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<document id="73C55955B1AA466218EFACBC2A3FC05F" ID-DOI="10.1206/3894.1" ID-ISSN="0003-0082" ID-Zenodo-Dep="5369712" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1630263992186" checkinUser="carolina" docAuthor="Emmons, Louise H. &amp; Fabre, Pierre-henri" docDate="2018" docId="039C2E2FFFCAD560FE4DFDBEFC16FA6C" docLanguage="en" docName="N3894.pdf" docOrigin="American Museum Novitates 2018 (3894)" docSource="http://www.bioone.org/doi/10.1206/3894.1" docStyle="DocumentStyle:26C3580A17D4BBE58BD8D8AC6AE09AC4.6:AmMusNovit.2010-.journal_article" docStyleId="26C3580A17D4BBE58BD8D8AC6AE09AC4" docStyleName="AmMusNovit.2010-.journal_article" docStyleVersion="6" docTitle="Pattonomys Emmons 2005" docType="treatment" docVersion="4" lastPageNumber="29" masterDocId="FFA55657FFD0D57CFFB9FFE4FFE5FFCB" masterDocTitle="A Review of the Pattonomys / Toromys Clade (Rodentia: Echimyidae), with Descriptions of a New Toromys Species and a New Genus" masterLastPageNumber="52" masterPageNumber="1" pageNumber="27" updateTime="1699203564564" updateUser="plazi" zenodo-license-document="CC-BY-4.0">
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<mods:title id="2377605CC02238237F080B6EEC7244F0">A Review of the Pattonomys / Toromys Clade (Rodentia: Echimyidae), with Descriptions of a New Toromys Species and a New Genus</mods:title>
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<mods:namePart id="822E48C891FCCC8CD85D6B02EB66FC49">Emmons, Louise H.</mods:namePart>
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<mods:namePart id="3F8873765DDCD651B0CC3DAEE56082CE">Fabre, Pierre-henri</mods:namePart>
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<mods:title id="A292951534CF25F0C304BE5915A77B24">American Museum Novitates</mods:title>
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<subSubSection id="C32FCCB2FFCAD566FE4DFDBEFCD6FDBF" box="[500,819,601,628]" pageId="26" pageNumber="27" type="nomenclature">
<paragraph id="8B8A9F39FFCAD566FE4DFDBEFCD6FDBF" blockId="26.[500,819,601,628]" box="[500,819,601,628]" pageId="26" pageNumber="27">
<heading id="D0C22855FFCAD566FE4DFDBEFCD6FDBF" box="[500,819,601,628]" centered="true" fontSize="11" level="2" pageId="26" pageNumber="27" reason="5">
<taxonomicName id="4C35E4BAFFCAD566FE4DFDBEFCD6FDBF" authority="Emmons, 2005" authorityName="Emmons" authorityYear="2005" box="[500,819,601,628]" class="Mammalia" family="Echimyidae" genus="Pattonomys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rodentia" pageId="26" pageNumber="27" phylum="Chordata" rank="genus">
<emphasis id="B941432BFFCAD566FE4DFDBEFD9EFDB8" box="[500,635,602,627]" italics="true" pageId="26" pageNumber="27">Pattonomys</emphasis>
<bibRefCitation id="EFA4E2C8FFCAD566FD3AFDBEFCD6FDBF" author="Emmons, L. H." box="[643,819,601,628]" pageId="26" pageNumber="27" pagination="247 - 309" refId="ref23229" refString="Emmons, L. H. 2005. A revision of the arboreal Echimyidae (Rodentia: Echimyidae, Echimyinae); with descriptions of two new genera. In E. A. Lacey, and P. Myers (editors), Mammalian diversification: from chromosomes to phylogeography: 247 - 309. Berkeley: University of California Press." type="book chapter" year="2005">Emmons, 2005</bibRefCitation>
</taxonomicName>
</heading>
</paragraph>
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<subSubSection id="C32FCCB2FFCAD566FF25FD76FD13FD1A" pageId="26" pageNumber="27" type="materials_examined">
<paragraph id="8B8A9F39FFCAD566FF25FD76FC75FD60" blockId="26.[108,1213,657,1767]" box="[156,912,657,684]" pageId="26" pageNumber="27">
<typeStatus id="548E219BFFCAD566FF25FD76FF32FD67" box="[156,215,658,684]" pageId="26" pageNumber="27">TYPE</typeStatus>
SPECIES:
<taxonomicName id="4C35E4BAFFCAD566FEF1FD75FD02FD60" authority="I. Geoffroy St." authorityName="I. Geoffroy St." box="[328,743,657,684]" class="Mammalia" family="Echimyidae" genus="Nelomys" kingdom="Animalia" order="Rodentia" pageId="26" pageNumber="27" phylum="Chordata" rank="species" species="semivillosus">
<emphasis id="B941432BFFCAD566FEF1FD75FDDCFD60" box="[328,569,657,683]" italics="true" pageId="26" pageNumber="27">Nelomys semivillosus</emphasis>
I. Geoffroy St.
</taxonomicName>
-Hilaire, 1838.
</paragraph>
<paragraph id="8B8A9F39FFCAD566FF25FD53FD13FD1A" blockId="26.[108,1213,657,1767]" box="[156,758,695,721]" pageId="26" pageNumber="27">CONTENTS: Four valid species, as diagnosed below.</paragraph>
</subSubSection>
<subSubSection id="C32FCCB2FFCAD566FF25FD38FC75FB7D" pageId="26" pageNumber="27" type="diagnosis">
<paragraph id="8B8A9F39FFCAD566FF25FD38FC75FB7D" blockId="26.[108,1213,657,1767]" pageId="26" pageNumber="27">
DIAGNOSIS: A genus of large, arboreally adapted
<taxonomicName id="4C35E4BAFFCAD566FD51FD38FC97FD3D" box="[744,882,732,758]" class="Mammalia" family="Echimyidae" kingdom="Animalia" order="Rodentia" pageId="26" pageNumber="27" phylum="Chordata" rank="family">Echimyidae</taxonomicName>
(
<tableCitation id="C6B7AA82FFCAD566FC3DFD38FC07FD3C" box="[900,994,732,759]" captionStart="TABLE 5" captionStartId="12.[108,182,224,248]" captionText="TABLE 5. External measurements (mm) of Leiuromys, Pattonomys, and Toromys.a" pageId="26" pageNumber="27">tables 5</tableCitation>
,
<tableCitation id="C6B7AA82FFCAD566FC48FD38FBE7FD3C" box="[1009,1026,732,759]" captionStart="TABLE 6" captionStartId="13.[109,175,238,259]" captionText="TABLE 6. Cranial measurements (mm) of Leiuromys, Pattonomys, and Toromys. Tabulated statistics include the sample mean and (in parentheses) the observed range and the sample size." pageId="26" pageNumber="27">6</tableCitation>
). Dorsal pelage with strong, wide, gray-based aristiform spines that narrow to sharp tips; some spines entirely black or dusky tipped, others with dark subterminal bands and contrasting white tips that impart a pronounced speckled appearance (
<figureCitation id="130E83BCFFCAD566FDD5FCA8FD59FCAD" box="[620,700,844,870]" captionStart="FIG" captionStartId="14.[108,149,1202,1223]" captionTargetBox="[180,1139,224,1174]" captionTargetId="figure-143@14.[180,1139,224,1174]" captionTargetPageId="14" captionText="FIG. 4. Pelage of dorsum at rump. A, Leiuromys occasius. FMNH 84529, high density of sharply pointed spines and fine, wavy red underhairs; B, Pattonomys semivillosus USNM 280204, dense, more tapered, white-tipped spines, abundant pale bristles, straight ochraceous underhairs; C, Makalata cf. didelphoides USNM 549593, agouti pelage, sparser narrowly buff-tipped spines, tapered hairlike spine tips; D, Toromys grandis AMNH 94036, thick bristles, no spines; E, T. rhipidurus AMNH 73273, fine-banded agouti bristles; F, T. sp. nov., narrow aristiforms and a few narrow spines with hairlike tips, abundant thin, agouti bristles. The resemblance between Toromys congeners is evident. Spine tips are more sharply tapered in A than in B." figureDoi="http://doi.org/10.5281/zenodo.5369718" httpUri="https://zenodo.org/record/5369718/files/figure.png" pageId="26" pageNumber="27">fig. 4B</figureCitation>
). Prominent pale postauricular patches (
<figureCitation id="130E83BCFFCAD566FB2FFCA8FF7BFC40" captionStart="FIG" captionStartId="25.[108,148,1514,1535]" captionTargetBox="[192,1128,224,1481]" captionTargetId="figure-144@25.[192,1128,224,1481]" captionTargetPageId="25" captionText="FIG. 14. Living Toromys and Pattonomys. A, Toromys grandis, lower Rio Purus, (fig. 1, locality 31). Note the elongate body, the longer black hair on the proximal tail, and the broad foot with narrow heel. There is a postauricular patch of pale skin. The anterior bright eyeshine spot seems to show an oval, vertical pupil, the other spot is a reflection of the flash (image by Vinicius Carvalho). B, Pattonomys semivillosus in a black mangrove tree (Avicennia germinans) in Parque Nacional Isla de Salamanca, Magdalena, Colombia. The white postauricular tufts above and behind the ear are striking (photograph by Fabrice Schmitt). C, Toromys rhipidurus, captured near Iquitos, Peru. The evident cream-colored postauricular patches are not visible on most museum skins. The camera-flash has accentuated the yellow midbody hues (image by Pamela Sánchez-Vendizú)." figureDoi="http://doi.org/10.5281/zenodo.5369738" httpUri="https://zenodo.org/record/5369738/files/figure.png" pageId="26" pageNumber="27">fig. 14B</figureCitation>
). Cranium generally like that of
<taxonomicName id="4C35E4BAFFCAD566FDAEFC95FD63FC40" authorityName="Emmons &amp; Fabre" authorityYear="2018" box="[535,646,881,907]" class="Mammalia" family="Echimyidae" genus="Makalata" kingdom="Animalia" order="Rodentia" pageId="26" pageNumber="27" phylum="Chordata" rank="genus">
<emphasis id="B941432BFFCAD566FDAEFC95FD63FC40" box="[535,646,881,907]" italics="true" pageId="26" pageNumber="27">Makalata</emphasis>
</taxonomicName>
and
<emphasis id="B941432BFFCAD566FD06FC95FCC1FC40" box="[703,804,881,907]" italics="true" pageId="26" pageNumber="27">
<taxonomicName id="4C35E4BAFFCAD566FD06FC95FCC5FC40" authorityName="G. Cuvier" authorityYear="1809" box="[703,800,881,907]" class="Mammalia" family="Echimyidae" genus="Echimys" kingdom="Animalia" order="Rodentia" pageId="26" pageNumber="27" phylum="Chordata" rank="genus">Echimys</taxonomicName>
,
</emphasis>
short and broad, with more or less expanded, winglike supraorbital shelves that curve upward from the frontal bones (
<figureCitation id="130E83BCFFCAD566FB80FC73FB5DFC7A" box="[1081,1208,919,945]" captionStart="FIG" captionStartId="17.[108,150,1500,1521]" captionTargetBox="[108,1212,224,1473]" captionTargetId="figure-7@17.[108,1212,224,1473]" captionTargetPageId="17" captionText="FIG. 7. Crania and mandibles, lateral view. A, Toromys grandis AMNH 93601; B, T. rhipidurus BMNH 28.7.21.89 (holotype); C, T. sp. nov., FMNH 55483 (holotype), D, Leiuromys occasius. AMNH 71897; E, Pattonomys punctatus BMNH 98.12.1.18 (holotype); F, P. flavidus USNM 63218 (holotype); G, P. carrikeri AMNH 130790; H, P. semivillosus USNM 280205. Black arrow in F indicates the deep mandible of P. flavidus relative to its congeners." figureDoi="http://doi.org/10.5281/zenodo.5369724" httpUri="https://zenodo.org/record/5369724/files/figure.png" pageId="26" pageNumber="27">figs. 7EH</figureCitation>
,
<figureCitation id="130E83BCFFCAD566FFD5FC58FF56FC1D" box="[108,179,956,982]" captionStart="FIG" captionStartId="19.[108,150,1560,1581]" captionTargetBox="[132,1188,224,1533]" captionTargetId="figure-6@19.[132,1188,224,1533]" captionTargetPageId="19" captionText="FIG. 9. Crania, dorsal view. A, Toromys grandis AMNH 93601; B, T. rhipidurus BMNH 28.7.21.89 (holotype); C, T. sp. nov., FMNH 55483 (holotype), D, Leiuromys occasius. AMNH 71897; E, Pattonomys punctatus BMNH 98.12.1.18 (holotype); F, P. flavidus USNM 63218 (holotype); G, P. carrikeri AMNH 130790; H, P. semivillosus USNM 280205. Black arrow in A indicates recurved parietal-interparietal suture, white arrow the expanded squamosal shelf." figureDoi="http://doi.org/10.5281/zenodo.5369728" httpUri="https://zenodo.org/record/5369728/files/figure.png" pageId="26" pageNumber="27">9EH</figureCitation>
,
<figureCitation id="130E83BCFFCAD566FF06FC58FEF1FC1D" box="[191,276,956,982]" captionStart="FIG" captionStartId="20.[108,149,1562,1583]" captionTargetBox="[132,1188,224,1534]" captionTargetId="figure-11@20.[132,1188,224,1534]" captionTargetPageId="20" captionText="FIG. 10. Crania, palatal view. A, Toromys grandis AMNH 93601; B, T. rhipidurus BMNH 28.7.21.89 (holotype); C, T. sp. nov., FMNH 55483 (holotype), D, Leiuromys occasius. AMNH 71897; E, Pattonomys punctatus BMNH 98.12.1.18 (holotype); F, P. flavidus USNM 63218 (holotype); G, P. carrikeri AMNH 130790; H, P. semivillosus USNM 280205." figureDoi="http://doi.org/10.5281/zenodo.5369730" httpUri="https://zenodo.org/record/5369730/files/figure.png" pageId="26" pageNumber="27">10EH</figureCitation>
; see also
<bibRefCitation id="EFA4E2C8FFCAD566FE3AFC59FDD2FC1C" author="Emmons, L. H." box="[387,567,956,983]" pageId="26" pageNumber="27" pagination="247 - 309" refId="ref23229" refString="Emmons, L. H. 2005. A revision of the arboreal Echimyidae (Rodentia: Echimyidae, Echimyinae); with descriptions of two new genera. In E. A. Lacey, and P. Myers (editors), Mammalian diversification: from chromosomes to phylogeography: 247 - 309. Berkeley: University of California Press." type="book chapter" year="2005">Emmons, 2005</bibRefCitation>
:
<figureCitation id="130E83BCFFCAD566FDFBFC58FD72FC1D" box="[578,663,956,982]" captionStart="FIG" captionStartId="21.[108,150,1406,1427]" captionTargetBox="[377,944,228,1379]" captionTargetId="figure-6@21.[377,944,227,1379]" captionTargetPageId="21" captionText="FIG. 11. Leiuromys occasius upper (top row) and lower (bottom row) occlusal patterns of cheek teeth at three states of toothwear. A, FMNH 84427; B, FMNH 84259; C, MCZ 37964, reversed. Note the trilophodont M3, the rounded posterior edges of the lower molars, the position of the mure of M12 on the lingual edges of the teeth, the quadralophodont lower premolar, and the M-shaped lower molars (viewed lingually)." figureDoi="http://doi.org/10.5281/zenodo.5369732" httpUri="https://zenodo.org/record/5369732/files/figure.png" pageId="26" pageNumber="27">figs. 11</figureCitation>
,
<figureCitation id="130E83BCFFCAD566FD1DFC58FD21FC1D" box="[676,708,956,982]" captionStart="FIG" captionStartId="25.[108,148,1514,1535]" captionTargetBox="[192,1128,224,1481]" captionTargetId="figure-144@25.[192,1128,224,1481]" captionTargetPageId="25" captionText="FIG. 14. Living Toromys and Pattonomys. A, Toromys grandis, lower Rio Purus, (fig. 1, locality 31). Note the elongate body, the longer black hair on the proximal tail, and the broad foot with narrow heel. There is a postauricular patch of pale skin. The anterior bright eyeshine spot seems to show an oval, vertical pupil, the other spot is a reflection of the flash (image by Vinicius Carvalho). B, Pattonomys semivillosus in a black mangrove tree (Avicennia germinans) in Parque Nacional Isla de Salamanca, Magdalena, Colombia. The white postauricular tufts above and behind the ear are striking (photograph by Fabrice Schmitt). C, Toromys rhipidurus, captured near Iquitos, Peru. The evident cream-colored postauricular patches are not visible on most museum skins. The camera-flash has accentuated the yellow midbody hues (image by Pamela Sánchez-Vendizú)." figureDoi="http://doi.org/10.5281/zenodo.5369738" httpUri="https://zenodo.org/record/5369738/files/figure.png" pageId="26" pageNumber="27">14</figureCitation>
). Upper cheekteeth with widely open flexi, greatly anteroposteriorly expanded protocone and posteroloph (probably metaloph and posteroloph combined), and a markedly oblique paraflexus on dP4 (
<figureCitation id="130E83BCFFCAD566FCDEFBE3FC0EFBEA" box="[871,1003,1031,1057]" captionStart="FIG" captionStartId="22.[108,150,1490,1511]" captionTargetBox="[115,1205,224,1460]" captionTargetId="figure-12@22.[115,1205,224,1460]" captionTargetPageId="22" captionText="FIG. 12. Occlusal patterns of upper (top row) and lower (bottom row) toothrows of Toromys and Pattonomys species, not to scale. A, Toromys sp. nov. FMNH 55483 holotype; B, T.sp. nov. LSUMZ 8904, an old individual; C, Toromys grandis AMNH 93604; D, Toromys rhipidurus AMNH 98667; E, Pattonomys carrikeri AMNH 31530; F, P. semivillosus USNM 280205; G, P. punctatus BMNH 9812118 (holotype). In Toromys maxillary teeth (AD) note laminar appearance of narrow, nearly parallel molar flexi, lophs of nearly uniform width, and mures lingually (left) of center of teeth. In Pattonomys (EG) note short, offset flexi, short, wide protoloph (arrow) especially pronounced on M2, mures about in midtooth; unequal loph sizes give irregular appearance. On mandibular teeth, note the straight (flat) posterior molar edges; the square (flat) lingual edge of central lophid (entoconid) in Toromys (AD, also compare with fig. 17)." figureDoi="http://doi.org/10.5281/zenodo.5369734" httpUri="https://zenodo.org/record/5369734/files/figure.png" pageId="26" pageNumber="27">fig. 12EG</figureCitation>
). These expanded lophs are accentuated with wear. The four lophs of dP4M2 are markedly unequal in length and width: the anteroloph is broad and squared lingually, tapering labially; and the posteroloph is pointed labially and strongly curved along the posterior margin of the tooth. Lower premolar tetralophodont, with two labial and two lingual flexids (
<figureCitation id="130E83BCFFCAD566FD46FB78FC66FB7D" box="[767,899,1180,1206]" captionStart="FIG" captionStartId="22.[108,150,1490,1511]" captionTargetBox="[115,1205,224,1460]" captionTargetId="figure-12@22.[115,1205,224,1460]" captionTargetPageId="22" captionText="FIG. 12. Occlusal patterns of upper (top row) and lower (bottom row) toothrows of Toromys and Pattonomys species, not to scale. A, Toromys sp. nov. FMNH 55483 holotype; B, T.sp. nov. LSUMZ 8904, an old individual; C, Toromys grandis AMNH 93604; D, Toromys rhipidurus AMNH 98667; E, Pattonomys carrikeri AMNH 31530; F, P. semivillosus USNM 280205; G, P. punctatus BMNH 9812118 (holotype). In Toromys maxillary teeth (AD) note laminar appearance of narrow, nearly parallel molar flexi, lophs of nearly uniform width, and mures lingually (left) of center of teeth. In Pattonomys (EG) note short, offset flexi, short, wide protoloph (arrow) especially pronounced on M2, mures about in midtooth; unequal loph sizes give irregular appearance. On mandibular teeth, note the straight (flat) posterior molar edges; the square (flat) lingual edge of central lophid (entoconid) in Toromys (AD, also compare with fig. 17)." figureDoi="http://doi.org/10.5281/zenodo.5369734" httpUri="https://zenodo.org/record/5369734/files/figure.png" pageId="26" pageNumber="27">fig. 12EG</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C32FCCB2FFCAD560FF25FB25FC16FA6C" lastPageId="28" lastPageNumber="29" pageId="26" pageNumber="27" type="description">
<paragraph id="8B8A9F39FFCAD560FF25FB25FD3CFC2D" blockId="26.[108,1213,657,1767]" lastBlockId="28.[108,1216,225,1447]" lastPageId="28" lastPageNumber="29" pageId="26" pageNumber="27">
DESCRIPTION: Spines wide, sharp, and stiff, with tips not drawn out into long hairlike processes but end in microscopic hairs (
<figureCitation id="130E83BCFFCAD566FDE4FB03FD57FACA" box="[605,690,1255,1281]" captionStart="FIG" captionStartId="14.[108,149,1202,1223]" captionTargetBox="[180,1139,224,1174]" captionTargetId="figure-143@14.[180,1139,224,1174]" captionTargetPageId="14" captionText="FIG. 4. Pelage of dorsum at rump. A, Leiuromys occasius. FMNH 84529, high density of sharply pointed spines and fine, wavy red underhairs; B, Pattonomys semivillosus USNM 280204, dense, more tapered, white-tipped spines, abundant pale bristles, straight ochraceous underhairs; C, Makalata cf. didelphoides USNM 549593, agouti pelage, sparser narrowly buff-tipped spines, tapered hairlike spine tips; D, Toromys grandis AMNH 94036, thick bristles, no spines; E, T. rhipidurus AMNH 73273, fine-banded agouti bristles; F, T. sp. nov., narrow aristiforms and a few narrow spines with hairlike tips, abundant thin, agouti bristles. The resemblance between Toromys congeners is evident. Spine tips are more sharply tapered in A than in B." figureDoi="http://doi.org/10.5281/zenodo.5369718" httpUri="https://zenodo.org/record/5369718/files/figure.png" pageId="26" pageNumber="27">fig. 4B</figureCitation>
). White-tipped spines are sprinkled singly among the lateral and dorsal pelage from shoulder or midback to rump, increasing posteriorly in number and in the length of pale tips. Pale-tipped spines sparsely and evenly distributed over the speckled rump. Dorsal spines mixed with fine, ochraceous yellow to orange setiform bristles and sparse dull reddish or yellow-ochraceous underhairs; pelage slate-gray based. Head, neck, and shoulders grizzled gray lined with black, in a mixture of long, narrow, dusky and whitish aristiforms (
<figureCitation id="130E83BCFFCAD566FD8AFA23FD7FFA2A" box="[563,666,1479,1505]" captionStart="FIG" captionStartId="25.[108,148,1514,1535]" captionTargetBox="[192,1128,224,1481]" captionTargetId="figure-144@25.[192,1128,224,1481]" captionTargetPageId="25" captionText="FIG. 14. Living Toromys and Pattonomys. A, Toromys grandis, lower Rio Purus, (fig. 1, locality 31). Note the elongate body, the longer black hair on the proximal tail, and the broad foot with narrow heel. There is a postauricular patch of pale skin. The anterior bright eyeshine spot seems to show an oval, vertical pupil, the other spot is a reflection of the flash (image by Vinicius Carvalho). B, Pattonomys semivillosus in a black mangrove tree (Avicennia germinans) in Parque Nacional Isla de Salamanca, Magdalena, Colombia. The white postauricular tufts above and behind the ear are striking (photograph by Fabrice Schmitt). C, Toromys rhipidurus, captured near Iquitos, Peru. The evident cream-colored postauricular patches are not visible on most museum skins. The camera-flash has accentuated the yellow midbody hues (image by Pamela Sánchez-Vendizú)." figureDoi="http://doi.org/10.5281/zenodo.5369738" httpUri="https://zenodo.org/record/5369738/files/figure.png" pageId="26" pageNumber="27">fig. 14B</figureCitation>
). Sides grayish from head to flank between gradually or sharply demarcated pale venter and yellowish or reddish-brown tinged middorsal pelage. Ear tips with an inconspicuous, sparse fringe of long fine, salient hairs. Dorsal pelage extends onto the tail base to about the hind feet of study skins; distal tail evenly covered above and below to its tip with fine, monocolored white to red-brown hairs that do not cover scales but curl outward, so that the tail looks distinctly hairy (
<figureCitation id="130E83BCFFCAD566FC58F965FBDCF957" box="[993,1081,1665,1692]" captionStart="FIG" captionStartId="15.[108,150,569,590]" captionTargetBox="[108,1212,224,540]" captionTargetId="figure-420@15.[108,1212,224,540]" captionTargetPageId="15" captionText="FIG. 5. Scales and hairs on dorsal tail base. A, Pattonomys carrikeri USNM 496483, irregular scales nearly square or slightly hexagonal, hairs unicolored, long, and curled outward; B, Leiuromys occasius FMNH 84259, short, wide hexagonal scales in spiral rows, short, adpressed, scalelike hairs in pairs; C, Toromys rhipidurus AMNH 276710, scales rounded posteriorly, about as long as wide, hairs robust, unicolor, black or rusty; D, Makalata macrura USNM 496480, short, rectangular scales in even rows, fine, salient hairs that are often banded dark at base, pale at tips. White scale bars 5 mm, fluid-fixed specimens except B." figureDoi="http://doi.org/10.5281/zenodo.5369720" httpUri="https://zenodo.org/record/5369720/files/figure.png" pageId="26" pageNumber="27">fig. 5A</figureCitation>
). Vibrissae robust, dense, and black, the longest reaching to shoulder when laid back. Two pairs of lateral mammae lie in the dorsal pelage field, one about midway between the limbs, the other just anterior to the flank. Hind feet, based only on
<taxonomicName id="4C35E4BAFFCBD567FD76FC3EFCB5FC38" box="[719,848,985,1011]" class="Mammalia" family="Echimyidae" genus="Pattonomys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rodentia" pageId="27" pageNumber="28" phylum="Chordata" rank="species" species="carrikeri">
<emphasis id="B941432BFFCBD567FD76FC3EFCB5FC38" box="[719,848,985,1011]" italics="true" pageId="27" pageNumber="28">P. carrikeri</emphasis>
</taxonomicName>
, with four small, ovoid digital pads and a fifth small outer pad budding laterally from D4. Hypothenar pad joined to large naked heel pad with only a slight depression between them (
<figureCitation id="130E83BCFFCBD567FCE4FBC0FC57FBF4" box="[861,946,1060,1087]" captionStart="FIG" captionStartId="16.[108,150,1542,1563]" captionTargetBox="[229,1091,224,1514]" captionTargetId="figure-11@16.[229,1091,224,1514]" captionTargetPageId="16" captionText="FIG. 6. Variation among genera in plantar surfaces of hindfeet, fluid-preserved specimens. A, Toromys rhipidurus AMNH 276710; B, Pattonomys carrikeri USNM 496501; C, Phyllomys pattoni, UFES 121; D, Makalata cf. didelphoides USNM 581981 (Bolivia); E, M. cf. macrura USNM 496480 (Venezuela); F, Echimys chrysurus, USNM 549839 (Rio Xingu). Note small digital pads, especially D3, and collars of small tubercles around pads in (AC); large digital pads and no collars of tubercles in (DF); merged D1 and thenar pad in (EF), leaving three separated digital pads; and differences in digital pad shapes (see text). Abbreviations: D, digital pads; Th, thenar pad; Hy, hypothenar pad. Arrow in A indicates supernumerary separated pad of Pattonomys spp. Figures not to scale. Photo panel C, Yuri Leite." figureDoi="http://doi.org/10.5281/zenodo.5369722" httpUri="https://zenodo.org/record/5369722/files/figure.png" pageId="27" pageNumber="28">fig. 6B</figureCitation>
); thenar pad elongate and slender, separated from the joined hypothenar and heel pads by a deep, narrow groove containing palmar tubercles. Plantar surfaces between pads with a few inconspicuous tubercles; digital pads bordered by distinct rings or collars of tubercles (
<figureCitation id="130E83BCFFCBD567FC25FB70FC15FB64" box="[924,1008,1172,1199]" captionStart="FIG" captionStartId="16.[108,150,1542,1563]" captionTargetBox="[229,1091,224,1514]" captionTargetId="figure-11@16.[229,1091,224,1514]" captionTargetPageId="16" captionText="FIG. 6. Variation among genera in plantar surfaces of hindfeet, fluid-preserved specimens. A, Toromys rhipidurus AMNH 276710; B, Pattonomys carrikeri USNM 496501; C, Phyllomys pattoni, UFES 121; D, Makalata cf. didelphoides USNM 581981 (Bolivia); E, M. cf. macrura USNM 496480 (Venezuela); F, Echimys chrysurus, USNM 549839 (Rio Xingu). Note small digital pads, especially D3, and collars of small tubercles around pads in (AC); large digital pads and no collars of tubercles in (DF); merged D1 and thenar pad in (EF), leaving three separated digital pads; and differences in digital pad shapes (see text). Abbreviations: D, digital pads; Th, thenar pad; Hy, hypothenar pad. Arrow in A indicates supernumerary separated pad of Pattonomys spp. Figures not to scale. Photo panel C, Yuri Leite." figureDoi="http://doi.org/10.5281/zenodo.5369722" httpUri="https://zenodo.org/record/5369722/files/figure.png" pageId="27" pageNumber="28">fig. 6B</figureCitation>
); forefoot digital pads also bordered by indistinct collars of tubercles, but the central palm with few or none. On the cranium, the frontal-squamosal suture at the back of the orbit is raised in a sharp ridge (
<figureCitation id="130E83BCFFCBD567FF07FAE0FED9FAD4" box="[190,316,1284,1311]" captionStart="FIG" captionStartId="17.[108,150,1500,1521]" captionTargetBox="[108,1212,224,1473]" captionTargetId="figure-7@17.[108,1212,224,1473]" captionTargetPageId="17" captionText="FIG. 7. Crania and mandibles, lateral view. A, Toromys grandis AMNH 93601; B, T. rhipidurus BMNH 28.7.21.89 (holotype); C, T. sp. nov., FMNH 55483 (holotype), D, Leiuromys occasius. AMNH 71897; E, Pattonomys punctatus BMNH 98.12.1.18 (holotype); F, P. flavidus USNM 63218 (holotype); G, P. carrikeri AMNH 130790; H, P. semivillosus USNM 280205. Black arrow in F indicates the deep mandible of P. flavidus relative to its congeners." figureDoi="http://doi.org/10.5281/zenodo.5369724" httpUri="https://zenodo.org/record/5369724/files/figure.png" pageId="27" pageNumber="28">fig. 7EH</figureCitation>
). Mastoid processes short, to the middle or lower edge of the auditory meatus, adpressed to the bulla, and enclosed to the tip by occipital bone (
<figureCitation id="130E83BCFFCBD567FC64FACDFBB6FA8F" box="[989,1107,1321,1348]" captionStart="FIG" captionStartId="17.[108,150,1500,1521]" captionTargetBox="[108,1212,224,1473]" captionTargetId="figure-7@17.[108,1212,224,1473]" captionTargetPageId="17" captionText="FIG. 7. Crania and mandibles, lateral view. A, Toromys grandis AMNH 93601; B, T. rhipidurus BMNH 28.7.21.89 (holotype); C, T. sp. nov., FMNH 55483 (holotype), D, Leiuromys occasius. AMNH 71897; E, Pattonomys punctatus BMNH 98.12.1.18 (holotype); F, P. flavidus USNM 63218 (holotype); G, P. carrikeri AMNH 130790; H, P. semivillosus USNM 280205. Black arrow in F indicates the deep mandible of P. flavidus relative to its congeners." figureDoi="http://doi.org/10.5281/zenodo.5369724" httpUri="https://zenodo.org/record/5369724/files/figure.png" pageId="27" pageNumber="28">fig. 7EH</figureCitation>
) and see also
<bibRefCitation id="EFA4E2C8FFCBD567FF1CFAABFE7FFAA2" author="Emmons, L. H." box="[165,410,1358,1385]" pageId="27" pageNumber="28" pagination="247 - 309" refId="ref23229" refString="Emmons, L. H. 2005. A revision of the arboreal Echimyidae (Rodentia: Echimyidae, Echimyinae); with descriptions of two new genera. In E. A. Lacey, and P. Myers (editors), Mammalian diversification: from chromosomes to phylogeography: 247 - 309. Berkeley: University of California Press." type="book chapter" year="2005">Emmons (2005: 263</bibRefCitation>
,
<figureCitation id="130E83BCFFCBD567FE11FAABFE02FAA2" box="[424,487,1358,1385]" captionStart="FIG" captionStartId="15.[108,150,569,590]" captionTargetBox="[108,1212,224,540]" captionTargetId="figure-420@15.[108,1212,224,540]" captionTargetPageId="15" captionText="FIG. 5. Scales and hairs on dorsal tail base. A, Pattonomys carrikeri USNM 496483, irregular scales nearly square or slightly hexagonal, hairs unicolored, long, and curled outward; B, Leiuromys occasius FMNH 84259, short, wide hexagonal scales in spiral rows, short, adpressed, scalelike hairs in pairs; C, Toromys rhipidurus AMNH 276710, scales rounded posteriorly, about as long as wide, hairs robust, unicolor, black or rusty; D, Makalata macrura USNM 496480, short, rectangular scales in even rows, fine, salient hairs that are often banded dark at base, pale at tips. White scale bars 5 mm, fluid-fixed specimens except B." figureDoi="http://doi.org/10.5281/zenodo.5369720" httpUri="https://zenodo.org/record/5369720/files/figure.png" pageId="27" pageNumber="28">fig. 5</figureCitation>
). Postglenoid foramen enclosed below by the petrosal bone in an anteriorly facing tube; the salient ventral lip of the auditory meatus is usually visible in dorsal aspect (
<figureCitation id="130E83BCFFCBD567FE85FA7DFE77FA78" box="[316,402,1433,1459]" captionStart="FIG" captionStartId="19.[108,150,1560,1581]" captionTargetBox="[132,1188,224,1533]" captionTargetId="figure-6@19.[132,1188,224,1533]" captionTargetPageId="19" captionText="FIG. 9. Crania, dorsal view. A, Toromys grandis AMNH 93601; B, T. rhipidurus BMNH 28.7.21.89 (holotype); C, T. sp. nov., FMNH 55483 (holotype), D, Leiuromys occasius. AMNH 71897; E, Pattonomys punctatus BMNH 98.12.1.18 (holotype); F, P. flavidus USNM 63218 (holotype); G, P. carrikeri AMNH 130790; H, P. semivillosus USNM 280205. Black arrow in A indicates recurved parietal-interparietal suture, white arrow the expanded squamosal shelf." figureDoi="http://doi.org/10.5281/zenodo.5369728" httpUri="https://zenodo.org/record/5369728/files/figure.png" pageId="27" pageNumber="28">fig. 9G</figureCitation>
), and is further extended in mature individuals by two overlapping additional bony rings that cup the meatus ventrally and laterally, but are incomplete dorsally (
<figureCitation id="130E83BCFFCBD567FFCCFA00FF3EFA34" box="[117,219,1508,1535]" captionStart="FIG" captionStartId="27.[109,149,846,867]" captionTargetBox="[132,1188,224,819]" captionTargetId="figure-319@27.[132,1188,224,819]" captionTargetPageId="27" captionText="FIG. 15. Pattonomys basicrania. A, Pattonomys punctatus USNM 374741; B, P. carrikeri USNM 456429. Note separate masticatory and buccinator foramina (arrows), robust extra bony rings below meatus, wide mesopterygoid fossa, and less inflated auditory tympanic bullae in A; joined foramina and more inflated bullae in B." figureDoi="http://doi.org/10.5281/zenodo.5369740" httpUri="https://zenodo.org/record/5369740/files/figure.png" pageId="27" pageNumber="28">fig. 15A</figureCitation>
). These are not fused and can be lost in skin preparation. Nasal bones in dorsal aspect flared distally, slightly pinched in medially, and about squared or slightly rounded posteriorly (
<figureCitation id="130E83BCFFCBD567FEBEF9CBFE63F982" box="[263,390,1583,1609]" captionStart="FIG" captionStartId="19.[108,150,1560,1581]" captionTargetBox="[132,1188,224,1533]" captionTargetId="figure-6@19.[132,1188,224,1533]" captionTargetPageId="19" captionText="FIG. 9. Crania, dorsal view. A, Toromys grandis AMNH 93601; B, T. rhipidurus BMNH 28.7.21.89 (holotype); C, T. sp. nov., FMNH 55483 (holotype), D, Leiuromys occasius. AMNH 71897; E, Pattonomys punctatus BMNH 98.12.1.18 (holotype); F, P. flavidus USNM 63218 (holotype); G, P. carrikeri AMNH 130790; H, P. semivillosus USNM 280205. Black arrow in A indicates recurved parietal-interparietal suture, white arrow the expanded squamosal shelf." figureDoi="http://doi.org/10.5281/zenodo.5369728" httpUri="https://zenodo.org/record/5369728/files/figure.png" pageId="27" pageNumber="28">fig. 9EH</figureCitation>
). Incisive foramina small, narrow, and slightly ovoid, with deep grooves and ridges leading onto the anterior palate to between the first or second cheekteeth (
<figureCitation id="130E83BCFFCBD567FFCCF99DFEE5F958" box="[117,256,1657,1683]" captionStart="FIG" captionStartId="20.[108,149,1562,1583]" captionTargetBox="[132,1188,224,1534]" captionTargetId="figure-11@20.[132,1188,224,1534]" captionTargetPageId="20" captionText="FIG. 10. Crania, palatal view. A, Toromys grandis AMNH 93601; B, T. rhipidurus BMNH 28.7.21.89 (holotype); C, T. sp. nov., FMNH 55483 (holotype), D, Leiuromys occasius. AMNH 71897; E, Pattonomys punctatus BMNH 98.12.1.18 (holotype); F, P. flavidus USNM 63218 (holotype); G, P. carrikeri AMNH 130790; H, P. semivillosus USNM 280205." figureDoi="http://doi.org/10.5281/zenodo.5369730" httpUri="https://zenodo.org/record/5369730/files/figure.png" pageId="27" pageNumber="28">fig. 10EH</figureCitation>
). Maxillary toothrows nearly parallel. Mandible with a low coronoid process only slightly shorter than the narrow condyloid process, and a shallow sigmoid notch (
<figureCitation id="130E83BCFFCBD567FB2CF97BFF5CF914" captionStart="FIG" captionStartId="17.[108,150,1500,1521]" captionTargetBox="[108,1212,224,1473]" captionTargetId="figure-7@17.[108,1212,224,1473]" captionTargetPageId="17" captionText="FIG. 7. Crania and mandibles, lateral view. A, Toromys grandis AMNH 93601; B, T. rhipidurus BMNH 28.7.21.89 (holotype); C, T. sp. nov., FMNH 55483 (holotype), D, Leiuromys occasius. AMNH 71897; E, Pattonomys punctatus BMNH 98.12.1.18 (holotype); F, P. flavidus USNM 63218 (holotype); G, P. carrikeri AMNH 130790; H, P. semivillosus USNM 280205. Black arrow in F indicates the deep mandible of P. flavidus relative to its congeners." figureDoi="http://doi.org/10.5281/zenodo.5369724" httpUri="https://zenodo.org/record/5369724/files/figure.png" pageId="27" pageNumber="28">fig. 7EH</figureCitation>
). Mandibular foramen in a fossa arising low on the condyloid ridge; masseteric crest not curved upward anteriorly, but forms a nearly straight line from the base of the angular process (
<figureCitation id="130E83BCFFCCD560FF6FFEE3FEAAFEEA" box="[214,335,263,289]" captionStart="FIG" captionStartId="17.[108,150,1500,1521]" captionTargetBox="[108,1212,224,1473]" captionTargetId="figure-7@17.[108,1212,224,1473]" captionTargetPageId="17" captionText="FIG. 7. Crania and mandibles, lateral view. A, Toromys grandis AMNH 93601; B, T. rhipidurus BMNH 28.7.21.89 (holotype); C, T. sp. nov., FMNH 55483 (holotype), D, Leiuromys occasius. AMNH 71897; E, Pattonomys punctatus BMNH 98.12.1.18 (holotype); F, P. flavidus USNM 63218 (holotype); G, P. carrikeri AMNH 130790; H, P. semivillosus USNM 280205. Black arrow in F indicates the deep mandible of P. flavidus relative to its congeners." figureDoi="http://doi.org/10.5281/zenodo.5369724" httpUri="https://zenodo.org/record/5369724/files/figure.png" pageId="28" pageNumber="29">fig. 7EH</figureCitation>
). Incisor enamel yellowish white, lowers tinged more orange than uppers; upper incisors opisthodont, robust, and strongly curved, with the base of the roots outside of, or level with, the maxillary base of the zygoma. Maxillary cheekteeth (
<figureCitation id="130E83BCFFCCD560FC66FEB5FB87FEA0" box="[991,1122,337,363]" captionStart="FIG" captionStartId="22.[108,150,1490,1511]" captionTargetBox="[115,1205,224,1460]" captionTargetId="figure-12@22.[115,1205,224,1460]" captionTargetPageId="22" captionText="FIG. 12. Occlusal patterns of upper (top row) and lower (bottom row) toothrows of Toromys and Pattonomys species, not to scale. A, Toromys sp. nov. FMNH 55483 holotype; B, T.sp. nov. LSUMZ 8904, an old individual; C, Toromys grandis AMNH 93604; D, Toromys rhipidurus AMNH 98667; E, Pattonomys carrikeri AMNH 31530; F, P. semivillosus USNM 280205; G, P. punctatus BMNH 9812118 (holotype). In Toromys maxillary teeth (AD) note laminar appearance of narrow, nearly parallel molar flexi, lophs of nearly uniform width, and mures lingually (left) of center of teeth. In Pattonomys (EG) note short, offset flexi, short, wide protoloph (arrow) especially pronounced on M2, mures about in midtooth; unequal loph sizes give irregular appearance. On mandibular teeth, note the straight (flat) posterior molar edges; the square (flat) lingual edge of central lophid (entoconid) in Toromys (AD, also compare with fig. 17)." figureDoi="http://doi.org/10.5281/zenodo.5369734" httpUri="https://zenodo.org/record/5369734/files/figure.png" pageId="28" pageNumber="29">fig. 12EG</figureCitation>
) squarish to rectangular, with rounded posterior lophs and an uneven appearance due to unequal loph sizes. DP4M3 always with two labial and two lingual flexi. Anterior lingual flexi (paraflexi) short, reaching only about half of the width of the tooth; the posterior lingual flexi (hypoflexi) longer; both labial flexi (para + hypoflexi) extend slightly more than half of the width of the tooth and are of the same length. DP4 and M1 always with a mure in the center of the tooth; M23 are with or without a mure, and all four flexi slant posteriorly from the edge to the middle of the tooth, so that labial and lingual flexi slant slightly in opposite directions. If without a mure, the joined flexi have a sharp elbow where the mure would be (
<figureCitation id="130E83BCFFCCD560FF49FD45FEB0FD70" box="[240,341,673,699]" captionStart="FIG" captionStartId="22.[108,150,1490,1511]" captionTargetBox="[115,1205,224,1460]" captionTargetId="figure-12@22.[115,1205,224,1460]" captionTargetPageId="22" captionText="FIG. 12. Occlusal patterns of upper (top row) and lower (bottom row) toothrows of Toromys and Pattonomys species, not to scale. A, Toromys sp. nov. FMNH 55483 holotype; B, T.sp. nov. LSUMZ 8904, an old individual; C, Toromys grandis AMNH 93604; D, Toromys rhipidurus AMNH 98667; E, Pattonomys carrikeri AMNH 31530; F, P. semivillosus USNM 280205; G, P. punctatus BMNH 9812118 (holotype). In Toromys maxillary teeth (AD) note laminar appearance of narrow, nearly parallel molar flexi, lophs of nearly uniform width, and mures lingually (left) of center of teeth. In Pattonomys (EG) note short, offset flexi, short, wide protoloph (arrow) especially pronounced on M2, mures about in midtooth; unequal loph sizes give irregular appearance. On mandibular teeth, note the straight (flat) posterior molar edges; the square (flat) lingual edge of central lophid (entoconid) in Toromys (AD, also compare with fig. 17)." figureDoi="http://doi.org/10.5281/zenodo.5369734" httpUri="https://zenodo.org/record/5369734/files/figure.png" pageId="28" pageNumber="29">fig. 12G</figureCitation>
, arrow). Lower premolar usually divided by one central flexid into two V-shaped lophs, but with wear forms a midtooth mure. In m12 the lingual flexids are parallel and of about equal length. Flexids of all mandibular cheekteeth wide. Hypoconid with a flat labial profile (
<figureCitation id="130E83BCFFCCD560FEFEFCF5FE35FCE0" box="[327,464,785,811]" captionStart="FIG" captionStartId="22.[108,150,1490,1511]" captionTargetBox="[115,1205,224,1460]" captionTargetId="figure-12@22.[115,1205,224,1460]" captionTargetPageId="22" captionText="FIG. 12. Occlusal patterns of upper (top row) and lower (bottom row) toothrows of Toromys and Pattonomys species, not to scale. A, Toromys sp. nov. FMNH 55483 holotype; B, T.sp. nov. LSUMZ 8904, an old individual; C, Toromys grandis AMNH 93604; D, Toromys rhipidurus AMNH 98667; E, Pattonomys carrikeri AMNH 31530; F, P. semivillosus USNM 280205; G, P. punctatus BMNH 9812118 (holotype). In Toromys maxillary teeth (AD) note laminar appearance of narrow, nearly parallel molar flexi, lophs of nearly uniform width, and mures lingually (left) of center of teeth. In Pattonomys (EG) note short, offset flexi, short, wide protoloph (arrow) especially pronounced on M2, mures about in midtooth; unequal loph sizes give irregular appearance. On mandibular teeth, note the straight (flat) posterior molar edges; the square (flat) lingual edge of central lophid (entoconid) in Toromys (AD, also compare with fig. 17)." figureDoi="http://doi.org/10.5281/zenodo.5369734" httpUri="https://zenodo.org/record/5369734/files/figure.png" pageId="28" pageNumber="29">fig. 12EG</figureCitation>
). Posterior borders of lower cheekteeth straight. Phallus of
<taxonomicName id="4C35E4BAFFCCD560FB11FCF6FF34FC9A" class="Mammalia" family="Echimyidae" genus="Pattonomys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rodentia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="carrikeri">
<emphasis id="B941432BFFCCD560FB11FCF6FF34FC9A" italics="true" pageId="28" pageNumber="29">P. carrikeri</emphasis>
</taxonomicName>
long and slender, with a long, pointed, bacular papilla, small urethral lappet, border of ventral crater wall a straight line without a U- or V-shaped ventral fold (
<bibRefCitation id="EFA4E2C8FFCCD560FC47FCB8FB5CFCBC" author="Emmons, L. H." box="[1022,1209,860,887]" pageId="28" pageNumber="29" pagination="247 - 309" refId="ref23229" refString="Emmons, L. H. 2005. A revision of the arboreal Echimyidae (Rodentia: Echimyidae, Echimyinae); with descriptions of two new genera. In E. A. Lacey, and P. Myers (editors), Mammalian diversification: from chromosomes to phylogeography: 247 - 309. Berkeley: University of California Press." type="book chapter" year="2005">Emmons, 2005</bibRefCitation>
:
<figureCitation id="130E83BCFFCCD560FFD7FC65FF2BFC50" box="[110,206,897,923]" captionStart="FIG" captionStartId="20.[108,149,1562,1583]" captionTargetBox="[132,1188,224,1534]" captionTargetId="figure-11@20.[132,1188,224,1534]" captionTargetPageId="20" captionText="FIG. 10. Crania, palatal view. A, Toromys grandis AMNH 93601; B, T. rhipidurus BMNH 28.7.21.89 (holotype); C, T. sp. nov., FMNH 55483 (holotype), D, Leiuromys occasius. AMNH 71897; E, Pattonomys punctatus BMNH 98.12.1.18 (holotype); F, P. flavidus USNM 63218 (holotype); G, P. carrikeri AMNH 130790; H, P. semivillosus USNM 280205." figureDoi="http://doi.org/10.5281/zenodo.5369730" httpUri="https://zenodo.org/record/5369730/files/figure.png" pageId="28" pageNumber="29">fig. 10C</figureCitation>
); those of other species are undescribed. Karyotypes of
<specimenCount id="9D3354B0FFCCD560FCD5FC65FBC3FC50" box="[876,1062,897,923]" pageId="28" pageNumber="29" type="generic">four specimens</specimenCount>
, which from their localities we assign to
<taxonomicName id="4C35E4BAFFCCD560FE7AFC43FDD8FC0A" box="[451,573,935,961]" class="Mammalia" family="Echimyidae" genus="Pattonomys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rodentia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="flavidus">
<emphasis id="B941432BFFCCD560FE7AFC43FDD8FC0A" box="[451,573,935,961]" italics="true" pageId="28" pageNumber="29">P. flavidus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C35E4BAFFCCD560FDC3FC43FD18FC0A" box="[634,765,935,961]" class="Mammalia" family="Echimyidae" genus="Pattonomys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rodentia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="carrikeri">
<emphasis id="B941432BFFCCD560FDC3FC43FD18FC0A" box="[634,765,935,961]" italics="true" pageId="28" pageNumber="29">P. carrikeri</emphasis>
</taxonomicName>
, were identical, 2
<emphasis id="B941432BFFCCD560FC6FFC43FC0EFC0B" box="[982,1003,935,960]" italics="true" pageId="28" pageNumber="29">N</emphasis>
= 94, among the highest found in
<taxonomicName id="4C35E4BAFFCCD560FE86FC28FD33FC2D" authority="(Aguilera et al., 1998)" baseAuthorityName="Aguilera" baseAuthorityYear="1998" box="[319,726,972,999]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="28" pageNumber="29" phylum="Chordata" rank="class">
Mammalia (
<bibRefCitation id="EFA4E2C8FFCCD560FE6BFC28FD29FC2D" author="Aguilera, M. &amp; N. Sangines &amp; A. Perez-Zapata" box="[466,716,972,999]" pageId="28" pageNumber="29" pagination="181 - 187" refId="ref22701" refString="Aguilera, M., N. Sangines, and A. Perez-Zapata. 1998. Echimys semivillosus, a rodent species with a very high chromosomal number. Caryologia 51: 181 - 187." type="journal article" year="1998">Aguilera et al., 1998</bibRefCitation>
)
</taxonomicName>
.
</paragraph>
<caption id="DF4ACFB1FFCBD567FFD4FCAAFB6EFC55" ID-DOI="http://doi.org/10.5281/zenodo.5369740" ID-Zenodo-Dep="5369740" httpUri="https://zenodo.org/record/5369740/files/figure.png" pageId="27" pageNumber="28" startId="27.[109,149,846,867]" targetBox="[132,1188,224,819]" targetPageId="27">
<paragraph id="8B8A9F39FFCBD567FFD4FCAAFB6EFC55" blockId="27.[108,1213,844,927]" pageId="27" pageNumber="28">
FIG. 15.
<taxonomicName id="4C35E4BAFFCBD567FF7DFCA9FED3FCAF" authorityName="Emmons" authorityYear="2005" box="[196,310,845,868]" class="Mammalia" family="Echimyidae" genus="Pattonomys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rodentia" pageId="27" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis id="B941432BFFCBD567FF7DFCA9FED3FCAF" box="[196,310,845,868]" italics="true" pageId="27" pageNumber="28">Pattonomys</emphasis>
</taxonomicName>
basicrania.
<emphasis id="B941432BFFCBD567FE08FCA8FE2DFCAF" bold="true" box="[433,456,844,868]" pageId="27" pageNumber="28">A,</emphasis>
<taxonomicName id="4C35E4BAFFCBD567FE68FCA9FD1CFCAF" authority="USNM" authorityName="USNM" box="[465,761,844,868]" class="Mammalia" family="Echimyidae" genus="Pattonomys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rodentia" pageId="27" pageNumber="28" phylum="Chordata" rank="species" species="punctatus">
<emphasis id="B941432BFFCBD567FE68FCA9FD4CFCAF" box="[465,681,845,868]" italics="true" pageId="27" pageNumber="28">Pattonomys punctatus</emphasis>
USNM
</taxonomicName>
374741;
<emphasis id="B941432BFFCBD567FCEFFCA8FC89FCAF" bold="true" box="[854,876,844,868]" pageId="27" pageNumber="28">B,</emphasis>
<taxonomicName id="4C35E4BAFFCBD567FCCDFCA9FBCAFCAF" authority="USNM" authorityName="USNM" box="[884,1071,844,868]" class="Mammalia" family="Echimyidae" genus="Pattonomys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rodentia" pageId="27" pageNumber="28" phylum="Chordata" rank="species" species="carrikeri">
<emphasis id="B941432BFFCBD567FCCDFCA9FC05FCAF" box="[884,992,844,868]" italics="true" pageId="27" pageNumber="28">P. carrikeri</emphasis>
USNM
</taxonomicName>
456429. Note separate masticatory and buccinator foramina (arrows), robust extra bony rings below meatus, wide mesopterygoid fossa, and less inflated auditory tympanic bullae in A; joined foramina and more inflated bullae in B.
</paragraph>
</caption>
<paragraph id="8B8A9F39FFCCD560FF25FC15FC16FA6C" blockId="28.[108,1216,225,1447]" pageId="28" pageNumber="29">
COMPARISONS WITH OTHER GENERA: A white-speckled, spiny pelage is unique to the genus, although
<taxonomicName id="4C35E4BAFFCCD560FF59FBF3FEBDFBFB" authorityName="Emmons &amp; Fabre" authorityYear="2018" box="[224,344,1047,1072]" class="Mammalia" family="Echimyidae" genus="Leiuromys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rodentia" pageId="28" pageNumber="29" phylum="Chordata" rank="genus">
<emphasis id="B941432BFFCCD560FF59FBF3FEBDFBFB" box="[224,344,1047,1072]" italics="true" pageId="28" pageNumber="29">Leiuromys</emphasis>
</taxonomicName>
,
<taxonomicName id="4C35E4BAFFCCD560FED3FBF3FE31FBFB" authorityName="Wagner" authorityYear="1845" box="[362,468,1047,1072]" class="Mammalia" family="Echimyidae" genus="Mesomys" kingdom="Animalia" order="Rodentia" pageId="28" pageNumber="29" phylum="Chordata" rank="genus">
<emphasis id="B941432BFFCCD560FED3FBF3FE31FBFB" box="[362,468,1047,1072]" italics="true" pageId="28" pageNumber="29">Mesomys</emphasis>
</taxonomicName>
,
<taxonomicName id="4C35E4BAFFCCD560FE5CFBF3FD97FBFA" authorityName="Thomas" authorityYear="1920" box="[485,626,1047,1073]" class="Mammalia" family="Echimyidae" genus="Lonchothrix" kingdom="Animalia" order="Rodentia" pageId="28" pageNumber="29" phylum="Chordata" rank="genus">
<emphasis id="B941432BFFCCD560FE5CFBF3FD97FBFA" box="[485,626,1047,1073]" italics="true" pageId="28" pageNumber="29">Lonchothrix</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C35E4BAFFCCD560FD02FBF3FC6BFBFB" box="[699,910,1047,1073]" class="Mammalia" family="Echimyidae" genus="Phyllomys" kingdom="Animalia" order="Rodentia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="pattoni">
<emphasis id="B941432BFFCCD560FD02FBF3FC6BFBFB" box="[699,910,1047,1073]" italics="true" pageId="28" pageNumber="29">Phyllomys pattoni</emphasis>
</taxonomicName>
have strong, buff-tipped spines. There is no geographic overlap of these genera with
<taxonomicName id="4C35E4BAFFCCD560FC98FBD9FC42FB9D" authorityName="Emmons" authorityYear="2005" box="[801,935,1085,1110]" class="Mammalia" family="Echimyidae" genus="Pattonomys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rodentia" pageId="28" pageNumber="29" phylum="Chordata" rank="genus">
<emphasis id="B941432BFFCCD560FC98FBD9FC42FB9D" box="[801,935,1085,1110]" italics="true" pageId="28" pageNumber="29">Pattonomys</emphasis>
</taxonomicName>
. The maxillary occlusal pattern of
<taxonomicName id="4C35E4BAFFCCD560FF55FB86FE2BFBB0" box="[236,462,1121,1147]" class="Mammalia" family="Echimyidae" genus="Pattonomys" kingdom="Animalia" order="Rodentia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B941432BFFCCD560FF55FB86FE96FBB0" box="[236,371,1122,1147]" italics="true" pageId="28" pageNumber="29">Pattonomys</emphasis>
species
</taxonomicName>
is unique for the family, and even single or worn upper teeth usually can be identified to genus. A quadralophodont lower premolar is shared only with
<taxonomicName id="4C35E4BAFFCCD560FFD5FB49FF06FB0D" authorityName="Emmons &amp; Fabre" authorityYear="2018" box="[108,227,1197,1222]" class="Mammalia" family="Echimyidae" genus="Leiuromys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rodentia" pageId="28" pageNumber="29" phylum="Chordata" rank="genus">
<emphasis id="B941432BFFCCD560FFD5FB49FF06FB0D" box="[108,227,1197,1222]" italics="true" pageId="28" pageNumber="29">Leiuromys</emphasis>
</taxonomicName>
among
<taxonomicName id="4C35E4BAFFCCD560FEFAFB48FE5AFB0D" box="[323,447,1196,1222]" class="Mammalia" family="Echimyidae" kingdom="Animalia" order="Rodentia" pageId="28" pageNumber="2" phylum="Chordata" rank="tribe" tribe="Echimyini">Echimyini</taxonomicName>
, but in that genus, the mure is near the lingual edge of the upper molars, and the posterior edges of the lower cheekteeth are curved, while the mure of
<taxonomicName id="4C35E4BAFFCCD560FBE0FB36FF17FADA" class="Mammalia" family="Echimyidae" genus="Pattonomys" kingdom="Animalia" order="Rodentia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B941432BFFCCD560FBE0FB36FF7CFADB" italics="true" pageId="28" pageNumber="29">Pattonomys</emphasis>
species
</taxonomicName>
is near the center of the upper molars, and the posterior edges of the lower molars are straight. The flexids of
<taxonomicName id="4C35E4BAFFCCD560FE13FAF9FDD4FAFD" authorityName="Emmons" authorityYear="2005" box="[426,561,1309,1334]" class="Mammalia" family="Echimyidae" genus="Pattonomys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rodentia" pageId="28" pageNumber="29" phylum="Chordata" rank="genus">
<emphasis id="B941432BFFCCD560FE13FAF9FDD4FAFD" box="[426,561,1309,1334]" italics="true" pageId="28" pageNumber="29">Pattonomys</emphasis>
</taxonomicName>
are wider than those of other
<taxonomicName id="4C35E4BAFFCCD560FC23FAF8FBF3FAFD" box="[922,1046,1308,1334]" class="Mammalia" family="Echimyidae" kingdom="Animalia" order="Rodentia" pageId="28" pageNumber="2" phylum="Chordata" rank="tribe" tribe="Echimyini">Echimyini</taxonomicName>
. Likewise, the long, thin, simple phallus of
<taxonomicName id="4C35E4BAFFCCD560FE07FAA6FDDEFA90" box="[446,571,1345,1371]" class="Mammalia" family="Echimyidae" genus="Pattonomys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rodentia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="carrikeri">
<emphasis id="B941432BFFCCD560FE07FAA6FDDEFA90" box="[446,571,1345,1371]" italics="true" pageId="28" pageNumber="29">P. carrikeri</emphasis>
</taxonomicName>
is thus far unique among the
<taxonomicName id="4C35E4BAFFCCD560FC24FAA5FBFCFA90" box="[925,1049,1345,1371]" class="Mammalia" family="Echimyidae" kingdom="Animalia" order="Rodentia" pageId="28" pageNumber="2" phylum="Chordata" rank="tribe" tribe="Echimyini">Echimyini</taxonomicName>
; those known for the other genera are short, thick, and with additional folds (
<bibRefCitation id="EFA4E2C8FFCCD560FCCBFA83FC1FFA4A" author="Leite, Y. L. R." box="[882,1018,1383,1409]" pageId="28" pageNumber="29" pagination="1 - 118" refId="ref24508" refString="Leite, Y. L. R. 2003. Evolution and systematics of the Atlantic tree rats, genus Phyllomys (Rodentia, Echimyidae), with description of two new species. University of California Publications in Zoology 132: 1 - 118 + xv." type="journal article" year="2003">Leite, 2003</bibRefCitation>
;
<bibRefCitation id="EFA4E2C8FFCCD560FBB1FA83FF4DFA6C" author="Emmons, L. H." pageId="28" pageNumber="29" pagination="247 - 309" refId="ref23229" refString="Emmons, L. H. 2005. A revision of the arboreal Echimyidae (Rodentia: Echimyidae, Echimyinae); with descriptions of two new genera. In E. A. Lacey, and P. Myers (editors), Mammalian diversification: from chromosomes to phylogeography: 247 - 309. Berkeley: University of California Press." type="book chapter" year="2005">Emmons et al., 2005</bibRefCitation>
:
<figureCitation id="130E83BCFFCCD560FF0AFA68FEF7FA6D" box="[179,274,1420,1446]" captionStart="FIG" captionStartId="20.[108,149,1562,1583]" captionTargetBox="[132,1188,224,1534]" captionTargetId="figure-11@20.[132,1188,224,1534]" captionTargetPageId="20" captionText="FIG. 10. Crania, palatal view. A, Toromys grandis AMNH 93601; B, T. rhipidurus BMNH 28.7.21.89 (holotype); C, T. sp. nov., FMNH 55483 (holotype), D, Leiuromys occasius. AMNH 71897; E, Pattonomys punctatus BMNH 98.12.1.18 (holotype); F, P. flavidus USNM 63218 (holotype); G, P. carrikeri AMNH 130790; H, P. semivillosus USNM 280205." figureDoi="http://doi.org/10.5281/zenodo.5369730" httpUri="https://zenodo.org/record/5369730/files/figure.png" pageId="28" pageNumber="29">fig. 10C</figureCitation>
), but descriptions are needed for other species in every genus.
</paragraph>
</subSubSection>
</treatment>
</document>