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<document id="8652AA27618AC769FFA7CF92ED70E82C" ID-DOI="10.1206/0003-0090(2000)244&lt;0001:MOTRJA&gt;2.0.CO;2" ID-ISSN="0003-0090" ID-Zenodo-Dep="5347311" IM.illustrations_approvedBy="felipe" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="felipe" checkinTime="1630203687125" checkinUser="felipe" docAuthor="PATTON, JAMES L., DA SILVA, MARIA NAZARETH F. &amp; MALCOLM, JAY R." docDate="2000" docId="039E01774BA3D942FCA9327AB46AFC57" docLanguage="en" docName="B244.pdf" docOrigin="Bulletin of the American Museum of Natural History 2000 (244)" docSource="http://www.bioone.org/perlserv/?request=get-abstract&amp;doi=10.1206%2F0003-0090(2000)244%3C0001%3AMOTRJA%3E2.0.CO%3B2" docStyle="DocumentStyle:2B34016C1EBFE58B07368F9517618743.3:BulAmeMusNatHis.2000-2010.journal_article.0cover.type1" docStyleId="2B34016C1EBFE58B07368F9517618743" docStyleName="BulAmeMusNatHis.2000-2010.journal_article.0cover.type1" docStyleVersion="3" docTitle="Proechimys steerei Goldman 1911" docType="treatment" docVersion="3" lastPageNumber="259" masterDocId="FFA7790F4B54D840FFB7335EB74EFFCF" masterDocTitle="Mammals Of The Rio Juruá And The Evolutionary And Ecological Diversification Of Amazonia" masterLastPageNumber="306" masterPageNumber="1" pageNumber="248" updateTime="1699183939665" updateUser="plazi" zenodo-license-document="CC-BY-4.0">
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<mods:title id="FC1A1EB5EE09F514404A88929A570C65">Mammals Of The Rio Juruá And The Evolutionary And Ecological Diversification Of Amazonia</mods:title>
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<mods:namePart id="95F30212344FA9677D7F48D864D933C3">DA SILVA, MARIA NAZARETH F.</mods:namePart>
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<subSubSection id="C32DE3EA4BA3D8B7FCA9327AB3F1FEF4" box="[798,1215,292,315]" pageId="247" pageNumber="248" type="nomenclature">
<paragraph id="8B88B0614BA3D8B7FCA9327AB3F1FEF4" blockId="247.[798,1215,292,315]" box="[798,1215,292,315]" pageId="247" pageNumber="248">
<heading id="D0C0070D4BA3D8B7FCA9327AB3F1FEF4" box="[798,1215,292,315]" centered="true" fontSize="9" level="2" pageId="247" pageNumber="248" reason="2">
<taxonomicName id="4C37CBE24BA3D8B7FCA9327AB3F1FEF4" ID-CoL="4MQLM" authority="Goldman, 1911" authorityName="Goldman" authorityYear="1911" box="[798,1215,292,315]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="247" pageNumber="248" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734BA3D8B7FCA9327AB4B0FEF4" box="[798,1022,292,315]" italics="true" pageId="247" pageNumber="248">Proechimys steerei</emphasis>
Goldman, 1911
</taxonomicName>
</heading>
</paragraph>
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<subSubSection id="C32DE3EA4BA3D8B7FCB33204B331FE27" pageId="247" pageNumber="248" type="materials_examined">
<paragraph id="8B88B0614BA3D8B7FCB33204B331FE27" blockId="247.[743,1272,346,1741]" pageId="247" pageNumber="248">
<materialsCitation id="3B5FBA3C4BA3D8B7FCB33204B470FE62" county="northwestern Brazil'" location="Rio Purus" municipality="Amazon" pageId="247" pageNumber="248" specimenCount="1" typeStatus="holotype">
<typeStatus id="548C0EC34BA3D8B7FCB33204B473FEBE" box="[772,829,346,369]" pageId="247" pageNumber="248">TYPE</typeStatus>
LOCALITY:
<location id="8EE8E6BA4BA3D8B7FC683204B32DFEBE" LSID="urn:lsid:plazi:treatment:039E01774BA3D942FCA9327AB46AFC57:8EE8E6BA4BA3D8B7FC683204B32DFEBE" box="[991,1123,346,369]" county="northwestern Brazil'" municipality="Amazon" name="Rio Purus" pageId="247" pageNumber="248">Rio Purus</location>
, a southern tributary of the
<collectingMunicipality id="6BEC2A1B4BA3D8B7FC0B3226B36FFE40" box="[956,1057,376,399]" pageId="247" pageNumber="248">Amazon</collectingMunicipality>
, in
<collectingCounty id="62E9C8ED4BA3D8B7FBD73226B470FE62" pageId="247" pageNumber="248">northwestern Brazil</collectingCounty>
</materialsCitation>
;
<materialsCitation id="3B5FBA3C4BA3D8B7FCE632C8B335FE27" country="Brazil" location="upper Rio Purus" pageId="247" pageNumber="248" specimenCount="1" stateProvince="Amazonas" typeStatus="holotype">
recorded as Hyutanahan,
<location id="8EE8E6BA4BA3D8B7FB3632C8B462FE05" LSID="urn:lsid:plazi:treatment:039E01774BA3D942FCA9327AB46AFC57:8EE8E6BA4BA3D8B7FB3632C8B462FE05" country="Brazil" name="upper Rio Purus" pageId="247" pageNumber="248" stateProvince="Amazonas">upper Rio Purus</location>
, Provincia Lábrea,
<collectingRegion id="49F37E834BA3D8B7FBB832EDB44CFE27" country="Brazil" name="Amazonas" pageId="247" pageNumber="248">Estado do Amazonas</collectingRegion>
,
<collectingCountry id="F320F0F14BA3D8B7FCA7328FB419FE27" box="[784,855,465,488]" name="Brazil" pageId="247" pageNumber="248">Brazil</collectingCountry>
by Moojen (1948: 338)
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="C32DE3EA4BA3D8BDFCB332B0B460F902" lastPageId="253" lastPageNumber="254" pageId="247" pageNumber="248" type="description">
<paragraph id="8B88B0614BA3D8B7FCB332B0B4E7FA86" blockId="247.[743,1272,346,1741]" pageId="247" pageNumber="248">
DESCRIPTION:
<taxonomicName id="4C37CBE24BA3D8B7FC0732B1B3D7FDC9" authorityName="Goldman" authorityYear="1911" box="[944,1177,495,518]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="247" pageNumber="248" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734BA3D8B7FC0732B1B3D7FDC9" box="[944,1177,495,518]" italics="true" pageId="247" pageNumber="248">Proechimys steerei</emphasis>
</taxonomicName>
is the largest species of spiny rats found in the Rio Jurua´, with individuals from true várzea sites during the flooding season maximally weighing nearly
<quantity id="4CCF1D844BA3D8B7FCDB313BB4E9FDB3" box="[876,935,613,636]" metricMagnitude="0" metricUnit="kg" metricValue="1.0" pageId="247" pageNumber="248" unit="kg" value="1.0">1 kg</quantity>
. The ears and hind feet are large, and the tail is proportionately short approximately two thirds that of the body (tables 60 and 64). The color of the tail is dark brown above and white to cream ventrally it is clothed by hair but the scales remain conspicuous to the eye. The color of the dorsal surface of the hind foot is characteristic of this species: a pale to dark brown outer band and whitish inner band along the length of the foot, from the tarsal joint to the end of the toes, in most individuals. Another distinctive feature is the narrow, short, and rath­ er lax aristiform hairs on the dorsum (da Silva, 1998: fig. 3), that contribute to a characteristically softer fur than is found in any other spiny rat, not only along the Rio Juruá but in all of western Amazonia. As is true of most species of
<taxonomicName id="4C37CBE24BA3D8B7FC1C3721B37AFB59" authorityName="Allen" authorityYear="1899" box="[939,1076,1151,1174]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="247" pageNumber="248" phylum="Chordata" rank="genus">
<emphasis id="B9436C734BA3D8B7FC1C3721B37AFB59" box="[939,1076,1151,1174]" italics="true" pageId="247" pageNumber="248">Proechimys</emphasis>
</taxonomicName>
, there is no lateral stripe and the reddish color of the sides of the body contrasts sharply with the pure white venter. The texture of the ventral fur in
<taxonomicName id="4C37CBE24BA3D8B7FCB237A9B43DFAC1" authorityName="Goldman" authorityYear="1911" box="[773,883,1270,1294]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="247" pageNumber="248" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734BA3D8B7FCB237A9B459FAC1" box="[773,791,1271,1294]" italics="true" pageId="247" pageNumber="248">P</emphasis>
.
<emphasis id="B9436C734BA3D8B7FC9137A9B43DFAC1" box="[806,883,1271,1294]" italics="true" pageId="247" pageNumber="248">steerei</emphasis>
</taxonomicName>
also seems thicker and distinctly more velvety to both the eye and touch than in other species.
</paragraph>
<paragraph id="8B88B0614BA3D8B7FCB3360EB4ACF9D5" blockId="247.[743,1272,346,1741]" pageId="247" pageNumber="248">
The baculum of our specimens of
<taxonomicName id="4C37CBE24BA3D8B7FB22360EB5B2FA4A" authorityName="Goldman" authorityYear="1911" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="247" pageNumber="248" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734BA3D8B7FB22360EB3E9FAA8" box="[1173,1191,1360,1383]" italics="true" pageId="247" pageNumber="248">P</emphasis>
.
<emphasis id="B9436C734BA3D8B7FB02360EB5B2FA4A" italics="true" pageId="247" pageNumber="248">steerei</emphasis>
</taxonomicName>
is similar to that described by Patton (1987) for his
<taxonomicName id="4C37CBE24BA3D8B7FC3936D2B491FA6C" authorityName="Thomas" authorityYear="1905" box="[910,991,1420,1443]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="247" pageNumber="248" phylum="Chordata" rank="species" species="goeldii">
<emphasis id="B9436C734BA3D8B7FC3936D2B491FA6C" box="[910,991,1420,1443]" italics="true" pageId="247" pageNumber="248">goeldii</emphasis>
</taxonomicName>
­ group; it is moderately long and narrow, especially when compared to bacula of
<taxonomicName id="4C37CBE24BA3D8B7FC3D3696B37AFA10" baseAuthorityName="sensu Patton" baseAuthorityYear="1987" box="[906,1076,1479,1503]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="247" pageNumber="248" phylum="Chordata" rank="species" species="brevicauda">
<emphasis id="B9436C734BA3D8B7FC3D3696B4D2FA10" box="[906,924,1480,1503]" italics="true" pageId="247" pageNumber="248">P</emphasis>
.
<emphasis id="B9436C734BA3D8B7FC063696B37AFA10" box="[945,1076,1480,1503]" italics="true" pageId="247" pageNumber="248">brevicauda</emphasis>
</taxonomicName>
and
<taxonomicName id="4C37CBE24BA3D8B7FBCE3696B3BEFA10" authorityName="Petter" authorityYear="1978" box="[1145,1264,1479,1503]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="247" pageNumber="248" phylum="Chordata" rank="species" species="cuvieri">
<emphasis id="B9436C734BA3D8B7FBCE3696B3C5FA10" box="[1145,1163,1480,1503]" italics="true" pageId="247" pageNumber="248">P</emphasis>
.
<emphasis id="B9436C734BA3D8B7FB283696B3BEFA10" box="[1183,1264,1480,1503]" italics="true" pageId="247" pageNumber="248">cuvieri</emphasis>
</taxonomicName>
although it is shorter and wider than that of
<taxonomicName id="4C37CBE24BA3D8B7FD50355DB42DF9D5" authorityName="Thomas" authorityYear="1900" box="[743,867,1539,1562]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="247" pageNumber="248" phylum="Chordata" rank="species" species="simonsi">
<emphasis id="B9436C734BA3D8B7FD50355DB5B7F9D5" box="[743,761,1539,1562]" italics="true" pageId="247" pageNumber="248">P</emphasis>
.
<emphasis id="B9436C734BA3D8B7FCBD355DB42DF9D5" box="[778,867,1539,1562]" italics="true" pageId="247" pageNumber="248">simonsi</emphasis>
</taxonomicName>
(fig. 137).
</paragraph>
<paragraph id="8B88B0614BA3D8B7FCB3357FB3B9F902" blockId="247.[743,1272,346,1741]" pageId="247" pageNumber="248">The skull is large, with a long and narrow rostrum (figs. 138 and 139) and well­developed supraorbital ledge (Patton, 1987: fig 21b). The incisive foramen is lyrate to oval in outline, with slightly to well­flanged posterolateral margins in the great majority of</paragraph>
<paragraph id="8B88B0614BACD8B8FF14339FB7A3FBE4" blockId="248.[163,692,193,1741]" pageId="248" pageNumber="249">
individuals (130 out of
<specimenCount id="9D317BE84BACD8B8FE74339FB53BFF17" box="[451,629,193,216]" pageId="248" pageNumber="249" type="generic">134 specimens</specimenCount>
) that form grooves extending onto the palate (fig. 140). The premaxillary portion of the septum is short, less than half the length of the opening, the maxillary portion is distinctly narrow, and both are in contact in most specimens (104 out of 135); the vomer is usually not visible (101 out of
<specimenCount id="9D317BE84BACD8B8FE7A32D0B5C8FE6A" box="[461,646,398,421]" pageId="248" pageNumber="249" type="generic">133 specimens</specimenCount>
). A groove is present on the floor of the infraorbital foramen, but development of the lateral flange is weak (of
<specimenCount id="9D317BE84BACD8B8FE1B32B8B514FE32" box="[428,602,486,509]" pageId="248" pageNumber="249" type="generic">134 specimens</specimenCount>
, 79 had a shallow groove but without lateral flanges and only four had a groove present with moderately developed lateral flanges; all 51 remaining specimens had a smooth floor without any groove). The mesopterygoid fossa is relatively broad, but penetrates the palate to a level between the posterior and anterior margins of M3 (fig. 141). Most specimens show three folds in PM4, M1, and M3 (118, 97, and 93 out of
<specimenCount id="9D317BE84BACD8B8FE673052B5CDFCEC" box="[464,643,780,803]" pageId="248" pageNumber="249" type="generic">134 specimens</specimenCount>
, respectively; the remaining individuals have four folds in those teeth) and four folds in M2 (91 out of
<specimenCount id="9D317BE84BACD8B8FEEA303AB55FFCB4" box="[349,529,868,891]" pageId="248" pageNumber="249" type="generic">134 specimens</specimenCount>
, with 3 folds in all others). Overall, these characters observed for specimens from the Rio Juruá compare with those described by Patton (1987) for specimens of his
<taxonomicName id="4C37CBE24BACD8B8FDB03087B516FC3F" authorityName="Thomas" authorityYear="1905" box="[519,600,985,1008]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="248" pageNumber="249" phylum="Chordata" rank="species" species="goeldii">
<emphasis id="B9436C734BACD8B8FDB03087B516FC3F" box="[519,600,985,1008]" italics="true" pageId="248" pageNumber="249">goeldii</emphasis>
</taxonomicName>
­ group from other localities outside the Rio Juruá Basin.
</paragraph>
<paragraph id="8B88B0614BACD8B8FF76376FB56FFB4C" blockId="248.[163,692,193,1741]" pageId="248" pageNumber="249">SELECTED MEASUREMENTS: Means and ranges of selected external and cranial measurements are given in table 64.</paragraph>
<paragraph id="8B88B0614BACD8B8FF7737D7B476FEDC" blockId="248.[163,692,193,1741]" lastBlockId="248.[739,1268,193,1741]" pageId="248" pageNumber="249">
COMPARISONS:
<taxonomicName id="4C37CBE24BACD8B8FEC437D7B518FB6F" authorityName="Goldman" authorityYear="1911" box="[371,598,1161,1184]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="248" pageNumber="249" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734BACD8B8FEC437D7B518FB6F" box="[371,598,1161,1184]" italics="true" pageId="248" pageNumber="249">Proechimys steerei</emphasis>
</taxonomicName>
is perhaps the most easily recognized species of spiny rat in the Rio Juruá basin, being distinguished from all other
<taxonomicName id="4C37CBE24BACD8B8FE6237BFB510FB37" authorityName="Allen" authorityYear="1899" box="[469,606,1249,1272]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="248" pageNumber="249" phylum="Chordata" rank="genus">
<emphasis id="B9436C734BACD8B8FE6237BFB510FB37" box="[469,606,1249,1272]" italics="true" pageId="248" pageNumber="249">Proechimys</emphasis>
</taxonomicName>
by the combination of very large body size, relatively short and bicolored tail, laterally bicolored dorsal surface of the hind foot, and the distinctly soft adult pelage covering the entire body, especially along the dorsum where the aristiforms are not stiff to the touch. The relatively short and narrow baculum is indicative of a short and thin phallus in the male, a structure that also easily separates this species from all other sympatric spiny rats throughout western Amazonia. Cranially,
<taxonomicName id="4C37CBE24BACD8B8FE90351FB6D1F997" authorityName="Goldman" authorityYear="1911" box="[295,415,1601,1624]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="248" pageNumber="249" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734BACD8B8FE90351FB677F997" box="[295,313,1601,1624]" italics="true" pageId="248" pageNumber="249">P</emphasis>
.
<emphasis id="B9436C734BACD8B8FEE5351FB6D1F997" box="[338,415,1601,1624]" italics="true" pageId="248" pageNumber="249">steerei</emphasis>
</taxonomicName>
can be distinguished from other sympatric species by the combination of its large size, typically four folds on M2, structure of the incisive foramina, and broad but relatively deep mesopterygoid fossa. Illustrations of these features are given in figures 139 and 140 as well as Patton (1987).
</paragraph>
<paragraph id="8B88B0614BACD8B8FCB73247B4D2F95C" blockId="248.[739,1268,193,1741]" pageId="248" pageNumber="249">
MOLECULAR PHYLOGEOGRAPHY: We have sampled 25 localities that span a substantial portion of the mapped range of the
<taxonomicName id="4C37CBE24BACD8B8FB2E320AB3A4FEA4" authorityName="Thomas" authorityYear="1905" box="[1177,1258,340,363]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="248" pageNumber="249" phylum="Chordata" rank="species" species="goeldii">
<emphasis id="B9436C734BACD8B8FB2E320AB3A4FEA4" box="[1177,1258,340,363]" italics="true" pageId="248" pageNumber="249">goeldii</emphasis>
</taxonomicName>
­ group of species, as diagnosed and mapped by Patton (1987: fig. 2). Included are samples of
<taxonomicName id="4C37CBE24BACD8B8FCB532F2B43EFE0C" authorityName="Goldman" authorityYear="1911" box="[770,880,428,451]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="248" pageNumber="249" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734BACD8B8FCB532F2B45AFE0C" box="[770,788,428,451]" italics="true" pageId="248" pageNumber="249">P</emphasis>
.
<emphasis id="B9436C734BACD8B8FC9432F2B43EFE0C" box="[803,880,428,451]" italics="true" pageId="248" pageNumber="249">steerei</emphasis>
</taxonomicName>
from 14 of the primary localities within the Rio Juruá basin (fig. 153; table 77). Up to five individuals were sequenced per population. Three major haplotype clades are recognizable, one of which is further divisible into two groups (fig. 154). Haplotypes from specimens allocated to
<taxonomicName id="4C37CBE24BACD8B8FB8A3102B3FFFDBC" authorityName="Thomas" authorityYear="1905" box="[1085,1201,604,627]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="248" pageNumber="249" phylum="Chordata" rank="species" species="goeldii">
<emphasis id="B9436C734BACD8B8FB8A3102B301FDBC" box="[1085,1103,604,627]" italics="true" pageId="248" pageNumber="249">P</emphasis>
.
<emphasis id="B9436C734BACD8B8FBD73102B3FFFDBC" box="[1120,1201,604,627]" italics="true" pageId="248" pageNumber="249">goeldii</emphasis>
</taxonomicName>
from two localities in Estado do Pará are so well differentiated from all others (13.2% relative to other members of the group) that it is basal to all other species of Amazonian
<taxonomicName id="4C37CBE24BACD8B8FB3A318FB45EFCC9" authorityName="Allen" authorityYear="1899" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="248" pageNumber="249" phylum="Chordata" rank="genus">
<emphasis id="B9436C734BACD8B8FB3A318FB45EFCC9" italics="true" pageId="248" pageNumber="249">Proechimys</emphasis>
</taxonomicName>
in analyses containing all taxa and geographic representatives (da Silva, 1998: fig 13). The other two clades separate samples from largely south of the Rio Solimões, but including those from the Rio Jaú west and south of the Rio Negro, from those in northern
<collectingCountry id="F320F0F14BACD8B8FCA730C0B40BFC7A" box="[784,837,926,949]" name="Peru" pageId="248" pageNumber="249">Perú</collectingCountry>
, southern
<collectingCountry id="F320F0F14BACD8B8FC7730C0B375FC7A" box="[960,1083,926,949]" name="Venezuela" pageId="248" pageNumber="249">Venezuela</collectingCountry>
, and
<collectingCountry id="F320F0F14BACD8B8FBCF30C0B3F1FC7A" box="[1144,1215,926,949]" name="Brazil" pageId="248" pageNumber="249">Brazil</collectingCountry>
east of the Rio Negro (fig. 153). These differ by an average of 10.9%. Although samples from the northern clade are sparse, sequence divergence is low, averaging only 2.6% across the 2000 kilometers between northern
<collectingCountry id="F320F0F14BACD8B8FB09376FB3BDFB87" box="[1214,1267,1073,1096]" name="Peru" pageId="248" pageNumber="249">Perú</collectingCountry>
and central
<collectingCountry id="F320F0F14BACD8B8FCC83710B488FBAA" box="[895,966,1102,1125]" name="Brazil" pageId="248" pageNumber="249">Brazil</collectingCountry>
north of Manaus. The southern clade, however, does exhibit substantial geographic variation, with the samples from north of the Rio Solimões along the Rio Jaú differing from those to the south by an average of 6.6%. A more extensive analysis of haplotype variation among localities within the Rio Juruá is in preparation for publication elsewhere (M. D. Matocq, J. L. Patton, and M. N. F. da Silva). A synopsis of these data is given below in the section on riverine barriers. However, from the tree in fig. 154 it is apparent that a reasonable degree of haplotype diversity is present along the river, as 801 bp haplotypes from 11 different localities differ by an average of more than 5%. Importantly, those haplotypes from the Headwaters form a monophyletic assemblage distinct from those from the other three sample regions.
</paragraph>
<paragraph id="8B88B0614BACD8B9FCB735C7B372FB14" blockId="248.[739,1268,193,1741]" lastBlockId="249.[743,1271,1161,1741]" lastPageId="249" lastPageNumber="250" pageId="248" pageNumber="249">
We consider the three clades identified in the mtDNA tree (fig. 154) to represent sep­ arate species. Based both on comparisons of specimens of each of these clades to
<typeStatus id="548C0EC34BADD8B9FDCE37F8B7AAFB14" pageId="249" pageNumber="250" type="holotype">holotypes</typeStatus>
of the various named forms that Patton (1987) allocated to his
<taxonomicName id="4C37CBE24BADD8B9FE6637BFB56CFB37" authorityName="Thomas" authorityYear="1905" box="[465,546,1249,1272]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="249" pageNumber="250" phylum="Chordata" rank="species" species="goeldii">
<emphasis id="B9436C734BADD8B9FE6637BFB56CFB37" box="[465,546,1249,1272]" italics="true" pageId="249" pageNumber="250">goeldii</emphasis>
</taxonomicName>
­ group, the eastern mtDNA clade represents
<taxonomicName id="4C37CBE24BADD8B9FDF737A1B61DFAFC" authority="Thomas, 1905" authorityName="Thomas" authorityYear="1905" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="249" pageNumber="250" phylum="Chordata" rank="species" species="goeldii">
<emphasis id="B9436C734BADD8B9FDF737A1B51CFAD9" box="[576,594,1279,1302]" italics="true" pageId="249" pageNumber="250">P</emphasis>
.
<emphasis id="B9436C734BADD8B9FDD137A1B5F9FAD9" box="[614,695,1279,1302]" italics="true" pageId="249" pageNumber="250">goeldii</emphasis>
Thomas, 1905
</taxonomicName>
, although the oldest name for the southern clade is
<taxonomicName id="4C37CBE24BADD8B9FE093667B7ABFAA2" authority="Goldman, 1911" authorityName="Goldman" authorityYear="1911" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="249" pageNumber="250" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734BADD8B9FE093667B69EFA9F" box="[446,464,1337,1360]" italics="true" pageId="249" pageNumber="250">P</emphasis>
.
<emphasis id="B9436C734BADD8B9FE503667B57AFA9F" box="[487,564,1337,1360]" italics="true" pageId="249" pageNumber="250">steerei</emphasis>
Goldman, 1911
</taxonomicName>
. The northern clade is
<taxonomicName id="4C37CBE24BADD8B9FE4B3609B6FCFA44" authority="Hershkovitz, 1948" authorityName="Hershkovitz" authorityYear="1948" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="249" pageNumber="250" phylum="Chordata" rank="species" species="quadruplicatus">
<emphasis id="B9436C734BADD8B9FE4B3609B540FAA1" box="[508,526,1367,1390]" italics="true" pageId="249" pageNumber="250">P</emphasis>
.
<emphasis id="B9436C734BADD8B9FDA83609B787FA44" italics="true" pageId="249" pageNumber="250">quadruplicatus</emphasis>
Hershkovitz, 1948
</taxonomicName>
, a name published in the same year but somewhat earlier than its junior synonym,
<taxonomicName id="4C37CBE24BADD8B9FEC036F1B7ABFA2C" authority="Moojen, 1948" authorityName="Moojen" authorityYear="1948" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="249" pageNumber="250" phylum="Chordata" rank="species" species="amphichoricus">
<emphasis id="B9436C734BADD8B9FEC036F1B6C7FA09" box="[375,393,1455,1478]" italics="true" pageId="249" pageNumber="250">P</emphasis>
.
<emphasis id="B9436C734BADD8B9FE2A36F1B504FA09" box="[413,586,1455,1478]" italics="true" pageId="249" pageNumber="250">amphichoricus</emphasis>
Moojen, 1948
</taxonomicName>
. Thorough analyses of morphological character variation for this group have yet to be performed, but the data summarized by Patton (1987) suggest that each clade is diagnosable by morphological traits as well as cytochrome­b sequences. For example, most individuals of
<taxonomicName id="4C37CBE24BADD8B9FEE13522B564F95C" authorityName="Hershkovitz" authorityYear="1948" box="[342,554,1660,1683]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="249" pageNumber="250" phylum="Chordata" rank="species" species="quadruplicatus">
<emphasis id="B9436C734BADD8B9FEE13522B626F95C" box="[342,360,1660,1683]" italics="true" pageId="249" pageNumber="250">P</emphasis>
.
<emphasis id="B9436C734BADD8B9FECD3522B564F95C" box="[378,554,1660,1683]" italics="true" pageId="249" pageNumber="250">quadruplicatus</emphasis>
</taxonomicName>
(more than 60%) possess four folds on all four upper cheekteeth, while this character state is much less frequent in
<taxonomicName id="4C37CBE24BADD8B9FC1C37D7B353FB6F" authorityName="Goldman" authorityYear="1911" box="[939,1053,1161,1184]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="249" pageNumber="250" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734BADD8B9FC1C37D7B4F3FB6F" box="[939,957,1161,1184]" italics="true" pageId="249" pageNumber="250">P</emphasis>
.
<emphasis id="B9436C734BADD8B9FC6737D7B353FB6F" box="[976,1053,1161,1184]" italics="true" pageId="249" pageNumber="250">steerei</emphasis>
</taxonomicName>
, and few individuals of
<taxonomicName id="4C37CBE24BADD8B9FCFC37F9B48AFB71" authorityName="Thomas" authorityYear="1905" box="[843,964,1190,1214]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="249" pageNumber="250" phylum="Chordata" rank="species" species="goeldii">
<emphasis id="B9436C734BADD8B9FCFC37F9B413FB71" box="[843,861,1191,1214]" italics="true" pageId="249" pageNumber="250">P</emphasis>
.
<emphasis id="B9436C734BADD8B9FCC437F9B48AFB71" box="[883,964,1191,1214]" italics="true" pageId="249" pageNumber="250">goeldii</emphasis>
</taxonomicName>
have four folds on any teeth (Patton, 1987: table 5).
</paragraph>
<caption id="DF48E0E94BADD8B9FF76308DB426FB9D" ID-DOI="http://doi.org/10.5281/zenodo.5347640" ID-Zenodo-Dep="5347640" httpUri="https://zenodo.org/record/5347640/files/figure.png" pageId="249" pageNumber="250" startId="249.[193,231,979,1000]" targetBox="[217,1218,188,956]" targetPageId="249">
<paragraph id="8B88B0614BADD8B9FF76308DB426FB9D" blockId="249.[167,1271,979,1107]" pageId="249" pageNumber="250">
Fig. 153. Map of the distribution of three species of the
<taxonomicName id="4C37CBE24BADD8B9FC8E308DB348FC27" authorityName="Thomas" authorityYear="1905" box="[825,1030,979,1000]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="249" pageNumber="250" phylum="Chordata" rank="species" species="goeldii">
<emphasis id="B9436C734BADD8B9FC8E308DB348FC27" box="[825,1030,979,1000]" italics="true" pageId="249" pageNumber="250">Proechimys goeldii</emphasis>
</taxonomicName>
group (modified from Patton, 1987) illustrating localities for which 801 bp of cytochrome­b sequence are available, lettered or numbered as in the tree (fig. 154). Localities from which individual specimens have been examined are identified by number (Rio Jurua´) or letter, and are listed in table 77. Solid triangles =
<taxonomicName id="4C37CBE24BADD8B9FB34377DB7B4FB9C" authorityName="Hershkovitz" authorityYear="1948" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="249" pageNumber="250" phylum="Chordata" rank="species" species="quadruplicatus">
<emphasis id="B9436C734BADD8B9FB34377DB3DDFBF7" box="[1155,1171,1059,1080]" italics="true" pageId="249" pageNumber="250">P</emphasis>
.
<emphasis id="B9436C734BADD8B9FB14377DB7B4FB9C" italics="true" pageId="249" pageNumber="250">quadruplicatus</emphasis>
</taxonomicName>
; solid circles =
<taxonomicName id="4C37CBE24BADD8B9FE183760B55BFB9C" authorityName="Goldman" authorityYear="1911" box="[431,533,1085,1107]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="249" pageNumber="250" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734BADD8B9FE183760B6F1FB9C" box="[431,447,1086,1107]" italics="true" pageId="249" pageNumber="250">P</emphasis>
.
<emphasis id="B9436C734BADD8B9FE783760B55BFB9C" box="[463,533,1086,1107]" italics="true" pageId="249" pageNumber="250">steerei</emphasis>
</taxonomicName>
; and open circles =
<taxonomicName id="4C37CBE24BADD8B9FD4E3760B42CFB9C" authorityName="Thomas" authorityYear="1905" box="[761,866,1085,1107]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="249" pageNumber="250" phylum="Chordata" rank="species" species="goeldii">
<emphasis id="B9436C734BADD8B9FD4E3760B447FB9C" box="[761,777,1086,1107]" italics="true" pageId="249" pageNumber="250">P</emphasis>
.
<emphasis id="B9436C734BADD8B9FCAE3760B42CFB9C" box="[793,866,1086,1107]" italics="true" pageId="249" pageNumber="250">goeldii</emphasis>
</taxonomicName>
.
</paragraph>
</caption>
<paragraph id="8B88B0614BADD8B9FCB337BFB3B9F902" blockId="249.[743,1271,1161,1741]" pageId="249" pageNumber="250">
MORPHOMETRIC VARIATION: As with
<taxonomicName id="4C37CBE24BADD8B9FB0037BFB462FAD9" authorityName="Thomas" authorityYear="1900" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="249" pageNumber="250" phylum="Chordata" rank="species" species="simonsi">
<emphasis id="B9436C734BADD8B9FB0037BFB387FB37" box="[1207,1225,1249,1272]" italics="true" pageId="249" pageNumber="250">P</emphasis>
.
<emphasis id="B9436C734BADD8B9FB6D37BFB462FAD9" italics="true" pageId="249" pageNumber="250">simonsi</emphasis>
</taxonomicName>
, above, we summarize the variation in mensural characters for adults (toothwear classes 8, 9, and 10) of
<taxonomicName id="4C37CBE24BADD8B9FBB43667B33DFA9F" authorityName="Goldman" authorityYear="1911" box="[1027,1139,1337,1360]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="249" pageNumber="250" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734BADD8B9FBB43667B35BFA9F" box="[1027,1045,1337,1360]" italics="true" pageId="249" pageNumber="250">P</emphasis>
.
<emphasis id="B9436C734BADD8B9FB913667B33DFA9F" box="[1062,1139,1337,1360]" italics="true" pageId="249" pageNumber="250">steerei</emphasis>
</taxonomicName>
that is due to locality, sex, and age effects by a nested ANOVA (table 78). No character exhibits significant interlocality differences, and the average contribution of this factor to character variation is only 5.6%. However, considerable differences due to both sexual dimorphism and age variation are apparent More than two thirds of the variables exhibit significant variant components for sex and age, with an average of 23.3% of the total pool of variation due to sex and 20.6% due to age. Hence, the degree of sexual dimorphism and the extent to which dimensions
</paragraph>
<paragraph id="8B88B0614BAED8BAFF143608B343FA0A" blockId="250.[163,692,1366,1742]" lastBlockId="250.[739,1267,1366,1741]" pageId="250" pageNumber="251">
continue to increase in older toothwear classes even in fully adult individuals is somewhat greater in this species than it is in
<taxonomicName id="4C37CBE24BAED8BAFD2D36CFB7B2FA09" authorityName="Thomas" authorityYear="1900" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="250" pageNumber="251" phylum="Chordata" rank="species" species="simonsi">
<emphasis id="B9436C734BAED8BAFD2D36CFB5E2FA67" box="[666,684,1425,1448]" italics="true" pageId="250" pageNumber="251">P</emphasis>
.
<emphasis id="B9436C734BAED8BAFF1436F1B7B2FA09" box="[163,252,1455,1478]" italics="true" pageId="250" pageNumber="251">simonsi</emphasis>
</taxonomicName>
, although the differences in patterns between the two species is not great (compare tables 72 and 78). When an analysis nested by sex and age is applied to our largest sample from Nova Empresa (locality
<emphasis id="B9436C734BAED8BAFD31357DB5DBF9F4" bold="true" box="[646,661,1571,1595]" pageId="250" pageNumber="251">8</emphasis>
, n = 41), seven cranial variables exhibit significant sexual dimorphism (
<emphasis id="B9436C734BAED8BAFE5A3501B6B2F9B9" box="[493,508,1631,1654]" italics="true" pageId="250" pageNumber="251">p</emphasis>
&lt;0.05: CIL, MB, RL, MPFW;
<emphasis id="B9436C734BAED8BAFECC3522B6C4F95C" box="[379,394,1660,1683]" italics="true" pageId="250" pageNumber="251">p</emphasis>
&lt;0.01: NL, D, and PL) and six show significant age differences (
<emphasis id="B9436C734BAED8BAFD1335C7B5FDF97F" box="[676,691,1689,1712]" italics="true" pageId="250" pageNumber="251">p</emphasis>
&lt;0.05: MB, RL, NL, MPFW;
<emphasis id="B9436C734BAED8BAFDAD35E9B567F901" box="[538,553,1719,1742]" italics="true" pageId="250" pageNumber="251">p</emphasis>
&lt;0.01: D, PL). Again, within­locality variance is partitioned somewhat differently than it is in
<taxonomicName id="4C37CBE24BAED8BAFB6D362AB472FA67" authorityName="Thomas" authorityYear="1900" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="250" pageNumber="251" phylum="Chordata" rank="species" species="simonsi">
<emphasis id="B9436C734BAED8BAFB6D362AB3A2FA44" box="[1242,1260,1396,1419]" italics="true" pageId="250" pageNumber="251">P</emphasis>
.
<emphasis id="B9436C734BAED8BAFD5436CFB472FA67" box="[739,828,1425,1448]" italics="true" pageId="250" pageNumber="251">simonsi</emphasis>
</taxonomicName>
, particularly in the degree of character sexual dimorphism.
</paragraph>
<paragraph id="8B88B0614BAED8BBFCB73692B477FCCA" blockId="250.[739,1267,1366,1741]" lastBlockId="251.[743,1272,193,1741]" lastPageId="251" lastPageNumber="252" pageId="250" pageNumber="251">
The small amount of morphometric variation attributable to interlocality differences contrasts somewhat with the mtDNA haplotype data, which indicates both a reasonable amount of sequence differentiation among haplotypes (over 5%) and some geographic structuring into two reciprocally monophyletic clades along the river. In order to examine geographic trends in morphology more thoroughly, we used both principal components and discriminant function analyses, and compared samples from the four sample regions as well as those from the left and right banks of the river. Despite negligible interlocality variation in univariate di­ mensions, limited regional effects are apparent. In a principal components analysis, samples from each of the four geographic regional sample areas overlap extensively in multivariate space in combinations of the first three axes, which combine to explain 76.1% of the total pool of variation (fig. 155 top). Nevertheless, there are significant differences between the regions in mean PC scores on each axis (PC­1:
<emphasis id="B9436C734BAFD8BBFB853297B30AFE2F" box="[1074,1092,457,480]" italics="true" pageId="251" pageNumber="252">F</emphasis>
<subScript id="17B3B2244BAFD8BBFBF4328BB324FE2C" attach="left" box="[1091,1130,469,483]" fontSize="6" pageId="251" pageNumber="252">3,161</subScript>
= 7.086,
<emphasis id="B9436C734BAFD8BBFB5F3297B3B9FE2F" box="[1256,1271,457,480]" italics="true" pageId="251" pageNumber="252">p</emphasis>
= 0.0002; PC­2:
<emphasis id="B9436C734BAFD8BBFC0232B9B489FE31" box="[949,967,487,510]" italics="true" pageId="251" pageNumber="252">F</emphasis>
<subScript id="17B3B2244BAFD8BBFC7132ADB4A3FDCE" attach="left" box="[966,1005,499,513]" fontSize="6" pageId="251" pageNumber="252">3,161</subScript>
= 9.111,
<emphasis id="B9436C734BAFD8BBFBDF32B9B339FE31" box="[1128,1143,487,510]" italics="true" pageId="251" pageNumber="252">p</emphasis>
= 0.0001 PC­3:
<emphasis id="B9436C734BAFD8BBFC8E315AB405FDD4" box="[825,843,516,539]" italics="true" pageId="251" pageNumber="252">F</emphasis>
<subScript id="17B3B2244BAFD8BBFCFC314EB43CFDD1" attach="left" box="[843,882,528,542]" fontSize="6" pageId="251" pageNumber="252">3,161</subScript>
= 15.410,
<emphasis id="B9436C734BAFD8BBFC4B315AB345FDD4" box="[1020,1035,516,539]" italics="true" pageId="251" pageNumber="252">p</emphasis>
= 0.0001, respectively). The first PC axis represents a general size axis, as indicated by high and positive factor coefficients (table 79) and by positive correlations of individual scores with their respective mensural variables. The correlation between PC­1 scores and individual values for logCIL, for example, is 0.976 (
<emphasis id="B9436C734BAFD8BBFB71318FB39BFD27" box="[1222,1237,721,744]" italics="true" pageId="251" pageNumber="252">p</emphasis>
&lt;0.001).
</paragraph>
<paragraph id="8B88B0614BAFD8BBFCB33052B47BFB72" blockId="251.[743,1272,193,1741]" pageId="251" pageNumber="252">
The trend in overall size in
<taxonomicName id="4C37CBE24BAFD8BBFBF43052B3FEFCEC" authorityName="Goldman" authorityYear="1911" box="[1091,1200,780,803]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="251" pageNumber="252" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734BAFD8BBFBF43052B31BFCEC" box="[1091,1109,780,803]" italics="true" pageId="251" pageNumber="252">P</emphasis>
.
<emphasis id="B9436C734BAFD8BBFBD43052B3FEFCEC" box="[1123,1200,780,803]" italics="true" pageId="251" pageNumber="252">steerei</emphasis>
</taxonomicName>
along the Rio Juruá from its headwaters to its mouth is more complex then in
<taxonomicName id="4C37CBE24BAFD8BBFBC43019B3BEFC91" authorityName="Thomas" authorityYear="1900" box="[1139,1264,838,862]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="251" pageNumber="252" phylum="Chordata" rank="species" species="simonsi">
<emphasis id="B9436C734BAFD8BBFBC43019B3CBFC91" box="[1139,1157,839,862]" italics="true" pageId="251" pageNumber="252">P</emphasis>
.
<emphasis id="B9436C734BAFD8BBFB203019B3BEFC91" box="[1175,1264,839,862]" italics="true" pageId="251" pageNumber="252">simonsi</emphasis>
</taxonomicName>
with parallel increases in size from the Headwaters Region to the Upper Central Region and from the Lower Central Region to the Mouth Region (fig. 156). The differences between samples from the Upper and Lower Central regions are significant, as are those between the Lower Central and Mouth (Duncans multiple range critical differences = 0.479 and 0.495, respectively, both
<emphasis id="B9436C734BAFD8BBFB083711B380FBA9" box="[1215,1230,1103,1126]" italics="true" pageId="251" pageNumber="252">p</emphasis>
&lt;0.05). This complex pattern parallels interlocality variation in karyotype, as discussed below.
</paragraph>
<caption id="DF48E0E94BAFD8BBFF76372DB603FA2D" ID-DOI="http://doi.org/10.5281/zenodo.5347642" ID-Zenodo-Dep="5347642" httpUri="https://zenodo.org/record/5347642/files/figure.png" pageId="251" pageNumber="252" startId="251.[193,231,1139,1160]" targetBox="[174,687,188,1117]" targetPageId="251">
<paragraph id="8B88B0614BAFD8BBFF76372DB603FA2D" blockId="251.[167,695,1139,1506]" pageId="251" pageNumber="252">
Fig. 154. Strict consensus maximum parsimony tree of 27 equally minimal length trees for 30 individual haplotypes of species of the
<taxonomicName id="4C37CBE24BAFD8BBFDCC37F6B60AFB17" authorityName="Thomas" authorityYear="1905" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="251" pageNumber="252" phylum="Chordata" rank="species" species="goeldii">
<emphasis id="B9436C734BAFD8BBFDCC37F6B60AFB17" italics="true" pageId="251" pageNumber="252">Proechimys goeldii</emphasis>
</taxonomicName>
­group from 23 localities in Peru´, Venezuela, and Brazil, as identified in the map (fig. 153). Length = 509 steps; CI = 0.672; RI = 0.814. Sequences of other species of
<taxonomicName id="4C37CBE24BAFD8BBFD8B364DB5F9FAE7" authorityName="Allen" authorityYear="1899" box="[572,695,1299,1320]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="251" pageNumber="252" phylum="Chordata" rank="genus">
<emphasis id="B9436C734BAFD8BBFD8B364DB5F9FAE7" box="[572,695,1299,1320]" italics="true" pageId="251" pageNumber="252">Proechimys</emphasis>
</taxonomicName>
and of
<taxonomicName id="4C37CBE24BAFD8BBFF433670B61BFA8C" authorityName="Wagner" authorityYear="1845" box="[244,341,1326,1347]" class="Mammalia" family="Echimyidae" genus="Mesomys" kingdom="Animalia" order="Rodentia" pageId="251" pageNumber="252" phylum="Chordata" rank="genus">
<emphasis id="B9436C734BAFD8BBFF433670B61BFA8C" box="[244,341,1326,1347]" italics="true" pageId="251" pageNumber="252">Mesomys</emphasis>
</taxonomicName>
were used to root the tree. Bold numbers at internal nodes are bootstrap values, based on 1000 replicates; percentages are average Kimura two­parameter distance. Provenance data and catalog numbers for each specimen can be found in table 77, listed in order from top to bottom in the tree.
</paragraph>
</caption>
<paragraph id="8B88B0614BAFD8BBFCB3379AB3B9F902" blockId="251.[743,1272,193,1741]" pageId="251" pageNumber="252">
As was the case for
<taxonomicName id="4C37CBE24BAFD8BBFBB5379AB331FB14" authorityName="Thomas" authorityYear="1900" box="[1026,1151,1220,1243]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="251" pageNumber="252" phylum="Chordata" rank="species" species="simonsi">
<emphasis id="B9436C734BAFD8BBFBB5379AB35AFB14" box="[1026,1044,1220,1243]" italics="true" pageId="251" pageNumber="252">P</emphasis>
.
<emphasis id="B9436C734BAFD8BBFB91379AB331FB14" box="[1062,1151,1220,1243]" italics="true" pageId="251" pageNumber="252">simonsi</emphasis>
</taxonomicName>
, although regional samples of
<taxonomicName id="4C37CBE24BAFD8BBFC6B37BFB302FB37" authorityName="Goldman" authorityYear="1911" box="[988,1100,1249,1272]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="251" pageNumber="252" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734BAFD8BBFC6B37BFB4A0FB37" box="[988,1006,1249,1272]" italics="true" pageId="251" pageNumber="252">P</emphasis>
.
<emphasis id="B9436C734BAFD8BBFC4837BFB302FB37" box="[1023,1100,1249,1272]" italics="true" pageId="251" pageNumber="252">steerei</emphasis>
</taxonomicName>
exhibit a pattern of size increase along the length of the Rio Jurua´, there is no differentiation between opposite­bank samples when characters are examined either in a univariate fashion or by multivariate principal components analysis For example, there is broad overlap in the bivariate plot of PC­1 and PC­2 scores for samples pooled by right and left bank localities (fig. 155, bottom), and one­way ANO­ VAs for PC scores were nonsignificant for each of the three axes of table 78: for PC­1
<emphasis id="B9436C734BAFD8BBFD50351FB5B7F997" box="[743,761,1601,1624]" italics="true" pageId="251" pageNumber="252">F</emphasis>
<subScript id="17B3B2244BAFD8BBFD4F3513B451F994" attach="left" box="[760,799,1613,1627]" fontSize="6" pageId="251" pageNumber="252">1,163</subScript>
= 1.943,
<emphasis id="B9436C734BAFD8BBFC26351FB4EEF997" box="[913,928,1601,1624]" italics="true" pageId="251" pageNumber="252">p</emphasis>
= 0.1652; for PC­2,
<emphasis id="B9436C734BAFD8BBFB14351FB3FBF997" box="[1187,1205,1601,1624]" italics="true" pageId="251" pageNumber="252">F</emphasis>
<subScript id="17B3B2244BAFD8BBFB033513B395F994" attach="left" box="[1204,1243,1613,1627]" fontSize="6" pageId="251" pageNumber="252">1,163</subScript>
= 0.133,
<emphasis id="B9436C734BAFD8BBFC8A3501B402F9B9" box="[829,844,1631,1654]" italics="true" pageId="251" pageNumber="252">p</emphasis>
= 0.7161; and for PC­3,
<emphasis id="B9436C734BAFD8BBFB293501B3FEF9B9" box="[1182,1200,1631,1654]" italics="true" pageId="251" pageNumber="252">F</emphasis>
<subScript id="17B3B2244BAFD8BBFB183535B398F9B6" attach="left" box="[1199,1238,1643,1657]" fontSize="6" pageId="251" pageNumber="252">1,163</subScript>
= 0.648,
<emphasis id="B9436C734BAFD8BBFC803522B408F95C" box="[823,838,1660,1683]" italics="true" pageId="251" pageNumber="252">p</emphasis>
= 0.4221. However, also as was true for
<taxonomicName id="4C37CBE24BAFD8BBFCA635C7B4C5F97F" authorityName="Thomas" authorityYear="1900" box="[785,907,1689,1712]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="251" pageNumber="252" phylum="Chordata" rank="species" species="simonsi">
<emphasis id="B9436C734BAFD8BBFCA635C7B46DF97F" box="[785,803,1689,1712]" italics="true" pageId="251" pageNumber="252">P</emphasis>
.
<emphasis id="B9436C734BAFD8BBFC8535C7B4C5F97F" box="[818,907,1689,1712]" italics="true" pageId="251" pageNumber="252">simonsi</emphasis>
</taxonomicName>
, if samples are pooled by river bank and subjected to a discriminant function
</paragraph>
<paragraph id="8B88B0614BA8D8BCFF1437D7B797FA67" blockId="252.[163,692,1161,1741]" pageId="252" pageNumber="253">
analysis (table 74), which maximizes between­group variance while minimizing that within groups, some segregation of oppositebank populations is apparent. A one­way ANOVA of individual scores on the first discriminant axis yields a significant river bank effect (
<emphasis id="B9436C734BA8D8BCFF403667B647FA9F" box="[247,265,1337,1360]" italics="true" pageId="252" pageNumber="253">F</emphasis>
<subScript id="17B3B2244BA8D8BCFEBE361BB67EFA9C" attach="left" box="[265,304,1349,1363]" fontSize="6" pageId="252" pageNumber="253">1,163</subScript>
= 20.660,
<emphasis id="B9436C734BA8D8BCFE013667B68BFA9F" box="[438,453,1337,1360]" italics="true" pageId="252" pageNumber="253">p</emphasis>
&lt;0.0001), and histograms of these scores (fig. 151, right) illustrate the slight differences among the samples.
</paragraph>
<paragraph id="8B88B0614BA8D8BCFF7736F0B344F97F" blockId="252.[163,692,1161,1741]" lastBlockId="252.[739,1268,1161,1741]" pageId="252" pageNumber="253">
We also examined the relationship between the morphometric and genetic distances among our samples of
<taxonomicName id="4C37CBE24BA8D8BCFE6B36B7B502F9CF" authorityName="Goldman" authorityYear="1911" box="[476,588,1513,1536]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="252" pageNumber="253" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734BA8D8BCFE6B36B7B6A0F9CF" box="[476,494,1513,1536]" italics="true" pageId="252" pageNumber="253">P</emphasis>
.
<emphasis id="B9436C734BA8D8BCFE4836B7B502F9CF" box="[511,588,1513,1536]" italics="true" pageId="252" pageNumber="253">steerei</emphasis>
</taxonomicName>
, as well as that between each of these variables and geographic distance. We used the Mahalanobis D
<superScript id="7C421D294BA8D8BCFF44351EB7B2F981" attach="left" box="[243,252,1600,1614]" fontSize="6" pageId="252" pageNumber="253">2</superScript>
matrix generated from the discriminant function analysis that specified localities as the a priori groups as a measure of morphometric distance, and a matrix of genetic similarities (Slatkins [1993] M­statistic) generated from the population cytochrome­b haplotypes by the AMOVA program of Excoffier et al. (1992). These were compared to the log
<subScript id="17B3B2244BA8D8BCFC8137B3B409FB34" attach="left" box="[822,839,1261,1275]" fontSize="6" pageId="252" pageNumber="253">10</subScript>
of the straight­line geographic distances among localities given in table 1. Morphometric distances increase significantly with an increase in geographic distance among locality pairs (fig. 157; Mantels matrix correlation coefficient
<emphasis id="B9436C734BA8D8BCFB9A362AB376FA44" box="[1069,1080,1396,1419]" italics="true" pageId="252" pageNumber="253">r</emphasis>
= 0.460,
<emphasis id="B9436C734BA8D8BCFB77362AB381FA44" box="[1216,1231,1396,1419]" italics="true" pageId="252" pageNumber="253">p</emphasis>
= 0.0015), and genetic similarity decreases sharply with geography (fig. 157;
<emphasis id="B9436C734BA8D8BCFB0836F1B384FA09" box="[1215,1226,1455,1478]" italics="true" pageId="252" pageNumber="253">r</emphasis>
= ‾0.804,
<emphasis id="B9436C734BA8D8BCFCFE3692B416FA2C" box="[841,856,1484,1507]" italics="true" pageId="252" pageNumber="253">p</emphasis>
&lt;0.0001). Not surprisingly, therefore, there is a significant, if weak, correlation between genetic similarity and morphometric distance (
<emphasis id="B9436C734BA8D8BCFC18357AB4F4F9F4" box="[943,954,1572,1595]" italics="true" pageId="252" pageNumber="253">r</emphasis>
= ‾0.275,
<emphasis id="B9436C734BA8D8BCFBE3357AB32DF9F4" box="[1108,1123,1572,1595]" italics="true" pageId="252" pageNumber="253">p</emphasis>
= 0.0192) Populations of
<taxonomicName id="4C37CBE24BA8D8BCFC24351FB34FF997" authorityName="Goldman" authorityYear="1911" box="[915,1025,1601,1624]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="252" pageNumber="253" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734BA8D8BCFC24351FB4EBF997" box="[915,933,1601,1624]" italics="true" pageId="252" pageNumber="253">P</emphasis>
.
<emphasis id="B9436C734BA8D8BCFC03351FB34FF997" box="[948,1025,1601,1624]" italics="true" pageId="252" pageNumber="253">steerei</emphasis>
</taxonomicName>
along the Rio Jurua´ as is true of
<taxonomicName id="4C37CBE24BA8D8BCFCCD3501B4B8F9B9" authorityName="Thomas" authorityYear="1900" box="[890,1014,1630,1654]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="252" pageNumber="253" phylum="Chordata" rank="species" species="simonsi">
<emphasis id="B9436C734BA8D8BCFCCD3501B4C2F9B9" box="[890,908,1631,1654]" italics="true" pageId="252" pageNumber="253">P</emphasis>
.
<emphasis id="B9436C734BA8D8BCFC2A3501B4B8F9B9" box="[925,1014,1631,1654]" italics="true" pageId="252" pageNumber="253">simonsi</emphasis>
</taxonomicName>
, exhibit a clinal, isolation­by­distance pattern in both morphometric and genetic traits.
</paragraph>
<paragraph id="8B88B0614BA8D8BDFCB735E8B460F902" blockId="252.[739,1268,1161,1741]" lastBlockId="253.[743,1271,1102,1741]" lastPageId="253" lastPageNumber="254" pageId="252" pageNumber="253">
REPRODUCTION: We obtained specimens of
<taxonomicName id="4C37CBE24BA9D8BDFF103711B655FBA9" authorityName="Goldman" authorityYear="1911" box="[167,283,1102,1126]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="253" pageNumber="254" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734BA9D8BDFF103711B7F7FBA9" box="[167,185,1103,1126]" italics="true" pageId="253" pageNumber="254">P</emphasis>
.
<emphasis id="B9436C734BA9D8BDFF793711B655FBA9" box="[206,283,1103,1126]" italics="true" pageId="253" pageNumber="254">steerei</emphasis>
</taxonomicName>
along the entire river and at all seasons during the year of our sampling. Of the
<specimenCount id="9D317BE84BA9D8BDFF6237D7B601FB6F" box="[213,335,1161,1184]" pageId="253" pageNumber="254" type="male">183 males</specimenCount>
we autopsied, 88 were reproductively active. These ranged in age from toothwear class 3 to 10, although 82% (68 of 83) were adults (age classes 8 to 10) and 17% (14 of 83) were subadults (age classes 6 and 7). Reproductively inactive males include both young and adult individuals (age classes 1 to 9), with 68% (59 or 87) individuals of age classes 3, 5, and 6, and 9% (8 of 87) full adults (age classes 8 and 9). We have reproductive data for
<specimenCount id="9D317BE84BA9D8BDFE1036F0B57BFA0A" box="[423,565,1454,1477]" pageId="253" pageNumber="254" type="female">179 females</specimenCount>
(table 75). Off these, 65% showed signs of current or previous pregnancy whereas 35% apparently had not yet reproduced. We caught pregnant, lactating, or postpartum females at all sites indicating that at least some females are breeding in all seasons of the year. Of the parous females, 83% (87 of 105) were adults, 16% subadults, and 1% young individuals. Seventy­five females were pregnant. The age class of these individuals ranged from 6 to 10; 87% (65 of 75) were fully adult (age classes 8 to 10) and the remaining subadults Seventeen percent (14 of 84) were both pregnant and lactating. Modal litter size is 3 range, 17. The great majority of nulliparous females are young (47 of 58) or subadults (10 of 58). Relative to other species of spiny rats, particularly
<taxonomicName id="4C37CBE24BA9D8BDFC0C3667B372FA9F" authorityName="Thomas" authorityYear="1900" box="[955,1084,1337,1360]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="253" pageNumber="254" phylum="Chordata" rank="species" species="simonsi">
<emphasis id="B9436C734BA9D8BDFC0C3667B483FA9F" box="[955,973,1337,1360]" italics="true" pageId="253" pageNumber="254">P</emphasis>
.
<emphasis id="B9436C734BA9D8BDFC543667B372FA9F" box="[995,1084,1337,1360]" italics="true" pageId="253" pageNumber="254">simonsi</emphasis>
</taxonomicName>
, which is also distributed along the entire river and for which we also have large samples,
<taxonomicName id="4C37CBE24BA9D8BDFB3E362AB3B9FA44" authorityName="Goldman" authorityYear="1911" box="[1161,1271,1396,1419]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="253" pageNumber="254" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734BA9D8BDFB3E362AB3D5FA44" box="[1161,1179,1396,1419]" italics="true" pageId="253" pageNumber="254">P</emphasis>
.
<emphasis id="B9436C734BA9D8BDFB1D362AB3B9FA44" box="[1194,1271,1396,1419]" italics="true" pageId="253" pageNumber="254">steerei</emphasis>
</taxonomicName>
exhibits reproductive characteristics tending towards a more
<emphasis id="B9436C734BA9D8BDFC1836F1B4F4FA09" box="[943,954,1455,1478]" italics="true" pageId="253" pageNumber="254">r</emphasis>
­selected life history, with somewhat earlier reproductive maturity, larg­ er litter sizes, larger percentage of young animals breeding, a larger percentage of postpartum estrous females, and a smaller proportion of nonbreeding adults (table 75) These features might be expected for a species that lives primarily in strongly seasonal habitats, such as the várzea forests of the Rio Jurua´.
</paragraph>
</subSubSection>
<caption id="DF48E0E94BA9D8BDFF7630C6B6EDFBD7" ID-DOI="http://doi.org/10.5281/zenodo.5347644" ID-Zenodo-Dep="5347644" httpUri="https://zenodo.org/record/5347644/files/figure.png" pageId="253" pageNumber="254" startId="253.[193,231,920,941]" targetBox="[220,1218,188,898]" targetPageId="253">
<paragraph id="8B88B0614BA9D8BDFF7630C6B6EDFBD7" blockId="253.[167,1271,920,1048]" pageId="253" pageNumber="254">
Fig. 155. Bivariate plots of the first and second principal components axes (
<emphasis id="B9436C734BA9D8BDFBA430C6B325FC62" bold="true" box="[1043,1131,920,941]" pageId="253" pageNumber="254">top, left</emphasis>
) and second and third principal components axes (
<emphasis id="B9436C734BA9D8BDFDFF30ECB5FBFC08" bold="true" box="[584,693,946,967]" pageId="253" pageNumber="254">top, right</emphasis>
) illustrating the morphometric relationships between samples of
<taxonomicName id="4C37CBE24BA9D8BDFE913090B6BCFC2C" authorityName="Goldman" authorityYear="1911" box="[294,498,974,995]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="253" pageNumber="254" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734BA9D8BDFE913090B6BCFC2C" box="[294,498,974,995]" italics="true" pageId="253" pageNumber="254">Proechimys steerei</emphasis>
</taxonomicName>
by geographic regions along the Rio Jurua´. (
<emphasis id="B9436C734BA9D8BDFC5E3093B372FC2D" bold="true" box="[1001,1084,973,994]" pageId="253" pageNumber="254">Bottom</emphasis>
) Plot of the first and second principal components axes in comparisons between samples of
<taxonomicName id="4C37CBE24BA9D8BDFC6430B6B376FC32" authorityName="Goldman" authorityYear="1911" box="[979,1080,1000,1021]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="253" pageNumber="254" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734BA9D8BDFC6430B6B4ADFC32" box="[979,995,1000,1021]" italics="true" pageId="253" pageNumber="254">P</emphasis>
.
<emphasis id="B9436C734BA9D8BDFC4530B6B376FC32" box="[1010,1080,1000,1021]" italics="true" pageId="253" pageNumber="254">steerei</emphasis>
</taxonomicName>
on opposite river banks of the Rio Jurua´.
</paragraph>
</caption>
<caption id="DF48E0E94BAAD8BEFD4A309BB4D8FB5A" ID-DOI="http://doi.org/10.5281/zenodo.5347646" ID-Zenodo-Dep="5347646" httpUri="https://zenodo.org/record/5347646/files/figure.png" pageId="254" pageNumber="255" startId="254.[765,803,965,986]" targetBox="[165,1267,188,947]" targetPageId="254">
<paragraph id="8B88B0614BAAD8BEFD4A309BB4D8FB5A" blockId="254.[739,1267,965,1173]" pageId="254" pageNumber="255">
Fig. 157. Bivariate relationship between morphometric distance (Mahalanobis D
<superScript id="7C421D294BAAD8BEFBD43081B325FC23" attach="left" box="[1123,1131,991,1004]" fontSize="5" pageId="254" pageNumber="255">2</superScript>
; above) and genetic similarity (Slatkins [1993] M­statistic below) and the log of the straight­line geographic distance between sample localities of
<taxonomicName id="4C37CBE24BAAD8BEFBCF376EB467FBAF" authorityName="Goldman" authorityYear="1911" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="254" pageNumber="255" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734BAAD8BEFBCF376EB467FBAF" italics="true" pageId="254" pageNumber="255">Proechimys steerei</emphasis>
</taxonomicName>
along the Rio Jurua´. Mantels matrix correlation coefficients and their significance are indicated for each.
</paragraph>
</caption>
<subSubSection id="C32DE3EA4BAAD941FF7730A8B65DFB14" lastPageId="257" lastPageNumber="258" pageId="254" pageNumber="255" type="distribution">
<paragraph id="8B88B0614BAAD8BEFF7730A8B5FDFBAA" blockId="254.[163,691,1014,1125]" pageId="254" pageNumber="255">
DISTRIBUTION AND HABITAT: Although distributional limits of
<taxonomicName id="4C37CBE24BAAD8BEFE23374AB54DFBE4" authorityName="Goldman" authorityYear="1911" box="[404,515,1044,1067]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="254" pageNumber="255" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734BAAD8BEFE23374AB6E8FBE4" box="[404,422,1044,1067]" italics="true" pageId="254" pageNumber="255">P</emphasis>
.
<emphasis id="B9436C734BAAD8BEFE01374AB54DFBE4" box="[438,515,1044,1067]" italics="true" pageId="254" pageNumber="255">steerei</emphasis>
</taxonomicName>
remain poorly understood, we have identified this species from localities from central
<collectingCountry id="F320F0F14BAAD8BEFE5B3710B56FFBAA" box="[492,545,1102,1125]" name="Peru" pageId="254" pageNumber="255">Perú</collectingCountry>
south of the
</paragraph>
<paragraph id="8B88B0614BAAD8BFFD54379AB601FAFC" blockId="254.[739,1268,1220,1741]" lastBlockId="255.[167,696,1190,1741]" lastPageId="255" lastPageNumber="256" pageId="254" pageNumber="255">
Río Marañón to northern
<collectingCountry id="F320F0F14BAAD8BEFB9E379AB331FB14" box="[1065,1151,1220,1243]" name="Bolivia" pageId="254" pageNumber="255">Bolivia</collectingCountry>
and east through western
<collectingCountry id="F320F0F14BAAD8BEFC1F37BFB4A1FB37" box="[936,1007,1249,1272]" name="Brazil" pageId="254" pageNumber="255">Brazil</collectingCountry>
in
<collectingRegion id="49F37E834BAAD8BEFBA237BFB303FB37" box="[1045,1101,1249,1272]" country="Brazil" name="Acre" pageId="254" pageNumber="255">Acre</collectingRegion>
and Amazonas states as far as the west bank of the Rio Negro, north of the Rio Solimões (fig. 152; da Silva and Patton, 1998: fig. 1). We found the species throughout the Rio Jurua´, although most specimens (74.6%) were collected in the extensive várzea forests of the Upper and Lower Central regions. The lower numbers of specimens collected at localities in the Mouth Region (only 15.5% of the total sample) likely reflect our sampling of that area during the rainy season when the habitat of
<taxonomicName id="4C37CBE24BAAD8BEFCB2351FB43BF997" authorityName="Goldman" authorityYear="1911" box="[773,885,1601,1624]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="254" pageNumber="255" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734BAAD8BEFCB2351FB459F997" box="[773,791,1601,1624]" italics="true" pageId="254" pageNumber="255">P</emphasis>
.
<emphasis id="B9436C734BAAD8BEFC9F351FB43BF997" box="[808,885,1601,1624]" italics="true" pageId="254" pageNumber="255">steerei</emphasis>
</taxonomicName>
was reduced by flooding, probably resulting in a seasonal decrease in population density. However, the relatively low percentage of animals found in the Headwaters Region (9.9%) might be due to other habitat features, or to competitive interactions with other species, since strict segregation by habitat is not evident in this region (table 63) and true, seasonally flooded várzea is not present.
</paragraph>
<caption id="DF48E0E94BAAD8BEFF0A36A3B674F902" ID-DOI="http://doi.org/10.5281/zenodo.5347648" ID-Zenodo-Dep="5347648" httpUri="https://zenodo.org/record/5347648/files/figure.png" pageId="254" pageNumber="255" startId="254.[189,227,1533,1554]" targetBox="[166,687,1180,1511]" targetPageId="254">
<paragraph id="8B88B0614BAAD8BEFF0A36A3B674F902" blockId="254.[163,691,1533,1741]" pageId="254" pageNumber="255">
Fig. 156. Geographic trend in overall size, as indexed by mean scores on the first principal components axis for samples of
<taxonomicName id="4C37CBE24BAAD8BEFE52356DB5FDF987" authorityName="Goldman" authorityYear="1911" box="[485,691,1587,1608]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="254" pageNumber="255" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734BAAD8BEFE52356DB5FDF987" box="[485,691,1587,1608]" italics="true" pageId="254" pageNumber="255">Proechimys steerei</emphasis>
</taxonomicName>
along the Rio Jurua´. Samples are positioned from left to right from the headwaters downriver to mouth localities. Solid circles represent population means; bars on either side represent 95% confidence limits.
</paragraph>
</caption>
<caption id="DF48E0E94BABD8BFFF7630BEB42DFBAF" ID-DOI="http://doi.org/10.5281/zenodo.5347650" ID-Zenodo-Dep="5347650" httpUri="https://zenodo.org/record/5347650/files/figure.png" pageId="255" pageNumber="256" startId="255.[193,231,992,1013]" targetBox="[262,1176,190,970]" targetPageId="255">
<paragraph id="8B88B0614BABD8BFFF7630BEB42DFBAF" blockId="255.[167,1271,992,1120]" pageId="255" pageNumber="256">
Fig. 158. Karyotypes of four specimens of
<taxonomicName id="4C37CBE24BABD8BFFD0330BEB4CFFC3A" authorityName="Goldman" authorityYear="1911" box="[692,897,992,1013]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="255" pageNumber="256" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734BABD8BFFD0330BEB4CFFC3A" box="[692,897,992,1013]" italics="true" pageId="255" pageNumber="256">Proechimys steerei</emphasis>
</taxonomicName>
:
<emphasis id="B9436C734BABD8BFFC2430BEB4E8FC3A" bold="true" box="[915,934,992,1013]" pageId="255" pageNumber="256">A</emphasis>
, Female; 2n = 24, FN = 40 MNFS 2096; Macaco, left bank Rio Jau´, Amazonas, Brazil.
<emphasis id="B9436C734BABD8BFFC8230A4B408FBC0" bold="true" box="[821,838,1018,1039]" pageId="255" pageNumber="256">B</emphasis>
, Female; 2n = 24, FN = 40; JUR 307 Vai­Quem­Quer (locality
<emphasis id="B9436C734BABD8BFFE0E374BB69DFBE5" bold="true" box="[441,467,1045,1066]" pageId="255" pageNumber="256">15</emphasis>
), right bank Rio Jurua´, Amazonas, Brazil.
<emphasis id="B9436C734BABD8BFFC11374BB4F7FBE5" bold="true" box="[934,953,1045,1066]" pageId="255" pageNumber="256">C</emphasis>
, Female; 2n = 24, FN = 41 Sacado (locality
<emphasis id="B9436C734BABD8BFFEED376EB629FB8A" bold="true" box="[346,359,1072,1093]" pageId="255" pageNumber="256">5</emphasis>
), right bank Rio Jurua´, Amazonas, Brazil. And,
<emphasis id="B9436C734BABD8BFFCC5376EB4CBFB8A" bold="true" box="[882,901,1072,1093]" pageId="255" pageNumber="256">D</emphasis>
, Male; 2n = 24, FN = 42; MNFS 1545; Nova Vida (locality
<emphasis id="B9436C734BABD8BFFE7F3714B69BFB90" bold="true" box="[456,469,1098,1119]" pageId="255" pageNumber="256">3</emphasis>
), right bank Rio Jurua´, Acre, Brazil.
</paragraph>
</caption>
<paragraph id="8B88B0614BABD8BFFF733667B5FFF95C" blockId="255.[167,696,1190,1741]" pageId="255" pageNumber="256">
While most common in seasonally flooded várzea,
<taxonomicName id="4C37CBE24BABD8BFFEB13609B636FAA1" authorityName="Goldman" authorityYear="1911" box="[262,376,1366,1390]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="255" pageNumber="256" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734BABD8BFFEB13609B656FAA1" box="[262,280,1367,1390]" italics="true" pageId="255" pageNumber="256">P</emphasis>
.
<emphasis id="B9436C734BABD8BFFE9C3609B636FAA1" box="[299,376,1367,1390]" italics="true" pageId="255" pageNumber="256">steerei</emphasis>
</taxonomicName>
was also found occasionally in secondary and disturbed terra firme forests, active and abandoned gardens, and margins of flooded grasslands (table 63). Elsewhere in Amazonia, we have taken this species in igapó (seasonally flooded black water) forests and along small creeks and riv­ er edge habitats within terra firme forest. The totality of our observations strongly suggest that
<taxonomicName id="4C37CBE24BABD8BFFF6B3501B602F9B9" authorityName="Goldman" authorityYear="1911" box="[220,332,1630,1654]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="255" pageNumber="256" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734BABD8BFFF6B3501B7A0F9B9" box="[220,238,1631,1654]" italics="true" pageId="255" pageNumber="256">P</emphasis>
.
<emphasis id="B9436C734BABD8BFFF483501B602F9B9" box="[255,332,1631,1654]" italics="true" pageId="255" pageNumber="256">steerei</emphasis>
</taxonomicName>
prefers flooded forests and riparian habitats within the lowland Amazon.
</paragraph>
<paragraph id="8B88B0614BABD8BFFF7335C7B314F95C" blockId="255.[167,696,1190,1741]" lastBlockId="255.[743,1272,1190,1741]" pageId="255" pageNumber="256">
We captured the species in the Tomahawk and Sherman traps in our standardized lines; in addition we also took specimens by hunting and with Victor and metal snap traps. Although the total number of each kind of traps varied, in our sample of specimens (n = 313 the majority (72%) was caught in Tomahawk traps and only 16% in Sherman traps, 4% were shot, 6% were caught with Victor snap traps, and 2% with the metal snap traps. Of the animals captured in live traps, most young and subadults (129 out of 178) were caught in Tomahawk traps and the remaining 49 individuals in Sherman traps; the same is true for adults with 129 (out of 135) caught in Tomahawk traps, and only six in the Sherman traps. Considering the large size (maximum weight
<quantity id="4CCF1D844BABD8BFFC3B3500B481F9BA" box="[908,975,1630,1653]" metricMagnitude="-1" metricUnit="kg" metricValue="8.0" pageId="255" pageNumber="256" unit="g" value="800.0">800 g</quantity>
) of individual
<taxonomicName id="4C37CBE24BABD8BFFB363501B3BEF9B9" authorityName="Goldman" authorityYear="1911" box="[1153,1264,1630,1654]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="255" pageNumber="256" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734BABD8BFFB363501B3DDF9B9" box="[1153,1171,1631,1654]" italics="true" pageId="255" pageNumber="256">P</emphasis>
.
<emphasis id="B9436C734BABD8BFFB143501B3BEF9B9" box="[1187,1264,1631,1654]" italics="true" pageId="255" pageNumber="256">steerei</emphasis>
</taxonomicName>
these results are not surprising.
</paragraph>
<paragraph id="8B88B0614BABD941FCB335C7B65DFB14" blockId="255.[743,1272,1190,1741]" lastBlockId="257.[167,696,956,1741]" lastPageId="257" lastPageNumber="258" pageId="255" pageNumber="256">
KARYOTYPE: 2n = 24; FN = 4042. We prepared karyotypes from 148 individuals, at least one from each locality from which
<taxonomicName id="4C37CBE24A54D940FD2D379AB7BEFB37" authorityName="Goldman" authorityYear="1911" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="256" pageNumber="257" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734A54D940FD2D379AB5E2FB14" box="[666,684,1220,1243]" italics="true" pageId="256" pageNumber="257">P</emphasis>
.
<emphasis id="B9436C734A54D940FF1437BFB7BEFB37" box="[163,240,1249,1272]" italics="true" pageId="256" pageNumber="257">steerei</emphasis>
</taxonomicName>
was collected. All specimens had an autosomal complement consisting of two pairs of large and seven pairs of medium to small meta and submetacentrics, one pair of large subtelocentrics, and one pair of small acrocentrics. Specimens from the Headwaters Region had a second, smaller pair of subtelocentrics (fig. 158D); those from the Mouth Region typically had this pair as a medium­sized acrocentric (fig. 158B). Finally, specimens from localities in the Upper and Lower Central Regions had one or the other of these two homozygous conditions, or were heterozygous (fig. 158C). The Xchromosome was a small acrocentric and the Y­chromosome an even smaller one in all individuals, regardless of autosomal comple­ ment. We caught heterozygous individuals at localities
<emphasis id="B9436C734A54D940FCEF37BFB429FB36" bold="true" box="[856,871,1249,1273]" pageId="256" pageNumber="257">9</emphasis>
,
<emphasis id="B9436C734A54D940FCCE37BFB4D8FB36" bold="true" box="[889,918,1249,1273]" pageId="256" pageNumber="257">10</emphasis>
, and
<emphasis id="B9436C734A54D940FC5737BFB4B3FB36" bold="true" box="[992,1021,1249,1273]" pageId="256" pageNumber="257">14</emphasis>
. At localities
<emphasis id="B9436C734A54D940FB1937BFB3F3FB36" bold="true" box="[1198,1213,1249,1273]" pageId="256" pageNumber="257">9</emphasis>
and
<emphasis id="B9436C734A54D940FD5437A0B44EFAD9" bold="true" box="[739,768,1278,1302]" pageId="256" pageNumber="257">14</emphasis>
, the parental homozygotes with mediumsized acrocentrics were more abundant (9 of 11 and 8 of 10 karyotyped individuals in each locality, respectively); the two remaining individuals from each locality were of both
<typeStatus id="548C0EC34A54D940FC9036CFB429FA67" box="[807,871,1425,1448]" pageId="256" pageNumber="257">types</typeStatus>
, the parental homozygote with medium­sized subtelocentric and the heterozygote. At locality
<emphasis id="B9436C734A54D940FC613695B4BDFA2C" bold="true" box="[982,1011,1483,1507]" pageId="256" pageNumber="257">10</emphasis>
, three of six karyotyped individuals were homozygotes with medium­sized acrocentrics, and three were heterozygotes. The downriver karyotype with the single pair of medium­sized acrocentrics also characterizes samples from the Rio Jau´, to the north of the Rio Solimões and west of the Rio Negro (fig. 158A). The upriver karyotype with the two pairs of subtel­ ocentrics is also found in samples from central, eastern, and southern
<collectingCountry id="F320F0F14A55D941FE423087B564FC3F" box="[501,554,985,1008]" name="Peru" pageId="257" pageNumber="258">Perú</collectingCountry>
(table 80). Despite the slight variation in the karyotype of
<taxonomicName id="4C37CBE24A55D941FF67374AB609FBE4" authorityName="Goldman" authorityYear="1911" box="[208,327,1044,1067]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="257" pageNumber="258" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734A55D941FF67374AB7ACFBE4" box="[208,226,1044,1067]" italics="true" pageId="257" pageNumber="258">P</emphasis>
.
<emphasis id="B9436C734A55D941FF4D374AB609FBE4" box="[250,327,1044,1067]" italics="true" pageId="257" pageNumber="258">steerei</emphasis>
</taxonomicName>
, it is readily distinguishable from that of
<taxonomicName id="4C37CBE24A55D941FEF5376FB6F8FB87" authorityName="Thomas" authorityYear="1905" box="[322,438,1073,1096]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="257" pageNumber="258" phylum="Chordata" rank="species" species="goeldii">
<emphasis id="B9436C734A55D941FEF5376FB61AFB87" box="[322,340,1073,1096]" italics="true" pageId="257" pageNumber="258">P</emphasis>
.
<emphasis id="B9436C734A55D941FED2376FB6F8FB87" box="[357,438,1073,1096]" italics="true" pageId="257" pageNumber="258">goeldii</emphasis>
</taxonomicName>
(2n = 24, FN = 42) from the Rio Xingu in Estado do Pará to the east, or to those of
<taxonomicName id="4C37CBE24A55D941FE253732B528FB4C" authorityName="Hershkovitz" authorityYear="1948" box="[402,614,1132,1155]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="257" pageNumber="258" phylum="Chordata" rank="species" species="quadruplicatus">
<emphasis id="B9436C734A55D941FE253732B6EAFB4C" box="[402,420,1132,1155]" italics="true" pageId="257" pageNumber="258">P</emphasis>
.
<emphasis id="B9436C734A55D941FE013732B528FB4C" box="[438,614,1132,1155]" italics="true" pageId="257" pageNumber="258">quadruplicatus</emphasis>
</taxonomicName>
(2n = 2628, FN = 4244) in northwestern and north­central Estado do Amazonia (table 75; fig. 159).
</paragraph>
</subSubSection>
<caption id="DF48E0E94A55D941FF7631A5B677FCB5" ID-DOI="http://doi.org/10.5281/zenodo.5347652" ID-Zenodo-Dep="5347652" httpUri="https://zenodo.org/record/5347652/files/figure.png" pageId="257" pageNumber="258" startId="257.[193,231,763,784]" targetBox="[260,1177,188,741]" targetPageId="257">
<paragraph id="8B88B0614A55D941FF7631A5B677FCB5" blockId="257.[167,1271,762,890]" pageId="257" pageNumber="258">
Fig. 159. Karyotypes of three specimens of
<taxonomicName id="4C37CBE24A55D941FD0831A5B474FCDF" authorityName="Allen" authorityYear="1899" box="[703,826,763,784]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="257" pageNumber="258" phylum="Chordata" rank="genus">
<emphasis id="B9436C734A55D941FD0831A5B474FCDF" box="[703,826,763,784]" italics="true" pageId="257" pageNumber="258">Proechimys</emphasis>
</taxonomicName>
:
<emphasis id="B9436C734A55D941FCF931A4B42FFCC0" bold="true" box="[846,865,762,783]" pageId="257" pageNumber="258">A</emphasis>
, a male
<taxonomicName id="4C37CBE24A55D941FC7E31A5B3D7FCDF" authorityName="Thomas" authorityYear="1905" box="[969,1177,763,784]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="257" pageNumber="258" phylum="Chordata" rank="species" species="goeldii">
<emphasis id="B9436C734A55D941FC7E31A5B3D7FCDF" box="[969,1177,763,784]" italics="true" pageId="257" pageNumber="258">Proechimys goeldii</emphasis>
</taxonomicName>
; USNM 549573; 2n=24, FN=42; 52 km S Altamira, Rio Xingu, Para´, Brazil.
<emphasis id="B9436C734A55D941FC14304BB4FAFCE5" bold="true" box="[931,948,789,810]" pageId="257" pageNumber="258">B</emphasis>
, A male
<taxonomicName id="4C37CBE24A55D941FBA93048B7B4FC8A" authorityName="Hershkovitz" authorityYear="1948" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="257" pageNumber="258" phylum="Chordata" rank="species" species="quadruplicatus">
<emphasis id="B9436C734A55D941FBA93048B7B4FC8A" italics="true" pageId="257" pageNumber="258">Proechimys quadruplicatus</emphasis>
</taxonomicName>
; MVZ 157860; 2n=28, FN=42; La Poza, Río Santiago, Amazonas, Peru´. And,
<emphasis id="B9436C734A55D941FBD8306EB3CCFC8A" bold="true" box="[1135,1154,816,837]" pageId="257" pageNumber="258">C</emphasis>
, A female
<taxonomicName id="4C37CBE24A55D941FF103015B687FCAF" authorityName="Hershkovitz" authorityYear="1948" box="[167,457,843,864]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="257" pageNumber="258" phylum="Chordata" rank="species" species="quadruplicatus">
<emphasis id="B9436C734A55D941FF103015B687FCAF" box="[167,457,843,864]" italics="true" pageId="257" pageNumber="258">Proechimys quadruplicatus</emphasis>
</taxonomicName>
; JLP 16794; 2n=28, FN=42; Lago Meduiním, left bank Rio Negro, Amazonas, Brazil.
</paragraph>
</caption>
<subSubSection id="C32DE3EA4A55D942FF7237BFB46AFC57" lastPageId="258" lastPageNumber="259" pageId="257" pageNumber="258" type="discussion">
<paragraph id="8B88B0614A55D941FF7237BFB3B9F902" blockId="257.[167,696,956,1741]" lastBlockId="257.[743,1271,956,1741]" pageId="257" pageNumber="258">
COMMENTS: Patton (1987) included
<taxonomicName id="4C37CBE24A55D941FD2A37BFB7BAFAD9" authorityName="Goldman" authorityYear="1911" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="257" pageNumber="258" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734A55D941FD2A37BFB5E1FB37" box="[669,687,1249,1272]" italics="true" pageId="257" pageNumber="258">P</emphasis>
.
<emphasis id="B9436C734A55D941FF1037A1B7BAFAD9" box="[167,244,1279,1302]" italics="true" pageId="257" pageNumber="258">steerei</emphasis>
</taxonomicName>
within his
<taxonomicName id="4C37CBE24A55D941FE3F37A1B697FAD9" authorityName="Thomas" authorityYear="1905" box="[392,473,1279,1302]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="257" pageNumber="258" phylum="Chordata" rank="species" species="goeldii">
<emphasis id="B9436C734A55D941FE3F37A1B697FAD9" box="[392,473,1279,1302]" italics="true" pageId="257" pageNumber="258">goeldii</emphasis>
</taxonomicName>
­ group, which he noted varied more over its geographic range than any other species group that he examined, and suggested the existence of an eastern (
<taxonomicName id="4C37CBE24A55D941FF53362AB612FA44" authorityName="Thomas" authorityYear="1905" box="[228,348,1396,1419]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="257" pageNumber="258" phylum="Chordata" rank="species" species="goeldii">
<emphasis id="B9436C734A55D941FF53362AB7B8FA44" box="[228,246,1396,1419]" italics="true" pageId="257" pageNumber="258">P</emphasis>
.
<emphasis id="B9436C734A55D941FEBC362AB612FA44" box="[267,348,1396,1419]" italics="true" pageId="257" pageNumber="258">goeldii</emphasis>
</taxonomicName>
) and a western (
<taxonomicName id="4C37CBE24A55D941FD8E362AB5E3FA44" authorityName="Goldman" authorityYear="1911" box="[569,685,1396,1419]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="257" pageNumber="258" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734A55D941FD8E362AB505FA44" box="[569,587,1396,1419]" italics="true" pageId="257" pageNumber="258">P</emphasis>
.
<emphasis id="B9436C734A55D941FDD7362AB5E3FA44" box="[608,685,1396,1419]" italics="true" pageId="257" pageNumber="258">steerei</emphasis>
</taxonomicName>
) species. The cytochrome­b analyses (fig. 154), however, subdivide western populations even further and, along with karyotypic data, suggest at least three species within the
<taxonomicName id="4C37CBE24A55D941FF103559B7B6F9D1" authorityName="Thomas" authorityYear="1905" box="[167,248,1543,1566]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="257" pageNumber="258" phylum="Chordata" rank="species" species="goeldii">
<emphasis id="B9436C734A55D941FF103559B7B6F9D1" box="[167,248,1543,1566]" italics="true" pageId="257" pageNumber="258">goeldii</emphasis>
</taxonomicName>
­ group. The senior names that would apply to each clade are: (1)
<taxonomicName id="4C37CBE24A55D941FE4B357AB6FCF997" authority="Hershkovitz, 1948" authorityName="Hershkovitz" authorityYear="1948" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="257" pageNumber="258" phylum="Chordata" rank="species" species="quadruplicatus">
<emphasis id="B9436C734A55D941FE4B357AB540F9F4" box="[508,526,1572,1595]" italics="true" pageId="257" pageNumber="258">P</emphasis>
.
<emphasis id="B9436C734A55D941FDA8357AB787F997" italics="true" pageId="257" pageNumber="258">quadruplicatus</emphasis>
Hershkovitz, 1948
</taxonomicName>
(with
<taxonomicName id="4C37CBE24A55D941FDBC351FB62DF9BA" authority="Moojen, 1948 a" authorityName="Moojen" authorityYear="1948" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="257" pageNumber="258" phylum="Chordata" rank="species" species="amphichoricus">
<emphasis id="B9436C734A55D941FDBC351FB5F6F997" box="[523,696,1601,1624]" italics="true" pageId="257" pageNumber="258">amphichoricus</emphasis>
Moojen, 1948 a
</taxonomicName>
junior synonym), which occurs from
<collectingCountry id="F320F0F14A55D941FE9A3522B62CF95C" box="[301,354,1660,1683]" name="Peru" pageId="257" pageNumber="258">Perú</collectingCountry>
north of the Rio Marañón through eastern
<collectingCountry id="F320F0F14A55D941FED035C7B686F97F" box="[359,456,1689,1712]" name="Ecuador" pageId="257" pageNumber="258">Ecuador</collectingCountry>
to southeastern
<collectingCountry id="F320F0F14A55D941FD3C35C7B7B6F902" name="Colombia" pageId="257" pageNumber="258">Colombia</collectingCountry>
and east across southern
<collectingCountry id="F320F0F14A55D941FD8835E8B5F9F902" box="[575,695,1718,1741]" name="Venezuela" pageId="257" pageNumber="258">Venezuela</collectingCountry>
and adjacent
<collectingCountry id="F320F0F14A55D941FC3430E2B484FC1C" box="[899,970,956,979]" name="Brazil" pageId="257" pageNumber="258">Brazil</collectingCountry>
north and east of the Rio Negro to at least the vicinity of Manaus Available geographic samples of this taxon are 2n = 26 (
<collectingCountry id="F320F0F14A55D941FC25374AB342FBE4" box="[914,1036,1044,1067]" name="Venezuela" pageId="257" pageNumber="258">Venezuela</collectingCountry>
[Reig and Useche 1976]) or 2n = 28 (
<collectingCountry id="F320F0F14A55D941FC78376FB348FB87" box="[975,1030,1073,1096]" name="Peru" pageId="257" pageNumber="258">Perú</collectingCountry>
and
<collectingCountry id="F320F0F14A55D941FBF6376FB3ECFB87" box="[1089,1186,1073,1096]" name="Ecuador" pageId="257" pageNumber="258">Ecuador</collectingCountry>
[Gardner and Emmons, 1984] and
<collectingCountry id="F320F0F14A55D941FBF03710B3C0FBAA" box="[1095,1166,1102,1125]" name="Brazil" pageId="257" pageNumber="258">Brazil</collectingCountry>
[this report]; see table 80 and fig. 159BC). We referred to this clade as
<taxonomicName id="4C37CBE24A55D941FBB537D7B39AFB6F" authorityName="Moojen" authorityYear="1948" box="[1026,1236,1161,1184]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="257" pageNumber="258" phylum="Chordata" rank="species" species="amphichoricus">
<emphasis id="B9436C734A55D941FBB537D7B35AFB6F" box="[1026,1044,1161,1184]" italics="true" pageId="257" pageNumber="258">P</emphasis>
.
<emphasis id="B9436C734A55D941FB9037D7B39AFB6F" box="[1063,1236,1161,1184]" italics="true" pageId="257" pageNumber="258">amphichoricus</emphasis>
</taxonomicName>
in earlier papers (e.g., da Silva and Patton 1998). (2)
<taxonomicName id="4C37CBE24A55D941FCDD379AB492FB14" authorityName="Goldman" authorityYear="1911" box="[874,988,1220,1243]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="257" pageNumber="258" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734A55D941FCDD379AB432FB14" box="[874,892,1220,1243]" italics="true" pageId="257" pageNumber="258">P</emphasis>
.
<emphasis id="B9436C734A55D941FC38379AB492FB14" box="[911,988,1220,1243]" italics="true" pageId="257" pageNumber="258">steerei</emphasis>
</taxonomicName>
(with
<emphasis id="B9436C734A55D941FB87379AB3C9FB14" box="[1072,1159,1220,1243]" italics="true" pageId="257" pageNumber="258">pachita</emphasis>
Thomas 1923, and
<emphasis id="B9436C734A55D941FCD337BFB4F0FB37" box="[868,958,1249,1272]" italics="true" pageId="257" pageNumber="258">rattinus</emphasis>
Thomas, 1926, clearly junior synonyms, and probably
<emphasis id="B9436C734A55D941FBE537A1B3EAFAD9" box="[1106,1188,1279,1302]" italics="true" pageId="257" pageNumber="258">kermiti</emphasis>
Allen 1915,
<emphasis id="B9436C734A55D941FC8C3642B439FAFC" box="[827,887,1308,1331]" italics="true" pageId="257" pageNumber="258">hilda</emphasis>
Thomas, 1924, and
<emphasis id="B9436C734A55D941FB263642B3B9FAFC" box="[1169,1271,1308,1331]" italics="true" pageId="257" pageNumber="258">liminalis</emphasis>
Moojen, 1948, as well) with 2n = 24, FN = 40 or 42 karyotype (table 80 and figs. 158) The range of this species is delimited above and in figure 153. And, (3)
<taxonomicName id="4C37CBE24A55D941FB8D36CFB3E1FA67" authorityName="Thomas" authorityYear="1905" box="[1082,1199,1425,1448]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="257" pageNumber="258" phylum="Chordata" rank="species" species="goeldii">
<emphasis id="B9436C734A55D941FB8D36CFB302FA67" box="[1082,1100,1425,1448]" italics="true" pageId="257" pageNumber="258">P</emphasis>
.
<emphasis id="B9436C734A55D941FBE936CFB3E1FA67" box="[1118,1199,1425,1448]" italics="true" pageId="257" pageNumber="258">goeldii</emphasis>
</taxonomicName>
(with
<emphasis id="B9436C734A55D941FD5036F1B476FA09" box="[743,824,1455,1478]" italics="true" pageId="257" pageNumber="258">hyleae</emphasis>
Moojen, 1948,
<emphasis id="B9436C734A55D941FBAE36F1B330FA09" box="[1049,1150,1455,1478]" italics="true" pageId="257" pageNumber="258">nesiotes</emphasis>
Moojen 1948, and
<emphasis id="B9436C734A55D941FCD83692B4A1FA2C" box="[879,1007,1484,1507]" italics="true" pageId="257" pageNumber="258">leioprimna</emphasis>
Moojen, 1948 synonyms), from Estado do Pará in the eastern Amazon Basin of
<collectingCountry id="F320F0F14A55D941FC0B3558B349F9D2" box="[956,1031,1542,1565]" name="Brazil" pageId="257" pageNumber="258">Brazil</collectingCountry>
. Our samples of this taxon from the lower Rio Xingu and the Serra Carajás are inadequate to do more than simply document the extensive degree of sequence divergence between
<taxonomicName id="4C37CBE24A55D941FB983522B3EFF95C" authorityName="Thomas" authorityYear="1905" box="[1071,1185,1660,1683]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="257" pageNumber="258" phylum="Chordata" rank="species" species="goeldii">
<emphasis id="B9436C734A55D941FB983522B30FF95C" box="[1071,1089,1660,1683]" italics="true" pageId="257" pageNumber="258">P</emphasis>
.
<emphasis id="B9436C734A55D941FBE73522B3EFF95C" box="[1104,1185,1660,1683]" italics="true" pageId="257" pageNumber="258">goeldii</emphasis>
</taxonomicName>
and either
<taxonomicName id="4C37CBE24A55D941FCAA35C7B4BEF97F" authorityName="Hershkovitz" authorityYear="1948" box="[797,1008,1689,1712]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="257" pageNumber="258" phylum="Chordata" rank="species" species="quadruplicatus">
<emphasis id="B9436C734A55D941FCAA35C7B461F97F" box="[797,815,1689,1712]" italics="true" pageId="257" pageNumber="258">P</emphasis>
.
<emphasis id="B9436C734A55D941FCF735C7B4BEF97F" box="[832,1008,1689,1712]" italics="true" pageId="257" pageNumber="258">quadruplicatus</emphasis>
</taxonomicName>
or
<taxonomicName id="4C37CBE24A55D941FBAC35C7B3C4F97F" authorityName="Goldman" authorityYear="1911" box="[1051,1162,1689,1712]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="257" pageNumber="258" phylum="Chordata" rank="species" species="steerei">
<emphasis id="B9436C734A55D941FBAC35C7B363F97F" box="[1051,1069,1689,1712]" italics="true" pageId="257" pageNumber="258">P</emphasis>
.
<emphasis id="B9436C734A55D941FB8A35C7B3C4F97F" box="[1085,1162,1689,1712]" italics="true" pageId="257" pageNumber="258">steerei</emphasis>
</taxonomicName>
(average from 13.2 to 14.4%). These samples also
</paragraph>
<paragraph id="8B88B0614A56D942FF14339FB683FE47" blockId="258.[163,691,193,920]" pageId="258" pageNumber="259">
have a different karyotype, with 2n = 24 and FN = 42 (fig. 159A; table 80). Additional sampling throughout the range of the
<taxonomicName id="4C37CBE24A56D942FDEE33A2B5E4FEDC" authorityName="Thomas" authorityYear="1905" box="[601,682,252,275]" class="Mammalia" family="Echimyidae" genus="Proechimys" kingdom="Animalia" order="Rodentia" pageId="258" pageNumber="259" phylum="Chordata" rank="species" species="goeldii">
<emphasis id="B9436C734A56D942FDEE33A2B5E4FEDC" box="[601,682,252,275]" italics="true" pageId="258" pageNumber="259">goeldii</emphasis>
</taxonomicName>
­ group is needed to further refine the geographic boundaries of the three species we identify here, as well as to determine if other taxa deserve recognition.
</paragraph>
<paragraph id="8B88B0614A56D942FF7732D0B46AFC57" blockId="258.[163,691,193,920]" lastBlockId="258.[739,1267,193,920]" pageId="258" pageNumber="259">
SPECIMENS EXAMINED (n = 453): (
<emphasis id="B9436C734A56D942FDE432D0B52CFE69" bold="true" box="[595,610,398,422]" pageId="258" pageNumber="259">2</emphasis>
) 1f — MNFS 1254; (
<emphasis id="B9436C734A56D942FEE732F5B611FE0C" bold="true" box="[336,351,427,451]" pageId="258" pageNumber="259">a</emphasis>
)
<quantity id="4CCF1D844A56D942FEC632F2B6D6FE0C" box="[369,408,428,451]" metricMagnitude="0" metricUnit="m" metricValue="7.0" pageId="258" pageNumber="259" unit="m" value="7.0">7m</quantity>
, 4f — MNFS 995997, 1032, 1046, 1056057, 10601063; (
<emphasis id="B9436C734A56D942FDE03297B52AFE2E" bold="true" box="[599,612,457,481]" pageId="258" pageNumber="259">c</emphasis>
) 1f — MNFS 1037; (
<emphasis id="B9436C734A56D942FEF832B8B610FE31" bold="true" box="[335,350,486,510]" pageId="258" pageNumber="259">3</emphasis>
)
<quantity id="4CCF1D844A56D942FEC732B8B6D6FE32" box="[368,408,486,509]" metricMagnitude="0" metricUnit="m" metricValue="4.0" pageId="258" pageNumber="259" unit="m" value="4.0">4m</quantity>
, 9f — JUR 206, MNFS 1521, 1543, 1545, 15471548, 15891592, 1633, 1655, 1682; (
<emphasis id="B9436C734A56D942FE27317FB6D1FDF6" bold="true" box="[400,415,545,569]" pageId="258" pageNumber="259">4</emphasis>
)
<quantity id="4CCF1D844A56D942FE04317FB694FDF7" box="[435,474,545,568]" metricMagnitude="0" metricUnit="m" metricValue="8.0" pageId="258" pageNumber="259" unit="m" value="8.0">8m</quantity>
, 13f — JUR 241, 245, MNFS 1430, 14451447, 1459, 1462, 14741477, 15061507, 1569, 1617, 1623 1625, 1643, 1662; (
<emphasis id="B9436C734A56D942FE3B3127B6D2FD5E" bold="true" box="[396,412,633,657]" pageId="258" pageNumber="259">d</emphasis>
)
<quantity id="4CCF1D844A56D942FE193127B69DFD5F" box="[430,467,633,656]" metricMagnitude="0" metricUnit="m" metricValue="1.0" pageId="258" pageNumber="259" unit="m" value="1.0">1m</quantity>
— JLP 15628; (
<emphasis id="B9436C734A56D942FD2C3127B5E4FD5E" bold="true" box="[667,682,633,657]" pageId="258" pageNumber="259">5</emphasis>
)
<quantity id="4CCF1D844A56D942FF1431C8B797FD62" box="[163,217,662,685]" metricMagnitude="1" metricUnit="m" metricValue="3.9" pageId="258" pageNumber="259" unit="m" value="39.0">39m</quantity>
, 31f, 1 unknown — JUR 118129, 133 134, 136143, 170, 172, MNFS 570576, 585, 594607, 614620, 626628, 641, 647650, 658667; (
<emphasis id="B9436C734A56D942FE0731B0B6F1FCC9" bold="true" box="[432,447,750,774]" pageId="258" pageNumber="259">6</emphasis>
)
<quantity id="4CCF1D844A56D942FE6C31B0B55FFCCA" box="[475,529,750,773]" metricMagnitude="1" metricUnit="m" metricValue="1.2" pageId="258" pageNumber="259" unit="m" value="12.0">12m</quantity>
, 7f — JLP 15558, 15614, 1569015692, 1569815700, 15705, 1570915712, 15722, 1573015732, JUR 176, MNFS 526; (
<emphasis id="B9436C734A56D942FE7E3018B696FC91" bold="true" box="[457,472,838,862]" pageId="258" pageNumber="259">7</emphasis>
)
<quantity id="4CCF1D844A56D942FE593018B56AFC92" box="[494,548,838,861]" metricMagnitude="1" metricUnit="m" metricValue="1.0" pageId="258" pageNumber="259" unit="m" value="10.0">10m</quantity>
, 8f — JLP 1524415245, 1525415255, 1526815269, 1545815459, MNFS 333334, 337338, 342, 346, 354, 472, 496497; (
<emphasis id="B9436C734A56D942FBD5339FB33FFF16" bold="true" box="[1122,1137,193,217]" pageId="258" pageNumber="259">8</emphasis>
)
<quantity id="4CCF1D844A56D942FB31339FB3F2FF17" box="[1158,1212,193,216]" metricMagnitude="1" metricUnit="m" metricValue="6.8" pageId="258" pageNumber="259" unit="m" value="68.0">68m</quantity>
, 61f — JLP 1537515382, 15386115391, 1539615401, JUR 1532, 44, 49, 5071, 80109, 111113, 115116, MNFS 443 463, 467469, 477483, 501; (
<emphasis id="B9436C734A56D942FBED3268B327FE81" bold="true" box="[1114,1129,310,334]" pageId="258" pageNumber="259">9</emphasis>
)
<quantity id="4CCF1D844A56D942FBCB3268B3EDFE82" box="[1148,1187,310,333]" metricMagnitude="0" metricUnit="m" metricValue="3.0" pageId="258" pageNumber="259" unit="m" value="3.0">3m</quantity>
, 2f — JLP 1592615927, 16019, 16053, MNFS 851; (
<emphasis id="B9436C734A56D942FC98322FB402FE46" bold="true" box="[815,844,369,393]" pageId="258" pageNumber="259">9a</emphasis>
)
<quantity id="4CCF1D844A56D942FCD3322FB4C2FE47" box="[868,908,369,392]" metricMagnitude="0" metricUnit="m" metricValue="5.0" pageId="258" pageNumber="259" unit="m" value="5.0">5m</quantity>
, 5f — DMN 1617, MNFS 926928, 939, 949, 958959, 963; (
<emphasis id="B9436C734A56D942FB3E32D0B3E8FE69" bold="true" box="[1161,1190,398,422]" pageId="258" pageNumber="259">10</emphasis>
)
<quantity id="4CCF1D844A56D942FB0F32D0B3A0FE6A" box="[1208,1262,398,421]" metricMagnitude="1" metricUnit="m" metricValue="1.5" pageId="258" pageNumber="259" unit="m" value="15.0">15m</quantity>
, 9f — DMN, 1213, MNFS 869877, 895, 898899, 915916, 919, 932933, 940, 945 947, 956; (
<emphasis id="B9436C734A56D942FCC332B8B4DFFE31" bold="true" box="[884,913,486,510]" pageId="258" pageNumber="259">11</emphasis>
)
<quantity id="4CCF1D844A56D942FC1C32B8B4AFFE32" box="[939,993,486,509]" metricMagnitude="1" metricUnit="m" metricValue="1.2" pageId="258" pageNumber="259" unit="m" value="12.0">12m</quantity>
, 17f — JLP 15749 15750, 1575315757, MNFS 679680, 688 691, 700701, 704, 709711, 714715, 720, 752753, 770771, 784785, 794; (
<emphasis id="B9436C734A56D942FB3E3160B3E8FD99" bold="true" box="[1161,1190,574,598]" pageId="258" pageNumber="259">12</emphasis>
)
<quantity id="4CCF1D844A56D942FB0F3160B3A0FD9A" box="[1208,1262,574,597]" metricMagnitude="1" metricUnit="m" metricValue="1.7" pageId="258" pageNumber="259" unit="m" value="17.0">17m</quantity>
, 13f — JLP 15789, 1579915800, 15835 15838, 1585615858, 1586015864, 15878 15880, 15889, 15915, MNFS 687, 730731, 733735, 780, 781, 782, 783; (
<emphasis id="B9436C734A56D942FBE031EDB33AFD04" bold="true" box="[1111,1140,691,715]" pageId="258" pageNumber="259">13</emphasis>
)
<quantity id="4CCF1D844A56D942FB3F31EAB3F0FD04" box="[1160,1214,692,715]" metricMagnitude="1" metricUnit="m" metricValue="1.9" pageId="258" pageNumber="259" unit="m" value="19.0">19m</quantity>
, 27f — JUR 252262, 264266, 274282, 293, 307314, 326330, 338341, 345, 347, 349 351; (
<emphasis id="B9436C734A56D942FC9C3055B406FCEC" bold="true" box="[811,840,779,803]" pageId="258" pageNumber="259">14</emphasis>
) 1f — MNFS 1777; (
<emphasis id="B9436C734A56D942FBEF3055B329FCEC" bold="true" box="[1112,1127,779,803]" pageId="258" pageNumber="259">o</emphasis>
)
<quantity id="4CCF1D844A56D942FBCC3052B3EDFCEC" box="[1147,1187,780,803]" metricMagnitude="0" metricUnit="m" metricValue="3.0" pageId="258" pageNumber="259" unit="m" value="3.0">3m</quantity>
, 2f — MNFS 17371738, 17701772; (
<emphasis id="B9436C734A56D942FBDD3077B3C9FC8E" bold="true" box="[1130,1159,809,833]" pageId="258" pageNumber="259">16</emphasis>
)
<quantity id="4CCF1D844A56D942FB2F3077B380FC8F" box="[1176,1230,809,832]" metricMagnitude="1" metricUnit="m" metricValue="1.2" pageId="258" pageNumber="259" unit="m" value="12.0">12m</quantity>
, 6f — JUR 477, 504, MNFS 17491750, 1752 1753, 17551756, 17631769, 1783, 1792 1793.
</paragraph>
</subSubSection>
</treatment>
</document>