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<document ENCODING="UTF-8" ID-DOI="https://doi.org/10.5281/zenodo.291971" ID-GBIF-Dataset="0bb2f033-2b79-4d87-803f-91694067c3f3" ID-ISBN="978-0-9506207-7-0" ID-Zenodo-Dep="291971" checkinTime="1588973351969" checkinUser="admin" docAuthor="Jarvis, Charlie" docDate="2007" docId="C34863FF91EFF6E5FD14D72E8578A629" docLanguage="en" docName="ch7.gg.C.xml" docOrigin="Order out of Chaos. Linnaean Plant Types and their Types, London: Linnaean Society of London in association with the Natural History Museum" docTitle="Coccoloba pubescens Linnaeus 1759" docType="treatment" docVersion="10" lastPageNumber="431" masterDocId="E213C9FA410C8D52CE14FE18C476F6F3" masterDocTitle="Chapter 7: Linnaean Plant Names and their Types (part C)" masterLastPageNumber="473" masterPageNumber="370" pageNumber="431" updateTime="1635912622391" updateUser="ExternalLinkService">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>Chapter 7: Linnaean Plant Names and their Types (part C)</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Jarvis, Charlie</mods:namePart>
<mods:affiliation>Department of Botany, Natural History Museum, Cromwell Road, London, UK</mods:affiliation>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
<mods:relatedItem type="host">
<mods:originInfo>
<mods:dateIssued>2007</mods:dateIssued>
<mods:publisher>Linnaean Society of London in association with the Natural History Museum</mods:publisher>
<mods:place>
<mods:placeTerm>London</mods:placeTerm>
</mods:place>
</mods:originInfo>
<mods:titleInfo>
<mods:title>Order out of Chaos. Linnaean Plant Types and their Types</mods:title>
</mods:titleInfo>
<mods:part>
<mods:extent unit="page">
<mods:start>370</mods:start>
<mods:end>473</mods:end>
</mods:extent>
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<mods:classification>book chapter</mods:classification>
<mods:identifier type="DOI">https://doi.org/10.5281/zenodo.291971</mods:identifier>
<mods:identifier type="ISBN">978-0-9506207-7-0</mods:identifier>
<mods:identifier type="Zenodo-Dep">291971</mods:identifier>
</mods:mods>
<treatment ID-DOI="http://doi.org/10.5281/zenodo.4379826" ID-GBIF-Taxon="164002697" ID-Zenodo-Dep="4379826" LSID="urn:lsid:plazi:treatment:C34863FF91EFF6E5FD14D72E8578A629" httpUri="http://treatment.plazi.org/id/C34863FF91EFF6E5FD14D72E8578A629" lastPageNumber="431" pageNumber="431">
<subSubSection pageNumber="431" type="reference_group">
<paragraph pageNumber="431">
<treatmentCitation author="Linnaeus" pageNumber="1007" year="1759">
<bibCitation author="Linnaeus" pagination="1007" title="Coccoloba pubescens" volumeTitle="Systema Naturae, ed. 10, 2" year="1759">
<taxonomicName authority="Linnaeus" authorityName="Linnaeus" authorityYear="1759" class="Magnoliopsida" family="Polygonaceae" genus="Coccoloba" kingdom="Plantae" order="Polygonales" phylum="Tracheophyta" rank="species" species="pubescens">
<emphasis bold="true" pageNumber="431">Coccoloba pubescens</emphasis>
Linnaeus
</taxonomicName>
,
<volumeTitle>
<emphasis italics="true" pageNumber="431">Systema Naturae</emphasis>
, ed. 10, 2
</volumeTitle>
: 1007. 1759
</bibCitation>
</treatmentCitation>
.
</paragraph>
</subSubSection>
<subSubSection pageNumber="431" type="distribution">
<paragraph pageNumber="431">[&quot;Habitat in America.&quot;] Sp. Pl. ed. 2, 1: 523 (1762). RCN: 2881.</paragraph>
</subSubSection>
<subSubSection pageNumber="431" type="materials_examined">
<paragraph pageNumber="431">
<typeStatus>Lectotype</typeStatus>
(Howard,
<emphasis italics="true" pageNumber="431">Fl. Lesser Antilles</emphasis>
4: 133. 1988): [icon] &quot;
<emphasis italics="true" pageNumber="431">Scortea arbor Americana, amplissimis foliis, aversa parte nervis extantibus hirsutie ferruginea refertis</emphasis>
&quot; in Plukenet, Phytographia: t. 222, f. 8. 1692; Almag. Bot.: 38. 1696. - Voucher: Herb. Sloane 98: 23; 101: 196 (BM-SL).
</paragraph>
</subSubSection>
<subSubSection pageNumber="431" type="nomenclature">
<paragraph pageNumber="431">
Current name:
<taxonomicName authority="L." authorityName="Linnaeus" authorityYear="1759" class="Magnoliopsida" family="Polygonaceae" genus="Coccoloba" kingdom="Plantae" order="Polygonales" phylum="Tracheophyta" rank="species" species="pubescens">
<emphasis italics="true" pageNumber="431">
<emphasis bold="true" italics="true" pageNumber="431">Coccoloba pubescens</emphasis>
</emphasis>
L.
</taxonomicName>
(
<taxonomicName class="Magnoliopsida" family="Polygonaceae" kingdom="Plantae" order="Polygonales" phylum="Tracheophyta" rank="family">Polygonaceae</taxonomicName>
).
</paragraph>
</subSubSection>
<subSubSection pageNumber="431" type="discussion">
<paragraph pageNumber="431">
<emphasis italics="true" pageNumber="431">Note:</emphasis>
Fawcett &amp; Rendle (in
<emphasis italics="true" pageNumber="431">J. Bot.</emphasis>
51: 123. 1913) provided an extensive discussion but did not indicate a type, but later (
<emphasis italics="true" pageNumber="431">Fl. Jamaica</emphasis>
3: 118. 1914) indicated a type in BM (presumably material in Herb. Plukenet that was not seen by Linnaeus and is not original material for the name). See discussion by Howard (in
<emphasis italics="true" pageNumber="431">J. Arnold Arbor.</emphasis>
38: 229. 1957).
</paragraph>
</subSubSection>
</treatment>
</document>

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<document ID-DOI="10.7717/peerj.6573" ID-GBIF-Dataset="08258b2a-72c1-4f13-a965-a986e4939e11" ID-PMC="PMC6404657" ID-PubMed="30863686" ID-Zenodo-Dep="3749254" approvalRequired="38" approvalRequired_for_bibRefs="2" approvalRequired_for_document="1" approvalRequired_for_textStreams="33" approvalRequired_for_treatments="2" checkinTime="1586705614621" checkinUser="jeremy" docAuthor="Joseph E. Peterson &amp; Karsen N. Daus" docDate="2019" docId="C34887A8FF99FFEEFDB4AD048E6962B7" docLanguage="en" docName="PetersonDaus2019.pdf.imf" docOrigin="PeerJ (e 6573) 7 (3)" docSource="http://dx.doi.org/10.7717/peerj.6573" docStyle="DocumentStyle:B6346867C9D49AB2DD3A1D607021D2E9:PeerJ.2019-.journal_article" docStyleId="B6346867C9D49AB2DD3A1D607021D2E9" docStyleName="PeerJ.2019-.journal_article" docTitle="Tyrannosaurus rex Osborn 1905" docType="treatment" docVersion="4" lastPageNumber="13" masterDocId="3F71FFD0FF98FFE2FFBFAF278A766608" masterDocTitle="Feeding traces attributable to juvenile Tyrannosaurus rex offer insight into ontogenetic dietary trends" masterLastPageNumber="17" masterPageNumber="1" pageNumber="2" updateTime="1668138954482" updateUser="ExternalLinkService">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:title>Feeding traces attributable to juvenile Tyrannosaurus rex offer insight into ontogenetic dietary trends</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Joseph E. Peterson</mods:namePart>
<mods:affiliation>Department of Geology, University of Wisconsin Oshkosh, Oshkosh, WI, United States of America</mods:affiliation>
<mods:nameIdentifier type="email">petersoj@uwosh.edu</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Karsen N. Daus</mods:namePart>
<mods:affiliation>Department of Geology, University of Wisconsin Oshkosh, Oshkosh, WI, United States of America</mods:affiliation>
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<mods:number>2019-03-04</mods:number>
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<mods:number>7</mods:number>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.3808813" ID-GBIF-Taxon="184587546" ID-Zenodo-Dep="3808813" LSID="urn:lsid:plazi:treatment:C34887A8FF99FFEEFDB4AD048E6962B7" httpUri="http://treatment.plazi.org/id/C34887A8FF99FFEEFDB4AD048E6962B7" lastPageId="12" lastPageNumber="13" pageId="1" pageNumber="2">
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<paragraph blockId="1.[491,1542,229,892]" pageId="1" pageNumber="2">
Here we report on the presence of feeding traces on the caudal vertebra of a hadrosaurid dinosaur (BMR P2007.4.1, ‘‘Constantine’’). Based on the shape and orientation of the traces, and the known fauna of the Hell Creek Formation, they are interpreted to be feeding traces of a large theropod dinosaur, such as
<taxonomicName authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[1100,1312,667,692]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="rex">
<emphasis box="[1100,1312,667,692]" italics="true" pageId="1" pageNumber="2">Tyrannosaurus rex</emphasis>
</taxonomicName>
(
<bibRefCitation author="Erickson GM &amp; Olson KH" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="1" pageNumber="2" pagination="175 - 178" part="16" refId="ref5340" refString="Erickson GM, Olson KH. 1996. Bite marks attributable to Tyrannosaurus rex: a preliminary description and implications. Journal of Vertebrate Paleontology 16 (1): 175 - 178 DOI 10.1080 / 02724634.1996.10011297." title="Bite marks attributable to Tyrannosaurus rex: a preliminary description and implications" type="journal article" year="1996">
<emphasis italics="true" pageId="1" pageNumber="2">Erickson &amp; Olson, 1996</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Horner JR &amp; Goodwin MB &amp; Myhrvold N." box="[561,974,706,733]" journalOrPublisher="PLOS ONE" pageId="1" pageNumber="2" pagination="e 16574" part="6" refId="ref5979" refString="Horner JR, Goodwin MB, Myhrvold N. 2011. Dinosaur census reveals abundant Tyrannosaurus and rare ontogenetic stages in the Upper Cretaceous Hell Creek Formation (Maastrichtian), Montana, USA. PLOS ONE 6 (2): e 16574 DOI 10.1371 / journal. pone. 0016574." title="Dinosaur census reveals abundant Tyrannosaurus and rare ontogenetic stages in the Upper Cretaceous Hell Creek Formation (Maastrichtian), Montana, USA" type="journal article" year="2011">
<emphasis box="[561,974,706,733]" italics="true" pageId="1" pageNumber="2">Horner, Goodwin &amp; Myhrvold, 2011</emphasis>
</bibRefCitation>
). By comparing the dimensions and spacing of the traces with the maxillae and dentaries of specimens of
<taxonomicName authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[1177,1390,747,772]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="rex">
<emphasis box="[1177,1390,747,772]" italics="true" pageId="1" pageNumber="2">Tyrannosaurus rex</emphasis>
</taxonomicName>
of different ontogenetic stages, we interpret these tooth marks to be feeding traces from a juvenile
<taxonomicName authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[491,704,827,852]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="rex">
<emphasis box="[491,704,827,852]" italics="true" pageId="1" pageNumber="2">Tyrannosaurus rex</emphasis>
</taxonomicName>
and discuss the insights the specimen provides for juvenile tyrannosaur feeding behavior.
</paragraph>
</subSubSection>
<subSubSection lastPageId="8" lastPageNumber="9" pageId="1" pageNumber="2" type="description">
<paragraph blockId="1.[491,1541,926,1793]" box="[491,741,926,956]" pageId="1" pageNumber="2">
<heading bold="true" box="[491,741,926,956]" fontSize="12" level="2" pageId="1" pageNumber="2" reason="0">
<emphasis bold="true" box="[491,741,926,956]" pageId="1" pageNumber="2">Geologic setting</emphasis>
</heading>
</paragraph>
<paragraph blockId="1.[491,1541,926,1793]" pageId="1" pageNumber="2">
Specimen BMR P2007.4.1 is a partial hadrosaurid skeleton collected from the Upper Cretaceous Hell Creek Formation of Carter County, southeastern Montana in the Powder River Basin (
<figureCitation box="[645,711,1050,1076]" captionStart="Figure 1" captionStartId="2.[507,571,676,698]" captionTargetBox="[491,1449,229,630]" captionTargetId="figure@2.[491,1452,229,631]" captionTargetPageId="2" captionText="Figure 1 Discovery location of BMR P2007.4.1. Locality map showing the geographic location of specimen BMR P2007.4.1 in Carter County, Montana. Full-size DOI: 10.7717/peerj.6573/fig-1" figureDoi="http://doi.org/10.5281/zenodo.3749256" httpUri="https://zenodo.org/record/3749256/files/figure.png" pageId="1" pageNumber="2">Fig. 1</figureCitation>
). This specimen was collected on public lands under BLM Permit # M96842-2007 issued to Northern Illinois University and is accessioned at the Burpee Museum of Natural History in Rockford, IL. Exact coordinates for the location are on file in the paleontology collections at the Burpee Museum of Natural History (BMR), where the specimen is reposited.
</paragraph>
<paragraph blockId="1.[491,1541,926,1793]" pageId="1" pageNumber="2">
The collection locality is composed of a 4 m fine-grained, gray-tan lenticular sandstone within a larger surrounding blocky mudstone unit (
<figureCitation box="[1095,1159,1289,1315]" captionStart="Figure 2" captionStartId="3.[507,571,1714,1736]" captionTargetBox="[492,921,232,1665]" captionTargetId="figure@3.[491,922,229,1669]" captionTargetPageId="3" captionText="Figure 2 Stratigraphic column of the ‘‘Constantine’’ Quarry. Stratigraphy of the BMR P2007.4.1 ‘‘Constantine’’ Quarry. Full-size DOI: 10.7717/peerj.6573/fig-2" figureDoi="http://doi.org/10.5281/zenodo.3749258" httpUri="https://zenodo.org/record/3749258/files/figure.png" pageId="1" pageNumber="2">Fig. 2</figureCitation>
). The sandstone lacks bedforms, resulting from either (a) rapid accumulation (resulting in a lack of sedimentary structures), or (b) sedimentary structures that were obliterated by later currents or bioturbation, and is rich in rounded and weathered microvertebrate remains. The site is stratigraphically positioned approximately 44 m above the underlying Fox Hills–Hell Creek contact
</paragraph>
<paragraph blockId="1.[491,1541,926,1793]" pageId="1" pageNumber="2">
and overlies 0.5 m of siderite, which sits above a 5 m blocky mudstone. Grains are subrounded to subangular. Microvertebrate and fragmented macrovertebrate fossils are abundant and heavily rounded and abraded (
<bibRefCitation author="Peterson JE &amp; Scherer RP &amp; Huffman KM" box="[1034,1433,1567,1594]" journalOrPublisher="Palaios" pageId="1" pageNumber="2" pagination="81 - 88" part="26" refId="ref6454" refString="Peterson JE, Scherer RP, Huffman KM. 2011. Methods of microvertebrate sam- pling and their influences on taphonomic interpretations. Palaios 26 (2): 81 - 88 DOI 10.2110 / palo. 2010. p 10 - 080 r." title="Methods of microvertebrate sam- pling and their influences on taphonomic interpretations" type="journal article" year="2011">
<emphasis box="[1034,1433,1567,1594]" italics="true" pageId="1" pageNumber="2">Peterson, Scherer &amp; Huffman, 2011</emphasis>
</bibRefCitation>
). The fine-grained composition suggests a channel-fill deposit, overlying a floodplain deposit (
<bibRefCitation author="Murphy EC &amp; Hoganson JW &amp; Johnson KR" box="[502,916,1647,1674]" editor="Hartman JH &amp; Johnson KR &amp; Nichols DJ" journalOrPublisher="Geological Society of America Special Papers 361, Boulder, Colorado: Geological Society of America" pageId="1" pageNumber="2" pagination="9 - 34" refId="ref6230" refString="Murphy EC, Hoganson JW, Johnson KR. 2002. Lithostratigraphy of the Hell Creek Formation in North Dakota. In: Hartman JH, Johnson KR, Nichols DJ, eds. The hell creek formation and the cretaceous-tertiary boundary in the Northern great plains: an integrated continental record of the end of the cretaceous. Geological Society of America Special Papers 361, Boulder, Colorado: Geological Society of America, 9 - 34." title="Lithostratigraphy of the Hell Creek Formation in North Dakota" type="book chapter" volumeTitle="The hell creek formation and the cretaceous-tertiary boundary in the Northern great plains: an integrated continental record of the end of the cretaceous" year="2002">
<emphasis box="[502,916,1647,1674]" italics="true" pageId="1" pageNumber="2">Murphy, Hoganson &amp; Johnson, 2002</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Peterson JE &amp; Scherer RP &amp; Huffman KM" box="[927,1326,1647,1674]" journalOrPublisher="Palaios" pageId="1" pageNumber="2" pagination="81 - 88" part="26" refId="ref6454" refString="Peterson JE, Scherer RP, Huffman KM. 2011. Methods of microvertebrate sam- pling and their influences on taphonomic interpretations. Palaios 26 (2): 81 - 88 DOI 10.2110 / palo. 2010. p 10 - 080 r." title="Methods of microvertebrate sam- pling and their influences on taphonomic interpretations" type="journal article" year="2011">
<emphasis box="[927,1326,1647,1674]" italics="true" pageId="1" pageNumber="2">Peterson, Scherer &amp; Huffman, 2011</emphasis>
</bibRefCitation>
). The taphonomic distribution of the elements and their stratigraphic position suggests the skeleton was subaerially exposed on a floodplain for a considerable period of time prior to burial, allowing for weathering, disarticulation, and removal of many skeletal elements.
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.3749256" ID-Zenodo-Dep="3749256" httpUri="https://zenodo.org/record/3749256/files/figure.png" pageId="2" pageNumber="3" startId="2.[507,571,676,698]" targetBox="[491,1449,229,630]" targetPageId="2">
<paragraph blockId="2.[507,1526,676,757]" pageId="2" pageNumber="3">
<emphasis bold="true" box="[507,979,676,698]" pageId="2" pageNumber="3">Figure 1 Discovery location of BMR P2007.4.1.</emphasis>
Locality map showing the geographic location of specimen BMR P2007.4.1 in Carter County, Montana. Full-size DOI: 10.7717/peerj.6573/fig-1
</paragraph>
</caption>
<paragraph blockId="2.[491,939,812,847]" box="[491,939,812,847]" pageId="2" pageNumber="3">
<heading allCaps="true" bold="true" box="[491,939,812,847]" fontSize="14" level="3" pageId="2" pageNumber="3" reason="0">
<emphasis bold="true" box="[491,939,812,847]" pageId="2" pageNumber="3">MATERIALS &amp; METHODS</emphasis>
</heading>
</paragraph>
<paragraph blockId="2.[491,1542,870,1892]" pageId="2" pageNumber="3">
Specimen BMR P2007.4.1 consists of weathered pelvic elements (sacrum, left and right ilia), three dorsal vertebrae, and two proximal caudal vertebrae (
<figureCitation box="[1234,1297,910,936]" captionStart="Figure 3" captionStartId="4.[507,571,1233,1255]" captionTargetBox="[493,1450,231,1186]" captionTargetId="figure@4.[491,1453,229,1188]" captionTargetPageId="4" captionText="Figure 3 Map of the BMR P2007.4.1 ‘‘Constantine’’ Quarry. Dorsal vertebrae (field numbers CON- 2007-010, CON-2007-011, and CON-2007-012) were too weathered for collection, though their relative locations were mapped. Note the relative association of dorsal and caudal vertebrae, and pelvic elements. Full-size DOI: 10.7717/peerj.6573/fig-3" figureDoi="http://doi.org/10.5281/zenodo.3749260" httpUri="https://zenodo.org/record/3749260/files/figure.png" pageId="2" pageNumber="3">Fig. 3</figureCitation>
,
<tableCitation box="[1310,1395,910,936]" captionStart="Table 1" captionStartId="5.[507,563,246,267]" captionTargetBox="[514,1515,328,786]" captionTargetPageId="5" captionText="Table 1 Skeletal elements from BMR P2007.4.1. Recovered and recorded skeletal elements from the ‘‘Constantine Quarry’’ (BMR P2007.4.1) and taphonomic condition." httpUri="http://table.plazi.org/id/1F9E6636FF9DFFE7FE44AFD18EFB6722" pageId="2" pageNumber="3" tableUuid="1F9E6636FF9DFFE7FE44AFD18EFB6722">Table 1</tableCitation>
). The dorsal vertebrae were too weathered for collection, though their dimensions and relative locations within the quarry assemblage were measured and documented. Additionally, a series of heavily-weathered bone fragments and a small shed theropod tooth (
<emphasis box="[1303,1520,1029,1055]" italics="true" pageId="2" pageNumber="3">
<taxonomicName box="[1303,1482,1029,1055]" class="Reptilia" family="Troodontidae" genus="Saurornithoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">Saurornithoides</taxonomicName>
sp.
</emphasis>
) were also collected.
</paragraph>
<paragraph blockId="2.[491,1542,870,1892]" pageId="2" pageNumber="3">
The ilium of BMR P2007.4.1 possesses a number of hadrosaurid characters such as (1) a shallow morphology, (2) a ∼ 23
<superScript attach="left" box="[862,873,1148,1162]" fontSize="8" pageId="2" pageNumber="3">◦</superScript>
preacetabular process in medial view relative to the main body, and (3) a well-developed supra-acetabular process caudal to the acetabulum. While these characters are common among hadrosaurids, the stratigraphic position of BMR P2007.4.1 suggests it is attributable to the Late Cretaceous hadrosaurid
<taxonomicName authorityName="Lambe" authorityYear="1917" box="[1339,1515,1268,1294]" class="Reptilia" family="Hadrosauridae" genus="Edmontosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis box="[1339,1515,1268,1294]" italics="true" pageId="2" pageNumber="3">Edmontosaurus</emphasis>
</taxonomicName>
(i.e.,
<bibRefCitation author="Brett-Surman MK &amp; Wagner JR" box="[550,900,1308,1334]" editor="Carpenter K" journalOrPublisher="Bloomington: Indiana University Press" pageId="2" pageNumber="3" pagination="135 - 169" refId="ref4904" refString="Brett-Surman MK, Wagner JR. 2007. Discussion of character analysis of the appendicular anatomy of Campanian and Maastrichtian North American hadrosauridsvariation and ontogeny. In: Carpenter K, ed. Horns and beaks-ceratopsian and ornithopod dinosaurs. Bloomington: Indiana University Press, 135 - 169." title="Discussion of character analysis of the appendicular anatomy of Campanian and Maastrichtian North American hadrosauridsvariation and ontogeny" type="book chapter" volumeTitle="Horns and beaks-ceratopsian and ornithopod dinosaurs" year="2007">
<emphasis box="[550,900,1308,1334]" italics="true" pageId="2" pageNumber="3">Brett-Surman &amp; Wagner, 2007</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Campione NE" box="[914,1100,1308,1334]" editor="Ebert DA &amp; Evans DE" journalOrPublisher="Bloomington and Indianapolis: Indiana University Press" pageId="2" pageNumber="3" pagination="208 - 244" refId="ref4954" refString="Campione NE. 2014. Postcranial anatomy of Edmontosaurus regalis (Hadrosauridae) from the Horseshoe Canyon Formation, Alberta, Canada. In: Ebert DA, Evans DE, eds. Hadrosaurs. Bloomington and Indianapolis: Indiana University Press, 208 - 244." title="Postcranial anatomy of Edmontosaurus regalis (Hadrosauridae) from the Horseshoe Canyon Formation, Alberta, Canada" type="book chapter" volumeTitle="Hadrosaurs" year="2014">
<emphasis box="[914,1100,1308,1334]" italics="true" pageId="2" pageNumber="3">Campione, 2014</emphasis>
</bibRefCitation>
).
</paragraph>
<paragraph blockId="2.[491,1542,870,1892]" pageId="2" pageNumber="3">
The centra of the two caudal vertebrae lack any evidence for hemal arch attachments, suggesting they are among the more cranial-positioned caudal vertebrae, such as C1–C4 (
<bibRefCitation author="Campione NE" box="[502,688,1427,1454]" editor="Ebert DA &amp; Evans DE" journalOrPublisher="Bloomington and Indianapolis: Indiana University Press" pageId="2" pageNumber="3" pagination="208 - 244" refId="ref4954" refString="Campione NE. 2014. Postcranial anatomy of Edmontosaurus regalis (Hadrosauridae) from the Horseshoe Canyon Formation, Alberta, Canada. In: Ebert DA, Evans DE, eds. Hadrosaurs. Bloomington and Indianapolis: Indiana University Press, 208 - 244." title="Postcranial anatomy of Edmontosaurus regalis (Hadrosauridae) from the Horseshoe Canyon Formation, Alberta, Canada" type="book chapter" volumeTitle="Hadrosaurs" year="2014">
<emphasis box="[502,688,1427,1454]" italics="true" pageId="2" pageNumber="3">Campione, 2014</emphasis>
</bibRefCitation>
). One of the caudal vertebrae possesses three v-shaped indentations on the ventral surface of the centrum (
<figureCitation box="[915,1054,1468,1494]" captionStart="Figure 4" captionStartId="5.[507,571,1775,1797]" captionTargetBox="[494,1438,838,1728]" captionTargetId="figure@5.[491,1441,835,1731]" captionTargetPageId="5" captionText="Figure 4 Punctured caudal vertebra of BMR P2007.4.1. BMR P2007.4.1 in anterior (A) posterior (B) and ventral (C), including the two elliptical punctures on the ventral surface of the centrum (D, E). Full-size DOI: 10.7717/peerj.6573/fig-4" figureDoi="http://doi.org/10.5281/zenodo.3749262" httpUri="https://zenodo.org/record/3749262/files/figure.png" pageId="2" pageNumber="3">Figs. 4A–4E</figureCitation>
;
<httpUri box="[1070,1247,1468,1494]" httpUri="https://doi.org/10.6084/m9.figshare.7424945.v2" pageId="2" pageNumber="3">Figs. S1 and S2</httpUri>
). These traces feature collapsed cortical bone within the indentation, producing puncture marks (
<emphasis italics="true" pageId="2" pageNumber="3">
sensu
<bibRefCitation author="Binford LR" bookContentInfo="320" journalOrPublisher="New York: Academic Press" pageId="2" pageNumber="3" refId="ref4883" refString="Binford LR. 1981. Bones: ancient men and modern myths. New York: Academic Press, 320." title="Bones: ancient men and modern myths" type="book" year="1981">Binford, 1981</bibRefCitation>
</emphasis>
). The punctures penetrate 5 mm deep, are spaced 68 mm apart from their apical centers, show no signs of healing, and are inferred to have been created postmortem as feeding traces (e.g.,
<bibRefCitation author="Noto CR &amp; Main DJ &amp; Drumheller SK" box="[727,1093,1627,1653]" journalOrPublisher="Palaios" pageId="2" pageNumber="3" pagination="105 - 115" part="27" refId="ref6310" refString="Noto CR, Main DJ, Drumheller SK. 2012. Feeding traces and paleobiology of a Cretaceous (Cenomanian) crocodyliform: example from the Woodbine Formation of Texas. Palaios 27 (2): 105 - 115 DOI 10.2110 / palo. 2011. p 11 - 052 r." title="Feeding traces and paleobiology of a Cretaceous (Cenomanian) crocodyliform: example from the Woodbine Formation of Texas" type="journal article" year="2012">
<emphasis box="[727,1093,1627,1653]" italics="true" pageId="2" pageNumber="3">Noto, Main &amp; Drumheller, 2012</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Hone DWE &amp; Tanke DH" box="[1109,1345,1627,1653]" journalOrPublisher="PeerJ" pageId="2" pageNumber="3" pagination="e 885" part="3" refId="ref5902" refString="Hone DWE, Tanke DH. 2015. Pre- and postmortem tyrannosaurid bite marks on the remains of Daspletosaurus (Tyrannosaurinae: Therpoda) from Dinosaur Provincial Park, Alberta, Canada. PeerJ 3: e 885 DOI 10.7717 / peerj. 885." title="Pre- and postmortem tyrannosaurid bite marks on the remains of Daspletosaurus (Tyrannosaurinae: Therpoda) from Dinosaur Provincial Park, Alberta, Canada" type="journal article" year="2015">
<emphasis box="[1109,1345,1627,1653]" italics="true" pageId="2" pageNumber="3">Hone &amp; Tanke, 2015</emphasis>
</bibRefCitation>
;
<bibRefCitation author="McLain MA &amp; Nelsen D &amp; Snyder K &amp; Griffin CT &amp; Siviero B &amp; Brand LR &amp; Chadwick AV" journalOrPublisher="Palaios" pageId="2" pageNumber="3" pagination="164 - 173" part="33" refId="ref6171" refString="McLain MA, Nelsen D, Snyder K, Griffin CT, Siviero B, Brand LR, Chadwick AV. 2018. Tyrannosaur cannibalism: a case of a tooth-traced tyrannosaurid bone in the Lance Formation (Maastrichtian), Wyoming. Palaios 33 (4): 164 - 173 DOI 10.2110 / palo. 2017.076." title="Tyrannosaur cannibalism: a case of a tooth-traced tyrannosaurid bone in the Lance Formation (Maastrichtian), Wyoming" type="journal article" year="2018">
<emphasis italics="true" pageId="2" pageNumber="3">McLain et al., 2018</emphasis>
</bibRefCitation>
). The v-shape preserved in each puncture indicates that the original teeth would have possessed a prominent keel, though no striations from serration marks are present in the traces (
<emphasis box="[577,1009,1746,1773]" italics="true" pageId="2" pageNumber="3">
sensu
<bibRefCitation author="D'Amore DC &amp; Blumensehine RJ" box="[644,1009,1746,1773]" journalOrPublisher="Paleobiology" pageId="2" pageNumber="3" pagination="525 - 552" part="35" refId="ref5148" refString="D'Amore DC, Blumensehine RJ. 2009. Komodo monitor (Varanus komodoensis) feeding behavior and dental function reflected through tooth marks on bone surfaces, and the application to ziphodont paleobiology. Paleobiology 35 (4): 525 - 552 DOI 10.1666 / 0094 - 8373 - 35.4.525." title="Komodo monitor (Varanus komodoensis) feeding behavior and dental function reflected through tooth marks on bone surfaces, and the application to ziphodont paleobiology" type="journal article" year="2009">D’Amore &amp; Blumensehine, 2009</bibRefCitation>
</emphasis>
).
</paragraph>
<paragraph blockId="2.[491,1542,870,1892]" lastBlockId="4.[491,1542,1397,1902]" lastPageId="4" lastPageNumber="5" pageId="2" pageNumber="3">
The large size and shape of the punctures suggests that they were produced by a largeto medium-bodied carnivore. Such carnivores from the Hell Creek Formation include tyrannosaurs such as
<taxonomicName authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[750,962,1866,1891]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="rex">
<emphasis box="[750,962,1866,1891]" italics="true" pageId="2" pageNumber="3">Tyrannosaurus rex</emphasis>
</taxonomicName>
(
<bibRefCitation author="Erickson GM &amp; Olson KH" box="[985,1250,1866,1892]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="2" pageNumber="3" pagination="175 - 178" part="16" refId="ref5340" refString="Erickson GM, Olson KH. 1996. Bite marks attributable to Tyrannosaurus rex: a preliminary description and implications. Journal of Vertebrate Paleontology 16 (1): 175 - 178 DOI 10.1080 / 02724634.1996.10011297." title="Bite marks attributable to Tyrannosaurus rex: a preliminary description and implications" type="journal article" year="1996">
<emphasis box="[985,1250,1866,1892]" italics="true" pageId="2" pageNumber="3">Erickson &amp; Olson, 1996</emphasis>
</bibRefCitation>
;
<emphasis box="[1266,1493,1866,1892]" italics="true" pageId="2" pageNumber="3">Horner, Goodwin &amp;</emphasis>
<emphasis box="[491,671,1397,1424]" italics="true" pageId="4" pageNumber="5">Myhrvold, 2011</emphasis>
), medium-sized dromaeosaurids such as
<taxonomicName authority="(DePalma et al., 2015)" baseAuthorityName="DePalma" baseAuthorityYear="2015" class="Reptilia" family="Dromaeosauridae" genus="Dakotaraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="steini">
<emphasis box="[1159,1379,1398,1424]" italics="true" pageId="4" pageNumber="5">Dakotaraptor steini</emphasis>
(
<bibRefCitation author="DePalma RA &amp; Burnham DA &amp; Martin LD &amp; Larson PT &amp; Bakker RT" journalOrPublisher="Paleontological Contributions" pageId="4" pageNumber="5" pagination="1 - 16" part="14" refId="ref5200" refString="DePalma RA, Burnham DA, Martin LD, Larson PT, Bakker RT. 2015. The first giant raptor (Theropoda: Dromaeosauridae) from the Hell Creek Formation. Paleontological Contributions 14: 1 - 16 DOI 10.17161 / paleo. 1808.18764." title="The first giant raptor (Theropoda: Dromaeosauridae) from the Hell Creek Formation" type="journal article" year="2015">
<emphasis italics="true" pageId="4" pageNumber="5">DePalma et al., 2015</emphasis>
</bibRefCitation>
)
</taxonomicName>
, and crocodylians such as
<taxonomicName baseAuthorityName="Gilmore" baseAuthorityYear="1910" box="[918,1202,1438,1464]" class="Reptilia" genus="Borealosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="sternbergii">
<emphasis box="[918,1202,1438,1464]" italics="true" pageId="4" pageNumber="5">Borealosuchus sternbergii</emphasis>
</taxonomicName>
,
<taxonomicName authorityName="Gilmore" authorityYear="1911" box="[1218,1504,1438,1464]" class="Reptilia" family="Crocodylidae" genus="Brachychampsa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="montana">
<emphasis box="[1218,1504,1438,1464]" italics="true" pageId="4" pageNumber="5">Brachychampsa montana</emphasis>
</taxonomicName>
, and
<taxonomicName authority="(Matsumoto &amp; Evans, 2010)" baseAuthorityName="Matsumoto &amp; Evans" baseAuthorityYear="2010" box="[542,1200,1477,1503]" class="Reptilia" family="Crocodylidae" genus="Thoracosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="neocesariensis">
<emphasis box="[542,870,1477,1503]" italics="true" pageId="4" pageNumber="5">Thoracosaurus neocesariensis</emphasis>
(
<bibRefCitation author="Matsumoto R &amp; Evans SE" box="[887,1188,1477,1503]" journalOrPublisher="Journal of Iberian Geology" pageId="4" pageNumber="5" pagination="253 - 274" part="36" refId="ref6128" refString="Matsumoto R, Evans SE. 2010. Choristoderes and the freshwater assemblages of Laurasia. Journal of Iberian Geology 36 (2): 253 - 274 DOI 10.5209 / rev _ JIGE. 2010. v 36. n 2.11." title="Choristoderes and the freshwater assemblages of Laurasia" type="journal article" year="2010">
<emphasis box="[887,1188,1477,1503]" italics="true" pageId="4" pageNumber="5">Matsumoto &amp; Evans, 2010</emphasis>
</bibRefCitation>
)
</taxonomicName>
. By comparing the shape and orientation of the traces to the teeth of these carnivores from the Hell Creek Formation, they are hypothesized to be bite marks from a large theropod dinosaur, such as
<taxonomicName authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="rex">
<emphasis italics="true" pageId="4" pageNumber="5">Tyrannosaurus rex</emphasis>
</taxonomicName>
(
<bibRefCitation author="Erickson GM &amp; Olson KH" box="[547,811,1597,1623]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="4" pageNumber="5" pagination="175 - 178" part="16" refId="ref5340" refString="Erickson GM, Olson KH. 1996. Bite marks attributable to Tyrannosaurus rex: a preliminary description and implications. Journal of Vertebrate Paleontology 16 (1): 175 - 178 DOI 10.1080 / 02724634.1996.10011297." title="Bite marks attributable to Tyrannosaurus rex: a preliminary description and implications" type="journal article" year="1996">
<emphasis box="[547,811,1597,1623]" italics="true" pageId="4" pageNumber="5">Erickson &amp; Olson, 1996</emphasis>
</bibRefCitation>
); crocodylian teeth are circular in cross-section and too small, and dromaeosaurid teeth—even large dromaeosaurids such as
<taxonomicName box="[1219,1313,1637,1662]" class="Reptilia" family="Dromaeosauridae" genus="Dakotaraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="steini">
<emphasis box="[1219,1313,1637,1662]" italics="true" pageId="4" pageNumber="5">D. steini</emphasis>
</taxonomicName>
—are too small and laterally-compressed to have produced the punctures observed on BMRP2007.4.1.
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.3749258" ID-Zenodo-Dep="3749258" httpUri="https://zenodo.org/record/3749258/files/figure.png" pageId="3" pageNumber="4" startId="3.[507,571,1714,1736]" targetBox="[492,921,232,1665]" targetPageId="3">
<paragraph blockId="3.[507,1526,1714,1795]" pageId="3" pageNumber="4">
<emphasis bold="true" box="[507,1114,1714,1736]" pageId="3" pageNumber="4">Figure 2 Stratigraphic column of the ‘‘Constantine’’ Quarry.</emphasis>
Stratigraphy of the BMR P2007.4.1 ‘‘Constantine’’ Quarry. Full-size DOI: 10.7717/peerj.6573/fig-2
</paragraph>
</caption>
<caption ID-DOI="http://doi.org/10.5281/zenodo.3749260" ID-Zenodo-Dep="3749260" httpUri="https://zenodo.org/record/3749260/files/figure.png" pageId="4" pageNumber="5" startId="4.[507,571,1233,1255]" targetBox="[493,1450,231,1186]" targetPageId="4">
<paragraph blockId="4.[507,1526,1233,1343]" pageId="4" pageNumber="5">
<emphasis bold="true" box="[507,1104,1233,1255]" pageId="4" pageNumber="5">Figure 3 Map of the BMR P2007.4.1 ‘‘Constantine’’ Quarry.</emphasis>
Dorsal vertebrae (field numbers CON- 2007-010, CON-2007-011, and CON-2007-012) were too weathered for collection, though their relative locations were mapped. Note the relative association of dorsal and caudal vertebrae, and pelvic elements. Full-size DOI: 10.7717/peerj.6573/fig-3
</paragraph>
</caption>
<paragraph blockId="4.[491,1542,1397,1902]" lastBlockId="6.[491,1541,1181,1885]" lastPageId="6" lastPageNumber="7" pageId="4" pageNumber="5">
To test this hypothesis, the punctures on the caudal vertebra of BMR P2007.4.1 were first coated in
<superScript attach="left" box="[714,746,1751,1770]" fontSize="8" pageId="4" pageNumber="5">ReboundTM</superScript>
25 platinum-cure silicone rubber (Smooth-On) in order to make a silicone peel of the punctures in order to better visualize the morphology and dimensions of the teeth responsible for the traces (
<figureCitation box="[946,1086,1836,1862]" captionStart="Figure 5" captionStartId="6.[507,571,1016,1038]" captionTargetBox="[493,1451,230,970]" captionTargetId="figure@6.[491,1452,229,971]" captionTargetPageId="6" captionText="Figure 5 Silicone peel produced from BMR P2007.4.1. Silicone peel produced from the ventral surface of the punctured caudal vertebra of BMR P2007.4.1 in vertical (A), and lateral (B) views. Note the traced outlines demonstrating the shape of the tooth casts. Full-size DOI: 10.7717/peerj.6573/fig-5" figureDoi="http://doi.org/10.5281/zenodo.3749264" httpUri="https://zenodo.org/record/3749264/files/figure.png" pageId="4" pageNumber="5">Figs. 5A–5B</figureCitation>
;
<httpUri box="[1101,1282,1836,1862]" httpUri="https://doi.org/10.6084/m9.figshare.7424945.v2" pageId="4" pageNumber="5">Figs. S3 and S4</httpUri>
). These ‘‘teeth’’ were then compared with the dental dimensions and spacing of two
<taxonomicName authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[1217,1388,1876,1901]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis box="[1217,1388,1876,1901]" italics="true" pageId="4" pageNumber="5">Tyrannosaurus</emphasis>
</taxonomicName>
maxillae and dentaries. To approximate the ontogenetic stage of the tyrannosaur, a late-stage juvenile specimen (
<materialsCitation ID-GBIF-Occurrence="3335984310" box="[617,899,1221,1247]" collectionCode="BMR" pageId="6" pageNumber="7" specimenCode="BMP P2002.4.1, &quot;Jane&quot;">BMR P2002.4.1, ‘‘Jane’’</materialsCitation>
) histologically determined to be approximately 11–12 years old at the time of death (
<bibRefCitation author="Erickson GM &amp; Currie PJ &amp; Inouye BD &amp; Winn AA" box="[859,1083,1261,1287]" journalOrPublisher="Science" pageId="6" pageNumber="7" pagination="213 - 217" part="313" refId="ref5301" refString="Erickson GM, Currie PJ, Inouye BD, Winn AA. 2006. Tyrannosaur life tables: An example of nonavian dinosaur population biology. Science 313: 213 - 217 DOI 10.1126 / science. 1125721." title="Tyrannosaur life tables: An example of nonavian dinosaur population biology" type="journal article" year="2006">
<emphasis box="[859,1083,1261,1287]" italics="true" pageId="6" pageNumber="7">Erickson et al., 2006</emphasis>
</bibRefCitation>
) that possesses laterally compressed, sharp crowns, and a mature specimen (
<materialsCitation ID-GBIF-Occurrence="3335984319" box="[949,1160,1301,1327]" collectionCode="BHI" pageId="6" pageNumber="7" specimenCode="BHI 3033 &quot;Stan&quot;">BHI 3033, ‘‘Stan’’</materialsCitation>
) with robust, blunt crowns were utilized.
</paragraph>
<caption ID-Table-UUID="1F9E6636FF9DFFE7FE44AFD18EFB6722" httpUri="http://table.plazi.org/id/1F9E6636FF9DFFE7FE44AFD18EFB6722" pageId="5" pageNumber="6" startId="5.[507,563,246,267]" targetBox="[514,1515,328,786]" targetIsTable="true" targetPageId="5">
<paragraph blockId="5.[507,1477,246,298]" pageId="5" pageNumber="6">
<emphasis bold="true" box="[507,985,246,268]" pageId="5" pageNumber="6">Table 1 Skeletal elements from BMR P2007.4.1.</emphasis>
Recovered and recorded skeletal elements from the ‘‘Constantine Quarry’’ (BMR P2007.4.1) and taphonomic condition.
</paragraph>
</caption>
<paragraph pageId="5" pageNumber="6">
<table box="[514,1515,328,786]" gridcols="3" gridrows="13" pageId="5" pageNumber="6">
<tr box="[514,1515,328,350]" gridrow="0" pageId="5" pageNumber="6">
<th box="[514,663,328,350]" gridcol="0" gridrow="0" pageId="5" pageNumber="6">
<emphasis bold="true" box="[514,648,328,350]" pageId="5" pageNumber="6">Field number</emphasis>
</th>
<th box="[790,1077,328,350]" gridcol="1" gridrow="0" pageId="5" pageNumber="6">
<emphasis bold="true" box="[790,872,328,349]" pageId="5" pageNumber="6">Element</emphasis>
</th>
<th box="[1204,1515,328,350]" gridcol="2" gridrow="0" pageId="5" pageNumber="6">
<emphasis bold="true" box="[1204,1364,328,350]" pageId="5" pageNumber="6">State/Condition</emphasis>
</th>
</tr>
<tr box="[514,1515,367,389]" gridrow="1" pageId="5" pageNumber="6">
<th box="[514,663,367,389]" gridcol="0" gridrow="1" pageId="5" pageNumber="6">CON-2007-001</th>
<td box="[790,1077,367,389]" gridcol="1" gridrow="1" pageId="5" pageNumber="6">Rib fragment</td>
<td box="[1204,1515,367,389]" gridcol="2" gridrow="1" pageId="5" pageNumber="6">Abraded</td>
</tr>
<tr box="[514,1515,403,425]" gridrow="2" pageId="5" pageNumber="6">
<th box="[514,663,403,425]" gridcol="0" gridrow="2" pageId="5" pageNumber="6">CON-2007-002</th>
<td box="[790,1077,403,425]" gridcol="1" gridrow="2" pageId="5" pageNumber="6">Left ilium</td>
<td box="[1204,1515,403,425]" gridcol="2" gridrow="2" pageId="5" pageNumber="6">Heavily weathered</td>
</tr>
<tr box="[514,1515,439,461]" gridrow="3" pageId="5" pageNumber="6">
<th box="[514,663,439,461]" gridcol="0" gridrow="3" pageId="5" pageNumber="6">CON-2007-003</th>
<td box="[790,1077,439,461]" gridcol="1" gridrow="3" pageId="5" pageNumber="6">Rib fragment</td>
<td box="[1204,1515,439,461]" gridcol="2" gridrow="3" pageId="5" pageNumber="6">Abraded</td>
</tr>
<tr box="[514,1515,475,497]" gridrow="4" pageId="5" pageNumber="6">
<th box="[514,663,475,497]" gridcol="0" gridrow="4" pageId="5" pageNumber="6">CON-2007-004</th>
<td box="[790,1077,475,497]" gridcol="1" gridrow="4" pageId="5" pageNumber="6">Sacrum and right ilium</td>
<td box="[1204,1515,475,497]" gridcol="2" gridrow="4" pageId="5" pageNumber="6">Heavy to moderate weathering</td>
</tr>
<tr box="[514,1515,511,533]" gridrow="5" pageId="5" pageNumber="6">
<th box="[514,663,511,533]" gridcol="0" gridrow="5" pageId="5" pageNumber="6">CON-2007-005</th>
<td box="[790,1077,511,533]" gridcol="1" gridrow="5" pageId="5" pageNumber="6">Neural arch</td>
<td box="[1204,1515,511,533]" gridcol="2" gridrow="5" pageId="5" pageNumber="6">Fractured, but mild weathering</td>
</tr>
<tr box="[514,1515,548,570]" gridrow="6" pageId="5" pageNumber="6">
<th box="[514,663,548,570]" gridcol="0" gridrow="6" pageId="5" pageNumber="6">CON-2007-006</th>
<td box="[790,1077,548,570]" gridcol="1" gridrow="6" pageId="5" pageNumber="6">Caudal vertebra</td>
<td box="[1204,1515,548,570]" gridcol="2" gridrow="6" pageId="5" pageNumber="6">Mild weathering</td>
</tr>
<tr box="[514,1515,584,606]" gridrow="7" pageId="5" pageNumber="6">
<th box="[514,663,584,606]" gridcol="0" gridrow="7" pageId="5" pageNumber="6">CON-2007-007</th>
<td box="[790,1077,584,606]" gridcol="1" gridrow="7" pageId="5" pageNumber="6">Caudal vertebra</td>
<td box="[1204,1515,584,606]" gridcol="2" gridrow="7" pageId="5" pageNumber="6">Mild weathering</td>
</tr>
<tr box="[514,1515,620,642]" gridrow="8" pageId="5" pageNumber="6">
<th box="[514,663,620,642]" gridcol="0" gridrow="8" pageId="5" pageNumber="6">CON-2007-008</th>
<td box="[790,1077,620,642]" gridcol="1" gridrow="8" pageId="5" pageNumber="6">Bone fragment</td>
<td box="[1204,1515,620,642]" gridcol="2" gridrow="8" pageId="5" pageNumber="6">Heavily abraded</td>
</tr>
<tr box="[514,1515,656,678]" gridrow="9" pageId="5" pageNumber="6">
<th box="[514,663,656,678]" gridcol="0" gridrow="9" pageId="5" pageNumber="6">CON-2007-009</th>
<td box="[790,1077,656,678]" gridcol="1" gridrow="9" pageId="5" pageNumber="6">
Shed
<emphasis box="[842,1021,656,677]" italics="true" pageId="5" pageNumber="6">
<taxonomicName box="[842,989,656,677]" class="Reptilia" family="Troodontidae" genus="Saurornithoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Saurornithoides</taxonomicName>
sp.
</emphasis>
tooth
</td>
<td box="[1204,1515,656,678]" gridcol="2" gridrow="9" pageId="5" pageNumber="6">No apparent abrasion</td>
</tr>
<tr box="[514,1515,692,714]" gridrow="10" pageId="5" pageNumber="6">
<th box="[514,663,692,714]" gridcol="0" gridrow="10" pageId="5" pageNumber="6">CON-2007-010</th>
<td box="[790,1077,692,714]" gridcol="1" gridrow="10" pageId="5" pageNumber="6">Dorsal vertebra</td>
<td box="[1204,1515,692,714]" gridcol="2" gridrow="10" pageId="5" pageNumber="6">Heavily weathered, not collected</td>
</tr>
<tr box="[514,1515,728,750]" gridrow="11" pageId="5" pageNumber="6">
<th box="[514,663,728,750]" gridcol="0" gridrow="11" pageId="5" pageNumber="6">CON-2007-011</th>
<td box="[790,1077,728,750]" gridcol="1" gridrow="11" pageId="5" pageNumber="6">Dorsal vertebra</td>
<td box="[1204,1515,728,750]" gridcol="2" gridrow="11" pageId="5" pageNumber="6">Heavily weathered, not collected</td>
</tr>
<tr box="[514,1515,764,786]" gridrow="12" pageId="5" pageNumber="6">
<th box="[514,663,764,786]" gridcol="0" gridrow="12" pageId="5" pageNumber="6">CON-2007-012</th>
<td box="[790,1077,764,786]" gridcol="1" gridrow="12" pageId="5" pageNumber="6">Dorsal vertebra</td>
<td box="[1204,1515,764,786]" gridcol="2" gridrow="12" pageId="5" pageNumber="6">Heavily weathered, not collected</td>
</tr>
</table>
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.3749262" ID-Zenodo-Dep="3749262" httpUri="https://zenodo.org/record/3749262/files/figure.png" pageId="5" pageNumber="6" startId="5.[507,571,1775,1797]" targetBox="[494,1438,838,1728]" targetPageId="5">
<paragraph blockId="5.[507,1525,1775,1857]" pageId="5" pageNumber="6">
<emphasis bold="true" box="[507,1057,1775,1797]" pageId="5" pageNumber="6">Figure 4 Punctured caudal vertebra of BMR P2007.4.1.</emphasis>
BMR P2007.4.1 in anterior (A) posterior (B) and ventral (C), including the two elliptical punctures on the ventral surface of the centrum (D, E). Full-size DOI: 10.7717/peerj.6573/fig-4
</paragraph>
</caption>
<caption ID-DOI="http://doi.org/10.5281/zenodo.3749264" ID-Zenodo-Dep="3749264" httpUri="https://zenodo.org/record/3749264/files/figure.png" pageId="6" pageNumber="7" startId="6.[507,571,1016,1038]" targetBox="[493,1451,230,970]" targetPageId="6">
<paragraph blockId="6.[507,1526,1016,1127]" pageId="6" pageNumber="7">
<emphasis bold="true" box="[507,1047,1016,1038]" pageId="6" pageNumber="7">Figure 5 Silicone peel produced from BMR P2007.4.1.</emphasis>
Silicone peel produced from the ventral surface of the punctured caudal vertebra of BMR P2007.4.1 in vertical (A), and lateral (B) views. Note the traced outlines demonstrating the shape of the tooth casts. Full-size DOI: 10.7717/peerj.6573/fig-5
</paragraph>
</caption>
<paragraph blockId="6.[491,1541,1181,1885]" pageId="6" pageNumber="7">
All specimens were digitized via triangulated laser texture scanning with a NextEngine 3D Laser Scanner, capturing data at seven scanning divisions in high-definition (2.0k points/in 2). The resulting digital models were built with the NextEngine ScanStudio HD Pro version 2.02, and finalized as STL models (
<httpUri box="[1046,1184,1500,1526]" httpUri="https://doi.org/10.6084/m9.figshare.7424945.v2" pageId="6" pageNumber="7">Figs. S1–S8</httpUri>
). Scanning was conducted at the Department of Geology at the University of Wisconsin-Oshkosh in Oshkosh, WI.
</paragraph>
<paragraph blockId="6.[491,1541,1181,1885]" pageId="6" pageNumber="7">
The tooth spacing of both adult and late-stage juvenile tyrannosaur maxillae and dentaries were measured for both immediately-adjacent teeth and teeth from alternating replacement positions (i.e.,
<emphasis box="[816,947,1659,1685]" italics="true" pageId="6" pageNumber="7">Zahnreihen</emphasis>
), and compared with the spacing of the punctures (
<figureCitation box="[502,642,1699,1725]" captionStart="Figure 6" captionStartId="7.[507,571,1084,1106]" captionTargetBox="[491,1450,229,1039]" captionTargetId="figure@7.[491,1450,229,1039]" captionTargetPageId="7" captionText="Figure 6 Casts of BMR P2002.4.1 maxilla (A) and dentary (B) to illustrate the tooth positions used for spacing measurements. Note the alternating replacement of teeth. Scale bars equal 10 cm. Full-size DOI: 10.7717/peerj.6573/fig-6" figureDoi="http://doi.org/10.5281/zenodo.3749266" httpUri="https://zenodo.org/record/3749266/files/figure.png" pageId="6" pageNumber="7">Figs. 6A–6B</figureCitation>
), similar to
<emphasis box="[786,861,1699,1725]" italics="true" pageId="6" pageNumber="7">Fahlke</emphasis>
’s (
<emphasis box="[897,952,1699,1725]" italics="true" pageId="6" pageNumber="7">2012</emphasis>
) investigation of likely
<taxonomicName authorityName="Harlan" authorityYear="1834" box="[1227,1368,1699,1725]" class="Mammalia" family="Basilosauridae" genus="Basilosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis box="[1227,1368,1699,1725]" italics="true" pageId="6" pageNumber="7">Basilosaurus</emphasis>
</taxonomicName>
bite marks on specimens the smaller whale
<taxonomicName authorityName="Gibbes" authorityYear="1845" box="[839,941,1739,1765]" class="Mammalia" family="Basilosauridae" genus="Dorudon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis box="[839,941,1739,1765]" italics="true" pageId="6" pageNumber="7">Dorudon</emphasis>
</taxonomicName>
. Furthermore, the cross-sectional morphology
</paragraph>
<paragraph blockId="6.[491,1541,1181,1885]" pageId="6" pageNumber="7">
of adult and late-stage juvenile tyrannosaur maxillae and dentaries were measured labiolingually and mesiodistally at a 5 mm apical depth for each tooth crown, and plotted with measurements from the punctures found on BMR P2007.4.1 (
<figureCitation box="[1277,1342,1859,1885]" captionStart="Figure 7" captionStartId="8.[507,571,1212,1234]" captionTargetBox="[494,1448,231,1164]" captionTargetId="figure@8.[491,1451,229,1167]" captionTargetPageId="8" captionText="Figure 7 Maxillary and dentary measurements for BMRP 2002.4.1 and BHI 3033 mesiodistal and labiolingual dimensions at 5 mm depth compared to the bite marks on BMR P2007.4.1. Full-size DOI: 10.7717/peerj.6573/fig-7" figureDoi="http://doi.org/10.5281/zenodo.3749268" httpUri="https://zenodo.org/record/3749268/files/figure.png" pageId="6" pageNumber="7">Fig. 7</figureCitation>
).
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.3749266" ID-Zenodo-Dep="3749266" httpUri="https://zenodo.org/record/3749266/files/figure.png" pageId="7" pageNumber="8" startId="7.[507,571,1084,1106]" targetBox="[491,1450,229,1039]" targetPageId="7">
<paragraph blockId="7.[507,1526,1084,1165]" pageId="7" pageNumber="8">
<emphasis bold="true" pageId="7" pageNumber="8">
Figure 6 Casts of
<materialsCitation ID-GBIF-Occurrence="3335984312" box="[690,842,1084,1105]" collectionCode="BMR" pageId="7" pageNumber="8" specimenCode="BMR P2002.4.1">BMR P2002.4.1</materialsCitation>
maxilla (A) and dentary (B) to illustrate the tooth positions used for spacing measurements.
</emphasis>
Note the alternating replacement of teeth. Scale bars equal 10 cm. Full-size DOI: 10.7717/peerj.6573/fig-6
</paragraph>
</caption>
<paragraph blockId="7.[491,661,1219,1254]" box="[491,661,1219,1254]" pageId="7" pageNumber="8">
<heading allCaps="true" bold="true" box="[491,661,1219,1254]" fontSize="14" level="3" pageId="7" pageNumber="8" reason="0">
<emphasis bold="true" box="[491,661,1219,1254]" pageId="7" pageNumber="8">RESULTS</emphasis>
</heading>
</paragraph>
<paragraph blockId="7.[491,1542,1278,1862]" pageId="7" pageNumber="8">
The mesiodistal width measurements from the silicone peel taken from BMR P2007.4.1 average 7.8 mm and the labiolingual depth average was 5.2 mm. Maxillary and dentary teeth of the adult
<taxonomicName authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[698,869,1358,1383]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
<emphasis box="[698,869,1358,1383]" italics="true" pageId="7" pageNumber="8">Tyrannosaurus</emphasis>
</taxonomicName>
(
<materialsCitation ID-GBIF-Occurrence="3335984301" box="[884,1002,1357,1383]" collectionCode="BHI" pageId="7" pageNumber="8" specimenCode="BHI 3033">BHI 3033</materialsCitation>
) were found to be too large and widely spaced to have produced the punctures (
<figureCitation box="[884,959,1397,1423]" captionStart="Figure 7" captionStartId="8.[507,571,1212,1234]" captionTargetBox="[494,1448,231,1164]" captionTargetId="figure@8.[491,1451,229,1167]" captionTargetPageId="8" captionText="Figure 7 Maxillary and dentary measurements for BMRP 2002.4.1 and BHI 3033 mesiodistal and labiolingual dimensions at 5 mm depth compared to the bite marks on BMR P2007.4.1. Full-size DOI: 10.7717/peerj.6573/fig-7" figureDoi="http://doi.org/10.5281/zenodo.3749268" httpUri="https://zenodo.org/record/3749268/files/figure.png" pageId="7" pageNumber="8">Figs. 7</figureCitation>
,
<figureCitation box="[974,1097,1397,1423]" captionStart="Figure 8" captionStartId="9.[507,571,666,688]" captionTargetBox="[491,1451,229,621]" captionTargetId="figure@9.[491,1451,229,621]" captionTargetPageId="9" captionText="Figure 8 Digitized comparisons between tyrannosaur maxillae and BMR P2007.4.1. Interactive manipulation of digitized NextEngine 3D scan of a cast of the right maxilla of BHI 3033 and BMR P2007.4.1 caudal vertebra. Full-size DOI: 10.7717/peerj.6573/fig-8" figureDoi="http://doi.org/10.5281/zenodo.3749270" httpUri="https://zenodo.org/record/3749270/files/figure.png" pageId="7" pageNumber="8">8A and 8B</figureCitation>
;
<httpUri box="[1112,1290,1397,1423]" httpUri="https://doi.org/10.6084/m9.figshare.7424945.v2" pageId="7" pageNumber="8">Figs. S4 and S5</httpUri>
;
<tableCitation box="[1303,1466,1397,1423]" captionStart="Table 2" captionStartId="10.[507,563,246,267]" captionTargetBox="[514,1517,351,628]" captionTargetPageId="10" captionText="Table 2 Measurements of tooth crowns of tyrannosaur specimens. Mesiodistal and labiolingual measurements of teeth at 5 mm depth from the crown apex for (A) BHI 3033, (B) BMR P2002.4.1, and (C) the inferred bite marks on BMR P2007.4.1. All measurements are in mm." httpUri="http://table.plazi.org/id/1F9E6636FF92FFE8FE44AFD18EE4674F" pageId="7" pageNumber="8" tableUuid="1F9E6636FF92FFE8FE44AFD18EE4674F">Tables 2A–2C</tableCitation>
). For
<materialsCitation ID-GBIF-Occurrence="3335984303" box="[491,606,1437,1463]" collectionCode="BHI" pageId="7" pageNumber="8" specimenCode="BHI 3033">BHI 3033</materialsCitation>
, the average dentary tooth crown mesiodistal width at 5 mm depth was 7.13 mm, and the average dentary tooth crown labiolingual depth at 5 mm was 4.10 mm. The average maxillary crown mesiodistal width at 5 mm were 7.72 mm, and the average maxillary crown labiolingual depth at 5 mm averaged to 4.21 mm.
</paragraph>
<paragraph blockId="7.[491,1542,1278,1862]" lastBlockId="8.[491,1541,1347,1413]" lastPageId="8" lastPageNumber="9" pageId="7" pageNumber="8">
However, the teeth of
<materialsCitation ID-GBIF-Occurrence="3335984315" box="[787,971,1597,1623]" collectionCode="BMR" pageId="7" pageNumber="8" specimenCode="BMR P2002.4.1">BMR P2002.4.1</materialsCitation>
produced similarly shaped punctures at 5 mm apical depth (
<figureCitation box="[653,728,1636,1662]" captionStart="Figure 7" captionStartId="8.[507,571,1212,1234]" captionTargetBox="[494,1448,231,1164]" captionTargetId="figure@8.[491,1451,229,1167]" captionTargetPageId="8" captionText="Figure 7 Maxillary and dentary measurements for BMRP 2002.4.1 and BHI 3033 mesiodistal and labiolingual dimensions at 5 mm depth compared to the bite marks on BMR P2007.4.1. Full-size DOI: 10.7717/peerj.6573/fig-7" figureDoi="http://doi.org/10.5281/zenodo.3749268" httpUri="https://zenodo.org/record/3749268/files/figure.png" pageId="7" pageNumber="8">Figs. 7</figureCitation>
and
<figureCitation box="[788,802,1636,1662]" captionStart="Figure 9" captionStartId="11.[507,571,1192,1214]" captionTargetBox="[492,1451,231,1146]" captionTargetId="figure@11.[491,1452,229,1147]" captionTargetPageId="11" captionText="Figure 9 Digitized comparisons between BMR P2002.4.1 and BMR P2007.4.1. Interactive manipulation of digitized NextEngine 3D scan of a cast of the right maxilla and dentary of BMR P2002.4.1, and BMR P2007.4.1 caudal vertebra. Full-size DOI: 10.7717/peerj.6573/fig-9" figureDoi="http://doi.org/10.5281/zenodo.3749274" httpUri="https://zenodo.org/record/3749274/files/figure.png" pageId="7" pageNumber="8">9</figureCitation>
;
<httpUri box="[817,995,1636,1662]" httpUri="https://doi.org/10.6084/m9.figshare.7424945.v2" pageId="7" pageNumber="8">Figs. S7 and S8</httpUri>
;
<tableCitation box="[1008,1169,1636,1662]" captionStart="Table 2" captionStartId="10.[507,563,246,267]" captionTargetBox="[514,1517,351,628]" captionTargetPageId="10" captionText="Table 2 Measurements of tooth crowns of tyrannosaur specimens. Mesiodistal and labiolingual measurements of teeth at 5 mm depth from the crown apex for (A) BHI 3033, (B) BMR P2002.4.1, and (C) the inferred bite marks on BMR P2007.4.1. All measurements are in mm." httpUri="http://table.plazi.org/id/1F9E6636FF92FFE8FE44AFD18EE4674F" pageId="7" pageNumber="8" tableUuid="1F9E6636FF92FFE8FE44AFD18EE4674F">Tables 2B–2C</tableCitation>
). The puncture measurements taken from the peel, BMR P2007.4.1 demonstrate a mesiodistal width and labiolingual depth consistent with the measurements taken from the maxillary and dentary teeth of the late-stage juvenile
<taxonomicName authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[756,927,1756,1781]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
<emphasis box="[756,927,1756,1781]" italics="true" pageId="7" pageNumber="8">Tyrannosaurus</emphasis>
</taxonomicName>
. When plotted against the mesiodistal width and labiolingual depth of the maxillary teeth, measurements from the peel taken from BMR P2007.4.1 fall well within the cluster radius created by the late-stage juvenile
<taxonomicName authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[1363,1534,1836,1861]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
<emphasis box="[1363,1534,1836,1861]" italics="true" pageId="7" pageNumber="8">Tyrannosaurus</emphasis>
</taxonomicName>
,
<materialsCitation ID-GBIF-Occurrence="3335984313" box="[491,674,1347,1373]" collectionCode="BMR" pageId="8" pageNumber="9" specimenCode="BMR P2002.4.1">BMR P2002.4.1</materialsCitation>
(
<figureCitation box="[692,756,1347,1373]" captionStart="Figure 7" captionStartId="8.[507,571,1212,1234]" captionTargetBox="[494,1448,231,1164]" captionTargetId="figure@8.[491,1451,229,1167]" captionTargetPageId="8" captionText="Figure 7 Maxillary and dentary measurements for BMRP 2002.4.1 and BHI 3033 mesiodistal and labiolingual dimensions at 5 mm depth compared to the bite marks on BMR P2007.4.1. Full-size DOI: 10.7717/peerj.6573/fig-7" figureDoi="http://doi.org/10.5281/zenodo.3749268" httpUri="https://zenodo.org/record/3749268/files/figure.png" pageId="8" pageNumber="9">Fig. 7</figureCitation>
). Furthermore, the inferred crown spacing of the punctures closely matched those of the late-stage juvenile tyrannosaur maxilla (
<tableCitation box="[1212,1373,1387,1413]" captionStart="Table 3" captionStartId="12.[507,563,246,267]" captionTargetBox="[514,1517,351,862]" captionTargetPageId="12" captionText="Table 3 Measurements of crown spacing in tyrannosaur specimens. Tooth crown spacing between maxillary (A) and dentary (B) teeth in the juvenile tyrannosaur BMR P2002.4.1. All measurements are in mm." httpUri="http://table.plazi.org/id/1F9E6636FF94FFEEFE44AFD1885F674F" pageId="8" pageNumber="9" tableUuid="1F9E6636FF94FFEEFE44AFD1885F674F">Tables 3A–3B</tableCitation>
).
</paragraph>
</subSubSection>
<caption ID-DOI="http://doi.org/10.5281/zenodo.3749268" ID-Zenodo-Dep="3749268" httpUri="https://zenodo.org/record/3749268/files/figure.png" pageId="8" pageNumber="9" startId="8.[507,571,1212,1234]" targetBox="[494,1448,231,1164]" targetPageId="8">
<paragraph blockId="8.[507,1526,1212,1292]" pageId="8" pageNumber="9">
Figure 7 Maxillary and dentary measurements for
<materialsCitation ID-GBIF-Occurrence="3335984316" box="[1015,1168,1212,1233]" collectionCode="BMR" pageId="8" pageNumber="9" specimenCode="BMR P2002.4.1">BMRP 2002.4.1</materialsCitation>
and
<materialsCitation ID-GBIF-Occurrence="3335984302" box="[1214,1308,1212,1233]" collectionCode="BHI" pageId="8" pageNumber="9" specimenCode="BHI 3033">BHI 3033</materialsCitation>
mesiodistal and labiolingual dimensions at 5 mm depth compared to the bite marks on BMR P2007.4.1. Full-size DOI: 10.7717/peerj.6573/fig-7
</paragraph>
</caption>
<subSubSection lastPageId="12" lastPageNumber="13" pageId="8" pageNumber="9" type="discussion">
<paragraph blockId="8.[491,1098,1462,1497]" box="[491,1098,1462,1497]" pageId="8" pageNumber="9">
<heading allCaps="true" bold="true" box="[491,1098,1462,1497]" centered="true" fontSize="14" level="1" pageId="8" pageNumber="9" reason="0">
<emphasis bold="true" box="[491,1098,1462,1497]" pageId="8" pageNumber="9">DISCUSSION AND CONCLUSIONS</emphasis>
</heading>
</paragraph>
<paragraph blockId="8.[491,1541,1520,1905]" pageId="8" pageNumber="9">
While feeding traces and bite marks attributed to mature tyrannosaurids are welldocumented in common Late Cretaceous taxa such as hadrosaurids and ceratopsians (i.e.,
<bibRefCitation author="Fiorillo AR" box="[551,702,1599,1626]" journalOrPublisher="Palaeogeography, Palaeoclimatology, Palaeoecology" pageId="8" pageNumber="9" pagination="157 - 166" part="88" refId="ref5517" refString="Fiorillo AR. 1991. Prey bone utilization by predatory dinosaurs. Palaeogeography, Palaeoclimatology, Palaeoecology 88 (3 - 4): 157 - 166 DOI 10.1016 / 0031 - 0182 (91) 90062 - V." title="Prey bone utilization by predatory dinosaurs" type="journal article" year="1991">
<emphasis box="[551,702,1599,1626]" italics="true" pageId="8" pageNumber="9">Fiorillo, 1991</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Erickson GM &amp; Olson KH" box="[717,942,1600,1626]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="8" pageNumber="9" pagination="175 - 178" part="16" refId="ref5340" refString="Erickson GM, Olson KH. 1996. Bite marks attributable to Tyrannosaurus rex: a preliminary description and implications. Journal of Vertebrate Paleontology 16 (1): 175 - 178 DOI 10.1080 / 02724634.1996.10011297." title="Bite marks attributable to Tyrannosaurus rex: a preliminary description and implications" type="journal article" year="1996">
<emphasis box="[717,942,1600,1626]" italics="true" pageId="8" pageNumber="9">Erickson et al., 1996</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Erickson GM &amp; Olson KH" box="[957,1221,1600,1626]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="8" pageNumber="9" pagination="175 - 178" part="16" refId="ref5340" refString="Erickson GM, Olson KH. 1996. Bite marks attributable to Tyrannosaurus rex: a preliminary description and implications. Journal of Vertebrate Paleontology 16 (1): 175 - 178 DOI 10.1080 / 02724634.1996.10011297." title="Bite marks attributable to Tyrannosaurus rex: a preliminary description and implications" type="journal article" year="1996">
<emphasis box="[957,1221,1600,1626]" italics="true" pageId="8" pageNumber="9">Erickson &amp; Olson, 1996</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Jacobsen AR" box="[1236,1401,1599,1626]" journalOrPublisher="Historical Biology" pageId="8" pageNumber="9" pagination="17 - 26" part="13" refId="ref6061" refString="Jacobsen AR. 1998. Feeding behavior in carnivorous dinosaurs determined by tooth marks on dinosaur bones. Historical Biology 13: 17 - 26 DOI 10.1080 / 08912969809386569." title="Feeding behavior in carnivorous dinosaurs determined by tooth marks on dinosaur bones" type="journal article" year="1998">
<emphasis box="[1236,1401,1599,1626]" italics="true" pageId="8" pageNumber="9">Jacobsen, 1998</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Farlow JO &amp; Holtz TR" editor="Kowalewski M &amp; Kelley PH" journalOrPublisher="New York: Cambridge University Press" pageId="8" pageNumber="9" pagination="251 - 265" part="8" refId="ref5475" refString="Farlow JO, Holtz TR. 2002. The fossil record of predation in dinosaurs. In: Kowalewski M, Kelley PH, eds. Paleontological Society Paper 8. New York: Cambridge University Press, 251 - 265." title="The fossil record of predation in dinosaurs" type="book chapter" volumeTitle="Paleontological Society Paper" year="2002">
<emphasis italics="true" pageId="8" pageNumber="9">Farlow &amp; Holtz, 2002</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Fowler DW &amp; Sullivan RM" box="[636,909,1640,1666]" journalOrPublisher="New Mexico Museum of Natural History and Science Bulletin" pageId="8" pageNumber="9" pagination="127 - 130" part="35" refId="ref5604" refString="Fowler DW, Sullivan RM. 2006. A ceratopsid pelvis with toothmarks from the Upper Cretaceous Kirtland Formation, New Mexico: evidence of late Campanian tyrannosaurid feeding behavior. New Mexico Museum of Natural History and Science Bulletin 35: 127 - 130." title="A ceratopsid pelvis with toothmarks from the Upper Cretaceous Kirtland Formation, New Mexico: evidence of late Campanian tyrannosaurid feeding behavior" type="journal article" year="2006">
<emphasis box="[636,909,1640,1666]" italics="true" pageId="8" pageNumber="9">Fowler &amp; Sullivan, 2006</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Peterson JE &amp; Henderson MD &amp; Scherer RP &amp; Vittore CP" box="[923,1148,1639,1666]" journalOrPublisher="Palaios" pageId="8" pageNumber="9" pagination="780 - 784" part="24" refId="ref6411" refString="Peterson JE, Henderson MD, Scherer RP, Vittore CP. 2009. Face biting on a juvenile tyrannosaurid and behavioral implications. Palaios 24: 780 - 784 DOI 10.2110 / palo. 2009. p 09 - 056 r." title="Face biting on a juvenile tyrannosaurid and behavioral implications" type="journal article" year="2009">
<emphasis box="[923,1148,1639,1666]" italics="true" pageId="8" pageNumber="9">Peterson et al., 2009</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Bell PR &amp; Currie PJ" box="[1162,1382,1639,1666]" journalOrPublisher="Lethaia" pageId="8" pageNumber="9" pagination="278 - 281" part="43" refId="ref4842" refString="Bell PR, Currie PJ. 2009. A tyrannosaur jaw bitten by a confamilial: scavenging or fatal agonism? Lethaia 43 (2): 278 - 281 DOI 10.1111 / j. 1502 - 3931.2009.00195. x." title="A tyrannosaur jaw bitten by a confamilial: scavenging or fatal agonism?" type="journal article" year="2009">
<emphasis box="[1162,1382,1639,1666]" italics="true" pageId="8" pageNumber="9">Bell &amp; Currie, 2009</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Fowler DW &amp; Scannella JB &amp; Goodwin MB &amp; Horner JR" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="8" pageNumber="9" pagination="32" part="S 96 A" refId="ref5557" refString="Fowler DW, Scannella JB, Goodwin MB, Horner JR. 2012. How to eat a Triceratops: large sample sample of toothmarks provides new insight into the feeding behavior of Tyrannosaurus [Abstract S 96 A]. Journal of Vertebrate Paleontology 32." title="How to eat a Triceratops: large sample sample of toothmarks provides new insight into the feeding behavior of Tyrannosaurus" type="journal article" year="2012">
<emphasis italics="true" pageId="8" pageNumber="9">Fowler et al., 2012</emphasis>
</bibRefCitation>
;
<bibRefCitation author="DePalma RA &amp; Burnham DA &amp; Martin LD &amp; Rothschild BM &amp; Larson PL" box="[561,796,1679,1706]" journalOrPublisher="Proceedings of the Na- tional Academy of Sciences of the United States of America" pageId="8" pageNumber="9" pagination="12560 - 12564" part="110" refId="ref5246" refString="DePalma RA, Burnham DA, Martin LD, Rothschild BM, Larson PL. 2013. Physical evidence of predatory behavior in Tyrannosaurus rex. Proceedings of the Na- tional Academy of Sciences of the United States of America 110 (31): 12560 - 12564 DOI 10.1073 / pnas. 1216534110." title="Physical evidence of predatory behavior in Tyrannosaurus rex" type="journal article" year="2013">
<emphasis box="[561,796,1679,1706]" italics="true" pageId="8" pageNumber="9">DePalma et al., 2013</emphasis>
</bibRefCitation>
;
<bibRefCitation author="McLain MA &amp; Nelsen D &amp; Snyder K &amp; Griffin CT &amp; Siviero B &amp; Brand LR &amp; Chadwick AV" box="[810,1027,1679,1706]" journalOrPublisher="Palaios" pageId="8" pageNumber="9" pagination="164 - 173" part="33" refId="ref6171" refString="McLain MA, Nelsen D, Snyder K, Griffin CT, Siviero B, Brand LR, Chadwick AV. 2018. Tyrannosaur cannibalism: a case of a tooth-traced tyrannosaurid bone in the Lance Formation (Maastrichtian), Wyoming. Palaios 33 (4): 164 - 173 DOI 10.2110 / palo. 2017.076." title="Tyrannosaur cannibalism: a case of a tooth-traced tyrannosaurid bone in the Lance Formation (Maastrichtian), Wyoming" type="journal article" year="2018">
<emphasis box="[810,1027,1679,1706]" italics="true" pageId="8" pageNumber="9">McLain et al., 2018</emphasis>
</bibRefCitation>
), the identification of juvenile tyrannosaur feeding traces adds insight into the role of juvenile theropods in Cretaceous ecosystems. The dimensions and spacing of the punctures closely matches the maxillary teeth of
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, a late-stage juvenile (11–12 yr old) tyrannosaur which incidentally itself possesses morphologically similar craniofacial lesions previously interpreted as a conspecific bite (
<bibRefCitation author="Peterson JE &amp; Henderson MD &amp; Scherer RP &amp; Vittore CP" box="[502,727,1878,1905]" journalOrPublisher="Palaios" pageId="8" pageNumber="9" pagination="780 - 784" part="24" refId="ref6411" refString="Peterson JE, Henderson MD, Scherer RP, Vittore CP. 2009. Face biting on a juvenile tyrannosaurid and behavioral implications. Palaios 24: 780 - 784 DOI 10.2110 / palo. 2009. p 09 - 056 r." title="Face biting on a juvenile tyrannosaurid and behavioral implications" type="journal article" year="2009">
<emphasis box="[502,727,1878,1905]" italics="true" pageId="8" pageNumber="9">Peterson et al., 2009</emphasis>
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).
</paragraph>
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<paragraph blockId="9.[507,1526,666,776]" pageId="9" pageNumber="10">
<emphasis bold="true" box="[507,1339,666,688]" pageId="9" pageNumber="10">Figure 8 Digitized comparisons between tyrannosaur maxillae and BMR P2007.4.1.</emphasis>
Interactive manipulation of digitized NextEngine 3D scan of a cast of the right maxilla of
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and BMR P2007.4.1 caudal vertebra. Full-size DOI: 10.7717/peerj.6573/fig-8
</paragraph>
</caption>
<paragraph blockId="9.[491,1542,830,1893]" pageId="9" pageNumber="10">
<bibRefCitation author="Longrich NR &amp; Horner JR &amp; Erickson GM &amp; Currie PJ" box="[523,766,830,857]" journalOrPublisher="PLOS ONE" pageId="9" pageNumber="10" pagination="e 13419" part="5" refId="ref6091" refString="Longrich NR, Horner JR, Erickson GM, Currie PJ. 2010. Cannibalism in Tyrannosaurus rex. PLOS ONE 5 (10): e 13419 DOI 10.1371 / journal / pone. 0013419." title="Cannibalism in Tyrannosaurus rex" type="journal article" year="2010">
<emphasis box="[523,766,830,857]" italics="true" pageId="9" pageNumber="10">Longrich et al. (2010)</emphasis>
</bibRefCitation>
reported on evidence of cannibalism in
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<emphasis box="[1239,1305,831,856]" italics="true" pageId="9" pageNumber="10">T. rex</emphasis>
</taxonomicName>
based on a number of bitten and scored remains of
<taxonomicName authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[861,1074,871,896]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="rex">
<emphasis box="[861,1074,871,896]" italics="true" pageId="9" pageNumber="10">Tyrannosaurus rex</emphasis>
</taxonomicName>
, some attributed to juvenile or subadult individuals. However, many of these traces resemble the ‘puncture and pull’ bite marks that have previously been attributed to
<taxonomicName authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[905,971,950,975]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="rex">
<emphasis box="[905,971,950,975]" italics="true" pageId="9" pageNumber="10">T. rex</emphasis>
</taxonomicName>
(
<bibRefCitation author="Erickson GM &amp; Olson KH" box="[993,1218,950,976]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="9" pageNumber="10" pagination="175 - 178" part="16" refId="ref5340" refString="Erickson GM, Olson KH. 1996. Bite marks attributable to Tyrannosaurus rex: a preliminary description and implications. Journal of Vertebrate Paleontology 16 (1): 175 - 178 DOI 10.1080 / 02724634.1996.10011297." title="Bite marks attributable to Tyrannosaurus rex: a preliminary description and implications" type="journal article" year="1996">
<emphasis box="[993,1218,950,976]" italics="true" pageId="9" pageNumber="10">Erickson et al., 1996</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Erickson GM &amp; Olson KH" box="[1233,1498,950,976]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="9" pageNumber="10" pagination="175 - 178" part="16" refId="ref5340" refString="Erickson GM, Olson KH. 1996. Bite marks attributable to Tyrannosaurus rex: a preliminary description and implications. Journal of Vertebrate Paleontology 16 (1): 175 - 178 DOI 10.1080 / 02724634.1996.10011297." title="Bite marks attributable to Tyrannosaurus rex: a preliminary description and implications" type="journal article" year="1996">
<emphasis box="[1233,1498,950,976]" italics="true" pageId="9" pageNumber="10">Erickson &amp; Olson, 1996</emphasis>
</bibRefCitation>
), and also include furrows and scores (
<emphasis box="[927,1149,990,1016]" italics="true" pageId="9" pageNumber="10">
sensu
<bibRefCitation author="Binford LR" bookContentInfo="320" box="[994,1149,990,1016]" journalOrPublisher="New York: Academic Press" pageId="9" pageNumber="10" refId="ref4883" refString="Binford LR. 1981. Bones: ancient men and modern myths. New York: Academic Press, 320." title="Bones: ancient men and modern myths" type="book" year="1981">Binford, 1981</bibRefCitation>
</emphasis>
).
</paragraph>
<paragraph blockId="9.[491,1542,830,1893]" pageId="9" pageNumber="10">
Correlating traces in bone, such as tooth marks, to specific taxa and ontogenetic stages usually requires direct comparisons (e.g.,
<bibRefCitation author="Peterson JE &amp; Henderson MD &amp; Scherer RP &amp; Vittore CP" box="[980,1205,1069,1096]" journalOrPublisher="Palaios" pageId="9" pageNumber="10" pagination="780 - 784" part="24" refId="ref6411" refString="Peterson JE, Henderson MD, Scherer RP, Vittore CP. 2009. Face biting on a juvenile tyrannosaurid and behavioral implications. Palaios 24: 780 - 784 DOI 10.2110 / palo. 2009. p 09 - 056 r." title="Face biting on a juvenile tyrannosaurid and behavioral implications" type="journal article" year="2009">
<emphasis box="[980,1205,1069,1096]" italics="true" pageId="9" pageNumber="10">Peterson et al., 2009</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Fahlke JM" bookContentInfo="16 p." box="[1220,1363,1069,1096]" journalOrPublisher="Palaeontologica Electronica" pageId="9" pageNumber="10" pagination="32 A" part="15" publicationUrl="palaeo-electronica.org/content/2012-issue-3-articles/339-archaeocete-predation" refId="ref5421" refString="Fahlke JM. 2012. Bite marks revisited-evidence for middle-to-late Eocene Basilosaurus isis predation on Dorudon atrox (both Cetacea, Basilosauridae). Palaeontologica Electronica 15 (3): 32 A, 16 p. palaeo-electronica. org / content / 2012 - issue- 3 - articles / 339 - archaeocete-predation." title="Bite marks revisited-evidence for middle-to-late Eocene Basilosaurus isis predation on Dorudon atrox (both Cetacea, Basilosauridae)" type="book" year="2012">
<emphasis box="[1220,1363,1069,1096]" italics="true" pageId="9" pageNumber="10">Fahlke, 2012</emphasis>
</bibRefCitation>
). However, in cases where direct comparisons are not available, estimates can be made for tooth size, morphology, and spacing based on ontogenetic trajectories. While bite marks and feeding traces attributable to younger juvenile and hatchling tyrannosaurs have not yet been identified, the punctures present on the caudal vertebra of BMR P2007.4.1 provide direct evidence that late-stage juvenile
<taxonomicName authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[870,1083,1269,1294]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="rex">
<emphasis box="[870,1083,1269,1294]" italics="true" pageId="9" pageNumber="10">Tyrannosaurus rex</emphasis>
</taxonomicName>
such as
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possessed—at least in part—a similar diet as adults.
</paragraph>
<paragraph blockId="9.[491,1542,830,1893]" pageId="9" pageNumber="10">
While bite marks resulting from active predation cannot easily be distinguished from postmortem feeding traces, the ventral position of the punctures in the caudal centrum of BMR P2007.4.1 suggests that the feeding was taking place postmortem with the hadrosaur already on its side (
<bibRefCitation author="Chure DJ &amp; Fiorillo AR &amp; Jacobsen A." box="[725,1093,1468,1494]" journalOrPublisher="Gaia" pageId="9" pageNumber="10" pagination="227 - 232" part="15" refId="ref5051" refString="Chure DJ, Fiorillo AR, Jacobsen A. 1998. Prey bone utilization by predatory dinosaur sin the Late Jurassic of North America, with comments on prey bone use by dinosaurs throughout the Mesozoic. Gaia 15: 227 - 232." title="Prey bone utilization by predatory dinosaur sin the Late Jurassic of North America, with comments on prey bone use by dinosaurs throughout the Mesozoic" type="journal article" year="1998">
<emphasis box="[725,1093,1468,1494]" italics="true" pageId="9" pageNumber="10">Chure, Fiorillo &amp; Jacobsen, 1998</emphasis>
</bibRefCitation>
). The afflicted vertebra is from the cranial-most part of the tail. Observations of the feeding behaviors of carnivoran mammals and birds indicate that in most cases, consumption of the axial skeleton occurs after limbs and viscera have been consumed (e.g.,
<bibRefCitation author="Hill AP" box="[947,1059,1587,1614]" editor="Behrensmeyer AK &amp; Hill AP" journalOrPublisher="Chicago: University of Chicago Press" pageId="9" pageNumber="10" pagination="131 - 152" refId="ref5823" refString="Hill AP. 1980. Early postmortem damage to the remains of some contemporary East African mammals. In: Behrensmeyer AK, Hill AP, eds. Fossils in the making. Vertebrate taphonomy and paleoecology. Chicago: University of Chicago Press, 131 - 152." title="Early postmortem damage to the remains of some contemporary East African mammals" type="book chapter" volumeTitle="Fossils in the making. Vertebrate taphonomy and paleoecology" year="1980">
<emphasis box="[947,1059,1587,1614]" italics="true" pageId="9" pageNumber="10">Hill, 1980</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Haglund WD" box="[1073,1242,1587,1614]" editor="Haglund WD &amp; Sorg MH" journalOrPublisher="Boca Raton: CRC Press" pageId="9" pageNumber="10" pagination="367 - 382" refId="ref5737" refString="Haglund WD. 1997. Dogs and coyotes: postermorem involvement with human remains. In: Haglund WD, Sorg MH, eds. Forensic taphonomy. Boca Raton: CRC Press, 367 - 382." title="Dogs and coyotes: postermorem involvement with human remains" type="book chapter" volumeTitle="Forensic taphonomy" year="1997">
<emphasis box="[1073,1242,1587,1614]" italics="true" pageId="9" pageNumber="10">Haglund, 1997</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Carson EA &amp; Stefan VH &amp; Powell JF" journalOrPublisher="Journal of Forensic Sciences" pageId="9" pageNumber="10" pagination="515 - 526" part="45" refId="ref5021" refString="Carson EA, Stefan VH, Powell JF. 2000. Skeletal manifestations of bear scavenging. Journal of Forensic Sciences 45 (3): 515 - 526." title="Skeletal manifestations of bear scavenging" type="journal article" year="2000">
<emphasis italics="true" pageId="9" pageNumber="10">Carson, Stefan &amp; Powell, 2000</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Behrensmeyer AK &amp; Stayton CT &amp; Chapman RE" box="[561,1034,1627,1654]" journalOrPublisher="Paleobiology" pageId="9" pageNumber="10" pagination="52 - 70" part="29" refId="ref4788" refString="Behrensmeyer AK, Stayton CT, Chapman RE. 2003. Taphonomy and ecology of modern avifaunal remains from Amboseli Park, Kenya. Paleobiology 29 (1): 52 - 70 DOI 10.1666 / 0094 - 8373 (2003) 029 &lt;0052: TAEOMA&gt; 2.0. CO; 2." title="Taphonomy and ecology of modern avifaunal remains from Amboseli Park, Kenya" type="journal article" year="2003">
<emphasis box="[561,1034,1627,1654]" italics="true" pageId="9" pageNumber="10">Behrensmeyer, Stayton &amp; Chapman, 2003</emphasis>
</bibRefCitation>
). Hadrosaur tails had substantial muscles such as m. ilio-ischocaudalis and m. caudofemoralis longus (
<bibRefCitation author="Persons WS &amp; Currie PJ" box="[1207,1469,1667,1693]" editor="Eberth DA &amp; Evans DE" journalOrPublisher="Bloomington and Indianapolis: Indiana University Press" pageId="9" pageNumber="10" pagination="449 - 458" refId="ref6363" refString="Persons WS, Currie PJ. 2014. Duckbills on the run: the cursorial abilities of hadrosaurs and implications for hadrosaur-avoidance strategies. In: Eberth DA, Evans DE, eds. Hadrosaurs. Bloomington and Indianapolis: Indiana University Press, 449 - 458." title="Duckbills on the run: the cursorial abilities of hadrosaurs and implications for hadrosaur-avoidance strategies" type="book chapter" volumeTitle="Hadrosaurs" year="2014">
<emphasis box="[1207,1469,1667,1693]" italics="true" pageId="9" pageNumber="10">Persons &amp; Currie, 2014</emphasis>
</bibRefCitation>
) that might be a target of early stage consumption. However, the ventral bite traces on BMR P2007.4.1 suggest that the tyrannosaur was feeding after the haemal complexes and most of the superficial hypaxial muscles and m. caudofemoralis longus had been removed. As such the punctures on BMR P2007.4.1 suggest later-stage carcass consumption and postmortem feeding behaviors.
</paragraph>
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<paragraph blockId="10.[507,1524,246,327]" pageId="10" pageNumber="11">
<emphasis bold="true" box="[507,1170,246,268]" pageId="10" pageNumber="11">Table 2 Measurements of tooth crowns of tyrannosaur specimens.</emphasis>
Mesiodistal and labiolingual measurements of teeth at 5 mm depth from the crown apex for (A)
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, (B)
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, and (C) the inferred bite marks on BMR P2007.4.1. All measurements are in mm.
</paragraph>
</caption>
<paragraph pageId="10" pageNumber="11">
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<emphasis bold="true" pageId="10" pageNumber="11">
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<th box="[514,607,351,403]" gridcol="0" gridrow="0" pageId="10" pageNumber="11">
A
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</th>
<th box="[855,1062,351,403]" gridcol="2" gridrow="0" pageId="10" pageNumber="11">Maxilla</th>
<th box="[1306,1517,351,403]" gridcol="4" gridrow="0" pageId="10" pageNumber="11">Dentary</th>
</tr>
<tr box="[514,1517,422,444]" gridrow="1" pageId="10" pageNumber="11" rowspan-0="1">
<td box="[714,829,422,444]" gridcol="1" gridrow="1" pageId="10" pageNumber="11">Mesiodistal</td>
<td box="[855,1062,422,444]" gridcol="2" gridrow="1" pageId="10" pageNumber="11">Labiolingual</td>
<td box="[1169,1284,422,444]" gridcol="3" gridrow="1" pageId="10" pageNumber="11">Mesiodistal</td>
<td box="[1306,1517,422,444]" gridcol="4" gridrow="1" pageId="10" pageNumber="11">Labiolingual</td>
</tr>
</emphasis>
<tr box="[514,1517,462,484]" gridrow="2" pageId="10" pageNumber="11" rowspan-0="1">
<td box="[714,829,462,484]" gridcol="1" gridrow="2" pageId="10" pageNumber="11">15</td>
<td box="[855,1062,462,484]" gridcol="2" gridrow="2" pageId="10" pageNumber="11">16.8</td>
<td box="[1169,1284,462,484]" gridcol="3" gridrow="2" pageId="10" pageNumber="11">9.3</td>
<td box="[1306,1517,462,484]" gridcol="4" gridrow="2" pageId="10" pageNumber="11">10.0</td>
</tr>
<tr box="[514,1517,498,520]" gridrow="3" pageId="10" pageNumber="11" rowspan-0="1">
<td box="[714,829,498,520]" gridcol="1" gridrow="3" pageId="10" pageNumber="11">11.4</td>
<td box="[855,1062,498,520]" gridcol="2" gridrow="3" pageId="10" pageNumber="11">8.2</td>
<td box="[1169,1284,498,520]" gridcol="3" gridrow="3" pageId="10" pageNumber="11">8.27</td>
<td box="[1306,1517,498,520]" gridcol="4" gridrow="3" pageId="10" pageNumber="11">10.27</td>
</tr>
<tr box="[514,1517,534,556]" gridrow="4" pageId="10" pageNumber="11" rowspan-0="1">
<td box="[714,829,534,556]" gridcol="1" gridrow="4" pageId="10" pageNumber="11">13.7</td>
<td box="[855,1062,534,556]" gridcol="2" gridrow="4" pageId="10" pageNumber="11">8.2</td>
<td box="[1169,1284,534,556]" gridcol="3" gridrow="4" pageId="10" pageNumber="11">6.8</td>
<td box="[1306,1517,534,556]" gridcol="4" gridrow="4" pageId="10" pageNumber="11">7.7</td>
</tr>
<tr box="[514,1517,570,592]" gridrow="5" pageId="10" pageNumber="11" rowspan-0="1" rowspan-1="1" rowspan-2="1">
<td box="[1169,1284,570,592]" gridcol="3" gridrow="5" pageId="10" pageNumber="11">9.4</td>
<td box="[1306,1517,570,592]" gridcol="4" gridrow="5" pageId="10" pageNumber="11">6.7</td>
</tr>
<tr box="[514,1517,606,628]" gridrow="6" pageId="10" pageNumber="11" rowspan-0="1" rowspan-1="1" rowspan-2="1">
<td box="[1169,1284,606,628]" gridcol="3" gridrow="6" pageId="10" pageNumber="11">9.7</td>
<td box="[1306,1517,606,628]" gridcol="4" gridrow="6" pageId="10" pageNumber="11">5.9</td>
</tr>
</table>
</paragraph>
<paragraph pageId="10" pageNumber="11">
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<emphasis bold="true" pageId="10" pageNumber="11">
<tr box="[514,1517,668,719]" gridrow="0" pageId="10" pageNumber="11">
<th box="[514,665,668,719]" gridcol="0" gridrow="0" pageId="10" pageNumber="11">
B
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</th>
<th box="[758,1091,668,719]" colspan="2" colspanRight="1" gridcol="1" gridrow="0" pageId="10" pageNumber="11">Maxilla</th>
<th box="[1183,1517,668,719]" colspan="2" colspanRight="1" gridcol="3" gridrow="0" pageId="10" pageNumber="11">Dentary</th>
</tr>
<tr box="[514,1517,739,761]" gridrow="1" pageId="10" pageNumber="11" rowspan-0="1">
<td box="[758,1091,739,761]" colspan="2" colspanRight="1" gridcol="1" gridrow="1" pageId="10" pageNumber="11">Mesiodistal Labiolingual</td>
<td box="[1183,1517,739,761]" colspan="2" colspanRight="1" gridcol="3" gridrow="1" pageId="10" pageNumber="11">Mesiodistal Labiolingual</td>
</tr>
</emphasis>
<tr box="[514,1517,778,800]" gridrow="2" pageId="10" pageNumber="11" rowspan-0="1">
<td box="[758,798,778,800]" gridcol="1" gridrow="2" pageId="10" pageNumber="11">6.77</td>
<td box="[965,1091,778,800]" gridcol="2" gridrow="2" pageId="10" pageNumber="11">4.4</td>
<td box="[1183,1223,778,800]" gridcol="3" gridrow="2" pageId="10" pageNumber="11">7.06</td>
<td box="[1391,1517,778,800]" gridcol="4" gridrow="2" pageId="10" pageNumber="11">4.39</td>
</tr>
<tr box="[514,1517,814,836]" gridrow="3" pageId="10" pageNumber="11" rowspan-0="1">
<td box="[758,798,814,836]" gridcol="1" gridrow="3" pageId="10" pageNumber="11">7.18</td>
<td box="[965,1091,814,836]" gridcol="2" gridrow="3" pageId="10" pageNumber="11">4.73</td>
<td box="[1183,1223,814,836]" gridcol="3" gridrow="3" pageId="10" pageNumber="11">6.54</td>
<td box="[1391,1517,814,836]" gridcol="4" gridrow="3" pageId="10" pageNumber="11">4.14</td>
</tr>
<tr box="[514,1517,850,872]" gridrow="4" pageId="10" pageNumber="11" rowspan-0="1">
<td box="[758,798,850,872]" gridcol="1" gridrow="4" pageId="10" pageNumber="11">7.35</td>
<td box="[965,1091,850,872]" gridcol="2" gridrow="4" pageId="10" pageNumber="11">3.9</td>
<td box="[1183,1223,850,872]" gridcol="3" gridrow="4" pageId="10" pageNumber="11">6.78</td>
<td box="[1391,1517,850,872]" gridcol="4" gridrow="4" pageId="10" pageNumber="11">3.77</td>
</tr>
<tr box="[514,1517,886,908]" gridrow="5" pageId="10" pageNumber="11" rowspan-0="1">
<td box="[758,798,886,908]" gridcol="1" gridrow="5" pageId="10" pageNumber="11">8.64</td>
<td box="[965,1091,886,908]" gridcol="2" gridrow="5" pageId="10" pageNumber="11">4.57</td>
<td box="[1183,1223,886,908]" gridcol="3" gridrow="5" pageId="10" pageNumber="11">7.25</td>
<td box="[1391,1517,886,908]" gridcol="4" gridrow="5" pageId="10" pageNumber="11">4.39</td>
</tr>
<tr box="[514,1517,923,945]" gridrow="6" pageId="10" pageNumber="11" rowspan-0="1">
<td box="[758,798,923,945]" gridcol="1" gridrow="6" pageId="10" pageNumber="11">7.91</td>
<td box="[965,1091,923,945]" gridcol="2" gridrow="6" pageId="10" pageNumber="11">4.19</td>
<td box="[1183,1223,923,945]" gridcol="3" gridrow="6" pageId="10" pageNumber="11">7.39</td>
<td box="[1391,1517,923,945]" gridcol="4" gridrow="6" pageId="10" pageNumber="11">4.47</td>
</tr>
<tr box="[514,1517,959,981]" gridrow="7" pageId="10" pageNumber="11" rowspan-0="1">
<td box="[758,798,959,981]" gridcol="1" gridrow="7" pageId="10" pageNumber="11">7.7</td>
<td box="[965,1091,959,981]" gridcol="2" gridrow="7" pageId="10" pageNumber="11">4.8</td>
<td box="[1183,1223,959,981]" gridcol="3" gridrow="7" pageId="10" pageNumber="11">7.48</td>
<td box="[1391,1517,959,981]" gridcol="4" gridrow="7" pageId="10" pageNumber="11">4.37</td>
</tr>
<tr box="[514,1517,995,1017]" gridrow="8" pageId="10" pageNumber="11" rowspan-0="1">
<td box="[758,798,995,1017]" gridcol="1" gridrow="8" pageId="10" pageNumber="11">8.74</td>
<td box="[965,1091,995,1017]" gridcol="2" gridrow="8" pageId="10" pageNumber="11">4.59</td>
<td box="[1183,1223,995,1017]" gridcol="3" gridrow="8" pageId="10" pageNumber="11">7.0</td>
<td box="[1391,1517,995,1017]" gridcol="4" gridrow="8" pageId="10" pageNumber="11">4.44</td>
</tr>
<tr box="[514,1517,1031,1053]" gridrow="9" pageId="10" pageNumber="11" rowspan-0="1">
<td box="[758,798,1031,1053]" gridcol="1" gridrow="9" pageId="10" pageNumber="11">8.52</td>
<td box="[965,1091,1031,1053]" gridcol="2" gridrow="9" pageId="10" pageNumber="11">4.21</td>
<td box="[1183,1223,1031,1053]" gridcol="3" gridrow="9" pageId="10" pageNumber="11">7.7</td>
<td box="[1391,1517,1031,1053]" gridcol="4" gridrow="9" pageId="10" pageNumber="11">4.24</td>
</tr>
<tr box="[514,1517,1067,1089]" gridrow="10" pageId="10" pageNumber="11" rowspan-0="1">
<td box="[758,798,1067,1089]" gridcol="1" gridrow="10" pageId="10" pageNumber="11">6.71</td>
<td box="[965,1091,1067,1089]" gridcol="2" gridrow="10" pageId="10" pageNumber="11">2.56</td>
<td box="[1183,1223,1067,1089]" gridcol="3" gridrow="10" pageId="10" pageNumber="11">7.19</td>
<td box="[1391,1517,1067,1089]" gridcol="4" gridrow="10" pageId="10" pageNumber="11">4.2</td>
</tr>
<tr box="[514,1517,1103,1125]" gridrow="11" pageId="10" pageNumber="11" rowspan-0="1" rowspan-1="1" rowspan-2="1">
<td box="[1183,1223,1103,1125]" gridcol="3" gridrow="11" pageId="10" pageNumber="11">6.34</td>
<td box="[1391,1517,1103,1125]" gridcol="4" gridrow="11" pageId="10" pageNumber="11">3.7</td>
</tr>
<tr box="[514,1517,1139,1161]" gridrow="12" pageId="10" pageNumber="11" rowspan-0="1" rowspan-1="1" rowspan-2="1">
<td box="[1183,1223,1139,1161]" gridcol="3" gridrow="12" pageId="10" pageNumber="11">7.7</td>
<td box="[1391,1517,1139,1161]" gridcol="4" gridrow="12" pageId="10" pageNumber="11">3.01</td>
</tr>
</table>
</paragraph>
<paragraph pageId="10" pageNumber="11">
<table box="[514,1517,1201,1328]" gridcols="3" gridrows="3" pageId="10" pageNumber="11">
<tr box="[514,1517,1201,1253]" gridrow="0" pageId="10" pageNumber="11">
<emphasis bold="true" box="[514,1517,1201,1253]" pageId="10" pageNumber="11">
<th box="[514,802,1201,1253]" gridcol="0" gridrow="0" pageId="10" pageNumber="11">C BMR P2007.4.1 ‘‘Bite Marks’’</th>
<th box="[1039,1154,1201,1253]" gridcol="1" gridrow="0" pageId="10" pageNumber="11">Mesiodistal</th>
<th box="[1391,1517,1201,1253]" gridcol="2" gridrow="0" pageId="10" pageNumber="11">Labiolingual</th>
</emphasis>
</tr>
<tr box="[514,1517,1270,1292]" gridrow="1" pageId="10" pageNumber="11" rowspan-0="1">
<td box="[1039,1154,1270,1292]" gridcol="1" gridrow="1" pageId="10" pageNumber="11">8.3</td>
<td box="[1391,1517,1270,1292]" gridcol="2" gridrow="1" pageId="10" pageNumber="11">4.31</td>
</tr>
<tr box="[514,1517,1306,1328]" gridrow="2" pageId="10" pageNumber="11" rowspan-0="1">
<td box="[1039,1154,1306,1328]" gridcol="1" gridrow="2" pageId="10" pageNumber="11">8.5</td>
<td box="[1391,1517,1306,1328]" gridcol="2" gridrow="2" pageId="10" pageNumber="11">4.3</td>
</tr>
</table>
</paragraph>
<paragraph blockId="10.[491,1541,1376,1880]" lastBlockId="11.[491,1542,1357,1901]" lastPageId="11" lastPageNumber="12" pageId="10" pageNumber="11">
The identification of penetrating bite marks attributable to not only
<taxonomicName authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[1322,1534,1376,1401]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="rex">
<emphasis box="[1322,1534,1376,1401]" italics="true" pageId="10" pageNumber="11">Tyrannosaurus rex</emphasis>
</taxonomicName>
, but an individual of 11–12 years of age can potentially allow for the determination of the ontogeny of bite force in
<taxonomicName authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[795,1007,1456,1481]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="rex">
<emphasis box="[795,1007,1456,1481]" italics="true" pageId="10" pageNumber="11">Tyrannosaurus rex</emphasis>
</taxonomicName>
and for comparison with other theropods (e.g.,
<bibRefCitation author="Barrett PM &amp; Rayfield EJ" box="[557,833,1496,1522]" journalOrPublisher="Trends in Ecology and Evolution" pageId="10" pageNumber="11" pagination="217 - 224" part="21" refId="ref4673" refString="Barrett PM, Rayfield EJ. 2006. Ecological and evolutionary implications of di- nosaur feeding behavior. Trends in Ecology and Evolution 21 (4): 217 - 224 DOI 10.1016 / j. tree. 2006.01.002." title="Ecological and evolutionary implications of di- nosaur feeding behavior" type="journal article" year="2006">
<emphasis box="[557,833,1496,1522]" italics="true" pageId="10" pageNumber="11">Barrett &amp; Rayfield, 2006</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Gignac PM &amp; Makovicky PJ &amp; Erickson GM &amp; Walsh RP" box="[848,1057,1495,1522]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="10" pageNumber="11" pagination="1169 - 1177" part="30" refId="ref5689" refString="Gignac PM, Makovicky PJ, Erickson GM, Walsh RP. 2010. A description of Deinonychus antirrhopus bite marks and estimates of bite force using tooth indentation simulations. Journal of Vertebrate Paleontology 30 (4): 1169 - 1177 DOI 10.1080 / 02724634.2010.483535." title="A description of Deinonychus antirrhopus bite marks and estimates of bite force using tooth indentation simulations" type="journal article" year="2010">
<emphasis box="[848,1057,1495,1522]" italics="true" pageId="10" pageNumber="11">Gignac et al., 2010</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Bates KT &amp; Falkingham PL" box="[1072,1371,1495,1522]" journalOrPublisher="Biology Letters" pageId="10" pageNumber="11" pagination="660 - 664" part="8" refId="ref4713" refString="Bates KT, Falkingham PL. 2012. Estimating maximum bite force in Tyrannosaurus rex using multi-body dynamics. Biology Letters 8: 660 - 664 DOI 10.1098 / rsbl. 2012.0056." title="Estimating maximum bite force in Tyrannosaurus rex using multi-body dynamics" type="journal article" year="2012">
<emphasis box="[1072,1371,1495,1522]" italics="true" pageId="10" pageNumber="11">Bates &amp; Falkingham, 2012</emphasis>
</bibRefCitation>
). Studies on the estimated bite forces of an adult
<taxonomicName authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[933,1146,1536,1561]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="rex">
<emphasis box="[933,1146,1536,1561]" italics="true" pageId="10" pageNumber="11">Tyrannosaurus rex</emphasis>
</taxonomicName>
have yielded a wide range of results. Estimates based on muscle volume proposed bite forces between 8,526 and 34,522 N, coupled with tooth pressures of 718–2,974 MPa, and a unique tooth morphology and arrangement to promote fine fragmentation of bone during osteophagy (
<bibRefCitation author="Gignac PM &amp; Erickson GM" journalOrPublisher="Scientific Reports" pageId="10" pageNumber="11" pagination="1 - 10" part="7" refId="ref5650" refString="Gignac PM, Erickson GM. 2017. The biomechanics behind extreme osteophagy in Tyrannosaurus rex. Scientific Reports 7 (2012): 1 - 10 DOI 10.1038 / s 41598 - 017 - 02161 - w." title="The biomechanics behind extreme osteophagy in Tyrannosaurus rex" type="journal article" year="2017">
<emphasis italics="true" pageId="10" pageNumber="11">Gignac &amp; Erickson, 2017</emphasis>
</bibRefCitation>
). However, estimates incorporating likely muscle fiber length produced results over 64,000 N for adult
<taxonomicName authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[864,931,1735,1760]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="rex">
<emphasis box="[864,931,1735,1760]" italics="true" pageId="10" pageNumber="11">T. rex</emphasis>
</taxonomicName>
(
<bibRefCitation author="Bates KT &amp; Falkingham PL" box="[952,1251,1734,1761]" journalOrPublisher="Journal of Anatomy" pageId="10" pageNumber="11" pagination="635 - 635" part="233" refId="ref4746" refString="Bates KT, Falkingham PL. 2018. The importance of muscle architecture in biomechanical reconstructures of extinct animals: a case study using Tyrannosaurus rex. Journal of Anatomy 233: 635 - 635 DOI 10.1111 / joa. 12874." title="The importance of muscle architecture in biomechanical reconstructures of extinct animals: a case study using Tyrannosaurus rex" type="journal article" year="2018">
<emphasis box="[952,1251,1734,1761]" italics="true" pageId="10" pageNumber="11">Bates &amp; Falkingham, 2018</emphasis>
</bibRefCitation>
). Juvenile
<taxonomicName authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[1381,1447,1735,1760]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="rex">
<emphasis box="[1381,1447,1735,1760]" italics="true" pageId="10" pageNumber="11">T. rex</emphasis>
</taxonomicName>
such as
<materialsCitation ID-GBIF-Occurrence="3335984320" box="[524,709,1775,1801]" collectionCode="BMR" pageId="10" pageNumber="11" specimenCode="BMR P2002.4.1">BMR P2002.4.1</materialsCitation>
have much narrower and blade-like tooth morphologies and were unlikely to have been able to withstand similar bite forces at this ontogenetic stage.
<bibRefCitation author="Bates KT &amp; Falkingham PL" journalOrPublisher="Biology Letters" pageId="10" pageNumber="11" pagination="660 - 664" part="8" refId="ref4713" refString="Bates KT, Falkingham PL. 2012. Estimating maximum bite force in Tyrannosaurus rex using multi-body dynamics. Biology Letters 8: 660 - 664 DOI 10.1098 / rsbl. 2012.0056." title="Estimating maximum bite force in Tyrannosaurus rex using multi-body dynamics" type="journal article" year="2012">
<emphasis italics="true" pageId="10" pageNumber="11">Bates &amp; Falkingham (2012)</emphasis>
</bibRefCitation>
estimated a maximum bite force for
<materialsCitation ID-GBIF-Occurrence="3335984304" box="[1175,1358,1854,1880]" collectionCode="BMR" pageId="10" pageNumber="11" specimenCode="BMR P2002.4.1">BMR P2002.4.1</materialsCitation>
at 2,400–3,850 N, and hypothesized that an increase in bite force during growth could indicate a change in feeding behavior and dietary partitioning while approaching adulthood.
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.3749274" ID-Zenodo-Dep="3749274" httpUri="https://zenodo.org/record/3749274/files/figure.png" pageId="11" pageNumber="12" startId="11.[507,571,1192,1214]" targetBox="[492,1451,231,1146]" targetPageId="11">
<paragraph blockId="11.[507,1526,1192,1303]" pageId="11" pageNumber="12">
<emphasis bold="true" box="[507,1279,1192,1214]" pageId="11" pageNumber="12">
Figure 9 Digitized comparisons between
<materialsCitation ID-GBIF-Occurrence="3335984306" box="[923,1075,1192,1214]" collectionCode="BMR" pageId="11" pageNumber="12" specimenCode="BMR P2002.4.1">BMR P2002.4.1</materialsCitation>
and BMR P2007.4.1.
</emphasis>
Interactive manipulation of digitized NextEngine 3D scan of a cast of the right maxilla and dentary of
<materialsCitation ID-GBIF-Occurrence="3335984317" box="[1236,1388,1222,1244]" collectionCode="BMR" pageId="11" pageNumber="12" specimenCode="BMR P2002.4.1">BMR P2002.4.1</materialsCitation>
, and BMR P2007.4.1 caudal vertebra. Full-size DOI: 10.7717/peerj.6573/fig-9
</paragraph>
</caption>
<paragraph blockId="11.[491,1542,1357,1901]" pageId="11" pageNumber="12">
Observations on extant crocodylians have documented a wide variety of dietary partitioning during ontogeny (e.g.,
<bibRefCitation author="Tucker AD &amp; Limpus CJ &amp; McCallum HI &amp; McDonald KR" box="[908,1116,1477,1503]" journalOrPublisher="Copeia" pageId="11" pageNumber="12" pagination="978 - 988" part="1996" refId="ref6621" refString="Tucker AD, Limpus CJ, McCallum HI, McDonald KR. 1996. Ontogenetic dietary partitioning by Crocodylus johnstoni during the dry season. Copeia 1996 (4): 978 - 988 DOI 10.2307 / 1447661." title="Ontogenetic dietary partitioning by Crocodylus johnstoni during the dry season" type="journal article" year="1996">
<emphasis box="[908,1116,1477,1503]" italics="true" pageId="11" pageNumber="12">Tucker et al., 1996</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Platt SG &amp; Rainwater TR &amp; Finger AG &amp; Thorbjarnarson JB &amp; Anderson TA &amp; McMurry ST" box="[1130,1314,1477,1503]" journalOrPublisher="Herpetological Journal" pageId="11" pageNumber="12" pagination="281 - 290" part="16" refId="ref6499" refString="Platt SG, Rainwater TR, Finger AG, Thorbjarnarson JB, Anderson TA, McMurry ST. 2006. Food habits, ontogenetic dietary partitioning and observations of foraging behaviour of Morelet's crocodile (Crocodylus moreletii) in northern Belize. Herpetological Journal 16: 281 - 290." title="Food habits, ontogenetic dietary partitioning and observations of foraging behaviour of Morelet's crocodile (Crocodylus moreletii) in northern Belize" type="journal article" year="2006">
<emphasis box="[1130,1314,1477,1503]" italics="true" pageId="11" pageNumber="12">Platt et al., 2006</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Platt SG &amp; Thorbjarnarson JB &amp; Rainwater TR &amp; Martin DR" box="[1329,1513,1476,1503]" journalOrPublisher="Journal of Herpetology" pageId="11" pageNumber="12" pagination="1 - 10" part="47" refId="ref6549" refString="Platt SG, Thorbjarnarson JB, Rainwater TR, Martin DR. 2013. Diet of the American crocodile (Crocodylus acutus) in marine environments of coastal Belize. Journal of Herpetology 47 (1): 1 - 10 DOI 10.1670 / 12 - 077." title="Diet of the American crocodile (Crocodylus acutus) in marine environments of coastal Belize" type="journal article" year="2013">
<emphasis box="[1329,1513,1476,1503]" italics="true" pageId="11" pageNumber="12">Platt et al., 2013</emphasis>
</bibRefCitation>
).
</paragraph>
<paragraph blockId="11.[491,1542,1357,1901]" pageId="11" pageNumber="12">
In the American crocodile (
<taxonomicName baseAuthorityName="Cuvier" baseAuthorityYear="1807" box="[812,1016,1516,1542]" class="Reptilia" family="Crocodylidae" genus="Crocodylus" kingdom="Animalia" order="Crocodylia" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="acutus">
<emphasis box="[812,1016,1516,1542]" italics="true" pageId="11" pageNumber="12">Crocodylus acutus</emphasis>
</taxonomicName>
), hatchling and small juveniles have a dietary overlap of over 80%, commonly feeding upon insects and crustaceans (
<bibRefCitation author="Platt SG &amp; Thorbjarnarson JB &amp; Rainwater TR &amp; Martin DR" box="[1330,1514,1556,1582]" journalOrPublisher="Journal of Herpetology" pageId="11" pageNumber="12" pagination="1 - 10" part="47" refId="ref6549" refString="Platt SG, Thorbjarnarson JB, Rainwater TR, Martin DR. 2013. Diet of the American crocodile (Crocodylus acutus) in marine environments of coastal Belize. Journal of Herpetology 47 (1): 1 - 10 DOI 10.1670 / 12 - 077." title="Diet of the American crocodile (Crocodylus acutus) in marine environments of coastal Belize" type="journal article" year="2013">
<emphasis box="[1330,1514,1556,1582]" italics="true" pageId="11" pageNumber="12">Platt et al., 2013</emphasis>
</bibRefCitation>
). Alternatively, larger juveniles, subadults, and adults possess a dietary overlap of over 75%, consisting of more birds, mammals, fish, and other reptiles (
<bibRefCitation author="Platt SG &amp; Thorbjarnarson JB &amp; Rainwater TR &amp; Martin DR" box="[1189,1374,1635,1662]" journalOrPublisher="Journal of Herpetology" pageId="11" pageNumber="12" pagination="1 - 10" part="47" refId="ref6549" refString="Platt SG, Thorbjarnarson JB, Rainwater TR, Martin DR. 2013. Diet of the American crocodile (Crocodylus acutus) in marine environments of coastal Belize. Journal of Herpetology 47 (1): 1 - 10 DOI 10.1670 / 12 - 077." title="Diet of the American crocodile (Crocodylus acutus) in marine environments of coastal Belize" type="journal article" year="2013">
<emphasis box="[1189,1374,1635,1662]" italics="true" pageId="11" pageNumber="12">Platt et al., 2013</emphasis>
</bibRefCitation>
). Comparable ontogenetic dietary partitions were also observed in Morelet’s crocodile (
<taxonomicName baseAuthorityName="Duméril &amp; Bibron" baseAuthorityYear="1851" class="Reptilia" family="Crocodylidae" genus="Crocodylus" kingdom="Animalia" order="Crocodylia" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="moreletii">
<emphasis italics="true" pageId="11" pageNumber="12">Crocodylus moreletii</emphasis>
</taxonomicName>
) (
<bibRefCitation author="Platt SG &amp; Rainwater TR &amp; Finger AG &amp; Thorbjarnarson JB &amp; Anderson TA &amp; McMurry ST" box="[619,803,1716,1742]" journalOrPublisher="Herpetological Journal" pageId="11" pageNumber="12" pagination="281 - 290" part="16" refId="ref6499" refString="Platt SG, Rainwater TR, Finger AG, Thorbjarnarson JB, Anderson TA, McMurry ST. 2006. Food habits, ontogenetic dietary partitioning and observations of foraging behaviour of Morelet's crocodile (Crocodylus moreletii) in northern Belize. Herpetological Journal 16: 281 - 290." title="Food habits, ontogenetic dietary partitioning and observations of foraging behaviour of Morelet's crocodile (Crocodylus moreletii) in northern Belize" type="journal article" year="2006">
<emphasis box="[619,803,1716,1742]" italics="true" pageId="11" pageNumber="12">Platt et al., 2006</emphasis>
</bibRefCitation>
), and in Australian freshwater crocodiles (
<taxonomicName box="[1295,1530,1716,1742]" class="Reptilia" family="Crocodylidae" genus="Crocodylus" kingdom="Animalia" order="Crocodylia" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="johnstoni">
<emphasis box="[1295,1530,1716,1742]" italics="true" pageId="11" pageNumber="12">Crocodylus johnstoni</emphasis>
</taxonomicName>
) (
<bibRefCitation author="Tucker AD &amp; Limpus CJ &amp; McCallum HI &amp; McDonald KR" box="[502,710,1755,1781]" journalOrPublisher="Copeia" pageId="11" pageNumber="12" pagination="978 - 988" part="1996" refId="ref6621" refString="Tucker AD, Limpus CJ, McCallum HI, McDonald KR. 1996. Ontogenetic dietary partitioning by Crocodylus johnstoni during the dry season. Copeia 1996 (4): 978 - 988 DOI 10.2307 / 1447661." title="Ontogenetic dietary partitioning by Crocodylus johnstoni during the dry season" type="journal article" year="1996">
<emphasis box="[502,710,1755,1781]" italics="true" pageId="11" pageNumber="12">Tucker et al., 1996</emphasis>
</bibRefCitation>
). However, crocodylians are less discriminant of food sources when scavenging (e.g.,
<bibRefCitation author="Antunes MT" box="[689,851,1795,1821]" journalOrPublisher="Anuario do Instituto de Geociencias" pageId="11" pageNumber="12" pagination="117 - 130" part="40" refId="ref4631" refString="Antunes MT. 2017. Huge miocene crocodilians from western Europe: predation, comparisons with the '' False Gharial' ' and size. Anuario do Instituto de Geociencias 40 (3): 117 - 130 DOI 10.11137 / 2017.3.117.130." title="Huge miocene crocodilians from western Europe: predation, comparisons with the '' False Gharial' ' and size" type="journal article" year="2017">
<emphasis box="[689,851,1795,1821]" italics="true" pageId="11" pageNumber="12">Antunes, 2017</emphasis>
</bibRefCitation>
). While the punctures present on BMR P2007.4.1 are likely from postmortem scavenging behaviors of a juvenile tyrannosaur, the degree of dietary overlap or partitioning between juvenile and adult tyrannosaurs remains unresolved.
</paragraph>
<caption ID-Table-UUID="1F9E6636FF94FFEEFE44AFD1885F674F" httpUri="http://table.plazi.org/id/1F9E6636FF94FFEEFE44AFD1885F674F" pageId="12" pageNumber="13" targetBox="[514,1517,351,862]" targetIsTable="true" targetPageId="12">
<paragraph blockId="12.[507,1507,246,327]" pageId="12" pageNumber="13">
<emphasis bold="true" box="[507,1182,246,268]" pageId="12" pageNumber="13">Table 3 Measurements of crown spacing in tyrannosaur specimens.</emphasis>
Tooth crown spacing between maxillary (A) and dentary (B) teeth in the juvenile tyrannosaur
<materialsCitation ID-GBIF-Occurrence="3335984321" box="[1115,1267,276,298]" collectionCode="BMR" pageId="12" pageNumber="13" specimenCode="BMR P2002.4.1">BMR P2002.4.1</materialsCitation>
. All measurements are in mm.
</paragraph>
</caption>
<paragraph pageId="12" pageNumber="13">
<table box="[514,1517,351,862]" gridcols="2" gridrows="12" pageId="12" pageNumber="13">
<tr box="[514,1517,351,403]" gridrow="0" pageId="12" pageNumber="13">
<th box="[514,661,351,403]" gridcol="0" gridrow="0" pageId="12" pageNumber="13">
<emphasis bold="true" box="[514,661,351,403]" pageId="12" pageNumber="13">A Crown spacing</emphasis>
</th>
<th box="[1359,1517,351,403]" gridcol="1" gridrow="0" pageId="12" pageNumber="13">
<emphasis bold="true" box="[1359,1517,381,403]" pageId="12" pageNumber="13">Maxillary (mm)</emphasis>
</th>
</tr>
<tr box="[514,1517,419,442]" gridrow="1" pageId="12" pageNumber="13">
<td box="[514,661,419,442]" gridcol="0" gridrow="1" pageId="12" pageNumber="13">
<emphasis bold="true" box="[514,549,419,440]" pageId="12" pageNumber="13">4–6</emphasis>
</td>
<td box="[1359,1517,419,442]" gridcol="1" gridrow="1" pageId="12" pageNumber="13">70.2</td>
</tr>
<tr box="[514,1517,456,478]" gridrow="2" pageId="12" pageNumber="13">
<td box="[514,661,456,478]" gridcol="0" gridrow="2" pageId="12" pageNumber="13">
<emphasis bold="true" box="[514,549,456,477]" pageId="12" pageNumber="13">6–8</emphasis>
</td>
<td box="[1359,1517,456,478]" gridcol="1" gridrow="2" pageId="12" pageNumber="13">73.3</td>
</tr>
<tr box="[514,1517,492,514]" gridrow="3" pageId="12" pageNumber="13">
<td box="[514,661,492,514]" gridcol="0" gridrow="3" pageId="12" pageNumber="13">
<emphasis bold="true" box="[514,561,492,513]" pageId="12" pageNumber="13">8–10</emphasis>
</td>
<td box="[1359,1517,492,514]" gridcol="1" gridrow="3" pageId="12" pageNumber="13">62.8</td>
</tr>
<tr box="[514,1517,528,550]" gridrow="4" pageId="12" pageNumber="13">
<td box="[514,661,528,550]" gridcol="0" gridrow="4" pageId="12" pageNumber="13">
<emphasis bold="true" box="[514,594,528,550]" pageId="12" pageNumber="13">Average</emphasis>
</td>
<td box="[1359,1517,528,550]" gridcol="1" gridrow="4" pageId="12" pageNumber="13">68.7</td>
</tr>
<tr box="[514,1517,591,642]" gridrow="5" pageId="12" pageNumber="13">
<td box="[514,661,591,642]" gridcol="0" gridrow="5" pageId="12" pageNumber="13">
<emphasis bold="true" box="[514,661,591,642]" pageId="12" pageNumber="13">B Crown spacing</emphasis>
</td>
<td box="[1359,1517,591,642]" gridcol="1" gridrow="5" pageId="12" pageNumber="13">
<emphasis bold="true" box="[1378,1516,620,642]" pageId="12" pageNumber="13">Dentary(mm)</emphasis>
</td>
</tr>
<tr box="[514,1517,658,681]" gridrow="6" pageId="12" pageNumber="13">
<td box="[514,661,658,681]" gridcol="0" gridrow="6" pageId="12" pageNumber="13">
<emphasis bold="true" box="[514,549,658,679]" pageId="12" pageNumber="13">4–6</emphasis>
</td>
<td box="[1359,1517,658,681]" gridcol="1" gridrow="6" pageId="12" pageNumber="13">53.3</td>
</tr>
<tr box="[514,1517,694,717]" gridrow="7" pageId="12" pageNumber="13">
<td box="[514,661,694,717]" gridcol="0" gridrow="7" pageId="12" pageNumber="13">
<emphasis bold="true" box="[514,549,694,715]" pageId="12" pageNumber="13">6–8</emphasis>
</td>
<td box="[1359,1517,694,717]" gridcol="1" gridrow="7" pageId="12" pageNumber="13">49.8</td>
</tr>
<tr box="[514,1517,731,753]" gridrow="8" pageId="12" pageNumber="13">
<td box="[514,661,731,753]" gridcol="0" gridrow="8" pageId="12" pageNumber="13">
<emphasis bold="true" box="[514,561,731,752]" pageId="12" pageNumber="13">8–10</emphasis>
</td>
<td box="[1359,1517,731,753]" gridcol="1" gridrow="8" pageId="12" pageNumber="13">39.2</td>
</tr>
<tr box="[514,1517,767,789]" gridrow="9" pageId="12" pageNumber="13">
<td box="[514,661,767,789]" gridcol="0" gridrow="9" pageId="12" pageNumber="13">
<emphasis bold="true" box="[514,572,767,788]" pageId="12" pageNumber="13">10–12</emphasis>
</td>
<td box="[1359,1517,767,789]" gridcol="1" gridrow="9" pageId="12" pageNumber="13">33.7</td>
</tr>
<tr box="[514,1517,803,825]" gridrow="10" pageId="12" pageNumber="13">
<td box="[514,661,803,825]" gridcol="0" gridrow="10" pageId="12" pageNumber="13">
<emphasis bold="true" box="[514,572,803,824]" pageId="12" pageNumber="13">12–14</emphasis>
</td>
<td box="[1359,1517,803,825]" gridcol="1" gridrow="10" pageId="12" pageNumber="13">33</td>
</tr>
<tr box="[514,1517,839,862]" gridrow="11" pageId="12" pageNumber="13">
<td box="[514,661,839,862]" gridcol="0" gridrow="11" pageId="12" pageNumber="13">
<emphasis bold="true" box="[514,594,839,861]" pageId="12" pageNumber="13">Average</emphasis>
</td>
<td box="[1359,1517,839,862]" gridcol="1" gridrow="11" pageId="12" pageNumber="13">41.8</td>
</tr>
</table>
</paragraph>
<paragraph blockId="12.[491,1542,910,1215]" pageId="12" pageNumber="13">
Despite not yet possessing the same feeding mechanisms of an adult
<taxonomicName authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="rex">
<emphasis italics="true" pageId="12" pageNumber="13">Tyrannosaurus rex</emphasis>
</taxonomicName>
(i.e., bone-crushing and osteophagy), the punctures present on BMR P2007.4.1 demonstrate that late-stage juvenile and subadult tyrannosaurs were already biomechanically capable of puncturing bone during feeding, and were doing so without the large, blunt dental crowns of adults. Further identification of tyrannosaur feeding traces from different ontogenetic stages coupled with experimental studies of the biomechanics of tyrannosaur bite forces may reveal more insight into dynamic dietary trends and ecological role of
<taxonomicName authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[575,787,1189,1214]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="rex">
<emphasis box="[575,787,1189,1214]" italics="true" pageId="12" pageNumber="13">Tyrannosaurus rex</emphasis>
</taxonomicName>
throughout ontogeny.
</paragraph>
</subSubSection>
</treatment>
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347
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<emphasis id="7995EAB5B333CD611BC1F0B2FEE4FF64" bold="true" box="[151,370,220,246]" italics="true" pageId="1" pageNumber="551">Zorotypus weiweii</emphasis>
Wang, Li &amp; Cai
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,
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, 28–30
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material
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.
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apterous male,
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:
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,
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,
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, 5˚26ʼ55.7ˮN, 116˚27ʼ08.6ˮE, rainforest, under bark of decaying log,
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, coll.
<collectorName id="E6145371B333CD611F78F103FB5BFE17" box="[1070,1229,364,389]" pageId="1" pageNumber="551">Wang Jianyun</collectorName>
&amp;
<collectorName id="E6145371B333CD611FA4F102FA0FFE17" box="[1266,1433,364,389]" pageId="1" pageNumber="551">Zhang Weiwei</collectorName>
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.
<materialsCitation id="FB893CFAB333CD611BC1F1FEFA0FFE5F" ID-GBIF-Occurrence="1306554733" collectingDate="2015-10-09" collectorName="Liang Feiyang &amp; Zhang Weiwei" country="Malaysia" elevation="962" location="Borneo Jungle Girl Camp" municipality="Keningau" pageId="1" pageNumber="551" specimenCount="2" stateProvince="Sabah" typeStatus="paratype">
<typeStatus id="945A8805B333CD611BC1F1FEFE91FE3A" box="[151,263,400,424]" pageId="1" pageNumber="551" type="paratype">Paratypes</typeStatus>
, 4 dealate males, 2 dealate females, same location data as
<typeStatus id="945A8805B333CD6118F0F1E1FB9DFE3A" box="[934,1035,399,424]" pageId="1" pageNumber="551" type="holotype">holotype</typeStatus>
; 2 apterous males, same rainforest, 5˚27ʼ03.9ˮN, 116˚26ʼ52.3ˮE, under bark of decaying log,
<elevation id="C0CCD194B333CD611849F1DAFCFEFE5E" box="[799,872,436,461]" metricMagnitude="2" metricUnit="m" metricValue="9.62" pageId="1" pageNumber="551" unit="m" value="962.0">
<quantity id="8C199B42B333CD611849F1DAFCFEFE5E" box="[799,872,436,461]" metricMagnitude="2" metricUnit="m" metricValue="9.62" pageId="1" pageNumber="551" unit="m" value="962.0">962 m</quantity>
</elevation>
,
<collectingDate id="2F1BE98FB333CD611825F1DAFC7DFE5F" box="[883,1003,436,461]" pageId="1" pageNumber="551" value="2015-10-09">
<date id="3F5F1067B333CD611825F1DAFC7DFE5F" box="[883,1003,436,461]" pageId="1" pageNumber="551" value="2015-10-09">9. X. 2015</date>
</collectingDate>
, coll.
<collectorName id="E6145371B333CD611F7BF1DAFB59FE5F" box="[1069,1231,436,461]" pageId="1" pageNumber="551">Liang Feiyang</collectorName>
&amp;
<collectorName id="E6145371B333CD611FA4F1DAFA0FFE5F" box="[1266,1433,436,461]" pageId="1" pageNumber="551">Zhang Weiwei</collectorName>
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="03FB652CB333CD611B91F1B9FE25FD36" pageId="1" pageNumber="551" type="diagnosis">
<paragraph id="4B5E36A7B333CD611B91F1B9FE25FD36" blockId="1.[151,1437,327,677]" pageId="1" pageNumber="551">
<emphasis id="7995EAB5B333CD611B91F1B9FED7FE62" bold="true" box="[199,321,471,496]" pageId="1" pageNumber="551">Diagnosis.</emphasis>
The new species is similar to
<taxonomicName id="8CE14D24B333CD6119FCF1B7FCA5FE62" box="[682,819,472,496]" class="Insecta" family="Zorotypidae" genus="Zorotypus" kingdom="Animalia" order="Zoraptera" pageId="1" pageNumber="551" phylum="Arthropoda" rank="species" species="impolitus">
<emphasis id="7995EAB5B333CD6119FCF1B7FCA5FE62" box="[682,819,472,496]" italics="true" pageId="1" pageNumber="551">Z. impolitus</emphasis>
</taxonomicName>
, but it can be easily separated from the latter by the following characters: curly setae denser on vertex of male, forming a “V” shape (
<emphasis id="7995EAB5B333CD611F6AF193FBC5FD86" box="[1084,1107,509,532]" italics="true" pageId="1" pageNumber="551">vs</emphasis>
. male vertex setae relatively sparse); number of stout bristles on ventral surface of metafemur ten (
<emphasis id="7995EAB5B333CD61189CF24FFC7FFDAA" box="[970,1001,545,568]" italics="true" pageId="1" pageNumber="551">vs.</emphasis>
eight to nine); male ninth abdominal tergum (T9) sclerotized (
<emphasis id="7995EAB5B333CD611AEFF22BFE46FDCE" box="[441,464,581,604]" italics="true" pageId="1" pageNumber="551">vs</emphasis>
. T9 scarcely sclerotized); male T11 with larger sclerotized area, extended to lateral margin (
<emphasis id="7995EAB5B333CD611BAEF207FE99FD12" box="[248,271,617,640]" italics="true" pageId="1" pageNumber="551">vs</emphasis>
. T11 laterally scarcely sclerotized); and the male genitalia relatively stronger, ventral sclerite broader (
<figureCitation id="D3DA2A22B333CD611BC9F2E2FEB0FD36" box="[159,294,652,677]" captionStart="FIGURES 2 – 10" captionStartId="2.[151,264,1904,1927]" captionTargetBox="[202,1379,202,1861]" captionTargetId="figure@2.[192,1388,193,1872]" captionTargetPageId="2" captionText="FIGURES 2 – 10. Zorotypus weiweii sp. nov., antennae and appendages removed. 2, head of apterous male; 3, head of alate female; 4, thorax of apterous male; 5, thorax of dealate female; 6, 7, abdomen of apterous male; 8, abdomen of dealate female; 9, terminalia of male; 10, terminalia of female. Figs. 2, 3, anterior view; 4 – 6, dorsal view; 7, 8, ventral view; 9, 10, posterior view. Scale bar: Figs. 2, 3, 9, 10 = 0.4 mm; 4 – 8 = 0.2 mm." httpUri="https://zenodo.org/record/256278/files/figure.png" pageId="1" pageNumber="551">Figs. 2, 6, 9</figureCitation>
,
<figureCitation id="D3DA2A22B333CD611A64F2E2FE31FD36" box="[306,423,652,677]" captionStart="FIGURES 16 – 23" captionStartId="4.[151,264,1890,1913]" captionTargetBox="[202,1412,207,1829]" captionTargetId="figure@4.[161,1446,193,1845]" captionTargetPageId="4" captionText="FIGURES 16 – 23. Apterous male of Zorotypus weiweii sp. nov.. 16, right metaleg; 17, 18, posterior segments of abdomen; 19, left cercus; 20 – 23, male genitalia. 16, posterior view; 17, lateral view, posterior to the bottom 18 – 21, dorsal view; 22, 23, different angles of male genitalia. Scale bar: Fig. 16 = 0.3 mm; 17, 18, 20 = 0.45 mm; 19 = 0.30 mm; 21 – 23 = 0.15 mm." httpUri="https://zenodo.org/record/256280/files/figure.png" pageId="1" pageNumber="551">16, 20–23</figureCitation>
).
</paragraph>
</subSubSection>
<caption id="1F9E662FB333CD611BC1F503FC6DFA11" box="[151,1019,1389,1411]" httpUri="https://zenodo.org/record/256277/files/figure.png" pageId="1" pageNumber="551" targetBox="[188,1412,783,1323]" targetPageId="1">
<paragraph id="4B5E36A7B333CD611BC1F503FC6DFA11" blockId="1.[151,1019,1389,1411]" box="[151,1019,1389,1411]" pageId="1" pageNumber="551">
<emphasis id="7995EAB5B333CD611BC1F503FE85FA11" bold="true" box="[151,275,1389,1411]" pageId="1" pageNumber="551">FIGURE 1.</emphasis>
<taxonomicName id="8CE14D24B333CD611A4FF500FE44FA11" box="[281,466,1390,1411]" class="Insecta" family="Zorotypidae" genus="Zorotypus" kingdom="Animalia" order="Zoraptera" pageId="1" pageNumber="551" phylum="Arthropoda" rank="species" species="weiweii" status="sp. nov.">
<emphasis id="7995EAB5B333CD611A4FF500FE44FA11" box="[281,466,1390,1411]" italics="true" pageId="1" pageNumber="551">Zorotypus weiweii</emphasis>
</taxonomicName>
<taxonomicNameLabel id="62A657CEB333CD611A8FF500FDBFFA11" box="[473,553,1390,1411]" pageId="1" pageNumber="551" rank="species">
<emphasis id="7995EAB5B333CD611A8FF500FDBFFA11" bold="true" box="[473,553,1390,1411]" pageId="1" pageNumber="551">sp. nov.</emphasis>
</taxonomicNameLabel>
Holotype, male, habitus. Scale bar = 0.5 mm.
</paragraph>
</caption>
<subSubSection id="03FB652CB333CD661B91F5C5FAE2F855" lastPageId="6" lastPageNumber="556" pageId="1" pageNumber="551" type="description">
<paragraph id="4B5E36A7B333CD611B91F5C5FD86F8BE" blockId="1.[151,1437,1451,2016]" pageId="1" pageNumber="551">
<emphasis id="7995EAB5B333CD611B91F5C5FD87FA56" bold="true" box="[199,529,1451,1476]" pageId="1" pageNumber="551">Description. Apterous male</emphasis>
(
<figureCitation id="D3DA2A22B333CD611977F5C5FDE6FA56" box="[545,624,1451,1476]" captionStart="FIGURE 1" captionStartId="1.[151,250,1389,1411]" captionTargetBox="[188,1412,783,1323]" captionTargetId="figure@1.[151,1436,743,1368]" captionTargetPageId="1" captionText="FIGURE 1. Zorotypus weiweii sp. nov. Holotype, male, habitus. Scale bar = 0.5 mm." httpUri="https://zenodo.org/record/256277/files/figure.png" pageId="1" pageNumber="551">Figs. 1</figureCitation>
,
<figureCitation id="D3DA2A22B333CD611929F5C2FC92FA56" box="[639,772,1451,1476]" captionStart="FIGURES 2 – 10" captionStartId="2.[151,264,1904,1927]" captionTargetBox="[202,1379,202,1861]" captionTargetId="figure@2.[192,1388,193,1872]" captionTargetPageId="2" captionText="FIGURES 2 – 10. Zorotypus weiweii sp. nov., antennae and appendages removed. 2, head of apterous male; 3, head of alate female; 4, thorax of apterous male; 5, thorax of dealate female; 6, 7, abdomen of apterous male; 8, abdomen of dealate female; 9, terminalia of male; 10, terminalia of female. Figs. 2, 3, anterior view; 4 – 6, dorsal view; 7, 8, ventral view; 9, 10, posterior view. Scale bar: Figs. 2, 3, 9, 10 = 0.4 mm; 4 – 8 = 0.2 mm." httpUri="https://zenodo.org/record/256278/files/figure.png" pageId="1" pageNumber="551">2, 4, 6, 7, 9</figureCitation>
,
<figureCitation id="D3DA2A22B333CD611844F5C2FCCFFA56" box="[786,857,1452,1476]" captionStart-0="FIGURES 11 – 15" captionStart-1="FIGURES 16 – 23" captionStartId-0="3.[151,264,1565,1588]" captionStartId-1="4.[151,264,1890,1913]" captionTargetBox-0="[234,1346,221,1528]" captionTargetBox-1="[202,1412,207,1829]" captionTargetId-0="figure@3.[214,1377,193,1541]" captionTargetId-1="figure@4.[161,1446,193,1845]" captionTargetPageId-0="3" captionTargetPageId-1="4" captionText-0="FIGURES 11 – 15. Apterous male of Zorotypus weiweii sp. nov.. 11, antenna; 12, left mandible; 13, right mandible; 14, maxillary palpus of apterous male; 15, labial palpus. Figs. 12 – 15, anterior view. Scale bar: Fig. 11 = 0.4 mm; 12, 13 = 0.1 mm; 14, 15 = 0.15 mm." captionText-1="FIGURES 16 – 23. Apterous male of Zorotypus weiweii sp. nov.. 16, right metaleg; 17, 18, posterior segments of abdomen; 19, left cercus; 20 – 23, male genitalia. 16, posterior view; 17, lateral view, posterior to the bottom 18 – 21, dorsal view; 22, 23, different angles of male genitalia. Scale bar: Fig. 16 = 0.3 mm; 17, 18, 20 = 0.45 mm; 19 = 0.30 mm; 21 – 23 = 0.15 mm." httpUri-0="https://zenodo.org/record/256279/files/figure.png" httpUri-1="https://zenodo.org/record/256280/files/figure.png" pageId="1" pageNumber="551">11–23</figureCitation>
, 28). Body length 2.3 mm, generally glossy brown except membranous regions, distal sections of antennae, tibiae, tarsi and cerci yellowish white to pale white. Head sub-triangular, wider than pronotum, vertex slightly concave, with curly setae grouped beside the midline, bases of those setae forming a “V” shape (
<figureCitation id="D3DA2A22B333CD61197DF679FDE2F9BD" box="[555,628,1559,1584]" captionStart="FIGURES 2 – 10" captionStartId="2.[151,264,1904,1927]" captionTargetBox="[202,1379,202,1861]" captionTargetId="figure@2.[192,1388,193,1872]" captionTargetPageId="2" captionText="FIGURES 2 – 10. Zorotypus weiweii sp. nov., antennae and appendages removed. 2, head of apterous male; 3, head of alate female; 4, thorax of apterous male; 5, thorax of dealate female; 6, 7, abdomen of apterous male; 8, abdomen of dealate female; 9, terminalia of male; 10, terminalia of female. Figs. 2, 3, anterior view; 4 – 6, dorsal view; 7, 8, ventral view; 9, 10, posterior view. Scale bar: Figs. 2, 3, 9, 10 = 0.4 mm; 4 – 8 = 0.2 mm." httpUri="https://zenodo.org/record/256278/files/figure.png" pageId="1" pageNumber="551">Fig. 2</figureCitation>
); eyes and ocelli absent; antennae about as long as head and thorax combined, nine-segmented; antennomere I slightly curved outward at basal half, with a faint constriction at the basal half, antennomere II shortest, about half length of antennomere III, antennomeres III–IX longer than wide, antennomere V longest (
<figureCitation id="D3DA2A22B333CD611AEDF6EDFD85F90E" box="[443,531,1667,1692]" captionStart="FIGURES 11 – 15" captionStartId="3.[151,264,1565,1588]" captionTargetBox="[234,1346,221,1528]" captionTargetId="figure@3.[214,1377,193,1541]" captionTargetPageId="3" captionText="FIGURES 11 – 15. Apterous male of Zorotypus weiweii sp. nov.. 11, antenna; 12, left mandible; 13, right mandible; 14, maxillary palpus of apterous male; 15, labial palpus. Figs. 12 – 15, anterior view. Scale bar: Fig. 11 = 0.4 mm; 12, 13 = 0.1 mm; 14, 15 = 0.15 mm." httpUri="https://zenodo.org/record/256279/files/figure.png" pageId="1" pageNumber="551">Fig. 11</figureCitation>
); left mandible with five teeth, prostheca inserted on the ventral surface of subapical tooth, molar region without dentes; right mandible with five teeth, the largest tooth (third tooth) with three tiny granulated dentes, molar region with fine dentes (
<figureCitation id="D3DA2A22B333CD611819F6A5FC4AF976" box="[847,988,1739,1764]" captionStart="FIGURES 11 – 15" captionStartId="3.[151,264,1565,1588]" captionTargetBox="[234,1346,221,1528]" captionTargetId="figure@3.[214,1377,193,1541]" captionTargetPageId="3" captionText="FIGURES 11 – 15. Apterous male of Zorotypus weiweii sp. nov.. 11, antenna; 12, left mandible; 13, right mandible; 14, maxillary palpus of apterous male; 15, labial palpus. Figs. 12 – 15, anterior view. Scale bar: Fig. 11 = 0.4 mm; 12, 13 = 0.1 mm; 14, 15 = 0.15 mm." httpUri="https://zenodo.org/record/256279/files/figure.png" pageId="1" pageNumber="551">Figs. 12, 13</figureCitation>
); maxillae symmetrical, distal divided, apex of inner part bifurcate, apex of outer part with dense hairs; maxillary palpus five-segmented (
<figureCitation id="D3DA2A22B333CD611FA2F681FADEF895" box="[1268,1352,1775,1800]" captionStart="FIGURES 11 – 15" captionStartId="3.[151,264,1565,1588]" captionTargetBox="[234,1346,221,1528]" captionTargetId="figure@3.[214,1377,193,1541]" captionTargetPageId="3" captionText="FIGURES 11 – 15. Apterous male of Zorotypus weiweii sp. nov.. 11, antenna; 12, left mandible; 13, right mandible; 14, maxillary palpus of apterous male; 15, labial palpus. Figs. 12 – 15, anterior view. Scale bar: Fig. 11 = 0.4 mm; 12, 13 = 0.1 mm; 14, 15 = 0.15 mm." httpUri="https://zenodo.org/record/256279/files/figure.png" pageId="1" pageNumber="551">Fig. 14</figureCitation>
); labial palpus three-segmented (
<figureCitation id="D3DA2A22B333CD611AF9F77DFD95F8BE" box="[431,515,1811,1836]" captionStart="FIGURES 11 – 15" captionStartId="3.[151,264,1565,1588]" captionTargetBox="[234,1346,221,1528]" captionTargetId="figure@3.[214,1377,193,1541]" captionTargetPageId="3" captionText="FIGURES 11 – 15. Apterous male of Zorotypus weiweii sp. nov.. 11, antenna; 12, left mandible; 13, right mandible; 14, maxillary palpus of apterous male; 15, labial palpus. Figs. 12 – 15, anterior view. Scale bar: Fig. 11 = 0.4 mm; 12, 13 = 0.1 mm; 14, 15 = 0.15 mm." httpUri="https://zenodo.org/record/256279/files/figure.png" pageId="1" pageNumber="551">Fig. 15</figureCitation>
).
</paragraph>
<paragraph id="4B5E36A7B333CD631B91F756FDB7F844" blockId="1.[151,1437,1451,2016]" lastBlockId="3.[151,1436,1693,2006]" lastPageId="3" lastPageNumber="553" pageId="1" pageNumber="551">
Pronotum (
<figureCitation id="D3DA2A22B333CD611A1EF759FE0FF8DD" box="[328,409,1847,1872]" captionStart="FIGURE 1" captionStartId="1.[151,250,1389,1411]" captionTargetBox="[188,1412,783,1323]" captionTargetId="figure@1.[151,1436,743,1368]" captionTargetPageId="1" captionText="FIGURE 1. Zorotypus weiweii sp. nov. Holotype, male, habitus. Scale bar = 0.5 mm." httpUri="https://zenodo.org/record/256277/files/figure.png" pageId="1" pageNumber="551">Figs. 1</figureCitation>
,
<figureCitation id="D3DA2A22B333CD611AFDF759FE2CF8DD" box="[427,442,1847,1871]" captionStart="FIGURES 2 – 10" captionStartId="2.[151,264,1904,1927]" captionTargetBox="[202,1379,202,1861]" captionTargetId="figure@2.[192,1388,193,1872]" captionTargetPageId="2" captionText="FIGURES 2 – 10. Zorotypus weiweii sp. nov., antennae and appendages removed. 2, head of apterous male; 3, head of alate female; 4, thorax of apterous male; 5, thorax of dealate female; 6, 7, abdomen of apterous male; 8, abdomen of dealate female; 9, terminalia of male; 10, terminalia of female. Figs. 2, 3, anterior view; 4 – 6, dorsal view; 7, 8, ventral view; 9, 10, posterior view. Scale bar: Figs. 2, 3, 9, 10 = 0.4 mm; 4 – 8 = 0.2 mm." httpUri="https://zenodo.org/record/256278/files/figure.png" pageId="1" pageNumber="551">4</figureCitation>
) sub-rectangular, posterior slightly narrowed; mesonotum trapezoidal, slightly shorter than pronotum, posterior broader than anterior; metanotum trapezoidal, shorter but wider than mesonotum, distinctly wider than long; thorax setose (
<figureCitation id="D3DA2A22B333CD61192BF711FD50F805" box="[637,710,1919,1944]" captionStart="FIGURES 2 – 10" captionStartId="2.[151,264,1904,1927]" captionTargetBox="[202,1379,202,1861]" captionTargetId="figure@2.[192,1388,193,1872]" captionTargetPageId="2" captionText="FIGURES 2 – 10. Zorotypus weiweii sp. nov., antennae and appendages removed. 2, head of apterous male; 3, head of alate female; 4, thorax of apterous male; 5, thorax of dealate female; 6, 7, abdomen of apterous male; 8, abdomen of dealate female; 9, terminalia of male; 10, terminalia of female. Figs. 2, 3, anterior view; 4 – 6, dorsal view; 7, 8, ventral view; 9, 10, posterior view. Scale bar: Figs. 2, 3, 9, 10 = 0.4 mm; 4 – 8 = 0.2 mm." httpUri="https://zenodo.org/record/256278/files/figure.png" pageId="1" pageNumber="551">Fig. 4</figureCitation>
). Legs with setae of moderate length, setae sparse on anterior surface of profemora and posterior surface of meso- and metafemora sparse, distal tarsomeres of all legs with short setae; posterior surface and dorsal surface of profemur covered with moderate-length setae, anterior surface and ventral surface with short setae; protibia with moderate-length setae, ventral surface of distal half with about 15 bristles, apically with two spurs; distal tarsomere about 2.5 times length of basal tarsomere. Mesofemur less strong than profemur, setae of mesofemur much like those of profemur; mesotibia with moderate-length setae and two small apical spurs. Metafemur (
<figureCitation id="D3DA2A22B331CD631957F767FDC2F8B0" box="[513,596,1801,1826]" captionStart="FIGURES 16 – 23" captionStartId="4.[151,264,1890,1913]" captionTargetBox="[202,1412,207,1829]" captionTargetId="figure@4.[161,1446,193,1845]" captionTargetPageId="4" captionText="FIGURES 16 – 23. Apterous male of Zorotypus weiweii sp. nov.. 16, right metaleg; 17, 18, posterior segments of abdomen; 19, left cercus; 20 – 23, male genitalia. 16, posterior view; 17, lateral view, posterior to the bottom 18 – 21, dorsal view; 22, 23, different angles of male genitalia. Scale bar: Fig. 16 = 0.3 mm; 17, 18, 20 = 0.45 mm; 19 = 0.30 mm; 21 – 23 = 0.15 mm." httpUri="https://zenodo.org/record/256280/files/figure.png" pageId="3" pageNumber="553">Fig. 16</figureCitation>
) swollen, much stronger than profemur or mesofemur, narrower apically, anterior and dorsal surface with moderate-length setae, apical half of ventral margin with three long, fine bristles on anterior surface, posterior surface nearly bare, with sparse, minute setae near the dorsal and ventral margin area, ventral surface with ten stout bristles, proximal first and third bristles longer than others; metatibia with moderatelength setae, dorsal surface subapically with one long bristle, ventral surface with two spurs apically; basal tarsomere with two spurs ventrally.
</paragraph>
<caption id="1F9E662FB330CD621BC1F71EFD70F871" httpUri="https://zenodo.org/record/256278/files/figure.png" pageId="2" pageNumber="552" targetBox="[202,1379,202,1861]" targetPageId="2">
<paragraph id="4B5E36A7B330CD621BC1F71EFD70F871" blockId="2.[151,1436,1904,2019]" pageId="2" pageNumber="552">
<emphasis id="7995EAB5B330CD621BC1F71EFEDFF814" bold="true" box="[151,329,1904,1927]" pageId="2" pageNumber="552">FIGURES 2–10.</emphasis>
<taxonomicName id="8CE14D24B330CD621A07F71CFD9AF815" box="[337,524,1906,1927]" class="Insecta" family="Zorotypidae" genus="Zorotypus" kingdom="Animalia" order="Zoraptera" pageId="2" pageNumber="552" phylum="Arthropoda" rank="species" species="weiweii" status="sp. nov.">
<emphasis id="7995EAB5B330CD621A07F71CFD9AF815" box="[337,524,1906,1927]" italics="true" pageId="2" pageNumber="552">Zorotypus weiweii</emphasis>
</taxonomicName>
<emphasis id="7995EAB5B330CD621943F71FFDF1F814" bold="true" box="[533,615,1905,1926]" pageId="2" pageNumber="552">
<taxonomicNameLabel id="62A657CEB330CD621943F71FFDF1F814" box="[533,615,1905,1926]" pageId="2" pageNumber="552" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
, antennae and appendages removed. 2, head of apterous male; 3, head of alate female; 4, thorax of apterous male; 5, thorax of dealate female; 6, 7, abdomen of apterous male; 8, abdomen of dealate female; 9, terminalia of male; 10, terminalia of female. Figs. 2, 3, anterior view; 4–6, dorsal view; 7, 8, ventral view; 9, 10, posterior view. Scale bar: Figs. 2, 3, 9, 10 = 0.4 mm; 4–8 = 0.2 mm.
</paragraph>
</caption>
<caption id="1F9E662FB331CD631BC1F673FEC4F9E3" httpUri="https://zenodo.org/record/256279/files/figure.png" pageId="3" pageNumber="553" targetBox="[234,1346,221,1528]" targetPageId="3">
<paragraph id="4B5E36A7B331CD631BC1F673FEC4F9E3" blockId="3.[151,1436,1565,1649]" pageId="3" pageNumber="553">
<emphasis id="7995EAB5B331CD631BC1F673FEC1F9A1" bold="true" box="[151,343,1565,1588]" pageId="3" pageNumber="553">FIGURES 11–15.</emphasis>
Apterous male of
<taxonomicName id="8CE14D24B331CD631973F670FD75F9A1" box="[549,739,1566,1587]" class="Insecta" family="Zorotypidae" genus="Zorotypus" kingdom="Animalia" order="Zoraptera" pageId="3" pageNumber="553" phylum="Arthropoda" rank="species" species="weiweii" status="sp. nov.">
<emphasis id="7995EAB5B331CD631973F670FD75F9A1" box="[549,739,1566,1587]" italics="true" pageId="3" pageNumber="553">Zorotypus weiweii</emphasis>
</taxonomicName>
<taxonomicNameLabel id="62A657CEB331CD6319B8F670FCD5F9A1" box="[750,835,1566,1587]" pageId="3" pageNumber="553" rank="species">
<emphasis id="7995EAB5B331CD6319B8F670FCD5F9A1" bold="true" box="[750,835,1566,1587]" pageId="3" pageNumber="553">sp. nov.</emphasis>
</taxonomicNameLabel>
. 11, antenna; 12, left mandible; 13, right mandible; 14, maxillary palpus of apterous male; 15, labial palpus. Figs. 12–15, anterior view. Scale bar: Fig. 11 = 0.4 mm; 12, 13 = 0.1 mm; 14, 15 = 0.15 mm.
</paragraph>
</caption>
<caption id="1F9E662FB336CD641BC1F70CFACEF824" httpUri="https://zenodo.org/record/256280/files/figure.png" pageId="4" pageNumber="554" targetBox="[202,1412,207,1829]" targetPageId="4">
<paragraph id="4B5E36A7B336CD641BC1F70CFACEF824" blockId="4.[151,1436,1890,1974]" pageId="4" pageNumber="554">
<emphasis id="7995EAB5B336CD641BC1F70CFEC2F8EA" bold="true" box="[151,340,1890,1913]" pageId="4" pageNumber="554">FIGURES 16–23.</emphasis>
Apterous male of
<taxonomicName id="8CE14D24B336CD641959F70AFD5FF8EB" box="[527,713,1892,1913]" class="Insecta" family="Zorotypidae" genus="Zorotypus" kingdom="Animalia" order="Zoraptera" pageId="4" pageNumber="554" phylum="Arthropoda" rank="species" species="weiweii" status="sp. nov.">
<emphasis id="7995EAB5B336CD641959F70AFD5FF8EB" box="[527,713,1892,1913]" italics="true" pageId="4" pageNumber="554">Zorotypus weiweii</emphasis>
</taxonomicName>
<taxonomicNameLabel id="62A657CEB336CD641999F70DFCB6F8EA" box="[719,800,1891,1912]" pageId="4" pageNumber="554" rank="species">
<emphasis id="7995EAB5B336CD641999F70DFCB6F8EA" bold="true" box="[719,800,1891,1912]" pageId="4" pageNumber="554">sp. nov.</emphasis>
</taxonomicNameLabel>
. 16, right metaleg; 17, 18, posterior segments of abdomen; 19, left cercus; 20–23, male genitalia. 16, posterior view; 17, lateral view, posterior to the bottom 18–21, dorsal view; 22, 23, different angles of male genitalia. Scale bar: Fig. 16 = 0.3 mm; 17, 18, 20 = 0.45 mm; 19 = 0.30 mm; 21–23 = 0.15 mm.
</paragraph>
</caption>
<caption id="1F9E662FB337CD651BC1F3A2FEB1FB93" httpUri="https://zenodo.org/record/256281/files/figure.png" pageId="5" pageNumber="555" targetBox="[253,1338,191,938]" targetPageId="5">
<paragraph id="4B5E36A7B337CD651BC1F3A2FEB1FB93" blockId="5.[151,1436,972,1025]" pageId="5" pageNumber="555">
<emphasis id="7995EAB5B337CD651BC1F3A2FEC2FC70" bold="true" box="[151,340,972,995]" pageId="5" pageNumber="555">FIGURES 24, 25.</emphasis>
Male genitalia of
<taxonomicName id="8CE14D24B337CD651958F3A3FC85FC70" authority="Karny" authorityName="Karny" box="[526,787,973,994]" class="Insecta" family="Zorotypidae" genus="Zorotypus" kingdom="Animalia" order="Zoraptera" pageId="5" pageNumber="555" phylum="Arthropoda" rank="species" species="caudelli">
<emphasis id="7995EAB5B337CD651958F3A3FD5BFC70" box="[526,717,973,994]" italics="true" pageId="5" pageNumber="555">Zorotypus caudelli</emphasis>
Karny
</taxonomicName>
, posterior to the bottom. 24, dorsal view; 25, lateral view. Scale bar = 0.2 mm.
</paragraph>
</caption>
<paragraph id="4B5E36A7B337CD651B91F443FF45FA64" blockId="5.[151,1437,1069,2030]" pageId="5" pageNumber="555">
First abdominal tergum (T1) with single transverse row of short setae along hind margin and a few small setae laterally; T2–7 with regular vestiture of short and moderate-length setae and a pair of longer setae along hind margin; T8 with short and moderate-length setae and two pairs of erect long setae along hind margin, inner side setae longer and stronger (
<figureCitation id="D3DA2A22B337CD651A95F4F4FD85FB21" box="[451,531,1178,1203]" captionStart="FIGURES 2 – 10" captionStartId="2.[151,264,1904,1927]" captionTargetBox="[202,1379,202,1861]" captionTargetId="figure@2.[192,1388,193,1872]" captionTargetPageId="2" captionText="FIGURES 2 – 10. Zorotypus weiweii sp. nov., antennae and appendages removed. 2, head of apterous male; 3, head of alate female; 4, thorax of apterous male; 5, thorax of dealate female; 6, 7, abdomen of apterous male; 8, abdomen of dealate female; 9, terminalia of male; 10, terminalia of female. Figs. 2, 3, anterior view; 4 – 6, dorsal view; 7, 8, ventral view; 9, 10, posterior view. Scale bar: Figs. 2, 3, 9, 10 = 0.4 mm; 4 – 8 = 0.2 mm." httpUri="https://zenodo.org/record/256278/files/figure.png" pageId="5" pageNumber="555">Figs. 6</figureCitation>
,
<figureCitation id="D3DA2A22B337CD651976F4F4FDFCFB20" box="[544,618,1178,1202]" captionStart="FIGURES 16 – 23" captionStartId="4.[151,264,1890,1913]" captionTargetBox="[202,1412,207,1829]" captionTargetId="figure@4.[161,1446,193,1845]" captionTargetPageId="4" captionText="FIGURES 16 – 23. Apterous male of Zorotypus weiweii sp. nov.. 16, right metaleg; 17, 18, posterior segments of abdomen; 19, left cercus; 20 – 23, male genitalia. 16, posterior view; 17, lateral view, posterior to the bottom 18 – 21, dorsal view; 22, 23, different angles of male genitalia. Scale bar: Fig. 16 = 0.3 mm; 17, 18, 20 = 0.45 mm; 19 = 0.30 mm; 21 – 23 = 0.15 mm." httpUri="https://zenodo.org/record/256280/files/figure.png" pageId="5" pageNumber="555">17, 18</figureCitation>
). T9 short, weakly sclerotized; T10 separated into anterior and posterior parts, anterior half sclerotized, with short and moderate-length setae laterally, posterior half mostly membranous, central region with medial spatula-like, upcurved projection; basal projection with several fine, short setae, apically with two setae (
<figureCitation id="D3DA2A22B337CD651A1DF56BFE0DFA8C" box="[331,411,1285,1310]" captionStart="FIGURES 2 – 10" captionStartId="2.[151,264,1904,1927]" captionTargetBox="[202,1379,202,1861]" captionTargetId="figure@2.[192,1388,193,1872]" captionTargetPageId="2" captionText="FIGURES 2 – 10. Zorotypus weiweii sp. nov., antennae and appendages removed. 2, head of apterous male; 3, head of alate female; 4, thorax of apterous male; 5, thorax of dealate female; 6, 7, abdomen of apterous male; 8, abdomen of dealate female; 9, terminalia of male; 10, terminalia of female. Figs. 2, 3, anterior view; 4 – 6, dorsal view; 7, 8, ventral view; 9, 10, posterior view. Scale bar: Figs. 2, 3, 9, 10 = 0.4 mm; 4 – 8 = 0.2 mm." httpUri="https://zenodo.org/record/256278/files/figure.png" pageId="5" pageNumber="555">Figs. 9</figureCitation>
,
<figureCitation id="D3DA2A22B337CD651AFEF568FE64FA8C" box="[424,498,1286,1310]" captionStart="FIGURES 16 – 23" captionStartId="4.[151,264,1890,1913]" captionTargetBox="[202,1412,207,1829]" captionTargetId="figure@4.[161,1446,193,1845]" captionTargetPageId="4" captionText="FIGURES 16 – 23. Apterous male of Zorotypus weiweii sp. nov.. 16, right metaleg; 17, 18, posterior segments of abdomen; 19, left cercus; 20 – 23, male genitalia. 16, posterior view; 17, lateral view, posterior to the bottom 18 – 21, dorsal view; 22, 23, different angles of male genitalia. Scale bar: Fig. 16 = 0.3 mm; 17, 18, 20 = 0.45 mm; 19 = 0.30 mm; 21 – 23 = 0.15 mm." httpUri="https://zenodo.org/record/256280/files/figure.png" pageId="5" pageNumber="555">17, 18</figureCitation>
); T11 with long, strongly upcurved medial projection centrally, bent in the middle, apically with a tiny hook (= male mating hook) (
<figureCitation id="D3DA2A22B337CD651997F544FCDCFAD0" box="[705,842,1322,1347]" captionStart="FIGURES 16 – 23" captionStartId="4.[151,264,1890,1913]" captionTargetBox="[202,1412,207,1829]" captionTargetId="figure@4.[161,1446,193,1845]" captionTargetPageId="4" captionText="FIGURES 16 – 23. Apterous male of Zorotypus weiweii sp. nov.. 16, right metaleg; 17, 18, posterior segments of abdomen; 19, left cercus; 20 – 23, male genitalia. 16, posterior view; 17, lateral view, posterior to the bottom 18 – 21, dorsal view; 22, 23, different angles of male genitalia. Scale bar: Fig. 16 = 0.3 mm; 17, 18, 20 = 0.45 mm; 19 = 0.30 mm; 21 – 23 = 0.15 mm." httpUri="https://zenodo.org/record/256280/files/figure.png" pageId="5" pageNumber="555">Figs. 17, 18</figureCitation>
), projection of T11 narrower and longer than that of T10 (
<figureCitation id="D3DA2A22B337CD651B8CF523FEC4FAF4" box="[218,338,1357,1382]" captionStart="FIGURES 2 – 10" captionStartId="2.[151,264,1904,1927]" captionTargetBox="[202,1379,202,1861]" captionTargetId="figure@2.[192,1388,193,1872]" captionTargetPageId="2" captionText="FIGURES 2 – 10. Zorotypus weiweii sp. nov., antennae and appendages removed. 2, head of apterous male; 3, head of alate female; 4, thorax of apterous male; 5, thorax of dealate female; 6, 7, abdomen of apterous male; 8, abdomen of dealate female; 9, terminalia of male; 10, terminalia of female. Figs. 2, 3, anterior view; 4 – 6, dorsal view; 7, 8, ventral view; 9, 10, posterior view. Scale bar: Figs. 2, 3, 9, 10 = 0.4 mm; 4 – 8 = 0.2 mm." httpUri="https://zenodo.org/record/256278/files/figure.png" pageId="5" pageNumber="555">Figs. 6, 9</figureCitation>
); base of T11 projection broadly sclerotized, narrower laterally, posterior region of broadly sclerotized margin bearing four moderate-length setae on each side (
<figureCitation id="D3DA2A22B337CD6518CBF51CFC74FA19" box="[925,994,1394,1419]" captionStart="FIGURES 2 – 10" captionStartId="2.[151,264,1904,1927]" captionTargetBox="[202,1379,202,1861]" captionTargetId="figure@2.[192,1388,193,1872]" captionTargetPageId="2" captionText="FIGURES 2 – 10. Zorotypus weiweii sp. nov., antennae and appendages removed. 2, head of apterous male; 3, head of alate female; 4, thorax of apterous male; 5, thorax of dealate female; 6, 7, abdomen of apterous male; 8, abdomen of dealate female; 9, terminalia of male; 10, terminalia of female. Figs. 2, 3, anterior view; 4 – 6, dorsal view; 7, 8, ventral view; 9, 10, posterior view. Scale bar: Figs. 2, 3, 9, 10 = 0.4 mm; 4 – 8 = 0.2 mm." httpUri="https://zenodo.org/record/256278/files/figure.png" pageId="5" pageNumber="555">Fig. 6</figureCitation>
); epiproct and paraproct unsclerotized; cercus (
<figureCitation id="D3DA2A22B337CD651BA7F5FBFED2FA3C" box="[241,324,1429,1454]" captionStart="FIGURES 16 – 23" captionStartId="4.[151,264,1890,1913]" captionTargetBox="[202,1412,207,1829]" captionTargetId="figure@4.[161,1446,193,1845]" captionTargetPageId="4" captionText="FIGURES 16 – 23. Apterous male of Zorotypus weiweii sp. nov.. 16, right metaleg; 17, 18, posterior segments of abdomen; 19, left cercus; 20 – 23, male genitalia. 16, posterior view; 17, lateral view, posterior to the bottom 18 – 21, dorsal view; 22, 23, different angles of male genitalia. Scale bar: Fig. 16 = 0.3 mm; 17, 18, 20 = 0.45 mm; 19 = 0.30 mm; 21 – 23 = 0.15 mm." httpUri="https://zenodo.org/record/256280/files/figure.png" pageId="5" pageNumber="555">Fig. 19</figureCitation>
) 1-segmented, conical, with one long apical seta, three subapical moderate-length setae, several short setae, and very long, fine setae; surface covered with numerous minute spicules (
<figureCitation id="D3DA2A22B337CD651F3DF5D4FB2BFA40" box="[1131,1213,1466,1491]" captionStart="FIGURES 16 – 23" captionStartId="4.[151,264,1890,1913]" captionTargetBox="[202,1412,207,1829]" captionTargetId="figure@4.[161,1446,193,1845]" captionTargetPageId="4" captionText="FIGURES 16 – 23. Apterous male of Zorotypus weiweii sp. nov.. 16, right metaleg; 17, 18, posterior segments of abdomen; 19, left cercus; 20 – 23, male genitalia. 16, posterior view; 17, lateral view, posterior to the bottom 18 – 21, dorsal view; 22, 23, different angles of male genitalia. Scale bar: Fig. 16 = 0.3 mm; 17, 18, 20 = 0.45 mm; 19 = 0.30 mm; 21 – 23 = 0.15 mm." httpUri="https://zenodo.org/record/256280/files/figure.png" pageId="5" pageNumber="555">Fig. 19</figureCitation>
) except at base and apex.
</paragraph>
<paragraph id="4B5E36A7B337CD651B91F66CFDC5F8A9" blockId="5.[151,1437,1069,2030]" pageId="5" pageNumber="555">
First abdominal sternum (S1) scarcely sclerotized; S2 weakly sclerotized centrally, with two or three short setae on each side; S3 with two transverse rows of short setae, basal row sparse; S4 with three transverse rows of short setae; S5–7 with short setae evenly scattered, S5 with pair of scarcely sclerotized circular areas; S8 and S9 fused, with shallow furrow, S8+9 subtriangular, with evenly scattered, moderate-length setae, posterior margin of S8 with a pair of longer setae; S10 invisible externally, beneath S8+9; S11 with two lateral sclerites, each with four short setae and two moderately long setae; on inner side setae longer, with a pair of tiny sensory setae on each side of inner membranous area (
<figureCitation id="D3DA2A22B337CD651A89F6B4FDC2F960" box="[479,596,1754,1779]" captionStart="FIGURES 2 – 10" captionStartId="2.[151,264,1904,1927]" captionTargetBox="[202,1379,202,1861]" captionTargetId="figure@2.[192,1388,193,1872]" captionTargetPageId="2" captionText="FIGURES 2 – 10. Zorotypus weiweii sp. nov., antennae and appendages removed. 2, head of apterous male; 3, head of alate female; 4, thorax of apterous male; 5, thorax of dealate female; 6, 7, abdomen of apterous male; 8, abdomen of dealate female; 9, terminalia of male; 10, terminalia of female. Figs. 2, 3, anterior view; 4 – 6, dorsal view; 7, 8, ventral view; 9, 10, posterior view. Scale bar: Figs. 2, 3, 9, 10 = 0.4 mm; 4 – 8 = 0.2 mm." httpUri="https://zenodo.org/record/256278/files/figure.png" pageId="5" pageNumber="555">Figs. 6, 9</figureCitation>
,
<figureCitation id="D3DA2A22B337CD651933F6B4FD38F960" box="[613,686,1754,1778]" captionStart="FIGURES 16 – 23" captionStartId="4.[151,264,1890,1913]" captionTargetBox="[202,1412,207,1829]" captionTargetId="figure@4.[161,1446,193,1845]" captionTargetPageId="4" captionText="FIGURES 16 – 23. Apterous male of Zorotypus weiweii sp. nov.. 16, right metaleg; 17, 18, posterior segments of abdomen; 19, left cercus; 20 – 23, male genitalia. 16, posterior view; 17, lateral view, posterior to the bottom 18 – 21, dorsal view; 22, 23, different angles of male genitalia. Scale bar: Fig. 16 = 0.3 mm; 17, 18, 20 = 0.45 mm; 19 = 0.30 mm; 21 – 23 = 0.15 mm." httpUri="https://zenodo.org/record/256280/files/figure.png" pageId="5" pageNumber="555">17–18</figureCitation>
). Genitalia asymmetrical (
<figureCitation id="D3DA2A22B337CD6518B3F6B4FBE5F960" box="[997,1139,1754,1779]" captionStart="FIGURES 16 – 23" captionStartId="4.[151,264,1890,1913]" captionTargetBox="[202,1412,207,1829]" captionTargetId="figure@4.[161,1446,193,1845]" captionTargetPageId="4" captionText="FIGURES 16 – 23. Apterous male of Zorotypus weiweii sp. nov.. 16, right metaleg; 17, 18, posterior segments of abdomen; 19, left cercus; 20 – 23, male genitalia. 16, posterior view; 17, lateral view, posterior to the bottom 18 – 21, dorsal view; 22, 23, different angles of male genitalia. Scale bar: Fig. 16 = 0.3 mm; 17, 18, 20 = 0.45 mm; 19 = 0.30 mm; 21 – 23 = 0.15 mm." httpUri="https://zenodo.org/record/256280/files/figure.png" pageId="5" pageNumber="555">Figs. 20–23</figureCitation>
), without elongate coiled flagellum or well-defined basal plate; dorsal sclerite twisted and elongated, slightly bent in middle; middle sclerite short, ventral sclerite broadly extended.
</paragraph>
<paragraph id="4B5E36A7B337CD651B91F72BFDB4F87C" blockId="5.[151,1437,1069,2030]" pageId="5" pageNumber="555">
<emphasis id="7995EAB5B337CD651B91F72BFE04F8CC" bold="true" box="[199,402,1861,1886]" pageId="5" pageNumber="555">Apterous female.</emphasis>
Generally as in male except as follows: head without curly setae on vertex. Abdominal T10 uniformly sclerotized, posterior margin with a pair of moderate length setae on each side, and with several short setae (
<figureCitation id="D3DA2A22B337CD651BB4F7E3FEAEF834" box="[226,312,1933,1958]" captionStart="FIGURES 2 – 10" captionStartId="2.[151,264,1904,1927]" captionTargetBox="[202,1379,202,1861]" captionTargetId="figure@2.[192,1388,193,1872]" captionTargetPageId="2" captionText="FIGURES 2 – 10. Zorotypus weiweii sp. nov., antennae and appendages removed. 2, head of apterous male; 3, head of alate female; 4, thorax of apterous male; 5, thorax of dealate female; 6, 7, abdomen of apterous male; 8, abdomen of dealate female; 9, terminalia of male; 10, terminalia of female. Figs. 2, 3, anterior view; 4 – 6, dorsal view; 7, 8, ventral view; 9, 10, posterior view. Scale bar: Figs. 2, 3, 9, 10 = 0.4 mm; 4 – 8 = 0.2 mm." httpUri="https://zenodo.org/record/256278/files/figure.png" pageId="5" pageNumber="555">Fig. 10</figureCitation>
); T11 without projection, mostly sclerotized, with several setae (
<figureCitation id="D3DA2A22B337CD651F7BF7E3FB12F834" box="[1069,1156,1933,1958]" captionStart="FIGURES 2 – 10" captionStartId="2.[151,264,1904,1927]" captionTargetBox="[202,1379,202,1861]" captionTargetId="figure@2.[192,1388,193,1872]" captionTargetPageId="2" captionText="FIGURES 2 – 10. Zorotypus weiweii sp. nov., antennae and appendages removed. 2, head of apterous male; 3, head of alate female; 4, thorax of apterous male; 5, thorax of dealate female; 6, 7, abdomen of apterous male; 8, abdomen of dealate female; 9, terminalia of male; 10, terminalia of female. Figs. 2, 3, anterior view; 4 – 6, dorsal view; 7, 8, ventral view; 9, 10, posterior view. Scale bar: Figs. 2, 3, 9, 10 = 0.4 mm; 4 – 8 = 0.2 mm." httpUri="https://zenodo.org/record/256278/files/figure.png" pageId="5" pageNumber="555">Fig. 10</figureCitation>
); abdominal sterna with more setae than male; S8 not fused with S9, S9 short and trapezoidal, with several short setae and a pair of moderate length setae (
<figureCitation id="D3DA2A22B337CD651ACCF7BBFD83F87C" box="[410,533,2005,2030]" captionStart="FIGURES 2 – 10" captionStartId="2.[151,264,1904,1927]" captionTargetBox="[202,1379,202,1861]" captionTargetId="figure@2.[192,1388,193,1872]" captionTargetPageId="2" captionText="FIGURES 2 – 10. Zorotypus weiweii sp. nov., antennae and appendages removed. 2, head of apterous male; 3, head of alate female; 4, thorax of apterous male; 5, thorax of dealate female; 6, 7, abdomen of apterous male; 8, abdomen of dealate female; 9, terminalia of male; 10, terminalia of female. Figs. 2, 3, anterior view; 4 – 6, dorsal view; 7, 8, ventral view; 9, 10, posterior view. Scale bar: Figs. 2, 3, 9, 10 = 0.4 mm; 4 – 8 = 0.2 mm." httpUri="https://zenodo.org/record/256278/files/figure.png" pageId="5" pageNumber="555">Figs. 8, 10</figureCitation>
).
</paragraph>
<paragraph id="4B5E36A7B334CD661BC1F685FEFFF8AD" pageId="6" pageNumber="556">
<table id="39E1C407B334329F1BC1F685FA0AF8AD" box="[151,1436,1771,1855]" gridcols="3" gridrows="3" pageId="6" pageNumber="556">
<tr id="F5D134E5B334329F1BC1F685FA0AF890" box="[151,1436,1771,1794]" gridrow="0" pageId="6" pageNumber="556">
<th id="B6005D99B334329F1BC1F685FC9FF890" box="[151,777,1771,1794]" gridcol="0" gridrow="0" pageId="6" pageNumber="556">
<emphasis id="7995EAB5B334CD661BC1F685FEC0F893" bold="true" box="[151,342,1771,1794]" pageId="6" pageNumber="556">FIGURES 26–30.</emphasis>
Habitat, environment and living status of
</th>
<th id="B6005D99B334329F1846F685FBA4F890" box="[784,1074,1771,1794]" gridcol="1" gridrow="0" pageId="6" pageNumber="556">
<taxonomicName id="8CE14D24B334CD661844F682FC58F893" box="[786,974,1772,1793]" class="Insecta" family="Zorotypidae" genus="Zorotypus" kingdom="Animalia" order="Zoraptera" pageId="6" pageNumber="556" phylum="Arthropoda" rank="species" species="weiweii" status="sp. nov.">
<emphasis id="7995EAB5B334CD661844F682FC58F893" box="[786,974,1772,1793]" italics="true" pageId="6" pageNumber="556">Zorotypus weiweii</emphasis>
</taxonomicName>
<taxonomicNameLabel id="62A657CEB334CD661881F682FBBCF893" box="[983,1066,1772,1793]" pageId="6" pageNumber="556" rank="species">
<emphasis id="7995EAB5B334CD661881F682FBBCF893" bold="true" box="[983,1066,1772,1793]" pageId="6" pageNumber="556">sp. nov.</emphasis>
</taxonomicNameLabel>
.
</th>
<th id="B6005D99B334329F1F6FF685FA0AF890" box="[1081,1436,1771,1794]" gridcol="2" gridrow="0" pageId="6" pageNumber="556">26, landscape of type locality; 27,</th>
</tr>
<tr id="F5D134E5B334329F1BC1F764FA0AF8B2" box="[151,1436,1802,1824]" gridrow="1" pageId="6" pageNumber="556">
<th id="B6005D99B334329F1BC1F764FC9FF8B2" box="[151,777,1802,1824]" gridcol="0" gridrow="1" pageId="6" pageNumber="556">
rotting logs where
<taxonomicName id="8CE14D24B334CD661A00F765FD86F8B2" box="[342,528,1803,1824]" class="Insecta" family="Zorotypidae" genus="Zorotypus" kingdom="Animalia" order="Zoraptera" pageId="6" pageNumber="556" phylum="Arthropoda" rank="species" species="weiweii" status="sp. nov.">
<emphasis id="7995EAB5B334CD661A00F765FD86F8B2" box="[342,528,1803,1824]" italics="true" pageId="6" pageNumber="556">Zorotypus weiweii</emphasis>
</taxonomicName>
<taxonomicNameLabel id="62A657CEB334CD661941F765FDFEF8B2" box="[535,616,1803,1824]" pageId="6" pageNumber="556" rank="species">
<emphasis id="7995EAB5B334CD661941F765FDFEF8B2" bold="true" box="[535,616,1803,1824]" pageId="6" pageNumber="556">sp. nov.</emphasis>
</taxonomicNameLabel>
were collected;
</th>
<td id="B6005D99B334329F1846F764FBA4F8B2" box="[784,1074,1802,1824]" gridcol="1" gridrow="1" pageId="6" pageNumber="556">28, living apterous male; 29,</td>
<td id="B6005D99B334329F1F6FF764FA0AF8B2" box="[1081,1436,1802,1824]" gridcol="2" gridrow="1" pageId="6" pageNumber="556">living alate female; 30, living alate</td>
</tr>
<tr id="F5D134E5B334329F1BC1F747FA0AF8AD" box="[151,1436,1833,1855]" gridrow="2" pageId="6" pageNumber="556" rowspan-1="1" rowspan-2="1">
<th id="B6005D99B334329F1BC1F747FC9FF8AD" box="[151,777,1833,1855]" gridcol="0" gridrow="2" pageId="6" pageNumber="556">nymph (final instar).</th>
</tr>
</table>
</paragraph>
<paragraph id="4B5E36A7B334CD661B91F708FAE2F855" blockId="6.[151,1437,1894,2027]" pageId="6" pageNumber="556">
<emphasis id="7995EAB5B334CD661B91F708FE9BF8ED" bold="true" box="[199,269,1894,1919]" pageId="6" pageNumber="556">Alate.</emphasis>
Body color generally darker than apteron (Fig. 29), color blackish brown. Compound eyes and three black ocelli present (
<figureCitation id="D3DA2A22B334CD661AD9F7E4FE4EF831" box="[399,472,1930,1955]" captionStart="FIGURES 2 – 10" captionStartId="2.[151,264,1904,1927]" captionTargetBox="[202,1379,202,1861]" captionTargetId="figure@2.[192,1388,193,1872]" captionTargetPageId="2" captionText="FIGURES 2 – 10. Zorotypus weiweii sp. nov., antennae and appendages removed. 2, head of apterous male; 3, head of alate female; 4, thorax of apterous male; 5, thorax of dealate female; 6, 7, abdomen of apterous male; 8, abdomen of dealate female; 9, terminalia of male; 10, terminalia of female. Figs. 2, 3, anterior view; 4 – 6, dorsal view; 7, 8, ventral view; 9, 10, posterior view. Scale bar: Figs. 2, 3, 9, 10 = 0.4 mm; 4 – 8 = 0.2 mm." httpUri="https://zenodo.org/record/256278/files/figure.png" pageId="6" pageNumber="556">Fig. 3</figureCitation>
). Pronotum wider than long, mesonotum and metanotum indistinctly divided into several areas: prescutum, scutum and scutellum (
<figureCitation id="D3DA2A22B334CD6619EAF7C0FC96F855" box="[700,768,1966,1991]" captionStart="FIGURES 2 – 10" captionStartId="2.[151,264,1904,1927]" captionTargetBox="[202,1379,202,1861]" captionTargetId="figure@2.[192,1388,193,1872]" captionTargetPageId="2" captionText="FIGURES 2 – 10. Zorotypus weiweii sp. nov., antennae and appendages removed. 2, head of apterous male; 3, head of alate female; 4, thorax of apterous male; 5, thorax of dealate female; 6, 7, abdomen of apterous male; 8, abdomen of dealate female; 9, terminalia of male; 10, terminalia of female. Figs. 2, 3, anterior view; 4 – 6, dorsal view; 7, 8, ventral view; 9, 10, posterior view. Scale bar: Figs. 2, 3, 9, 10 = 0.4 mm; 4 – 8 = 0.2 mm." httpUri="https://zenodo.org/record/256278/files/figure.png" pageId="6" pageNumber="556">Fig. 5</figureCitation>
). Wings clothed and fringed with numerous microsetae.
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<emphasis id="7995EAB5B334CD661B91F7BCFEB1F879" bold="true" box="[199,295,2002,2027]" pageId="6" pageNumber="556">Biology.</emphasis>
Apterons, alates and dealates, as well as juveniles, were found living together under rotting bark of rain forest (Figs. 26, 27). Zorapterans mainly feed on fungal spores, organic matter or perhaps small arthropods. Alates and dealates were easily distinguished by nimble movements and compound eyes, even in juvenile stages (Figs. 28–30). We did not find any ecdysing juvenile individuals but zorapterans spent much of their time grooming almost every part of their bodies; thus, we speculate that they may be grooming to clear off fragments of exuviae. The antennae of
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are nine-segmented, the last segment being developed at the apex of the second segment before the body color turns brown.
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<emphasis id="7995EAB5B335CD671B91F101FED8FE1A" bold="true" box="[199,334,367,392]" pageId="7" pageNumber="557">Etymology.</emphasis>
The specific epithet refers to the Chinese entomologist Zhang Weiwei, one of the collectors of this new species, in honor of his great contributions to entomology.
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<emphasis id="7995EAB5B335CD671B91F1D9FEF6FE42" bold="true" box="[199,352,439,464]" pageId="7" pageNumber="557">Distribution.</emphasis>
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, East
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(
<figureCitation id="D3DA7BE8FFFF6343EEC4C231FEC2051F" box="[159,267,1121,1146]" captionStart-0="FIGURE 1" captionStart-1="FIGURE 2" captionStart-2="FIGURE 3" captionStart-3="FIGURE 4" captionStart-4="FIGURE 5" captionStartId-0="3.[151,250,1935,1957]" captionStartId-1="4.[151,250,1696,1718]" captionStartId-2="5.[151,250,1194,1216]" captionStartId-3="6.[151,250,1952,1974]" captionStartId-4="7.[151,250,1660,1682]" captionTargetBox-0="[151,1436,481,1912]" captionTargetBox-1="[349,1239,204,1666]" captionTargetBox-2="[349,1236,184,1162]" captionTargetBox-3="[384,1184,211,1923]" captionTargetBox-4="[151,1436,193,1637]" captionTargetId-0="figure@3.[151,1436,481,1914]" captionTargetId-1="figure@4.[340,1247,193,1675]" captionTargetId-2="figure@5.[340,1247,173,1173]" captionTargetId-3="figure@6.[376,1206,193,1942]" captionTargetId-4="figure@7.[151,1436,193,1639]" captionTargetPageId-0="3" captionTargetPageId-1="4" captionTargetPageId-2="5" captionTargetPageId-3="6" captionTargetPageId-4="7" captionText-0="FIGURE 1. Stiphodon palawanensis immediately after fixation. a, WPU-PPC-P 5, holotype (male, 62.2 mm SL); b, URM-P 48659, paratype (male, 51.1 mm SL); c, URM-P 48666, paratype (female, 58.5 mm SL); d, URM-P 48661, paratype (female, 41.1 mm SL)." captionText-1="FIGURE 2. Number of premaxillary and horizontal dentary teeth of Stiphodon palawanensis (red circles), S. alcedo (pink circles), S. atratus (black diamonds), S. imperiorientis (light blue triangles), S. maculidorsalis (light green triangles), S. multisquamus (deep blue circles), S. ornatus (orange triangles), S. pelewensis (gray diamonds), S. pulchellus (deep green triangles), and S. weberi (purple diamonds)." captionText-2="FIGURE 3. Dorsal scalation on head and nape in Stiphodon palawanensis. a, male (43.1 mm SL, CMK 11966); b, female (41.9 mm SL, CMK 11966)." captionText-3="FIGURE 4. Diagrammatic illustration of head showing arrangement of cephalic sensory pores and cutaneous sensory papillae in Stiphodon palawanensis (41.9 mm SL, CMK 11966). a, dorsal view; b, lateral view; c, ventral view. AN, anterior nostril; PN, posterior nostril." captionText-4="FIGURE 5. Males (a – d) and females (e, f) of live Stiphodon palawanensis observed in Balsahan Stream, Puerto Princesa City (b, e, f, 18 May 2015; a, c, 19 May 2015) and Barake Stream, Aborlan (d, 15 May 2015)." httpUri-0="https://zenodo.org/record/232900/files/figure.png" httpUri-1="https://zenodo.org/record/232901/files/figure.png" httpUri-2="https://zenodo.org/record/232902/files/figure.png" httpUri-3="https://zenodo.org/record/232903/files/figure.png" httpUri-4="https://zenodo.org/record/232904/files/figure.png" pageId="1" pageNumber="382">Figs. 1–5</figureCitation>
; Table 1)
</paragraph>
</subSubSection>
<subSubSection id="03FB34E6FFFF6343EECCC2F9FDF1062F" pageId="1" pageNumber="382" type="materials_examined">
<paragraph id="4B5E676DFFFF6343EECCC2F9FED40583" blockId="1.[151,1437,1193,2010]" pageId="1" pageNumber="382">
<emphasis id="7995BB7FFFFF6343EECCC2F9FE4E05A7" bold="true" box="[151,391,1193,1218]" pageId="1" pageNumber="382">Material examined.</emphasis>
<specimenCount id="5DE7ACE4FFFF6343EFCFC2F9FDFA05A7" box="[404,563,1193,1218]" pageId="1" pageNumber="382" type="generic">57 specimens</specimenCount>
(
<specimenCount id="5DE7ACE4FFFF6343EC12C2FAFD7C05A7" box="[585,693,1193,1218]" pageId="1" pageNumber="382" type="male">24 males</specimenCount>
and
<specimenCount id="5DE7ACE4FFFF6343ECA1C2FAFCB705A7" box="[762,894,1193,1218]" pageId="1" pageNumber="382" type="female">33 females</specimenCount>
, 27.0–
<quantity id="8C19CA88FFFF6343ED94C2F9FBF005A4" box="[975,1081,1193,1218]" metricMagnitude="-2" metricUnit="m" metricValue="6.39" pageId="1" pageNumber="382" unit="mm" value="63.9">63.9 mm</quantity>
SL) collected from Palawan,
<collectingCountry id="33F627FDFFFF6343EECCC29DFED00583" box="[151,281,1229,1254]" name="Philippines" pageId="1" pageNumber="382">Philippines</collectingCountry>
.
</paragraph>
<paragraph id="4B5E676DFFFF6343EE9CC2A1FC6B044B" blockId="1.[151,1437,1193,2010]" pageId="1" pageNumber="382">
<emphasis id="7995BB7FFFFF6343EE9CC2A1FEF2046F" bold="true" box="[199,315,1265,1290]" pageId="1" pageNumber="382">
<typeStatus id="945AD9CFFFFF6343EE9CC2A1FEFF046F" box="[199,310,1265,1290]" pageId="1" pageNumber="382" type="holotype">Holotype</typeStatus>
.
</emphasis>
WPU-PPC-P 5, male (
<quantity id="8C19CA88FFFF6343EC65C2A1FD6D046C" box="[574,676,1265,1290]" metricMagnitude="-2" metricUnit="m" metricValue="6.22" pageId="1" pageNumber="382" unit="mm" value="62.2">62.2 mm</quantity>
SL), Balsahan Stream in the Iwahig Prison and Penal Farm, Puerto Princesa City, Palawan,
<date id="3F5F41ADFFFF6343EFFDC346FDF7044B" box="[422,574,1301,1326]" pageId="1" pageNumber="382" value="2015-05-18">18 May 2015</date>
, coll. K. Maeda and H. P. Palla.
</paragraph>
<paragraph id="4B5E676DFFFF6343EE9CC369FDF1062F" blockId="1.[151,1437,1193,2010]" pageId="1" pageNumber="382">
<emphasis id="7995BB7FFFFF6343EE9CC369FE8F0437" bold="true" box="[199,326,1337,1362]" pageId="1" pageNumber="382">
<typeStatus id="945AD9CFFFFF6343EE9CC369FE880437" box="[199,321,1337,1362]" pageId="1" pageNumber="382" type="paratype">Paratypes</typeStatus>
.
</emphasis>
CMK 11966,
<specimenCount id="5DE7ACE4FFFF6343EFB3C36AFDF60437" box="[488,575,1337,1362]" pageId="1" pageNumber="382" type="male">2 males</specimenCount>
(37.3 and
<quantity id="8C19CA88FFFF6343ECE3C36AFCD40434" box="[696,797,1338,1362]" metricMagnitude="-2" metricUnit="m" metricValue="4.31" pageId="1" pageNumber="382" unit="mm" value="43.1">43.1 mm</quantity>
SL) and
<specimenCount id="5DE7ACE4FFFF6343EDDDC36AFC3B0437" box="[902,1010,1337,1362]" pageId="1" pageNumber="382" type="female">8 females</specimenCount>
(
<quantity id="8C19CA88FFFF6343EA59C369FB6E0434" box="[1026,1191,1337,1362]" metricMagnitude="-2" metricUnit="m" metricValue="3.785" metricValueMax="4.72" metricValueMin="2.85" pageId="1" pageNumber="382" unit="mm" value="37.85" valueMax="47.2" valueMin="28.5">28.5–47.2 mm</quantity>
SL), Malatgao River, Narra, Palawan,
<date id="3F5F41ADFFFF6343EF0EC30EFDFC0413" box="[341,565,1373,1398]" pageId="1" pageNumber="382" value="1994-09-29">29 September 1994</date>
, coll. J. Margraf; CMK 11974,
<specimenCount id="5DE7ACE4FFFF6343EDFBC30EFBC70413" box="[928,1038,1373,1398]" pageId="1" pageNumber="382" type="female">3 females</specimenCount>
(
<quantity id="8C19CA88FFFF6343EA45C30DFB0C0410" box="[1054,1221,1373,1398]" metricMagnitude="-2" metricUnit="m" metricValue="4.52" metricValueMax="4.85" metricValueMin="4.1899999999999995" pageId="1" pageNumber="382" unit="mm" value="45.2" valueMax="48.5" valueMin="41.9">41.9–48.5 mm</quantity>
SL), Estrella Falls (tributary of Malatgao River), Narra, Palawan,
<date id="3F5F41ADFFFF6343ECEBC3D2FC5904FF" box="[688,912,1409,1434]" pageId="1" pageNumber="382" value="1994-09-29">29 September 1994</date>
, coll. J. Margraf; NSMT-P 45091, 45092, and 45094,
<specimenCount id="5DE7ACE4FFFF6343EEB0C3F6FE8804DB" box="[235,321,1445,1470]" pageId="1" pageNumber="382" type="male">2 males</specimenCount>
(27.7 and
<quantity id="8C19CA88FFFF6343EFE3C3F6FDD504D8" box="[440,540,1446,1470]" metricMagnitude="-2" metricUnit="m" metricValue="3.88" pageId="1" pageNumber="382" unit="mm" value="38.8">38.8 mm</quantity>
SL) and female (
<quantity id="8C19CA88FFFF6343EC85C3F5FC8A04D8" box="[734,835,1445,1470]" metricMagnitude="-2" metricUnit="m" metricValue="3.66" pageId="1" pageNumber="382" unit="mm" value="36.6">36.6 mm</quantity>
SL), Iwahig River, Puerto Princesa City, Palawan,
<date id="3F5F41ADFFFF6343EBDBC3F6FE980487" pageId="1" pageNumber="382" value="1988-11-13">13 November 1988</date>
, coll. K. Matsuura; URM-P 31438, female (27.0 mm SL), Nagsagoiri River, Palawan,
<date id="3F5F41ADFFFF6343EB6FC39AFF1B0763" pageId="1" pageNumber="382" value="1985-08-09">9 August 1985</date>
, coll. S. Shokita; URM-P 31439,
<specimenCount id="5DE7ACE4FFFF6343EC3BC3BDFD730763" box="[608,698,1517,1542]" pageId="1" pageNumber="382" type="male">6 males</specimenCount>
(
<quantity id="8C19CA88FFFF6343EC97C3BDFCBD0760" box="[716,884,1517,1542]" metricMagnitude="-2" metricUnit="m" metricValue="3.545" metricValueMax="3.6799999999999997" metricValueMin="3.41" pageId="1" pageNumber="382" unit="mm" value="35.45" valueMax="36.8" valueMin="34.1">34.1–36.8 mm</quantity>
SL) and
<specimenCount id="5DE7ACE4FFFF6343EDBDC3BEFB9F0763" box="[998,1110,1517,1542]" pageId="1" pageNumber="382" type="female">7 females</specimenCount>
(
<quantity id="8C19CA88FFFF6343EA33C3BDFAD90760" box="[1128,1296,1517,1542]" metricMagnitude="-2" metricUnit="m" metricValue="3.4299999999999997" metricValueMax="3.62" metricValueMin="3.2399999999999998" pageId="1" pageNumber="382" unit="mm" value="34.3" valueMax="36.2" valueMin="32.4">32.4–36.2 mm</quantity>
SL), Papait River, Palawan,
<date id="3F5F41ADFFFF6343EF08C042FE37074F" box="[339,510,1553,1578]" pageId="1" pageNumber="382" value="1985-08-09">9 August 1985</date>
, coll. S. Shokita; URM-P 31440,
<specimenCount id="5DE7ACE4FFFF6343EDD9C042FC39074F" box="[898,1008,1553,1578]" pageId="1" pageNumber="382" type="female">2 females</specimenCount>
(38.6 and
<quantity id="8C19CA88FFFF6343EA36C042FB1A074C" box="[1133,1235,1554,1578]" metricMagnitude="-2" metricUnit="m" metricValue="4.07" pageId="1" pageNumber="382" unit="mm" value="40.7">40.7 mm</quantity>
SL),
<collectingCountry id="33F627FDFFFF6343EB4DC042FA99074C" box="[1302,1360,1554,1577]" name="Iran" pageId="1" pageNumber="382">Iraan</collectingCountry>
River, Palawan,
<date id="3F5F41ADFFFF6343EF5EC065FE79072B" box="[261,432,1589,1614]" pageId="1" pageNumber="382" value="1985-08-05">5 August 1985</date>
, coll. S. Shokita; URM-P 31441,
<specimenCount id="5DE7ACE4FFFF6343ED69C065FC43072B" box="[818,906,1589,1614]" pageId="1" pageNumber="382" type="male">6 males</specimenCount>
(
<quantity id="8C19CA88FFFF6343EDC0C066FB880728" box="[923,1089,1590,1614]" metricMagnitude="-2" metricUnit="m" metricValue="3.635" metricValueMax="4.14" metricValueMin="3.13" pageId="1" pageNumber="382" unit="mm" value="36.35" valueMax="41.4" valueMin="31.3">31.3–41.4 mm</quantity>
SL) and
<specimenCount id="5DE7ACE4FFFF6343EAF6C066FAD3072B" box="[1197,1306,1589,1614]" pageId="1" pageNumber="382" type="female">3 females</specimenCount>
(
<quantity id="8C19CA88FFFF6343EB71C066FF0A0714" metricMagnitude="-2" metricUnit="m" metricValue="3.4899999999999998" metricValueMax="3.91" metricValueMin="3.07" pageId="1" pageNumber="382" unit="mm" value="34.9" valueMax="39.1" valueMin="30.7">30.7–39.1 mm</quantity>
SL), Tagbariri, Palawan,
<date id="3F5F41ADFFFF6343EFBCC00AFD580717" box="[487,657,1625,1650]" pageId="1" pageNumber="382" value="1985-08-09">9 August 1985</date>
, coll. S. Shokita; URM-P 48659–48662,
<specimenCount id="5DE7ACE4FFFF6343EA3EC00AFB740717" box="[1125,1213,1625,1650]" pageId="1" pageNumber="382" type="male">2 males</specimenCount>
(46.2 and
<quantity id="8C19CA88FFFF6343EB6CC009FA540714" box="[1335,1437,1625,1650]" metricMagnitude="-2" metricUnit="m" metricValue="5.11" pageId="1" pageNumber="382" unit="mm" value="51.1">51.1 mm</quantity>
SL) and
<specimenCount id="5DE7ACE4FFFF6343EEA6C02EFEA207F3" box="[253,363,1661,1686]" pageId="1" pageNumber="382" type="female">2 females</specimenCount>
(41.1 and
<quantity id="8C19CA88FFFF6343EFB7C02EFD9B07F0" box="[492,594,1662,1686]" metricMagnitude="-2" metricUnit="m" metricValue="4.72" pageId="1" pageNumber="382" unit="mm" value="47.2">47.2 mm</quantity>
SL), Barake Stream (tributary of Aborlan River), Barangay Magbabadil, Aborlan, Palawan,
<date id="3F5F41ADFFFF6343EF2AC0F1FDC407DF" box="[369,525,1697,1722]" pageId="1" pageNumber="382" value="2015-05-15">15 May 2015</date>
, coll. K. Maeda and H. P. Palla; URM-P 48663–48666,
<specimenCount id="5DE7ACE4FFFF6343EACEC0F2FB2407DF" box="[1173,1261,1697,1722]" pageId="1" pageNumber="382" type="male">2 males</specimenCount>
(59.7 and
<quantity id="8C19CA88FFFF6343EB30C0F1FF0A07B8" metricMagnitude="-2" metricUnit="m" metricValue="6.39" pageId="1" pageNumber="382" unit="mm" value="63.9">63.9 mm</quantity>
SL) and
<specimenCount id="5DE7ACE4FFFF6343EF76C096FE5307BB" box="[301,410,1733,1758]" pageId="1" pageNumber="382" type="female">2 females</specimenCount>
(58.5 and
<quantity id="8C19CA88FFFF6343EC4DC095FDB307B8" box="[534,634,1733,1758]" metricMagnitude="-2" metricUnit="m" metricValue="6.36" pageId="1" pageNumber="382" unit="mm" value="63.6">63.6 mm</quantity>
SL), same data as
<typeStatus id="945AD9CFFFFF6343ED08C095FC7107BB" box="[851,952,1733,1758]" pageId="1" pageNumber="382" type="holotype">holotype</typeStatus>
; WPU-PPC-P 2–4,
<specimenCount id="5DE7ACE4FFFF6343EAC2C096FB3907BB" box="[1177,1264,1733,1758]" pageId="1" pageNumber="382" type="male">2 males</specimenCount>
(52.6 and
<quantity id="8C19CA88FFFF6343EB30C095FF0A0664" metricMagnitude="-2" metricUnit="m" metricValue="5.92" pageId="1" pageNumber="382" unit="mm" value="59.2">59.2 mm</quantity>
SL) and female (
<quantity id="8C19CA88FFFF6343EFD0C0B9FE3B0664" box="[395,498,1769,1794]" metricMagnitude="-2" metricUnit="m" metricValue="4.63" pageId="1" pageNumber="382" unit="mm" value="46.3">46.3 mm</quantity>
SL), Barake Stream (tributary of Aborlan River), Barangay Magbabadil, Aborlan, Palawan,
<date id="3F5F41ADFFFF6343EF58C15DFE530643" box="[259,410,1805,1830]" pageId="1" pageNumber="382" value="2015-05-15">15 May 2015</date>
, coll. K. Maeda and H. P. Palla; WPU-PPC-P 6–9, male (
<quantity id="8C19CA88FFFF6343EA4CC15DFBB50640" box="[1047,1148,1805,1830]" metricMagnitude="-2" metricUnit="m" metricValue="5.9399999999999995" pageId="1" pageNumber="382" unit="mm" value="59.4">59.4 mm</quantity>
SL) and
<specimenCount id="5DE7ACE4FFFF6343EAB9C15EFA840643" box="[1250,1357,1805,1830]" pageId="1" pageNumber="382" type="female">3 females</specimenCount>
(
<quantity id="8C19CA88FFFF6343EB00C15DFF32062C" metricMagnitude="-2" metricUnit="m" metricValue="5.9799999999999995" metricValueMax="6.21" metricValueMin="5.75" pageId="1" pageNumber="382" unit="mm" value="59.8" valueMax="62.1" valueMin="57.5">57.5– 62.1 mm</quantity>
SL), same data as
<typeStatus id="945AD9CFFFFF6343EF95C161FDFD062F" box="[462,564,1841,1866]" pageId="1" pageNumber="382" type="holotype">holotype</typeStatus>
.
</paragraph>
</subSubSection>
<subSubSection id="03FB34E6FFFF6340EE9CC105FD68019D" lastPageId="2" lastPageNumber="383" pageId="1" pageNumber="382" type="diagnosis">
<paragraph id="4B5E676DFFFF6340EE9CC105FD68019D" blockId="1.[151,1437,1193,2010]" lastBlockId="2.[151,1437,151,536]" lastPageId="2" lastPageNumber="383" pageId="1" pageNumber="382">
<emphasis id="7995BB7FFFFF6343EE9CC105FE88060B" bold="true" box="[199,321,1877,1902]" pageId="1" pageNumber="382">Diagnosis.</emphasis>
Second dorsal fin usually with one spine and nine segmented rays, pectoral fin usually with 15 segmented rays; first dorsal fin pointed in male; relatively large caudal fin (26–34% of SL) in male; premaxilla with 45–71 tricuspid teeth, dentary with 2–6 (male) or 1–4 (female) symphyseal teeth and 45–79 unicuspid horizontal teeth; nape and posterior part of occipital region covered by cycloid scales; 9–11 dusky transverse bars laterally on trunk and tail; first dorsal fin gray or dusky after preservation (orange or reddish brown in life) without any distinct markings, having a line of black blotches (male) or a black band (female) on distal part of second dorsal fin; pectoral fin without clear markings.
</paragraph>
</subSubSection>
<subSubSection id="03FB34E6FFFC634AEE9CC754FE5C0305" lastPageId="8" lastPageNumber="389" pageId="2" pageNumber="383" type="description">
<paragraph id="4B5E676DFFFC6340EE9CC754FBDC037D" blockId="2.[151,1437,151,536]" pageId="2" pageNumber="383">
<emphasis id="7995BB7FFFFC6340EE9CC754FE900078" bold="true" box="[199,345,260,285]" pageId="2" pageNumber="383">Description.</emphasis>
Morphometric measurements given in Table 1. Body elongate, cylindrical anteriorly and somewhat compressed posteriorly. Head somewhat depressed with a round snout protruding beyond upper lip. Anterior nostril tubular and short, posterior nostril round or oblong, not tubular. Mouth inferior with upper jaw projecting beyond lower jaw. Upper lip thick with small medial cleft. Premaxillary teeth 45–71, fine and tricuspid. Dentary with canine-like symphyseal teeth (number of teeth
<quantity id="8C19CA88FFFC6340ED16C7C4FC5F00C8" box="[845,918,404,429]" metricMagnitude="-1" metricUnit="m" metricValue="1.016" metricValueMax="1.524" metricValueMin="0.508" pageId="2" pageNumber="383" unit="in" value="4.0" valueMax="6.0" valueMin="2.0">2–6 in</quantity>
male,
<quantity id="8C19CA88FFFC6340EDBEC7C4FBE400C8" box="[997,1069,404,429]" metricMagnitude="-2" metricUnit="m" metricValue="6.35" metricValueMax="10.16" metricValueMin="2.54" pageId="2" pageNumber="383" unit="in" value="2.5" valueMax="4.0" valueMin="1.0">1–4 in</quantity>
female) and a row of unicuspid horizontal teeth enclosed in fleshy sheath (number of teeth 45–79). Larger fish having more premaxillary and horizontal teeth (
<figureCitation id="D3DA7BE8FFFC6340EF39C78CFE670091" box="[354,430,476,501]" captionStart="FIGURE 2" captionStartId="4.[151,250,1696,1718]" captionTargetBox="[349,1239,204,1666]" captionTargetId="figure@4.[340,1247,193,1675]" captionTargetPageId="4" captionText="FIGURE 2. Number of premaxillary and horizontal dentary teeth of Stiphodon palawanensis (red circles), S. alcedo (pink circles), S. atratus (black diamonds), S. imperiorientis (light blue triangles), S. maculidorsalis (light green triangles), S. multisquamus (deep blue circles), S. ornatus (orange triangles), S. pelewensis (gray diamonds), S. pulchellus (deep green triangles), and S. weberi (purple diamonds)." httpUri="https://zenodo.org/record/232901/files/figure.png" pageId="2" pageNumber="383">Fig. 2</figureCitation>
). Urogenital papilla in male rounded with one cleft at posterior tip; that in female rectangular, bearing one small projection at each corner of the posterior margin.
</paragraph>
<paragraph id="4B5E676DFFFC6340EECCC418FC1C03E4" blockId="2.[151,1436,583,646]" pageId="2" pageNumber="383">
<emphasis id="7995BB7FFFFC6340EECCC418FEC3033A" bold="true" box="[151,266,584,607]" pageId="2" pageNumber="383">TABLE 1.</emphasis>
Morphometrics of
<taxonomicName id="8CE11CEEFFFC6340EF81C417FD14033A" box="[474,733,583,607]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="383" phylum="Chordata" rank="species" species="palawanensis">
<emphasis id="7995BB7FFFFC6340EF81C417FD14033A" box="[474,733,583,607]" italics="true" pageId="2" pageNumber="383">Stiphodon palawanensis</emphasis>
</taxonomicName>
, expressed as a percentage of standard length. D1, first dorsal fin; D2, second dorsal fin; A, anal fin; C, caudal fin; P1, pectoral fin; P2, pelvic fin.
</paragraph>
<paragraph id="4B5E676DFFFC6340EEC4C499FACC064C" pageId="2" pageNumber="383">
<table id="39E195CDFFFC9CBDEECCC4F0FA55064C" box="[151,1436,672,1833]" gridcols="4" gridrows="8" pageId="2" pageNumber="383">
<tr id="F5D1652FFFFC9CBDEECCC4F0FA55026D" box="[151,1436,672,776]" gridrow="0" pageId="2" pageNumber="383">
<th id="B6000C53FFFC9CBDEECCC4F0FE39026D" box="[151,496,672,776]" gridcol="0" gridrow="0" pageId="2" pageNumber="383">Sex Number of specimens measured</th>
<th id="B6000C53FFFC9CBDECF1C4F0FCC1026D" box="[682,776,672,776]" gridcol="1" gridrow="0" pageId="2" pageNumber="383">Holotype Male 1</th>
<th id="B6000C53FFFC9CBDEDC3C4F0FC37026D" box="[920,1022,672,776]" gridcol="2" gridrow="0" pageId="2" pageNumber="383">Paratype Male 23</th>
<th id="B6000C53FFFC9CBDEAFBC4F0FA55026D" box="[1184,1436,672,776]" gridcol="3" gridrow="0" pageId="2" pageNumber="383">Female 33</th>
</tr>
<tr id="F5D1652FFFFC9CBDEECCC54CFA5502E1" box="[151,1436,796,900]" gridrow="1" pageId="2" pageNumber="383">
<th id="B6000C53FFFC9CBDEECCC54CFE3902E1" box="[151,496,796,900]" gridcol="0" gridrow="1" pageId="2" pageNumber="383">Standard length (mm) Head length Snout length</th>
<td id="B6000C53FFFC9CBDECF1C54CFCC102E1" box="[682,776,796,900]" gridcol="1" gridrow="1" pageId="2" pageNumber="383">62.2 22.8 9.2</td>
<td id="B6000C53FFFC9CBDEDC3C54CFC3702E1" box="[920,1022,796,900]" gridcol="2" gridrow="1" pageId="2" pageNumber="383">27.7–63.9 21.5–23.9 6.8–9.0</td>
<td id="B6000C53FFFC9CBDEAFBC54CFA5502E1" box="[1184,1436,796,900]" gridcol="3" gridrow="1" pageId="2" pageNumber="383">27.0–63.6 20.8–25.2 6.8–9.2</td>
</tr>
<tr id="F5D1652FFFFC9CBDEECCC5C8FA550565" box="[151,1436,920,1024]" gridrow="2" pageId="2" pageNumber="383">
<th id="B6000C53FFFC9CBDEECCC5C8FE390565" box="[151,496,920,1024]" gridcol="0" gridrow="2" pageId="2" pageNumber="383">Eye diameter Postorbital length of head Upper jaw length</th>
<td id="B6000C53FFFC9CBDECF1C5C8FCC10565" box="[682,776,920,1024]" gridcol="1" gridrow="2" pageId="2" pageNumber="383">4.3 10.9 10.3</td>
<td id="B6000C53FFFC9CBDEDC3C5C8FC370565" box="[920,1022,920,1024]" gridcol="2" gridrow="2" pageId="2" pageNumber="383">4.1–6.0 9.8–12.0 9.0–11.2</td>
<td id="B6000C53FFFC9CBDEAFBC5C8FA550565" box="[1184,1436,920,1024]" gridcol="3" gridrow="2" pageId="2" pageNumber="383">4.4–7.0 9.4–12.0 9.1–10.8</td>
</tr>
<tr id="F5D1652FFFFC9CBDEECCC244FA550530" box="[151,1436,1044,1109]" gridrow="3" pageId="2" pageNumber="383">
<th id="B6000C53FFFC9CBDEECCC244FE390530" box="[151,496,1044,1109]" gridcol="0" gridrow="3" pageId="2" pageNumber="383">Body depth at P2 origin Body depth at A origin</th>
<td id="B6000C53FFFC9CBDECF1C244FCC10530" box="[682,776,1044,1109]" gridcol="1" gridrow="3" pageId="2" pageNumber="383">15.4 16.2</td>
<td id="B6000C53FFFC9CBDEDC3C244FC370530" box="[920,1022,1044,1109]" gridcol="2" gridrow="3" pageId="2" pageNumber="383">12.9–15.2 14.9–17.2</td>
<td id="B6000C53FFFC9CBDEAFBC244FA550530" box="[1184,1436,1044,1109]" gridcol="3" gridrow="3" pageId="2" pageNumber="383">13.0–15.6 15.3–17.9</td>
</tr>
<tr id="F5D1652FFFFC9CBDEECCC239FA5505CD" box="[151,1436,1129,1192]" gridrow="4" pageId="2" pageNumber="383">
<th id="B6000C53FFFC9CBDEECCC239FE3905CD" box="[151,496,1129,1192]" gridcol="0" gridrow="4" pageId="2" pageNumber="383">Depth of caudal peduncle Length of caudal peduncle</th>
<td id="B6000C53FFFC9CBDECF1C239FCC105CD" box="[682,776,1129,1192]" gridcol="1" gridrow="4" pageId="2" pageNumber="383">13.0 19.3</td>
<td id="B6000C53FFFC9CBDEDC3C239FC3705CD" box="[920,1022,1129,1192]" gridcol="2" gridrow="4" pageId="2" pageNumber="383">11.3–12.6 18.2–21.1</td>
<td id="B6000C53FFFC9CBDEAFBC239FA5505CD" box="[1184,1436,1129,1192]" gridcol="3" gridrow="4" pageId="2" pageNumber="383">10.5–12.3 18.3–21.4</td>
</tr>
<tr id="F5D1652FFFFC9CBDEECCC2ECFA550732" box="[151,1436,1212,1623]" gridrow="5" pageId="2" pageNumber="383">
<th id="B6000C53FFFC9CBDEECCC2ECFE390732" box="[151,496,1212,1623]" gridcol="0" gridrow="5" pageId="2" pageNumber="383">Predorsal length Length of D1 base D1 length Length of longest spine of D1 Interval between D1 and D2 bases Length of D2 base D2 length Length of longest ray of D2 Preanal length Length of A base</th>
<td id="B6000C53FFFC9CBDECF1C2ECFCC10732" box="[682,776,1212,1623]" gridcol="1" gridrow="5" pageId="2" pageNumber="383">32.0 18.6 35.4 30.5 3.2 26.7 45.3 21.1 51.4 29.4</td>
<td id="B6000C53FFFC9CBDEDC3C2ECFC370732" box="[920,1022,1212,1623]" gridcol="2" gridrow="5" pageId="2" pageNumber="383">31.9–35.0 16.6–21.7 25.3–41.7 18.4–35.9 1.7–5.4 23.6–27.2 37.2–46.8 14.4–21.5 49.5–54.0 26.5–30.3</td>
<td id="B6000C53FFFC9CBDEAFBC2ECFA550732" box="[1184,1436,1212,1623]" gridcol="3" gridrow="5" pageId="2" pageNumber="383">33.0–37.0 17.0–23.0 18.1–23.0 13.4–17.8 2.5–7.2 21.4–25.7 28.5–36.7 12.7–16.7 49.4–56.0 18.6–27.5</td>
</tr>
<tr id="F5D1652FFFFC9CBDEECCC03BFA5507B6" box="[151,1436,1643,1747]" gridrow="6" pageId="2" pageNumber="383">
<th id="B6000C53FFFC9CBDEECCC03BFE3907B6" box="[151,496,1643,1747]" gridcol="0" gridrow="6" pageId="2" pageNumber="383">A length Length of longest ray of A Length from anus to A</th>
<td id="B6000C53FFFC9CBDECF1C03BFCC107B6" box="[682,776,1643,1747]" gridcol="1" gridrow="6" pageId="2" pageNumber="383">43.9 18.2 3.5</td>
<td id="B6000C53FFFC9CBDEDC3C03BFC3707B6" box="[920,1022,1643,1747]" gridcol="2" gridrow="6" pageId="2" pageNumber="383">38.6–47.0 13.7–18.3 2.9–4.2</td>
<td id="B6000C53FFFC9CBDEAFBC03BFA5507B6" box="[1184,1436,1643,1747]" gridcol="3" gridrow="6" pageId="2" pageNumber="383">31.6–37.7 11.1–14.4 3.0–5.6</td>
</tr>
<tr id="F5D1652FFFFC9CBDEECCC0B7FA55064C" box="[151,1436,1767,1833]" gridrow="7" pageId="2" pageNumber="383">
<th id="B6000C53FFFC9CBDEECCC0B7FE39064C" box="[151,496,1767,1833]" gridcol="0" gridrow="7" pageId="2" pageNumber="383">Length of longest ray of P1 C length</th>
<td id="B6000C53FFFC9CBDECF1C0B7FCC1064C" box="[682,776,1767,1833]" gridcol="1" gridrow="7" pageId="2" pageNumber="383">20.7 31.4</td>
<td id="B6000C53FFFC9CBDEDC3C0B7FC37064C" box="[920,1022,1767,1833]" gridcol="2" gridrow="7" pageId="2" pageNumber="383">18.2–23.7 26.4–34.2</td>
<td id="B6000C53FFFC9CBDEAFBC0B7FA55064C" box="[1184,1436,1767,1833]" gridcol="3" gridrow="7" pageId="2" pageNumber="383">17.9–21.9 21.3–26.1</td>
</tr>
</table>
</paragraph>
<paragraph id="4B5E676DFFFC6341EE9CC13DFE1300C8" blockId="2.[151,1436,1901,2034]" lastBlockId="3.[151,1436,151,429]" lastPageId="3" lastPageNumber="384" pageId="2" pageNumber="383">First dorsal fin with six spines, except two specimens with five spines; second dorsal fin usually with one spine and nine segmented rays (one specimen with one spine and eight segmented rays). In female, first dorsal fin rounded, almost semicircular, usually second and/or third spines longest, but its posteriormost tip never extending to origin of second dorsal fin. In male, first dorsal fin spines elongate (usually fourth spine longest) and posteriormost point of first dorsal fin (tip of fourth spine) extending to base of first to sixth segmented ray of second dorsal fin when depressed. Anal fin with one spine and ten segmented rays. In female, usually first or second and second or third segmented rays longest in second dorsal and anal fins, respectively; in male, posterior rays longer than anterior rays (last ray and/or penultimate ray usually longest). Caudal fin usually with 17 segmented rays, including 13 branched rays, posterior margin rounded; caudal fin relatively larger in male than in female (caudal-fin length 26–34% of SL in male, 21–26% of SL in female). Pectoral fin with 14 (n=7), 15 (n=47), or 16 (n=3) rays. Pelvic fin with one spine and five segmented rays; pelvic fins joined together to form strong, cuplike disk with fleshy frenum.</paragraph>
<caption id="1F9E37E5FFFD6341EECCC1DFFEED0687" httpUri="https://zenodo.org/record/232900/files/figure.png" pageId="3" pageNumber="384" targetBox="[151,1436,481,1912]" targetPageId="3">
<paragraph id="4B5E676DFFFD6341EECCC1DFFEED0687" blockId="3.[151,1436,1935,2018]" pageId="3" pageNumber="384">
<emphasis id="7995BB7FFFFD6341EECCC1DFFEDD06C1" bold="true" box="[151,276,1935,1957]" pageId="3" pageNumber="384">FIGURE 1.</emphasis>
<taxonomicName id="8CE11CEEFFFD6341EF40C1DFFDDA06C0" box="[283,531,1935,1957]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="3" pageNumber="384" phylum="Chordata" rank="species" species="palawanensis">
<emphasis id="7995BB7FFFFD6341EF40C1DFFDDA06C0" box="[283,531,1935,1957]" italics="true" pageId="3" pageNumber="384">Stiphodon palawanensis</emphasis>
</taxonomicName>
immediately after fixation. a, WPU-PPC-P 5, holotype (male, 62.2 mm SL); b, URM-P 48659, paratype (male, 51.1 mm SL); c, URM-P 48666, paratype (female, 58.5 mm SL); d, URM-P 48661, paratype (female, 41.1 mm SL).
</paragraph>
</caption>
<caption id="1F9E37E5FFFA6346EECCC0F0FD9D0677" httpUri="https://zenodo.org/record/232901/files/figure.png" pageId="4" pageNumber="385" targetBox="[349,1239,204,1666]" targetPageId="4">
<paragraph id="4B5E676DFFFA6346EECCC0F0FD9D0677" blockId="4.[151,1436,1696,1810]" pageId="4" pageNumber="385">
<emphasis id="7995BB7FFFFA6346EECCC0F0FEDC07D0" bold="true" box="[151,277,1696,1718]" pageId="4" pageNumber="385">FIGURE 2.</emphasis>
Number of premaxillary and horizontal dentary teeth of
<taxonomicName id="8CE11CEEFFFA6346ED34C0F0FBA107D3" box="[879,1128,1696,1718]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="4" pageNumber="385" phylum="Chordata" rank="species" species="palawanensis">
<emphasis id="7995BB7FFFFA6346ED34C0F0FBA107D3" box="[879,1128,1696,1718]" italics="true" pageId="4" pageNumber="385">Stiphodon palawanensis</emphasis>
</taxonomicName>
(red circles),
<taxonomicName id="8CE11CEEFFFA6346EAA6C0F1FA9707D3" box="[1277,1374,1696,1718]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="4" pageNumber="385" phylum="Chordata" rank="species" species="alcedo">
<emphasis id="7995BB7FFFFA6346EAA6C0F1FA9707D3" box="[1277,1374,1696,1718]" italics="true" pageId="4" pageNumber="385">S. alcedo</emphasis>
</taxonomicName>
(pink circles),
<taxonomicName id="8CE11CEEFFFA6346EEA8C090FE9307B0" box="[243,346,1728,1749]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="4" pageNumber="385" phylum="Chordata" rank="species" species="atratus">
<emphasis id="7995BB7FFFFA6346EEA8C090FE9307B0" box="[243,346,1728,1749]" italics="true" pageId="4" pageNumber="385">S. atratus</emphasis>
</taxonomicName>
(black diamonds),
<taxonomicName id="8CE11CEEFFFA6346EC77C090FD1207B0" box="[556,731,1728,1749]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="4" pageNumber="385" phylum="Chordata" rank="species" species="imperiorientis">
<emphasis id="7995BB7FFFFA6346EC77C090FD1207B0" box="[556,731,1728,1749]" italics="true" pageId="4" pageNumber="385">S. imperiorientis</emphasis>
</taxonomicName>
(light blue triangles),
<taxonomicName id="8CE11CEEFFFA6346ED8AC090FB4C07B0" box="[977,1157,1727,1749]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="4" pageNumber="385" phylum="Chordata" rank="species" species="maculidorsalis">
<emphasis id="7995BB7FFFFA6346ED8AC090FB4C07B0" box="[977,1157,1727,1749]" italics="true" pageId="4" pageNumber="385">S. maculidorsalis</emphasis>
</taxonomicName>
(light green triangles),
<taxonomicName id="8CE11CEEFFFA6346EBD2C090FEED0796" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="4" pageNumber="385" phylum="Chordata" rank="species" species="multisquamus">
<emphasis id="7995BB7FFFFA6346EBD2C090FEED0796" italics="true" pageId="4" pageNumber="385">S. multisquamus</emphasis>
</taxonomicName>
(deep blue circles),
<taxonomicName id="8CE11CEEFFFA6346EC59C08EFDA40791" box="[514,621,1758,1780]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="4" pageNumber="385" phylum="Chordata" rank="species" species="ornatus">
<emphasis id="7995BB7FFFFA6346EC59C08EFDA40791" box="[514,621,1758,1780]" italics="true" pageId="4" pageNumber="385">S. ornatus</emphasis>
</taxonomicName>
(orange triangles),
<taxonomicName id="8CE11CEEFFFA6346ED1BC08EFC020796" box="[832,971,1757,1779]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="4" pageNumber="385" phylum="Chordata" rank="species" species="pelewensis">
<emphasis id="7995BB7FFFFA6346ED1BC08EFC020796" box="[832,971,1757,1779]" italics="true" pageId="4" pageNumber="385">S. pelewensis</emphasis>
</taxonomicName>
(gray diamonds),
<taxonomicName id="8CE11CEEFFFA6346EACAC08EFADF0796" box="[1169,1302,1757,1779]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="4" pageNumber="385" phylum="Chordata" rank="species" species="pulchellus">
<emphasis id="7995BB7FFFFA6346EACAC08EFADF0796" box="[1169,1302,1757,1779]" italics="true" pageId="4" pageNumber="385">S. pulchellus</emphasis>
</taxonomicName>
(deep green triangles), and
<taxonomicName id="8CE11CEEFFFA6346EF75C0ADFE450677" box="[302,396,1788,1810]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="4" pageNumber="385" phylum="Chordata" rank="species" species="weberi">
<emphasis id="7995BB7FFFFA6346EF75C0ADFE450677" box="[302,396,1788,1810]" italics="true" pageId="4" pageNumber="385">S. weberi</emphasis>
</taxonomicName>
(purple diamonds).
</paragraph>
</caption>
<caption id="1F9E37E5FFFB6347EECCC2FAFEB405BA" httpUri="https://zenodo.org/record/232902/files/figure.png" pageId="5" pageNumber="386" targetBox="[349,1236,184,1162]" targetPageId="5">
<paragraph id="4B5E676DFFFB6347EECCC2FAFEB405BA" blockId="5.[151,1436,1194,1247]" pageId="5" pageNumber="386">
<emphasis id="7995BB7FFFFB6347EECCC2FAFEDB05DA" bold="true" box="[151,274,1194,1216]" pageId="5" pageNumber="386">FIGURE 3.</emphasis>
Dorsal scalation on head and nape in
<taxonomicName id="8CE11CEEFFFB6347ECD4C2FAFC4C05A5" box="[655,901,1194,1216]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="5" pageNumber="386" phylum="Chordata" rank="species" species="palawanensis">
<emphasis id="7995BB7FFFFB6347ECD4C2FAFC4C05A5" box="[655,901,1194,1216]" italics="true" pageId="5" pageNumber="386">Stiphodon palawanensis</emphasis>
</taxonomicName>
. a, male (43.1 mm SL, CMK 11966); b, female (41.9 mm SL, CMK 11966).
</paragraph>
</caption>
<paragraph id="4B5E676DFFFB6347EE9CC35BFD3B0745" blockId="5.[151,1437,1291,2036]" pageId="5" pageNumber="386">
Scales in a longitudinal series 29 (n=4), 30 (n=13), 31 (n=19), 32 (n=15, including
<typeStatus id="945AD9CFFFFB6347EAC9C35BFB300441" box="[1170,1273,1291,1316]" pageId="5" pageNumber="386" type="holotype">holotype</typeStatus>
), or 33 (n=6); scales in a transverse series 9 (n=1), 10 (n=3), 11 (n=52, including
<typeStatus id="945AD9CFFFFB6347ED25C37FFC2D042D" box="[894,996,1327,1352]" pageId="5" pageNumber="386" type="holotype">holotype</typeStatus>
), 12 (n=1); circumpeduncular scales 15 (n=1) or 16 (n=55) (not including one broken specimen). Ctenoid scales covering tail, sides and dorsum of posterior trunk. Pectoral-fin base naked. Small naked area behind pectoral-fin base; some anteriormost scales on lateral sides of trunk cycloid. Belly covered with cycloid scales. Nape and posterior part of occipital region covered by cycloid scales (
<figureCitation id="D3DA7BE8FFFB6347EF37C3EFFE7804BD" box="[364,433,1471,1496]" captionStart="FIGURE 3" captionStartId="5.[151,250,1194,1216]" captionTargetBox="[349,1236,184,1162]" captionTargetId="figure@5.[340,1247,173,1173]" captionTargetPageId="5" captionText="FIGURE 3. Dorsal scalation on head and nape in Stiphodon palawanensis. a, male (43.1 mm SL, CMK 11966); b, female (41.9 mm SL, CMK 11966)." httpUri="https://zenodo.org/record/232902/files/figure.png" pageId="5" pageNumber="386">Fig. 3</figureCitation>
); in female, scaled area slightly exceeding middle of occipital region; male similar, but sometimes not exceeding the middle. Cycloid scales also occurring on first and second dorsal-fin bases, anal-fin base, caudal-fin base, and proximal part of caudal fin.
</paragraph>
<paragraph id="4B5E676DFFFB6347EE9CC07BFE6D07E9" blockId="5.[151,1437,1291,2036]" pageId="5" pageNumber="386">
Cephalic sensory pore system always with A´, B, C, D(S), F, H´, K´, L´, N´, and
<collectingCountry id="33F627FDFFFB6347EA35C07CFB4D0721" box="[1134,1156,1580,1604]" name="Iceland" pageId="5" pageNumber="386">O</collectingCountry>
´(
<figureCitation id="D3DA7BE8FFFB6347EAC6C07BFB2D0721" box="[1181,1252,1579,1604]" captionStart="FIGURE 4" captionStartId="6.[151,250,1952,1974]" captionTargetBox="[384,1184,211,1923]" captionTargetId="figure@6.[376,1206,193,1942]" captionTargetPageId="6" captionText="FIGURE 4. Diagrammatic illustration of head showing arrangement of cephalic sensory pores and cutaneous sensory papillae in Stiphodon palawanensis (41.9 mm SL, CMK 11966). a, dorsal view; b, lateral view; c, ventral view. AN, anterior nostril; PN, posterior nostril." httpUri="https://zenodo.org/record/232903/files/figure.png" pageId="5" pageNumber="386">Fig. 4</figureCitation>
). Oculoscapular canal interrupted between pores H´and K´. Cutaneous sensory papillae developed over dorsal, lateral, and ventral surface of head (
<figureCitation id="D3DA7BE8FFFB6347EF0AC023FE5F07E9" box="[337,406,1651,1676]" captionStart="FIGURE 4" captionStartId="6.[151,250,1952,1974]" captionTargetBox="[384,1184,211,1923]" captionTargetId="figure@6.[376,1206,193,1942]" captionTargetPageId="6" captionText="FIGURE 4. Diagrammatic illustration of head showing arrangement of cephalic sensory pores and cutaneous sensory papillae in Stiphodon palawanensis (41.9 mm SL, CMK 11966). a, dorsal view; b, lateral view; c, ventral view. AN, anterior nostril; PN, posterior nostril." httpUri="https://zenodo.org/record/232903/files/figure.png" pageId="5" pageNumber="386">Fig. 4</figureCitation>
).
</paragraph>
<paragraph id="4B5E676DFFFB6347EE9CC0C7FEED07B1" blockId="5.[151,1437,1291,2036]" pageId="5" pageNumber="386">
The largest male and female specimens were 63.9 and
<quantity id="8C19CA88FFFB6347ED0CC0C7FC0907CA" box="[855,960,1687,1712]" metricMagnitude="-2" metricUnit="m" metricValue="6.36" pageId="5" pageNumber="386" unit="mm" value="63.6">63.6 mm</quantity>
SL (82.9 and
<quantity id="8C19CA88FFFB6347EA29C0C8FB1207CA" box="[1138,1243,1688,1712]" metricMagnitude="-2" metricUnit="m" metricValue="7.79" pageId="5" pageNumber="386" unit="mm" value="77.9">77.9 mm</quantity>
in total length), respectively.
</paragraph>
<paragraph id="4B5E676DFFFB6347EE9CC08FFE9E0691" blockId="5.[151,1437,1291,2036]" pageId="5" pageNumber="386">
<emphasis id="7995BB7FFFFB6347EE9CC08FFE0E0792" box="[199,455,1759,1783]" italics="true" pageId="5" pageNumber="386">Color in preservative.</emphasis>
In male, background brown; 9–11 (usually 10) dusky transverse bars laterally and dorsally on trunk and tail. First dorsal fin gray or dusky without distinct markings. Second dorsal fin gray or dusky; each ray having a black blotch surrounded by a translucent margin, these black blotches forming a line along second dorsal fin margin, but posterior part of the fin sometimes lacking this black blotch. Anal fin gray or dusky. Caudal fin with dusky longitudinal band on upper part; dorsal to this band, gray; ventrally along this band translucent; middle and lower parts of caudal fin gray. Pectoral fin pale gray without clear markings, but sometimes with one to four obscure, dusky spots on each of middle rays. Pelvic disk dusky, but margin of anterior and middle parts translucent.
</paragraph>
<caption id="1F9E37E5FFF86344EECCC1F0FEF60691" httpUri="https://zenodo.org/record/232903/files/figure.png" pageId="6" pageNumber="387" targetBox="[384,1184,211,1923]" targetPageId="6">
<paragraph id="4B5E676DFFF86344EECCC1F0FEF60691" blockId="6.[151,1436,1952,2036]" pageId="6" pageNumber="387">
<emphasis id="7995BB7FFFF86344EECCC1F0FEDA06D0" bold="true" box="[151,275,1952,1974]" pageId="6" pageNumber="387">FIGURE 4.</emphasis>
Diagrammatic illustration of head showing arrangement of cephalic sensory pores and cutaneous sensory papillae in
<taxonomicName id="8CE11CEEFFF86344EEEBC1EFFE6F06B0" box="[176,422,1983,2005]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="6" pageNumber="387" phylum="Chordata" rank="species" species="palawanensis">
<emphasis id="7995BB7FFFF86344EEEBC1EFFE6F06B0" box="[176,422,1983,2005]" italics="true" pageId="6" pageNumber="387">Stiphodon palawanensis</emphasis>
</taxonomicName>
(41.9 mm SL, CMK 11966). a, dorsal view; b, lateral view; c, ventral view. AN, anterior nostril; PN, posterior nostril.
</paragraph>
</caption>
<caption id="1F9E37E5FFF96345EECCC02CFBD707D4" httpUri="https://zenodo.org/record/232904/files/figure.png" pageId="7" pageNumber="388" targetBox="[151,1436,193,1637]" targetPageId="7">
<paragraph id="4B5E676DFFF96345EECCC02CFBD707D4" blockId="7.[151,1436,1660,1713]" pageId="7" pageNumber="388">
<emphasis id="7995BB7FFFF96345EECCC02CFEDA07F7" bold="true" box="[151,275,1660,1682]" pageId="7" pageNumber="388">FIGURE 5.</emphasis>
Males (a–d) and females (e, f) of live
<taxonomicName id="8CE11CEEFFF96345ECC0C02CFC5807F7" box="[667,913,1660,1682]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="7" pageNumber="388" phylum="Chordata" rank="species" species="palawanensis">
<emphasis id="7995BB7FFFF96345ECC0C02CFC5807F7" box="[667,913,1660,1682]" italics="true" pageId="7" pageNumber="388">Stiphodon palawanensis</emphasis>
</taxonomicName>
observed in Balsahan Stream, Puerto Princesa City (b, e, f, 18 May 2015; a, c, 19 May 2015) and Barake Stream, Aborlan (d, 15 May 2015).
</paragraph>
</caption>
<paragraph id="4B5E676DFFF96345EE9CC08EFA540697" blockId="7.[151,1437,1757,2034]" pageId="7" pageNumber="388">In female, background brown or yellowish brown; blackish longitudinal band extending from snout and upper lip to below eye and to middle of pectoral-fin base, band continuing from behind pectoral-fin base to posterior end of caudal peduncle through lateral midline; nine or ten (usually ten) dusky transverse bars laterally on trunk and tail intersecting with the mid-lateral longitudinal band. The longitudinal band often unclear if the transverse bars accentuated, and transverse bars often unclear if the longitudinal band accentuated. Additional blackish longitudinal band above the mid-lateral band from dorsoposterior edge of eye to base of upper procurrent caudalfin rays, but often obscure. First dorsal-fin gray without distinct markings. Second dorsal and anal fins gray with a black submarginal band and translucent margin; the black band on second dorsal fin thicker than that on anal fin.</paragraph>
<paragraph id="4B5E676DFFF6634AEECCC6C7FF1D0025" blockId="8.[151,1437,151,861]" pageId="8" pageNumber="389">Black blotch on middle of proximal part of caudal fin; other part of caudal fin translucent with a black band (upside-down “L” shape) along dorsal and posterior margins and one to four black vertical stripes on middle part. Pectoral fin pale gray without clear markings, but sometimes with one or two obscure, dusky spots along middle rays. Pelvic fin translucent, but middle parts of rays, membranes, and frenum often dusky, forming a ring in ventral view.</paragraph>
<paragraph id="4B5E676DFFF6634AEE9CC71CFE5C0305" blockId="8.[151,1437,151,861]" pageId="8" pageNumber="389">
<emphasis id="7995BB7FFFF6634AEE9CC71CFE900001" box="[199,345,332,356]" italics="true" pageId="8" pageNumber="389">Color in life.</emphasis>
Body and fin markings of male and female similar to those of preserved specimens, but in male (
<figureCitation id="D3DA7BE8FFF6634AEEC4C73FFF2D00ED" box="[159,228,367,392]" captionStart="FIGURE 5" captionStartId="7.[151,250,1660,1682]" captionTargetBox="[151,1436,193,1637]" captionTargetId="figure@7.[151,1436,193,1639]" captionTargetPageId="7" captionText="FIGURE 5. Males (a – d) and females (e, f) of live Stiphodon palawanensis observed in Balsahan Stream, Puerto Princesa City (b, e, f, 18 May 2015; a, c, 19 May 2015) and Barake Stream, Aborlan (d, 15 May 2015)." httpUri="https://zenodo.org/record/232904/files/figure.png" pageId="8" pageNumber="389">Fig. 5</figureCitation>
a–d), background of body grayish, purplish, or yellowish brown; first and second dorsal fins orange or reddish brown; black blotches on second dorsal fin surrounded by white or yellow margins; anal fin brown, gray, or orange; and dorsal part of caudal fin reddish or yellowish with bluish-white margin. In female (
<figureCitation id="D3DA7BE8FFF6634AEAB1C7E7FAE600B5" box="[1258,1327,439,464]" captionStart="FIGURE 5" captionStartId="7.[151,250,1660,1682]" captionTargetBox="[151,1436,193,1637]" captionTargetId="figure@7.[151,1436,193,1639]" captionTargetPageId="7" captionText="FIGURE 5. Males (a – d) and females (e, f) of live Stiphodon palawanensis observed in Balsahan Stream, Puerto Princesa City (b, e, f, 18 May 2015; a, c, 19 May 2015) and Barake Stream, Aborlan (d, 15 May 2015)." httpUri="https://zenodo.org/record/232904/files/figure.png" pageId="8" pageNumber="389">Fig. 5</figureCitation>
e, f), first dorsal fin reddish brown; second dorsal fin reddish brown with a black submarginal band and bluish white margin; white spots often lining proximal side of this band; anal fin brown with a black submarginal band and bluish white margin; upper and middle parts of caudal fin fringed by bluish white margin; area between the white margin and the black band orange.
</paragraph>
</subSubSection>
<subSubSection id="03FB34E6FFF6634BEE9CC43CFE5100B8" lastPageId="9" lastPageNumber="390" pageId="8" pageNumber="389" type="distribution">
<paragraph id="4B5E676DFFF6634AEE9CC43CFC140238" blockId="8.[151,1437,151,861]" pageId="8" pageNumber="389">
<emphasis id="7995BB7FFFF6634AEE9CC43CFEA903E0" bold="true" box="[199,352,620,645]" pageId="8" pageNumber="389">Distribution.</emphasis>
The new species is currently known only from Palawan, in the western
<collectingCountry id="33F627FDFFF6634AEACCC43CFAD003E0" box="[1175,1305,620,645]" name="Philippines" pageId="8" pageNumber="389">Philippines</collectingCountry>
. All known habitats are streams flowing into the Sulu Sea in the central part of the island (
<figureCitation id="D3DA7BE8FFF6634AEA6EC4DFFBB403CD" box="[1077,1149,655,680]" captionStart="FIGURE 6" captionStartId="8.[151,250,1866,1888]" captionTargetBox="[345,1240,917,1839]" captionTargetId="figure@8.[340,1248,913,1845]" captionTargetPageId="8" captionText="FIGURE 6. Location of Palawan in the Philippines (inset map) and sites where Stiphodon species have been found on Palawan. Red circles, S. palawanensis; green circles, S. pulchellus; blue circle, S. atropurpureus; and orange circle, S. percnopterygionus." httpUri="https://zenodo.org/record/232905/files/figure.png" pageId="8" pageNumber="389">Fig. 6</figureCitation>
). When we explored the Iwahig River where it flows into the South
<collectingCountry id="33F627FDFFF6634AECEEC4E4FD3103A8" box="[693,760,692,717]" name="China" pageId="8" pageNumber="389">China</collectingCountry>
Sea at Quezon (
<figureCitation id="D3DA7BE8FFF6634AED97C4E4FBDF03A8" box="[972,1046,692,717]" captionStart="FIGURE 6" captionStartId="8.[151,250,1866,1888]" captionTargetBox="[345,1240,917,1839]" captionTargetId="figure@8.[340,1248,913,1845]" captionTargetPageId="8" captionText="FIGURE 6. Location of Palawan in the Philippines (inset map) and sites where Stiphodon species have been found on Palawan. Red circles, S. palawanensis; green circles, S. pulchellus; blue circle, S. atropurpureus; and orange circle, S. percnopterygionus." httpUri="https://zenodo.org/record/232905/files/figure.png" pageId="8" pageNumber="389">Fig. 6</figureCitation>
) for two days in
<date id="3F5F41ADFFF6634AEAA3C4E5FABC03A8" box="[1272,1397,692,717]" pageId="8" pageNumber="389" value="2015-05">May 2015</date>
,
<taxonomicName id="8CE11CEEFFF6634AEBDCC4E4FEF90395" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="8" pageNumber="389" phylum="Chordata" rank="species" species="palawanensis">
<emphasis id="7995BB7FFFF6634AEBDCC4E4FEF90395" italics="true" pageId="8" pageNumber="389">S. palawanensis</emphasis>
</taxonomicName>
was not found, but other areas, such as the northern and southern parts of the island, have not yet been explored. Given its amphidromous life cycle with the high dispersal ability of the larvae suggested in this genus (
<bibRefCitation id="2F701A9CFFF6634AEEB7C54FFE3A025D" author="Yamasaki" box="[236,499,799,824]" pageId="8" pageNumber="389" refString="Yamasaki, N., Maeda, K. &amp; Tachihara, K. (2007) Pelagic larval duration and morphology at recruitment of Stiphodon percnopterygionus (Gobiidae: Sicydiinae). Raffles Bulletin of Zoology, 14 (Supplement), 209 - 214." type="journal article" year="2007">
Yamasaki
<emphasis id="7995BB7FFFF6634AEF3CC571FE6C025D" box="[359,421,800,824]" italics="true" pageId="8" pageNumber="389">et al.</emphasis>
, 2007
</bibRefCitation>
;
<bibRefCitation id="2F701A9CFFF6634AEC5FC54FFD31025D" author="Maeda" box="[516,760,799,824]" pageId="8" pageNumber="389" refString="Maeda, K., Mukai, T. &amp; Tachihara, K. (2012 b) A new species of amphidromous goby, Stiphodon alcedo, from the Ryukyu Archipelago (Gobiidae: Sicydiinae). Cybium, 35, 285 - 298." type="journal article" year="2012" yearSuffix="b">
Maeda
<emphasis id="7995BB7FFFF6634AEC06C571FD53025D" box="[605,666,800,824]" italics="true" pageId="8" pageNumber="389">et al.</emphasis>
, 2012b
</bibRefCitation>
), and the geographically limited survey made to date,
<taxonomicName id="8CE11CEEFFF6634AEBDCC570FEF90239" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="8" pageNumber="389" phylum="Chordata" rank="species" species="palawanensis">
<emphasis id="7995BB7FFFF6634AEBDCC570FEF90239" italics="true" pageId="8" pageNumber="389">S. palawanensis</emphasis>
</taxonomicName>
is expected to be found elsewhere in the region in the future.
</paragraph>
<caption id="1F9E37E5FFF6634AEECCC11AFE9306F8" httpUri="https://zenodo.org/record/232905/files/figure.png" pageId="8" pageNumber="389" targetBox="[345,1240,917,1839]" targetPageId="8">
<paragraph id="4B5E676DFFF6634AEECCC11AFE9306F8" blockId="8.[151,1436,1866,1950]" pageId="8" pageNumber="389">
<emphasis id="7995BB7FFFF6634AEECCC11AFEDE063A" bold="true" box="[151,279,1866,1888]" pageId="8" pageNumber="389">FIGURE 6.</emphasis>
Location of Palawan in the Philippines (inset map) and sites where
<taxonomicName id="8CE11CEEFFF6634AEA59C11AFBA10605" box="[1026,1128,1866,1888]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="8" pageNumber="389" phylum="Chordata" rank="genus">
<emphasis id="7995BB7FFFF6634AEA59C11AFBA10605" box="[1026,1128,1866,1888]" italics="true" pageId="8" pageNumber="389">Stiphodon</emphasis>
</taxonomicName>
species have been found on Palawan. Red circles,
<taxonomicName id="8CE11CEEFFF6634AEFD3C13AFDF8061A" box="[392,561,1897,1919]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="8" pageNumber="389" phylum="Chordata" rank="species" species="palawanensis">
<emphasis id="7995BB7FFFF6634AEFD3C13AFDF8061A" box="[392,561,1897,1919]" italics="true" pageId="8" pageNumber="389">S. palawanensis</emphasis>
</taxonomicName>
; green circles,
<taxonomicName id="8CE11CEEFFF6634AEC86C13AFCAD061A" box="[733,868,1897,1919]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="8" pageNumber="389" phylum="Chordata" rank="species" species="pulchellus">
<emphasis id="7995BB7FFFF6634AEC86C13AFCAD061A" box="[733,868,1897,1919]" italics="true" pageId="8" pageNumber="389">S. pulchellus</emphasis>
</taxonomicName>
; blue circle,
<taxonomicName id="8CE11CEEFFF6634AEDA1C13AFB60061A" box="[1018,1193,1898,1919]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="8" pageNumber="389" phylum="Chordata" rank="species" species="atropurpureus">
<emphasis id="7995BB7FFFF6634AEDA1C13AFB60061A" box="[1018,1193,1898,1919]" italics="true" pageId="8" pageNumber="389">S. atropurpureus</emphasis>
</taxonomicName>
; and orange circle,
<taxonomicName id="8CE11CEEFFF6634AEBD2C13AFE9D06FB" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="8" pageNumber="389" phylum="Chordata" rank="species" species="percnopterygionus">
<emphasis id="7995BB7FFFF6634AEBD2C13AFE9D06FB" italics="true" pageId="8" pageNumber="389">S. percnopterygionus</emphasis>
</taxonomicName>
.
</paragraph>
</caption>
<paragraph id="4B5E676DFFF7634BEE9CC6C7FE5100B8" blockId="9.[151,1437,151,1943]" pageId="9" pageNumber="390">
<emphasis id="7995BB7FFFF7634BEE9CC6C7FEE201D5" bold="true" box="[199,299,151,176]" pageId="9" pageNumber="390">Ecology.</emphasis>
The new species was one of the dominant fish species observed in the middle reaches of Balsahan Stream (Puerto Princesa City) and Barake Stream (Aborlan) in
<date id="3F5F41ADFFF7634BED38C6EAFC1301B7" box="[867,986,185,210]" pageId="9" pageNumber="390" value="2015-05">May 2015</date>
.
<taxonomicName id="8CE11CEEFFF7634BEDBEC6E9FB3F01B4" box="[997,1270,185,209]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="palawanensis">
<emphasis id="7995BB7FFFF7634BEDBEC6E9FB3F01B4" box="[997,1270,185,209]" italics="true" pageId="9" pageNumber="390">Stiphodon palawanensis</emphasis>
</taxonomicName>
inhabits pools with a substrate that is a mixture of boulders, gravel, and pebbles with exposed bedrock in some places. It also inhabits rapids. The water was clear. The fish were clinging to the rocks while feeding on algae. When disturbed, they swam to nearby rocks or hid in crevices under or between rocks. The maximum water depth of the sites was
<quantity id="8C19CA88FFF7634BEECCC76EFF1E0033" box="[151,215,318,343]" metricMagnitude="0" metricUnit="m" metricValue="1.5" pageId="9" pageNumber="390" unit="m" value="1.5">1.5 m</quantity>
in May, but it could reach 3.0 m during the wet season. Normally, that occurs from June to December, while the dry season runs from January to May. In these two sites sampled, two cyprinid species
<taxonomicName id="8CE11CEEFFF7634BEACCC730FED700FF" authority="Boulenger" authorityName="Boulenger" class="Actinopterygii" family="Cyprinidae" genus="Barbodes" kingdom="Animalia" order="Cypriniformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="palavanensis">
<emphasis id="7995BB7FFFF7634BEACCC730FA55001D" box="[1175,1436,352,376]" italics="true" pageId="9" pageNumber="390">Barbodes palavanensis</emphasis>
(Boulenger)
</taxonomicName>
and
<taxonomicName id="8CE11CEEFFF7634BEF0CC7D1FD4400FF" authority="Boulenger" authorityName="Boulenger" box="[343,653,385,410]" class="Actinopterygii" family="Cyprinidae" genus="Rasbora" kingdom="Animalia" order="Cypriniformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="everetti">
<emphasis id="7995BB7FFFF7634BEF0CC7D1FDD800FC" box="[343,529,385,409]" italics="true" pageId="9" pageNumber="390">Rasbora everetti</emphasis>
Boulenger
</taxonomicName>
were very abundant with
<taxonomicName id="8CE11CEEFFF7634BEDECC7D1FBA400FC" box="[951,1133,385,409]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="palawanensis">
<emphasis id="7995BB7FFFF7634BEDECC7D1FBA400FC" box="[951,1133,385,409]" italics="true" pageId="9" pageNumber="390">S. palawanensis</emphasis>
</taxonomicName>
. Other common species at these sites were two gobioids,
<taxonomicName id="8CE11CEEFFF7634BEFB0C7F3FCB400DE" authority="Hoese &amp; Allen" authorityName="Hoese &amp; Allen" box="[491,893,418,443]" class="Actinopterygii" family="Gobiidae" genus="Glossogobius" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="illimis">
<emphasis id="7995BB7FFFF7634BEFB0C7F3FD1900DE" box="[491,720,419,443]" italics="true" pageId="9" pageNumber="390">Glossogobius illimis</emphasis>
Hoese &amp; Allen
</taxonomicName>
and
<taxonomicName id="8CE11CEEFFF7634BEDE8C7F3FBF800DE" box="[947,1073,419,443]" class="Actinopterygii" family="Gobiidae" genus="Redigobius" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="genus">
<emphasis id="7995BB7FFFF7634BEDE8C7F3FBF800DE" box="[947,1073,419,443]" italics="true" pageId="9" pageNumber="390">Redigobius</emphasis>
</taxonomicName>
sp., and a halfbeak
<taxonomicName id="8CE11CEEFFF7634BEB55C7F3FE5100B8" authority="Meisner." authorityName="Meisner." class="Actinopterygii" family="Zenarchopteridae" genus="Dermogenys" kingdom="Animalia" order="Beloniformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="palawanensis">
<emphasis id="7995BB7FFFF7634BEB55C7F3FEF900B9" italics="true" pageId="9" pageNumber="390">Dermogenys palawanensis</emphasis>
Meisner.
</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection id="03FB34E6FFF76348EE9CC7B5FEB106EB" lastPageId="10" lastPageNumber="391" pageId="9" pageNumber="390" type="etymology">
<paragraph id="4B5E676DFFF7634BEE9CC7B5FF16037B" blockId="9.[151,1437,151,1943]" pageId="9" pageNumber="390">
<emphasis id="7995BB7FFFF7634BEE9CC7B5FE87009B" bold="true" box="[199,334,485,510]" pageId="9" pageNumber="390">Etymology.</emphasis>
The name of the new species is derived from Palawan, the
<typeStatus id="945AD9CFFFF7634BEA4DC7B6FB8F009B" box="[1046,1094,486,510]" pageId="9" pageNumber="390">type</typeStatus>
locality, and the Latin suffix
<emphasis id="7995BB7FFFF7634BEECCC457FF11037B" box="[151,216,519,542]" italics="true" pageId="9" pageNumber="390">-ensis</emphasis>
.
</paragraph>
<paragraph id="4B5E676DFFF7634BEE9CC477FE0C02B5" blockId="9.[151,1437,151,1943]" pageId="9" pageNumber="390">
<emphasis id="7995BB7FFFF7634BEE9CC477FEA80325" bold="true" box="[199,353,551,576]" pageId="9" pageNumber="390">Comparison.</emphasis>
The new species was compared with congeners sharing the second dorsal- and pectoral-fin ray counts (one spine and nine segmented rays in the second dorsal fin, and usually 15 or 16 rays in the pectoral fin).
<taxonomicName id="8CE11CEEFFF7634BEECCC43BFE6003E6" box="[151,425,619,643]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="palawanensis">
<emphasis id="7995BB7FFFF7634BEECCC43BFE6003E6" box="[151,425,619,643]" italics="true" pageId="9" pageNumber="390">Stiphodon palawanensis</emphasis>
</taxonomicName>
shares 9–11 dusky transverse bars laterally on the trunk and tail with
<taxonomicName id="8CE11CEEFFF7634BEA8CC43BFEF803C1" authority="Maeda &amp; Tan" authorityName="Maeda &amp; Tan" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="maculidorsalis">
<emphasis id="7995BB7FFFF7634BEA8CC43BFA5203E6" box="[1239,1435,619,643]" italics="true" pageId="9" pageNumber="390">S. maculidorsalis</emphasis>
Maeda &amp; Tan
</taxonomicName>
and
<taxonomicName id="8CE11CEEFFF7634BEF33C4DCFD5803C0" authority="Wu &amp; Ni" authorityName="Wu &amp; Ni" box="[360,657,652,677]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="multisquamus">
<emphasis id="7995BB7FFFF7634BEF33C4DCFDE903C1" box="[360,544,652,676]" italics="true" pageId="9" pageNumber="390">S. multisquamus</emphasis>
Wu &amp; Ni
</taxonomicName>
, but it differs from
<taxonomicName id="8CE11CEEFFF7634BED36C4DCFBF903C1" box="[877,1072,652,676]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="maculidorsalis">
<emphasis id="7995BB7FFFF7634BED36C4DCFBF903C1" box="[877,1072,652,676]" italics="true" pageId="9" pageNumber="390">S. maculidorsalis</emphasis>
</taxonomicName>
in the lack of clear markings on the pectoral fin (vs. having fine black spots along rays), dorsal markings (black spots scattered dorsally on the head and tail of
<taxonomicName id="8CE11CEEFFF7634BEF55C49FFE060383" box="[270,463,718,742]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="maculidorsalis">
<emphasis id="7995BB7FFFF7634BEF55C49FFE060383" box="[270,463,718,742]" italics="true" pageId="9" pageNumber="390">S. maculidorsalis</emphasis>
</taxonomicName>
, but not on
<taxonomicName id="8CE11CEEFFF7634BEC0FC49FFCC10383" box="[596,776,718,742]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="palawanensis">
<emphasis id="7995BB7FFFF7634BEC0FC49FFCC10383" box="[596,776,718,742]" italics="true" pageId="9" pageNumber="390">S. palawanensis</emphasis>
</taxonomicName>
), and predorsal scalation (the posterior part of the occipital region is scaled in
<taxonomicName id="8CE11CEEFFF7634BEF34C4A0FDED026D" box="[367,548,752,776]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="palawanensis">
<emphasis id="7995BB7FFFF7634BEF34C4A0FDED026D" box="[367,548,752,776]" italics="true" pageId="9" pageNumber="390">S. palawanensis</emphasis>
</taxonomicName>
,
<figureCitation id="D3DA7BE8FFF7634BEC6FC4BFFDB0026D" box="[564,633,751,776]" captionStart="FIGURE 3" captionStartId="5.[151,250,1194,1216]" captionTargetBox="[349,1236,184,1162]" captionTargetId="figure@5.[340,1247,173,1173]" captionTargetPageId="5" captionText="FIGURE 3. Dorsal scalation on head and nape in Stiphodon palawanensis. a, male (43.1 mm SL, CMK 11966); b, female (41.9 mm SL, CMK 11966)." httpUri="https://zenodo.org/record/232902/files/figure.png" pageId="9" pageNumber="390">Fig. 3</figureCitation>
, vs. an occipital region that is almost naked in
<taxonomicName id="8CE11CEEFFF7634BEACDC4A0FA93026D" box="[1174,1370,752,776]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="maculidorsalis">
<emphasis id="7995BB7FFFF7634BEACDC4A0FA93026D" box="[1174,1370,752,776]" italics="true" pageId="9" pageNumber="390">S. maculidorsalis</emphasis>
</taxonomicName>
); and from
<taxonomicName id="8CE11CEEFFF7634BEE8FC542FE42024C" box="[212,395,785,809]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="multisquamus">
<emphasis id="7995BB7FFFF7634BEE8FC542FE42024C" box="[212,395,785,809]" italics="true" pageId="9" pageNumber="390">S. multisquamus</emphasis>
</taxonomicName>
by the lack of distinct markings on the first dorsal and pectoral fins (vs. having a black blotch on the posterior part of the first dorsal fin in males, thick dusky lines along spines of the first dorsal fin in females, and fine black spots along the pectoral-fin rays in both sexes), having a line of black blotches (in males) or a black band (in females) on the distal part of the second dorsal fin (vs. no such black markings), and predorsal scalation of males (posterior part of the occipital region scaled in
<taxonomicName id="8CE11CEEFFF7634BECAAC5C7FC6E02CB" box="[753,935,918,942]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="palawanensis">
<emphasis id="7995BB7FFFF7634BECAAC5C7FC6E02CB" box="[753,935,918,942]" italics="true" pageId="9" pageNumber="390">S. palawanensis</emphasis>
</taxonomicName>
,
<figureCitation id="D3DA7BE8FFF7634BEDEEC5C6FC3002CA" box="[949,1017,918,943]" captionStart="FIGURE 3" captionStartId="5.[151,250,1194,1216]" captionTargetBox="[349,1236,184,1162]" captionTargetId="figure@5.[340,1247,173,1173]" captionTargetPageId="5" captionText="FIGURE 3. Dorsal scalation on head and nape in Stiphodon palawanensis. a, male (43.1 mm SL, CMK 11966); b, female (41.9 mm SL, CMK 11966)." httpUri="https://zenodo.org/record/232902/files/figure.png" pageId="9" pageNumber="390">Fig. 3</figureCitation>
, vs. an occipital region that is almost naked in
<taxonomicName id="8CE11CEEFFF7634BEEA6C5E8FE7C02B5" box="[253,437,952,976]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="multisquamus">
<emphasis id="7995BB7FFFF7634BEEA6C5E8FE7C02B5" box="[253,437,952,976]" italics="true" pageId="9" pageNumber="390">S. multisquamus</emphasis>
</taxonomicName>
).
</paragraph>
<paragraph id="4B5E676DFFF7634BEE9CC589FDD90512" blockId="9.[151,1437,151,1943]" pageId="9" pageNumber="390">
<taxonomicName id="8CE11CEEFFF7634BEE9CC589FD8B0297" authority="Maeda" authorityName="Maeda" box="[199,578,985,1010]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="niraikanaiensis">
<emphasis id="7995BB7FFFF7634BEE9CC589FE240294" box="[199,493,985,1009]" italics="true" pageId="9" pageNumber="390">Stiphodon niraikanaiensis</emphasis>
Maeda
</taxonomicName>
males have black longitudinal bands on the second dorsal and caudal fins, but females lack them. Although a line of black blotches on the second dorsal fin in
<taxonomicName id="8CE11CEEFFF7634BEACEC5ABFA860576" box="[1173,1359,1019,1043]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="palawanensis">
<emphasis id="7995BB7FFFF7634BEACEC5ABFA860576" box="[1173,1359,1019,1043]" italics="true" pageId="9" pageNumber="390">S. palawanensis</emphasis>
</taxonomicName>
males resembles a black band when the fin is not fully open,
<taxonomicName id="8CE11CEEFFF7634BED5FC24CFC700551" box="[772,953,1052,1076]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="palawanensis">
<emphasis id="7995BB7FFFF7634BED5FC24CFC700551" box="[772,953,1052,1076]" italics="true" pageId="9" pageNumber="390">S. palawanensis</emphasis>
</taxonomicName>
differs from
<taxonomicName id="8CE11CEEFFF7634BEA14C24CFAD00551" box="[1103,1305,1052,1076]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="niraikanaiensis">
<emphasis id="7995BB7FFFF7634BEA14C24CFAD00551" box="[1103,1305,1052,1076]" italics="true" pageId="9" pageNumber="390">S. niraikanaiensis</emphasis>
</taxonomicName>
also by the lack of conspicuous markings on the first dorsal and pectoral fins (vs. having black spots), and in the mode of the pectoral-fin ray count (15 vs. 16).
</paragraph>
<paragraph id="4B5E676DFFF7634BEE9CC2D0FB14074D" blockId="9.[151,1437,151,1943]" pageId="9" pageNumber="390">
<taxonomicName id="8CE11CEEFFF7634BEE9CC2D0FE1E05FD" box="[199,471,1152,1176]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="palawanensis">
<emphasis id="7995BB7FFFF7634BEE9CC2D0FE1E05FD" box="[199,471,1152,1176]" italics="true" pageId="9" pageNumber="390">Stiphodon palawanensis</emphasis>
</taxonomicName>
differs from
<taxonomicName id="8CE11CEEFFF7634BEC35C2D0FBDC05FD" authority="Maeda, Mukai &amp; Tachihara" authorityName="Maeda, Mukai &amp; Tachihara" box="[622,1045,1151,1176]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="alcedo">
<emphasis id="7995BB7FFFF7634BEC35C2D0FD1F05FD" box="[622,726,1152,1176]" italics="true" pageId="9" pageNumber="390">S. alcedo</emphasis>
Maeda, Mukai &amp; Tachihara
</taxonomicName>
by having a line of black blotches (in males) or a black band (in females) on the distal part of the second dorsal fin (vs. no such markings), having more premaxillary and horizontal teeth (
<figureCitation id="D3DA7BE8FFF7634BEC33C292FD6705BE" box="[616,686,1218,1243]" captionStart="FIGURE 2" captionStartId="4.[151,250,1696,1718]" captionTargetBox="[349,1239,204,1666]" captionTargetId="figure@4.[340,1247,193,1675]" captionTargetPageId="4" captionText="FIGURE 2. Number of premaxillary and horizontal dentary teeth of Stiphodon palawanensis (red circles), S. alcedo (pink circles), S. atratus (black diamonds), S. imperiorientis (light blue triangles), S. maculidorsalis (light green triangles), S. multisquamus (deep blue circles), S. ornatus (orange triangles), S. pelewensis (gray diamonds), S. pulchellus (deep green triangles), and S. weberi (purple diamonds)." httpUri="https://zenodo.org/record/232901/files/figure.png" pageId="9" pageNumber="390">Fig. 2</figureCitation>
), and the mode of the pectoral-fin ray count (15 vs. 16); from
<taxonomicName id="8CE11CEEFFF7634BEBDCC293FEB10599" authority="Watson" authorityName="Watson" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="martenstyni">
<emphasis id="7995BB7FFFF7634BEBDCC293FED20599" italics="true" pageId="9" pageNumber="390">S. martenstyni</emphasis>
Watson
</taxonomicName>
(of which only the male
<typeStatus id="945AD9CFFFF7634BECC1C2B4FD350598" box="[666,764,1252,1277]" pageId="9" pageNumber="390" type="holotype">holotype</typeStatus>
is known) by lack of clear markings on the first dorsal fin (vs. having two to five dusky spots along each spine) and having a line of black blotches on the second dorsal fin (vs. lack of it); from
<taxonomicName id="8CE11CEEFFF7634BEF25C377FD8F045B" authority="Watson" authorityName="Watson" box="[382,582,1319,1342]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="atratus">
<emphasis id="7995BB7FFFF7634BEF25C377FE20045B" box="[382,489,1319,1342]" italics="true" pageId="9" pageNumber="390">S. atratus</emphasis>
Watson
</taxonomicName>
,
<taxonomicName id="8CE11CEEFFF7634BEC0AC377FC0F045A" authority="Watson &amp; Chen" authorityName="Watson &amp; Chen" box="[593,966,1318,1343]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="imperiorientis">
<emphasis id="7995BB7FFFF7634BEC0AC377FCC2045B" box="[593,779,1319,1342]" italics="true" pageId="9" pageNumber="390">S. imperiorientis</emphasis>
Watson &amp; Chen
</taxonomicName>
,
<taxonomicName id="8CE11CEEFFF7634BED8AC377FB66045A" authority="Meinken" authorityName="Meinken" box="[977,1199,1318,1343]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="ornatus">
<emphasis id="7995BB7FFFF7634BED8AC377FB8B045B" box="[977,1090,1319,1342]" italics="true" pageId="9" pageNumber="390">S. ornatus</emphasis>
Meinken
</taxonomicName>
,
<taxonomicName id="8CE11CEEFFF7634BEAE1C377FA51045B" authority="Herre" authorityName="Herre" box="[1210,1432,1318,1342]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="pelewensis">
<emphasis id="7995BB7FFFF7634BEAE1C377FA87045B" box="[1210,1358,1318,1342]" italics="true" pageId="9" pageNumber="390">S. pelewensis</emphasis>
Herre
</taxonomicName>
,
<taxonomicName id="8CE11CEEFFF7634BEECCC318FEEE0405" box="[151,295,1352,1376]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="pulchellus">
<emphasis id="7995BB7FFFF7634BEECCC318FEEE0405" box="[151,295,1352,1376]" italics="true" pageId="9" pageNumber="390">S. pulchellus</emphasis>
</taxonomicName>
, and
<taxonomicName id="8CE11CEEFFF7634BEF3CC318FD300405" authority="Watson, Allen &amp; Kottelat" authorityName="Watson, Allen &amp; Kottelat" box="[359,761,1351,1376]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="weberi">
<emphasis id="7995BB7FFFF7634BEF3CC318FE190405" box="[359,464,1352,1376]" italics="true" pageId="9" pageNumber="390">S. weberi</emphasis>
Watson, Allen &amp; Kottelat
</taxonomicName>
by lack of distinct markings on the pectoral fin (vs. having many black spots along rays), having a line of black blotches (in males) or a black band (in females) on the second dorsal fin (vs. lack of it), and having more premaxillary and horizontal teeth (
<figureCitation id="D3DA7BE8FFF7634BEDACC3DAFBF504C6" box="[1015,1084,1418,1443]" captionStart="FIGURE 2" captionStartId="4.[151,250,1696,1718]" captionTargetBox="[349,1239,204,1666]" captionTargetId="figure@4.[340,1247,193,1675]" captionTargetPageId="4" captionText="FIGURE 2. Number of premaxillary and horizontal dentary teeth of Stiphodon palawanensis (red circles), S. alcedo (pink circles), S. atratus (black diamonds), S. imperiorientis (light blue triangles), S. maculidorsalis (light green triangles), S. multisquamus (deep blue circles), S. ornatus (orange triangles), S. pelewensis (gray diamonds), S. pulchellus (deep green triangles), and S. weberi (purple diamonds)." httpUri="https://zenodo.org/record/232901/files/figure.png" pageId="9" pageNumber="390">Fig. 2</figureCitation>
); and from
<taxonomicName id="8CE11CEEFFF7634BEAE5C3DBFAB004C7" box="[1214,1401,1419,1442]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="atropurpureus">
<emphasis id="7995BB7FFFF7634BEAE5C3DBFAB004C7" box="[1214,1401,1419,1442]" italics="true" pageId="9" pageNumber="390">S. atropurpureus</emphasis>
</taxonomicName>
,
<taxonomicName id="8CE11CEEFFF7634BEBDCC3DBFEF204A1" authority="Watson" authorityName="Watson" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="carisa">
<emphasis id="7995BB7FFFF7634BEBDCC3DBFF1404A1" italics="true" pageId="9" pageNumber="390">S. carisa</emphasis>
Watson
</taxonomicName>
,
<taxonomicName id="8CE11CEEFFF7634BEF13C3FCFD3804A1" authority="Keith, Marquet &amp; Watson" authorityName="Keith, Marquet &amp; Watson" box="[328,753,1452,1477]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="kalfatak">
<emphasis id="7995BB7FFFF7634BEF13C3FCFE7604A1" box="[328,447,1452,1476]" italics="true" pageId="9" pageNumber="390">S. kalfatak</emphasis>
Keith, Marquet &amp; Watson
</taxonomicName>
,
<taxonomicName id="8CE11CEEFFF7634BECA6C3FCFC0804A1" authority="Watson" authorityName="Watson" box="[765,961,1452,1476]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="larson">
<emphasis id="7995BB7FFFF7634BECA6C3FCFCAA04A1" box="[765,867,1452,1476]" italics="true" pageId="9" pageNumber="390">S. larson</emphasis>
Watson
</taxonomicName>
, and
<taxonomicName id="8CE11CEEFFF7634BEDA5C3FCFB7504A0" authority="Weber" authorityName="Weber" box="[1022,1212,1452,1477]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="semoni">
<emphasis id="7995BB7FFFF7634BEDA5C3FCFBA204A1" box="[1022,1131,1452,1476]" italics="true" pageId="9" pageNumber="390">S. semoni</emphasis>
Weber
</taxonomicName>
by having a line of black blotches (in males) or a black band (in females) on the second dorsal fin (vs. lack of it), a pointed first dorsal fin in males (vs. rounded, except
<taxonomicName id="8CE11CEEFFF7634BEC4CC3BFFDB30763" box="[535,634,1519,1542]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="carisa">
<emphasis id="7995BB7FFFF7634BEC4CC3BFFDB30763" box="[535,634,1519,1542]" italics="true" pageId="9" pageNumber="390">S. carisa</emphasis>
</taxonomicName>
males having a pointed fin), and the predorsal scalation of males (the posterior part of the occipital region is scaled,
<figureCitation id="D3DA7BE8FFF7634BECC4C05FFD2B074D" box="[671,738,1551,1576]" captionStart="FIGURE 3" captionStartId="5.[151,250,1194,1216]" captionTargetBox="[349,1236,184,1162]" captionTargetId="figure@5.[340,1247,173,1173]" captionTargetPageId="5" captionText="FIGURE 3. Dorsal scalation on head and nape in Stiphodon palawanensis. a, male (43.1 mm SL, CMK 11966); b, female (41.9 mm SL, CMK 11966)." httpUri="https://zenodo.org/record/232902/files/figure.png" pageId="9" pageNumber="390">Fig. 3</figureCitation>
, vs. an occipital region that is almost naked).
</paragraph>
<paragraph id="4B5E676DFFF7634BEE9CC061FB4107E9" blockId="9.[151,1437,151,1943]" pageId="9" pageNumber="390">
Briefly, the new species is most easily distinguished by its unique fin markings: any
<taxonomicName id="8CE11CEEFFF7634BEAD0C061FB35072C" box="[1163,1276,1585,1609]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="genus">
<emphasis id="7995BB7FFFF7634BEAD0C061FB35072C" box="[1163,1276,1585,1609]" italics="true" pageId="9" pageNumber="390">Stiphodon</emphasis>
</taxonomicName>
with a line of black blotches (in males) or a black band (in females) on the distal part of the second dorsal fin, and without distinct markings on the first dorsal and pectoral fins can be identified as
<taxonomicName id="8CE11CEEFFF7634BED97C024FB4907E9" box="[972,1152,1652,1676]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="palawanensis">
<emphasis id="7995BB7FFFF7634BED97C024FB4907E9" box="[972,1152,1652,1676]" italics="true" pageId="9" pageNumber="390">S. palawanensis</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="4B5E676DFFF7634BEE9CC0C5FC5706F2" blockId="9.[151,1437,151,1943]" pageId="9" pageNumber="390">
<emphasis id="7995BB7FFFF7634BEE9CC0C5FD0207CB" bold="true" box="[199,715,1684,1710]" pageId="9" pageNumber="390">
Other
<taxonomicName id="8CE11CEEFFF7634BEF4CC0C4FE4207C8" box="[279,395,1684,1709]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="genus">
<emphasis id="7995BB7FFFF7634BEF4CC0C4FE4207C8" bold="true" box="[279,395,1684,1709]" italics="true" pageId="9" pageNumber="390">Stiphodon</emphasis>
</taxonomicName>
species found in Palawan.
</emphasis>
During our two-day exploration in the
<typeStatus id="945AD9CFFFF7634BEAD5C0C6FB7707CB" box="[1166,1214,1686,1710]" pageId="9" pageNumber="390">type</typeStatus>
locality of the new species (Balsahan Stream in the Iwahig Prison and Penal Farm, Sulu Sea side of Puerto Princesa City) in
<date id="3F5F41ADFFF7634BEB32C0E7FF1B0795" pageId="9" pageNumber="390" value="2015-05">May 2015</date>
, two other
<taxonomicName id="8CE11CEEFFF7634BEF38C088FE1D0795" box="[355,468,1752,1776]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="genus">
<emphasis id="7995BB7FFFF7634BEF38C088FE1D0795" box="[355,468,1752,1776]" italics="true" pageId="9" pageNumber="390">Stiphodon</emphasis>
</taxonomicName>
species were found; three female individuals of
<taxonomicName id="8CE11CEEFFF7634BEA72C088FB250795" box="[1065,1260,1752,1776]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="atropurpureus">
<emphasis id="7995BB7FFFF7634BEA72C088FB250795" box="[1065,1260,1752,1776]" italics="true" pageId="9" pageNumber="390">S. atropurpureus</emphasis>
</taxonomicName>
and one male individual of
<taxonomicName id="8CE11CEEFFF7634BEF69C0AAFD280677" authority="Watson &amp; Chen" authorityName="Watson &amp; Chen" box="[306,737,1785,1810]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="percnopterygionus">
<emphasis id="7995BB7FFFF7634BEF69C0AAFDE80674" box="[306,545,1786,1809]" italics="true" pageId="9" pageNumber="390">S. percnopterygionus</emphasis>
Watson &amp; Chen
</taxonomicName>
(
<figureCitation id="D3DA7BE8FFF7634BECA8C0A9FCF00677" box="[755,825,1785,1810]" captionStart="FIGURE 6" captionStartId="8.[151,250,1866,1888]" captionTargetBox="[345,1240,917,1839]" captionTargetId="figure@8.[340,1248,913,1845]" captionTargetPageId="8" captionText="FIGURE 6. Location of Palawan in the Philippines (inset map) and sites where Stiphodon species have been found on Palawan. Red circles, S. palawanensis; green circles, S. pulchellus; blue circle, S. atropurpureus; and orange circle, S. percnopterygionus." httpUri="https://zenodo.org/record/232905/files/figure.png" pageId="9" pageNumber="390">Fig. 6</figureCitation>
). Two of the three
<taxonomicName id="8CE11CEEFFF7634BEA42C0AAFB1E0674" box="[1049,1239,1786,1809]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="atropurpureus">
<emphasis id="7995BB7FFFF7634BEA42C0AAFB1E0674" box="[1049,1239,1786,1809]" italics="true" pageId="9" pageNumber="390">S. atropurpureus</emphasis>
</taxonomicName>
individuals were collected (
<figureCitation id="D3DA7BE8FFF7634BEF50C14AFE840656" box="[267,333,1818,1843]" captionStart="FIGURE 7" captionStartId="10.[151,250,1241,1263]" captionTargetBox="[161,1420,201,1188]" captionTargetId="figure@10.[151,1436,193,1220]" captionTargetPageId="10" captionText="FIGURE 7. Stiphodon pulchellus (a, WPU-PPC-P 29, male, 45.9 mm SL; b, URM-P 48681, female, 39.6 mm SL) and Stiphodon atropurpureus (c, WPU-PPC-P 10, female, 43.7 mm SL) specimens collected from Palawan immediately after fixation." httpUri="https://zenodo.org/record/232906/files/figure.png" pageId="9" pageNumber="390">Fig. 7</figureCitation>
c) and listed as comparative material in this paper.
<taxonomicName id="8CE11CEEFFF7634BEDDFC14BFB050657" box="[900,1228,1819,1843]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="percnopterygionus">
<emphasis id="7995BB7FFFF7634BEDDFC14BFB050657" box="[900,1228,1819,1843]" italics="true" pageId="9" pageNumber="390">Stiphodon percnopterygionus</emphasis>
</taxonomicName>
was not collected, however a wild individual was identified on site by snorkel based observation of characteristic markings of the body and fins (
<figureCitation id="D3DA7BE8FFF7634BEF1EC10DFE430613" box="[325,394,1885,1910]" captionStart="FIGURE 8" captionStartId="11.[151,250,1139,1161]" captionTargetBox="[151,1436,179,1117]" captionTargetId="figure@11.[151,1436,153,1117]" captionTargetPageId="11" captionText="FIGURE 8. Stiphodon percnopterygionus (a) with Stiphodon palawanensis observed in Balsahan Stream, Puerto Princesa City, Palawan, on 18 May 2015." httpUri="https://zenodo.org/record/232907/files/figure.png" pageId="9" pageNumber="390">Fig. 8</figureCitation>
) and its unique, high triangular first dorsal fin. While
<taxonomicName id="8CE11CEEFFF7634BEDA4C10EFB7F0610" box="[1023,1206,1885,1909]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="palawanensis">
<emphasis id="7995BB7FFFF7634BEDA4C10EFB7F0610" box="[1023,1206,1885,1909]" italics="true" pageId="9" pageNumber="390">S. palawanensis</emphasis>
</taxonomicName>
was very abundant, both
<taxonomicName id="8CE11CEEFFF7634BEE94C12FFE4206F3" box="[207,395,1919,1942]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="atropurpureus">
<emphasis id="7995BB7FFFF7634BEE94C12FFE4206F3" box="[207,395,1919,1942]" italics="true" pageId="9" pageNumber="390">S. atropurpureus</emphasis>
</taxonomicName>
and
<taxonomicName id="8CE11CEEFFF7634BEF99C12FFD6606F3" box="[450,687,1919,1942]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="390" phylum="Chordata" rank="species" species="percnopterygionus">
<emphasis id="7995BB7FFFF7634BEF99C12FFD6606F3" box="[450,687,1919,1942]" italics="true" pageId="9" pageNumber="390">S. percnopterygionus</emphasis>
</taxonomicName>
were rare at this site.
</paragraph>
<caption id="1F9E37E5FFF46348EECCC289FF240449" httpUri="https://zenodo.org/record/232906/files/figure.png" pageId="10" pageNumber="391" targetBox="[161,1420,201,1188]" targetPageId="10">
<paragraph id="4B5E676DFFF46348EECCC289FF240449" blockId="10.[151,1435,1241,1324]" pageId="10" pageNumber="391">
<emphasis id="7995BB7FFFF46348EECCC289FEDE058A" bold="true" box="[151,279,1241,1263]" pageId="10" pageNumber="391">FIGURE 7.</emphasis>
<taxonomicName id="8CE11CEEFFF46348EF79C289FE33058A" box="[290,506,1241,1263]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="10" pageNumber="391" phylum="Chordata" rank="species" species="pulchellus">
<emphasis id="7995BB7FFFF46348EF79C289FE33058A" box="[290,506,1241,1263]" italics="true" pageId="10" pageNumber="391">Stiphodon pulchellus</emphasis>
</taxonomicName>
(a, WPU-PPC-P 29, male, 45.9 mm SL; b, URM-P 48681, female, 39.6 mm SL) and
<taxonomicName id="8CE11CEEFFF46348EECCC2A8FE51046B" box="[151,408,1272,1294]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="10" pageNumber="391" phylum="Chordata" rank="species" species="atropurpureus">
<emphasis id="7995BB7FFFF46348EECCC2A8FE51046B" box="[151,408,1272,1294]" italics="true" pageId="10" pageNumber="391">Stiphodon atropurpureus</emphasis>
</taxonomicName>
(c, WPU-PPC-P 10, female, 43.7 mm SL) specimens collected from Palawan immediately after fixation.
</paragraph>
</caption>
<paragraph id="4B5E676DFFF46348EE9CC30AFED80767" blockId="10.[151,1437,1369,1934]" pageId="10" pageNumber="391">
Four
<taxonomicName id="8CE11CEEFFF46348EF59C309FEBA0414" box="[258,371,1369,1393]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="10" pageNumber="391" phylum="Chordata" rank="genus">
<emphasis id="7995BB7FFFF46348EF59C309FEBA0414" box="[258,371,1369,1393]" italics="true" pageId="10" pageNumber="391">Stiphodon</emphasis>
</taxonomicName>
specimens collected in the Iwahig River at Puerto Princesa City in 1988 have been deposited in the National Museum of Nature and Science (Ibaraki,
<collectingCountry id="33F627FDFFF46348ED4BC32EFC9C04F3" box="[784,853,1406,1430]" name="Japan" pageId="10" pageNumber="391">Japan</collectingCountry>
). The Iwahig is a relatively large river next to the Balsahan Stream. The specimens are composed of three
<taxonomicName id="8CE11CEEFFF46348ED49C3F2FC0E04DC" box="[786,967,1441,1465]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="10" pageNumber="391" phylum="Chordata" rank="species" species="palawanensis">
<emphasis id="7995BB7FFFF46348ED49C3F2FC0E04DC" box="[786,967,1441,1465]" italics="true" pageId="10" pageNumber="391">S. palawanensis</emphasis>
</taxonomicName>
(NSMT-P 45091, 45092, and 45094) and one
<taxonomicName id="8CE11CEEFFF46348EE9CC396FE9104B8" box="[199,344,1477,1501]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="10" pageNumber="391" phylum="Chordata" rank="species" species="pulchellus">
<emphasis id="7995BB7FFFF46348EE9CC396FE9104B8" box="[199,344,1477,1501]" italics="true" pageId="10" pageNumber="391">S. pulchellus</emphasis>
</taxonomicName>
(NSMT-P 45093), but the latter species was not found in our exploration in Balsahan Stream in
<date id="3F5F41ADFFF46348EECCC3BAFEC40767" box="[151,269,1513,1538]" pageId="10" pageNumber="391" value="2015-05">May 2015</date>
.
</paragraph>
<paragraph id="4B5E676DFFF46348EE9CC05EFCB0070B" blockId="10.[151,1437,1369,1934]" pageId="10" pageNumber="391">
In Barake Stream (Sulu Sea side of Aborlan), the only
<taxonomicName id="8CE11CEEFFF46348ED72C05DFC530740" box="[809,922,1549,1573]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="10" pageNumber="391" phylum="Chordata" rank="genus">
<emphasis id="7995BB7FFFF46348ED72C05DFC530740" box="[809,922,1549,1573]" italics="true" pageId="10" pageNumber="391">Stiphodon</emphasis>
</taxonomicName>
species found during our one-day exploration was
<taxonomicName id="8CE11CEEFFF46348EE97C062FE4B072C" box="[204,386,1585,1609]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="10" pageNumber="391" phylum="Chordata" rank="species" species="palawanensis">
<emphasis id="7995BB7FFFF46348EE97C062FE4B072C" box="[204,386,1585,1609]" italics="true" pageId="10" pageNumber="391">S. palawanensis</emphasis>
</taxonomicName>
. All collections from Aborlan and Narra, examined in the present study (CMK and URM), also comprised entirely of individuals of
<taxonomicName id="8CE11CEEFFF46348EC38C006FCD10708" box="[611,792,1621,1645]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="10" pageNumber="391" phylum="Chordata" rank="species" species="palawanensis">
<emphasis id="7995BB7FFFF46348EC38C006FCD10708" box="[611,792,1621,1645]" italics="true" pageId="10" pageNumber="391">S. palawanensis</emphasis>
</taxonomicName>
(
<figureCitation id="D3DA7BE8FFF46348ED7DC005FCA5070B" box="[806,876,1621,1646]" captionStart="FIGURE 6" captionStartId="8.[151,250,1866,1888]" captionTargetBox="[345,1240,917,1839]" captionTargetId="figure@8.[340,1248,913,1845]" captionTargetPageId="8" captionText="FIGURE 6. Location of Palawan in the Philippines (inset map) and sites where Stiphodon species have been found on Palawan. Red circles, S. palawanensis; green circles, S. pulchellus; blue circle, S. atropurpureus; and orange circle, S. percnopterygionus." httpUri="https://zenodo.org/record/232905/files/figure.png" pageId="10" pageNumber="391">Fig. 6</figureCitation>
).
</paragraph>
<paragraph id="4B5E676DFFF46348EE9CC02AFF1F07BF" blockId="10.[151,1437,1369,1934]" pageId="10" pageNumber="391">
We also explored another river bearing the name Iwahig at Quezon, on the South
<collectingCountry id="33F627FDFFF46348EA3AC029FB6D07F7" box="[1121,1188,1657,1682]" name="China" pageId="10" pageNumber="391">China</collectingCountry>
Sea side of the island (
<figureCitation id="D3DA7BE8FFF46348EEC4C0CDFF2C07D3" box="[159,229,1693,1718]" captionStart="FIGURE 6" captionStartId="8.[151,250,1866,1888]" captionTargetBox="[345,1240,917,1839]" captionTargetId="figure@8.[340,1248,913,1845]" captionTargetPageId="8" captionText="FIGURE 6. Location of Palawan in the Philippines (inset map) and sites where Stiphodon species have been found on Palawan. Red circles, S. palawanensis; green circles, S. pulchellus; blue circle, S. atropurpureus; and orange circle, S. percnopterygionus." httpUri="https://zenodo.org/record/232905/files/figure.png" pageId="10" pageNumber="391">Fig. 6</figureCitation>
) in
<date id="3F5F41ADFFF46348EF43C0CEFE5807D3" box="[280,401,1693,1718]" pageId="10" pageNumber="391" value="2015-05">May 2015</date>
.
<taxonomicName id="8CE11CEEFFF46348EFC5C0CDFD4507D0" box="[414,652,1693,1717]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="10" pageNumber="391" phylum="Chordata" rank="species" species="pulchellus">
<emphasis id="7995BB7FFFF46348EFC5C0CDFD4507D0" box="[414,652,1693,1717]" italics="true" pageId="10" pageNumber="391">Stiphodon pulchellus</emphasis>
</taxonomicName>
(
<figureCitation id="D3DA7BE8FFF46348ECC4C0CDFD2C07D0" box="[671,741,1693,1718]" captionStart="FIGURE 7" captionStartId="10.[151,250,1241,1263]" captionTargetBox="[161,1420,201,1188]" captionTargetId="figure@10.[151,1436,193,1220]" captionTargetPageId="10" captionText="FIGURE 7. Stiphodon pulchellus (a, WPU-PPC-P 29, male, 45.9 mm SL; b, URM-P 48681, female, 39.6 mm SL) and Stiphodon atropurpureus (c, WPU-PPC-P 10, female, 43.7 mm SL) specimens collected from Palawan immediately after fixation." httpUri="https://zenodo.org/record/232906/files/figure.png" pageId="10" pageNumber="391">Fig. 7</figureCitation>
a, b) was abundant, but
<taxonomicName id="8CE11CEEFFF46348EDA5C0CEFB7C07D0" box="[1022,1205,1693,1717]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="10" pageNumber="391" phylum="Chordata" rank="species" species="palawanensis">
<emphasis id="7995BB7FFFF46348EDA5C0CEFB7C07D0" box="[1022,1205,1693,1717]" italics="true" pageId="10" pageNumber="391">S. palawanensis</emphasis>
</taxonomicName>
could not be found there.
</paragraph>
<paragraph id="4B5E676DFFF46348EE9CC0B5FEB106EB" blockId="10.[151,1437,1369,1934]" pageId="10" pageNumber="391">
Thus, the species composition of any
<taxonomicName id="8CE11CEEFFF46348EC26C0B5FD270798" box="[637,750,1765,1789]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="10" pageNumber="391" phylum="Chordata" rank="genus">
<emphasis id="7995BB7FFFF46348EC26C0B5FD270798" box="[637,750,1765,1789]" italics="true" pageId="10" pageNumber="391">Stiphodon</emphasis>
</taxonomicName>
assemblage seems to depend on the location on the island.
<taxonomicName id="8CE11CEEFFF46348EECCC159FE630644" box="[151,426,1801,1825]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="10" pageNumber="391" phylum="Chordata" rank="species" species="palawanensis">
<emphasis id="7995BB7FFFF46348EECCC159FE630644" box="[151,426,1801,1825]" italics="true" pageId="10" pageNumber="391">Stiphodon palawanensis</emphasis>
</taxonomicName>
may be the dominant species in some areas, whereas
<taxonomicName id="8CE11CEEFFF46348EA40C15AFB650644" box="[1051,1196,1801,1825]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="10" pageNumber="391" phylum="Chordata" rank="species" species="pulchellus">
<emphasis id="7995BB7FFFF46348EA40C15AFB650644" box="[1051,1196,1801,1825]" italics="true" pageId="10" pageNumber="391">S. pulchellus</emphasis>
</taxonomicName>
may be dominant in others. However, because only a small survey effort has been applied at each site and only a small part of the island has been explored so far, comprehensive surveys will be required to understand the distribution of
<taxonomicName id="8CE11CEEFFF46348EB70C101FA55060C" box="[1323,1436,1873,1897]" class="Actinopterygii" family="Gobiidae" genus="Stiphodon" kingdom="Animalia" order="Perciformes" pageId="10" pageNumber="391" phylum="Chordata" rank="genus">
<emphasis id="7995BB7FFFF46348EB70C101FA55060C" box="[1323,1436,1873,1897]" italics="true" pageId="10" pageNumber="391">Stiphodon</emphasis>
</taxonomicName>
species on Palawan.
</paragraph>
</subSubSection>
</treatment>
</document>