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<document id="D068295A017E76E106EC4630EE5B6B81" ID-CLB-Dataset="3674" ID-DOI="10.1206/0003-0082(2006)3509[1:ANMSOS]2.0.CO;2" ID-GBIF-Dataset="2f8c7065-6e4b-46b1-89af-809f51afdcf2" ID-ISSN="0003-0082" ID-Zenodo-Dep="4735318" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1620068761546" checkinUser="felipe" docAuthor="Anderson, Robert P. &amp; Timm, Robert M." docDate="2006" docId="03DA87BDFF952560FF66FD380653FA66" docLanguage="en" docName="AmMusNovit.2006.3509.1-38.pdf.imf" docOrigin="American Museum Novitates 3509 (1)" docSource="http://www.bioone.org/perlserv/?request=get-abstract&amp;doi=10.1206%2F0003-0082(2006)3509%5B1%3AANMSOS%5D2.0.CO%3B2" docStyle="DocumentStyle:70DDF1BDF6846024993F05262BAE3001.3:AmMusNovit.2000-2010.journal_article.type1" docStyleId="70DDF1BDF6846024993F05262BAE3001" docStyleName="AmMusNovit.2000-2010.journal_article.type1" docStyleVersion="3" docTitle="Heteromys DESMARESTIANUS 1817" docType="treatment" docVersion="9" lastPageNumber="25" masterDocId="FFE3FFC5FF832578FFFDFFEC024AFF9A" masterDocTitle="A New Montane Species of Spiny Pocket Mouse (Rodentia: Heteromyidae: Heteromys) from Northwestern Costa Rica" masterLastPageNumber="39" masterPageNumber="1" pageNumber="23" updateTime="1698938181996" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0">
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<mods:title id="A9EBAF383C2D9D0B6D8004A3A1095449">A New Montane Species of Spiny Pocket Mouse (Rodentia: Heteromyidae: Heteromys) from Northwestern Costa Rica</mods:title>
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<paragraph id="8BCC36ABFF95256EFF66FD380390FC93" blockId="22.[155,626,724,778]" pageId="22" pageNumber="23">
TAXONOMIC NOTES ON OTHER
<taxonomicName id="4C734D28FF95256EFE0DFD390038FD71" ID-CoL="4WKC" authority="IN" authorityName="DESMARESTIANUS" authorityYear="1817" box="[496,626,725,749]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="22" pageNumber="23" phylum="Chordata" rank="genus">
<emphasis id="B907EAB9FF95256EFE0DFD390038FD71" box="[496,626,725,749]" italics="true" pageId="22" pageNumber="23">HETEROMYS</emphasis>
</taxonomicName>
IN
<collectingCountry id="F364763BFF95256EFEAEFD1E0390FC93" box="[339,474,754,778]" name="Costa Rica" pageId="22" pageNumber="23">COSTA RICA</collectingCountry>
</paragraph>
</subSubSection>
<subSubSection id="C3696520FF95256EFF67FCCA012DFCA4" pageId="22" pageNumber="23" type="reference_group">
<paragraph id="8BCC36ABFF95256EFF67FCCA00C0F942" blockId="22.[128,654,806,1753]" pageId="22" pageNumber="23">
<taxonomicName id="4C734D28FF95256EFF67FCCA03D1FCA7" ID-CoL="4WKC" authority="ORESTERUS" authorityName="ORESTERUS" box="[154,411,806,830]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="22" pageNumber="23" phylum="Chordata" rank="genus">
<emphasis id="B907EAB9FF95256EFF67FCCA03D1FCA7" box="[154,411,806,830]" italics="true" pageId="22" pageNumber="23">HETEROMYS ORESTERUS</emphasis>
</taxonomicName>
: Our examinations indicate that
<taxonomicName id="4C734D28FF95256EFED9FCA8005BFCC6" ID-CoL="7VD2T" box="[292,529,836,860]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="oresterus">
<emphasis id="B907EAB9FF95256EFED9FCA8005BFCC6" box="[292,529,836,860]" italics="true" pageId="22" pageNumber="23">Heteromys oresterus</emphasis>
</taxonomicName>
is known definitively from only three localities in the western portion of the Cordillera de Talamanca in central
<collectingCountry id="F364763BFF95256EFE69FC710057FC2F" box="[404,541,925,949]" name="Costa Rica" pageId="22" pageNumber="23">Costa Rica</collectingCountry>
(fig. 10; appendix 2). In the original description,
<bibRefCitation id="EFE24B5AFF95256EFDB9FC57028DFC6A" author="Harris, W. P., Jr." pageId="22" pageNumber="23" pagination="1 - 6" refId="ref19177" refString="Harris, W. P., Jr. 1932. Four new mammals from Costa Rica. Occasional Papers of the Museum of Zoology, University of Michigan 248: 1 - 6." type="journal article" year="1932">Harris (1932)</bibRefCitation>
named
<taxonomicName id="4C734D28FF95256EFEDAFC3503F8FC6B" ID-CoL="7VD2T" box="[295,434,985,1009]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="oresterus">
<emphasis id="B907EAB9FF95256EFEDAFC3503F8FC6B" box="[295,434,985,1009]" italics="true" pageId="22" pageNumber="23">H. oresterus</emphasis>
</taxonomicName>
from El Copey de Dota,
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[
<quantity id="4C8B9B4EFF95256EFED8FC1A0331FB94" box="[293,379,1014,1038]" metricMagnitude="3" metricUnit="m" metricValue="1.829" pageId="22" pageNumber="23" unit="m" value="1829.0">1829 m</quantity>
] in the
<collectingRegion id="49B7F849FF95256EFE32FC1A02FAFBB6" country="Costa Rica" name="San Jose" pageId="22" pageNumber="23">Provincia de San José</collectingRegion>
[9
<emphasis id="B907EAB9FF95256EFF33FBF80292FBB6" bold="true" box="[206,216,1044,1068]" pageId="22" pageNumber="23">°</emphasis>
399N, 83
<emphasis id="B907EAB9FF95256EFEC1FBF8030CFBB6" bold="true" box="[316,326,1044,1068]" pageId="22" pageNumber="23">°</emphasis>
559W,
<bibRefCitation id="EFE24B5AFF95256EFE68FBF8002BFBB6" author="McPherson, A. B." box="[405,609,1044,1068]" pageId="22" pageNumber="23" pagination="97 - 273" refId="ref20231" refString="McPherson, A. B. 1985. A biogeographical analysis of factors influencing the distribution of Costa Rican rodents. Brenesia 23: 97 - 273." type="journal article" year="1985">McPherson, 1985</bibRefCitation>
]. In addition to the
<typeStatus id="54C88809FF95256EFECDFBDE03DDFBD0" box="[304,407,1074,1098]" pageId="22" pageNumber="23" type="holotype">holotype</typeStatus>
, he made reference to eight other adult topotypes. The full series collected by Austin Smith in 1931 includes those
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plus one juvenal specimen (UMMZ 6402664034, 66478) collected at elevations ranging from
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[
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]. In 1933, Smith obtained another specimen (UMMZ 67306) from a nearby locality on the Caribbean side of the continental divide in the
<collectingRegion id="49B7F849FF95256EFE7EFAD100C3FACF" box="[387,649,1341,1365]" country="Costa Rica" name="Cartago" pageId="22" pageNumber="23">Provincia de Cartago</collectingRegion>
: El Muñeco, Río Navarro,
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S of
<collectingRegion id="49B7F849FF95256EFF7DFA9502AAFA0B" box="[128,224,1401,1425]" country="Costa Rica" name="Cartago" pageId="22" pageNumber="23">Cartago</collectingRegion>
,
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[
<quantity id="4C8B9B4EFF95256EFEA8FA9503E6FA0B" box="[341,428,1401,1425]" metricMagnitude="3" metricUnit="m" metricValue="1.311" pageId="22" pageNumber="23" unit="m" value="1311.0">1311 m</quantity>
; 9
<emphasis id="B907EAB9FF95256EFE36FA95039FFA0B" bold="true" box="[459,469,1401,1425]" pageId="22" pageNumber="23">°</emphasis>
489N, 83
<emphasis id="B907EAB9FF95256EFDC3FA950002FA0B" bold="true" box="[574,584,1401,1425]" pageId="22" pageNumber="23">°</emphasis>
539W,
<bibRefCitation id="EFE24B5AFF95256EFF7DFA7B0300FA35" author="McPherson, A. B." box="[128,330,1431,1455]" pageId="22" pageNumber="23" pagination="97 - 273" refId="ref20231" refString="McPherson, A. B. 1985. A biogeographical analysis of factors influencing the distribution of Costa Rican rodents. Brenesia 23: 97 - 273." type="journal article" year="1985">McPherson, 1985</bibRefCitation>
]. A half century later, Duke S. Rogers and colleagues collected additional specimens in 1981 and 1982 from the Provincia de San Jose´: 2.2 km E (by road) La
<collectingCountry id="F364763BFF95256EFF7DF9E202AEF9BC" box="[128,228,1550,1574]" name="Trinidad and Tobago" pageId="22" pageNumber="23">Trinidad</collectingCountry>
de Dota,
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[9
<emphasis id="B907EAB9FF95256EFE32F9E20393F9BC" bold="true" box="[463,473,1550,1574]" pageId="22" pageNumber="23">°</emphasis>
409N, 83
<emphasis id="B907EAB9FF95256EFDC3F9E20002F9BC" bold="true" box="[574,584,1550,1574]" pageId="22" pageNumber="23">°</emphasis>
529W,
<bibRefCitation id="EFE24B5AFF95256EFF7DF9C0035FF9DE" author="IGCR" box="[128,277,1580,1604]" pageId="22" pageNumber="23" refId="ref19434" refString="IGCR. 1963. Hoja Tapanti', escala 1: 50,000. San Jose', Costa Rica: Instituto Geografico de Costa Rica." type="book" year="1963">IGCR, 1963</bibRefCitation>
; MVZ 164860164864, 165786;
<bibRefCitation id="EFE24B5AFF95256EFF7DF9A50396F9FB" author="Mascarello, J. T. &amp; D. S. Rogers" box="[128,476,1609,1633]" pageId="22" pageNumber="23" pagination="126 - 130" refId="ref20059" refString="Mascarello, J. T., and D. S. Rogers. 1988. Banded chromosomes of Liomys salvini, Heteromys oresterus, and H. desmarestianus. Journal of Mammalogy 69: 126 - 130." type="journal article" year="1988">Mascarello and Rogers, 1988</bibRefCitation>
; Rogers, 1989, 1990; Rogers and Rogers, 1992]. We have examined the specimens from these three localities and concur that they all represent
<taxonomicName id="4C734D28FF95256EFF7DF92D0340F943" ID-CoL="7VD2T" box="[128,266,1729,1753]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="oresterus">
<emphasis id="B907EAB9FF95256EFF7DF92D0340F943" box="[128,266,1729,1753]" italics="true" pageId="22" pageNumber="23">H. oresterus</emphasis>
</taxonomicName>
(appendix 2; see also figs. 7, 10).
</paragraph>
<paragraph id="8BCC36ABFF95256EFD25FF3F012DFCA4" blockId="22.[701,1228,211,1753]" pageId="22" pageNumber="23">
However, other specimens from the
<collectingRegion id="49B7F849FF95256EFD40FF1D018BFE93" box="[701,961,241,265]" country="Costa Rica" name="San Jose" pageId="22" pageNumber="23">Provincia de San José</collectingRegion>
reported in the literature as
<taxonomicName id="4C734D28FF95256EFCE5FEE2064AFEBC" ID-CoL="7VD2T" box="[792,1024,270,294]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="oresterus">
<emphasis id="B907EAB9FF95256EFCE5FEE2064AFEBC" box="[792,1024,270,294]" italics="true" pageId="22" pageNumber="23">Heteromys oresterus</emphasis>
</taxonomicName>
do not represent that species, but rather individuals of the
<taxonomicName id="4C734D28FF95256EFB53FEC0012EFEF8" ID-CoL="7VD2N" authorityName="Gray" authorityYear="1868" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="desmarestianus">
<emphasis id="B907EAB9FF95256EFB53FEC0012EFEF8" italics="true" pageId="22" pageNumber="23">H. desmarestianus</emphasis>
</taxonomicName>
species complex (fig. 10; appendix 2). These include a series collected by Alfred L. Gardner in 1967 from Fila la Máquina, ca. 7.5 km E El Canaan,
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[
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; 9
<emphasis id="B907EAB9FF95256EFCCBFE2D010AFE43" bold="true" box="[822,832,449,473]" pageId="22" pageNumber="23">°</emphasis>
289N, 83
<emphasis id="B907EAB9FF95256EFC59FE2D01E4FE43" bold="true" box="[932,942,449,473]" pageId="22" pageNumber="23">°</emphasis>
349W;
<bibRefCitation id="EFE24B5AFF95256EFC01FE2D068DFE43" author="McPherson, A. B." box="[1020,1223,449,473]" pageId="22" pageNumber="23" pagination="97 - 273" refId="ref20231" refString="McPherson, A. B. 1985. A biogeographical analysis of factors influencing the distribution of Costa Rican rodents. Brenesia 23: 97 - 273." type="journal article" year="1985">McPherson, 1985</bibRefCitation>
; LSUMZ 1313213138] considered to be
<taxonomicName id="4C734D28FF95256EFB53FE33016BFD8F" ID-CoL="7VD2T" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="oresterus">
<emphasis id="B907EAB9FF95256EFB53FE33016BFD8F" italics="true" pageId="22" pageNumber="23">H. oresterus</emphasis>
</taxonomicName>
by
<bibRefCitation id="EFE24B5AFF95256EFCA6FE100679FD8E" author="McPherson, A. B." box="[859,1075,508,532]" pageId="22" pageNumber="23" pagination="97 - 273" refId="ref20231" refString="McPherson, A. B. 1985. A biogeographical analysis of factors influencing the distribution of Costa Rican rodents. Brenesia 23: 97 - 273." type="journal article" year="1985">McPherson (1985)</bibRefCitation>
and Rogers and Rogers (1992). Furthermore, we tentatively reidentify a single specimen collected by R.M.T. in 1986 from
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S,
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E Ojo de Agua,
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[9
<emphasis id="B907EAB9FF95256EFC75FD9F01D8FD11" bold="true" box="[904,914,627,651]" pageId="22" pageNumber="23">°</emphasis>
359N, 83
<emphasis id="B907EAB9FF95256EFC01FD9F064CFD11" bold="true" box="[1020,1030,627,651]" pageId="22" pageNumber="23">°</emphasis>
489W;
<bibRefCitation id="EFE24B5AFF95256EFBA4FD9F0112FD33" author="Ashe, J. S. &amp; R. M. Timm" pageId="22" pageNumber="23" pagination="429 - 437" refId="ref17538" refString="Ashe, J. S., and R. M. Timm. 1987 a. Predation by and activity patterns of ' parasitic' beetles of the genus Amblyopinus (Coleoptera: Staphylinidae). Journal of Zoology (London) 212: 429 - 437." type="journal article" year="1987">Ashe and Timm, 1987a</bibRefCitation>
; FMNH 128459] reported as
<taxonomicName id="4C734D28FF95256EFB53FD7D016BFD5D" ID-CoL="7VD2T" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="oresterus">
<emphasis id="B907EAB9FF95256EFB53FD7D016BFD5D" italics="true" pageId="22" pageNumber="23">H. oresterus</emphasis>
</taxonomicName>
by Rogers and Rogers (1992). These specimens lack the narrow (and dorsally flared) rostrum of
<taxonomicName id="4C734D28FF95256EFC4EFD07060DFC99" ID-CoL="7VD2T" box="[947,1095,747,771]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="oresterus">
<emphasis id="B907EAB9FF95256EFC4EFD07060DFC99" box="[947,1095,747,771]" italics="true" pageId="22" pageNumber="23">H. oresterus</emphasis>
</taxonomicName>
and have a wider interorbital constriction than that species (fig. 7).
</paragraph>
</subSubSection>
<subSubSection id="C3696520FF952560FD25FCA80653FA66" lastPageId="24" lastPageNumber="25" pageId="22" pageNumber="23" type="description">
<paragraph id="8BCC36ABFF95256FFD25FCA8005CF905" blockId="22.[701,1228,211,1753]" lastBlockId="23.[88,614,1466,1754]" lastPageId="23" lastPageNumber="24" pageId="22" pageNumber="23">
The subgeneric placement of
<taxonomicName id="4C734D28FF95256EFBAFFCA8016BFCE0" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="oresterus">
<emphasis id="B907EAB9FF95256EFBAFFCA8016BFCE0" italics="true" pageId="22" pageNumber="23">Heteromys oresterus</emphasis>
</taxonomicName>
remains controversial and unclear. Only two subgenera (
<taxonomicName id="4C734D28FF95256EFC4DFC6C0660FC02" box="[944,1066,896,920]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="22" pageNumber="23" phylum="Chordata" rank="genus">
<emphasis id="B907EAB9FF95256EFC4DFC6C0660FC02" box="[944,1066,896,920]" italics="true" pageId="22" pageNumber="23">Heteromys</emphasis>
</taxonomicName>
and
<taxonomicName id="4C734D28FF95256EFB9CFC6C0688FC02" authorityName="Merriam" authorityYear="1902" box="[1121,1218,896,920]" class="Mammalia" family="Heteromyidae" genus="Xylomys" kingdom="Animalia" order="Rodentia" pageId="22" pageNumber="23" phylum="Chordata" rank="genus">
<emphasis id="B907EAB9FF95256EFB9CFC6C0688FC02" box="[1121,1218,896,920]" italics="true" pageId="22" pageNumber="23">Xylomys</emphasis>
</taxonomicName>
) have been proposed for the genus
<taxonomicName id="4C734D28FF95256EFBB7FC72068EFC2C" box="[1098,1220,926,950]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="22" pageNumber="23" phylum="Chordata" rank="genus">
<emphasis id="B907EAB9FF95256EFBB7FC72068EFC2C" box="[1098,1220,926,950]" italics="true" pageId="22" pageNumber="23">Heteromys</emphasis>
</taxonomicName>
.
<taxonomicName id="4C734D28FF95256EFD40FC500154FC4E" authorityName="Merriam" authorityYear="1902" box="[701,798,956,980]" class="Mammalia" family="Heteromyidae" genus="Xylomys" kingdom="Animalia" order="Rodentia" pageId="22" pageNumber="23" phylum="Chordata" rank="genus">
<emphasis id="B907EAB9FF95256EFD40FC500154FC4E" box="[701,798,956,980]" italics="true" pageId="22" pageNumber="23">Xylomys</emphasis>
</taxonomicName>
was initially described by
<bibRefCitation id="EFE24B5AFF95256EFB9BFC57014FFC6B" author="Merriam, C. H." pageId="22" pageNumber="23" pagination="41 - 50" refId="ref20292" refString="Merriam, C. H. 1902. Twenty new pocket mice (Heteromys and Liomys) from Mexico. Proceedings of the Biological Society of Washington 15: 41 - 50." type="journal article" year="1902">Merriam (1902)</bibRefCitation>
for the distinctive species
<taxonomicName id="4C734D28FF95256EFBAEFC350681FC6B" box="[1107,1227,985,1009]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="nelsoni">
<emphasis id="B907EAB9FF95256EFBAEFC350681FC6B" box="[1107,1227,985,1009]" italics="true" pageId="22" pageNumber="23">H. nelsoni</emphasis>
</taxonomicName>
(which is the largest in the genus and shows extremely soft dorsal pelage).
<bibRefCitation id="EFE24B5AFF95256EFBECFBF80681FBB6" author="Goldman, E. A." box="[1041,1227,1044,1068]" pageId="22" pageNumber="23" pagination="1 - 70" refId="ref18761" refString="Goldman, E. A. 1911. Revision of the spiny pocket mice (genera Heteromys and Liomys). North American Fauna 34: 1 - 70." type="journal article" year="1911">Goldman (1911)</bibRefCitation>
listed several additional cranial characters that appeared to distinguish
<taxonomicName id="4C734D28FF95256EFC2CFBBC0678FBF2" authorityName="Merriam" authorityYear="1902" box="[977,1074,1104,1128]" class="Mammalia" family="Heteromyidae" genus="Xylomys" kingdom="Animalia" order="Rodentia" pageId="22" pageNumber="23" phylum="Chordata" rank="genus">
<emphasis id="B907EAB9FF95256EFC2CFBBC0678FBF2" box="[977,1074,1104,1128]" italics="true" pageId="22" pageNumber="23">Xylomys</emphasis>
</taxonomicName>
based on the material available at the time. Subsequently,
<bibRefCitation id="EFE24B5AFF95256EFD40FB6001A7FB3E" author="Hall, E. R. &amp; K. R. Kelson" box="[701,1005,1164,1188]" pageId="22" pageNumber="23" refId="ref19145" refString="Hall, E. R., and K. R. Kelson. 1959. The mammals of North America, vol. 1. New York: Ronald Press." type="book" year="1959">Hall and Kelson (1959)</bibRefCitation>
and
<bibRefCitation id="EFE24B5AFF95256EFBC2FB600681FB3E" author="Hall, E. R." box="[1087,1227,1164,1188]" pageId="22" pageNumber="23" refId="ref19117" refString="Hall, E. R. 1981. The mammals of North America, 2 nd ed., vol. 1. New York: Wiley." type="book" year="1981">Hall (1981)</bibRefCitation>
considered
<taxonomicName id="4C734D28FF95256EFCB4FB450197FB5B" box="[841,989,1193,1217]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="oresterus">
<emphasis id="B907EAB9FF95256EFCB4FB450197FB5B" box="[841,989,1193,1217]" italics="true" pageId="22" pageNumber="23">H. oresterus</emphasis>
</taxonomicName>
a member of the subgenus
<taxonomicName id="4C734D28FF95256EFCC7FB2B01D1FB45" authorityName="Merriam" authorityYear="1902" box="[826,923,1223,1247]" class="Mammalia" family="Heteromyidae" genus="Xylomys" kingdom="Animalia" order="Rodentia" pageId="22" pageNumber="23" phylum="Chordata" rank="genus">
<emphasis id="B907EAB9FF95256EFCC7FB2B01D1FB45" box="[826,923,1223,1247]" italics="true" pageId="22" pageNumber="23">Xylomys</emphasis>
</taxonomicName>
. However, more recent work has shown that the only morphological character that unites
<taxonomicName id="4C734D28FF95256EFC24FAEF0612FA81" box="[985,1112,1283,1307]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="nelsoni">
<emphasis id="B907EAB9FF95256EFC24FAEF0612FA81" box="[985,1112,1283,1307]" italics="true" pageId="22" pageNumber="23">H. nelsoni</emphasis>
</taxonomicName>
and
<taxonomicName id="4C734D28FF95256EFB53FAEF016BFAA3" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="oresterus">
<emphasis id="B907EAB9FF95256EFB53FAEF016BFAA3" italics="true" pageId="22" pageNumber="23">H. oresterus</emphasis>
</taxonomicName>
is their possession of soft dorsal pelage (Rogers, 1986; Rogers and Rogers, 1992; see
<bibRefCitation id="EFE24B5AFF95256EFCCAFAB006D1FAEE" author="Homan, J. A. &amp; H. H. Genoways" box="[823,1179,1372,1396]" pageId="22" pageNumber="23" pagination="740 - 760" refId="ref19337" refString="Homan, J. A., and H. H. Genoways. 1978. An analysis of hair structure and its phylogenetic implications among heteromyid rodents. Journal of Mammalogy 59: 740 - 760." type="journal article" year="1978">Homan and Genoways [1978]</bibRefCitation>
for morphology of heteromyid hair). Unfortunately, spininess often varies clinally with elevation within a species of
<taxonomicName id="4C734D28FF95256EFBCAFA5906FBFA57" box="[1079,1201,1461,1485]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="22" pageNumber="23" phylum="Chordata" rank="genus">
<emphasis id="B907EAB9FF95256EFBCAFA5906FBFA57" box="[1079,1201,1461,1485]" italics="true" pageId="22" pageNumber="23">Heteromys</emphasis>
</taxonomicName>
— spinier in the lowlands and softer at higher elevations (
<bibRefCitation id="EFE24B5AFF95256EFCB9FA1C0675F992" author="Anderson, R. P." box="[836,1087,1520,1544]" pageId="22" pageNumber="23" pagination="613 - 630" refId="ref17183" refString="Anderson, R. P. 1999 [2000]. Preliminary review of the systematics and biogeography of the spiny pocket mice (Heteromys) of Colombia. Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales 23 (suplemento especial): 613 - 630." type="book chapter" year="1999">Anderson, 1999: 619</bibRefCitation>
,
<bibRefCitation id="EFE24B5AFF95256EFBACFA1C068DF992" author="Anderson, R. P." box="[1105,1223,1520,1544]" pageId="22" pageNumber="23" pagination="1 - 43" refId="ref17272" refString="Anderson, R. P. 2003 b. Taxonomy, distribution, and natural history of the genus Heteromys (Rodentia: Heteromyidae) in western Venezuela, with the description of a dwarf species from the Peninsula de Paraguana. American Museum Novitates 3396: 1 - 43." type="journal article" year="2003">2003b: 23</bibRefCitation>
, 26; Anderson and Jarrín-V., 2002: 12). Hence, we agree with Rogers (1986) and Rogers and Rogers (1992) in doubting the taxonomic utility of this character. No modern phylogenetic study of heteromyines has succeeded in determining the evolutionary relationships of the species of
<taxonomicName id="4C734D28FF95256EFC9EF92D0197F943" box="[867,989,1729,1753]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="22" pageNumber="23" phylum="Chordata" rank="genus">
<emphasis id="B907EAB9FF95256EFC9EF92D0197F943" box="[867,989,1729,1753]" italics="true" pageId="22" pageNumber="23">Heteromys</emphasis>
</taxonomicName>
, but morphological, karyological, and genetic data suggest that
<taxonomicName id="4C734D28FF94256FFDB4FA5702F6FA6A" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="oresterus">
<emphasis id="B907EAB9FF94256FFDB4FA5702F6FA6A" italics="true" pageId="23" pageNumber="24">H. oresterus</emphasis>
</taxonomicName>
is probably more closely related to members of the
<taxonomicName id="4C734D28FF94256FFED2FA19004CF997" authorityName="Gray" authorityYear="1868" box="[303,518,1525,1549]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="desmarestianus">
<emphasis id="B907EAB9FF94256FFED2FA19004CF997" box="[303,518,1525,1549]" italics="true" pageId="23" pageNumber="24">H. desmarestianus</emphasis>
</taxonomicName>
species complex than to
<taxonomicName id="4C734D28FF94256FFECDF9FE03E3F9B0" box="[304,425,1554,1578]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="nelsoni">
<emphasis id="B907EAB9FF94256FFECDF9FE03E3F9B0" box="[304,425,1554,1578]" italics="true" pageId="23" pageNumber="24">H. nelsoni</emphasis>
</taxonomicName>
(Rogers, 1986: 186191, 1989, 1990;
<bibRefCitation id="EFE24B5AFF94256FFEAFF9C302D8F9FE" author="Mascarello, J. T. &amp; D. S. Rogers" pageId="23" pageNumber="24" pagination="126 - 130" refId="ref20059" refString="Mascarello, J. T., and D. S. Rogers. 1988. Banded chromosomes of Liomys salvini, Heteromys oresterus, and H. desmarestianus. Journal of Mammalogy 69: 126 - 130." type="journal article" year="1988">Mascarello and Rogers, 1988</bibRefCitation>
). We conclude that recognition of defensible subgenera in
<taxonomicName id="4C734D28FF94256FFE9EF9860397F918" box="[355,477,1642,1666]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="23" pageNumber="24" phylum="Chordata" rank="genus">
<emphasis id="B907EAB9FF94256FFE9EF9860397F918" box="[355,477,1642,1666]" italics="true" pageId="23" pageNumber="24">Heteromys</emphasis>
</taxonomicName>
awaits the results of future phylogenetic analyses.
</paragraph>
<caption id="DF0C6623FF94256FFF8DFB1D013DFA3D" ID-DOI="http://doi.org/10.5281/zenodo.4735336" ID-Zenodo-Dep="4735336" httpUri="https://zenodo.org/record/4735336/files/figure.png" pageId="23" pageNumber="24" startId="23.[112,150,1265,1287]" targetBox="[88,1187,210,1231]" targetPageId="23">
<paragraph id="8BCC36ABFF94256FFF8DFB1D013DFA3D" blockId="23.[88,1188,1265,1447]" pageId="23" pageNumber="24">
Fig. 10. Map showing selected collection localities for
<taxonomicName id="4C734D28FF94256FFD67FB1D0168FA9D" authority="in" authorityName="DESMARESTIANUS" authorityYear="1817" box="[666,802,1265,1287]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="23" pageNumber="23" phylum="Chordata" rank="genus">
<emphasis id="B907EAB9FF94256FFD67FB1D014DFA9C" box="[666,775,1265,1286]" italics="true" pageId="23" pageNumber="24">Heteromys</emphasis>
in
</taxonomicName>
Costa Rica. Localities are plotted for voucher specimens from karyological and genetic studies (
<bibRefCitation id="EFE24B5AFF94256FFD55FAE00185FAB8" author="Mascarello, J. T. &amp; D. S. Rogers" box="[680,975,1292,1314]" pageId="23" pageNumber="24" pagination="126 - 130" refId="ref20059" refString="Mascarello, J. T., and D. S. Rogers. 1988. Banded chromosomes of Liomys salvini, Heteromys oresterus, and H. desmarestianus. Journal of Mammalogy 69: 126 - 130." type="journal article" year="1988">Mascarello and Rogers 1988</bibRefCitation>
; Rogers 1989, 1990). In addition, localities are plotted for confirmed specimens of
<taxonomicName id="4C734D28FF94256FFD39FACB0174FAA6" box="[708,830,1319,1340]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="oresterus">
<emphasis id="B907EAB9FF94256FFD39FACB0174FAA6" box="[708,830,1319,1340]" italics="true" pageId="23" pageNumber="24">H. oresterus</emphasis>
</taxonomicName>
and specimens previously reported as that species (
<bibRefCitation id="EFE24B5AFF94256FFEE5FAAD0392FACD" author="McPherson, A. B." box="[280,472,1345,1367]" pageId="23" pageNumber="24" pagination="97 - 273" refId="ref20231" refString="McPherson, A. B. 1985. A biogeographical analysis of factors influencing the distribution of Costa Rican rodents. Brenesia 23: 97 - 273." type="journal article" year="1985">McPherson, 1985</bibRefCitation>
; Rogers and Rogers, 1992). Circles represent localities of
<taxonomicName id="4C734D28FF94256FFB74FAAD02A4FAEB" authorityName="Gray" authorityYear="1868" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="desmarestianus">
<emphasis id="B907EAB9FF94256FFB74FAAD02A4FAEB" italics="true" pageId="23" pageNumber="24">H. desmarestianus</emphasis>
</taxonomicName>
, the triangle denotes the locality of
<taxonomicName id="4C734D28FF94256FFD7DFAB00147FAEB" baseAuthorityName="Rogers" baseAuthorityYear="1989" box="[640,781,1372,1393]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="nubicolens">
<emphasis id="B907EAB9FF94256FFD7DFAB00147FAEB" box="[640,781,1372,1393]" italics="true" pageId="23" pageNumber="24">H. nubicolens</emphasis>
</taxonomicName>
, and squares indicate localities of
<taxonomicName id="4C734D28FF94256FFB74FAB002F8FA11" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="oresterus">
<emphasis id="B907EAB9FF94256FFB74FAB002F8FA11" italics="true" pageId="23" pageNumber="24">H. oresterus</emphasis>
</taxonomicName>
. See appendices 1 and 2 for full provenience and specimen numbers. Gray shading denotes regions higher than 1000 m in elevation, and areas shown in black lie above 2000 m.
</paragraph>
</caption>
<paragraph id="8BCC36ABFF942560FF8FF949061BFEBC" blockId="23.[88,614,1466,1754]" lastBlockId="24.[701,1228,211,1532]" lastPageId="24" lastPageNumber="25" pageId="23" pageNumber="24">
<taxonomicName id="4C734D28FF94256FFF8FF9490051F921" box="[114,539,1701,1725]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B907EAB9FF94256FFF8FF94903F3F921" box="[114,441,1701,1725]" italics="true" pageId="23" pageNumber="24">HETEROMYS DESMARESTIANUS</emphasis>
SPECIES
</taxonomicName>
COMPLEX: Even after recognizing
<emphasis id="B907EAB9FF94256FFE40F92E00A4FA49" box="[445,750,1467,1754]" italics="true" pageId="23" pageNumber="24">
<taxonomicName id="4C734D28FF94256FFE40F92E007DF940" authorityName="DESMARESTIANUS" authorityYear="1817" box="[445,567,1730,1754]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="23" pageNumber="24" phylum="Chordata" rank="genus">Heteromys</taxonomicName>
nubicolens
</emphasis>
as a distinct species,
<taxonomicName id="4C734D28FF94256FFC0DFA5700F1FA6A" authorityName="Gray" authorityYear="1868" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="desmarestianus">
<emphasis id="B907EAB9FF94256FFC0DFA5700F1FA6A" italics="true" pageId="23" pageNumber="24">H. desmarestianus</emphasis>
</taxonomicName>
remains a confusing complex of morphologically similar species that range from
<collectingCountry id="F364763BFF94256FFD6BF9FE00A1F9B0" box="[662,747,1554,1578]" name="Mexico" pageId="23" pageNumber="24">Mexico</collectingCountry>
to
<collectingCountry id="F364763BFF94256FFCD4F9FE01D0F9B0" box="[809,922,1554,1578]" name="Colombia" pageId="23" pageNumber="24">Colombia</collectingCountry>
and vary widely in karyology and protein allozymes. Except for
<taxonomicName id="4C734D28FF94256FFD6BF9A10159F9FF" authorityName="J. A. Allen &amp; Chapman" authorityYear="1897" box="[662,787,1613,1637]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="gaumeri">
<emphasis id="B907EAB9FF94256FFD6BF9A10159F9FF" box="[662,787,1613,1637]" italics="true" pageId="23" pageNumber="24">H. gaumeri</emphasis>
</taxonomicName>
(2n 5 56) and
<taxonomicName id="4C734D28FF94256FFC46F9A10661F9FF" box="[955,1067,1613,1637]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="nelsoni">
<emphasis id="B907EAB9FF94256FFC46F9A10661F9FF" box="[955,1067,1613,1637]" italics="true" pageId="23" pageNumber="24">H. nelsoni</emphasis>
</taxonomicName>
(2n 5 42), all
<taxonomicName id="4C734D28FF94256FFD42F9860173F918" authorityName="DESMARESTIANUS" authorityYear="1817" box="[703,825,1642,1666]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="23" pageNumber="24" phylum="Chordata" rank="genus">
<emphasis id="B907EAB9FF94256FFD42F9860173F918" box="[703,825,1642,1666]" italics="true" pageId="23" pageNumber="24">Heteromys</emphasis>
</taxonomicName>
with known karyotypes have a diploid number of 2n 5 60 (
<bibRefCitation id="EFE24B5AFF94256FFC11F96B0084F927" author="Engstrom, M. D. &amp; H. H. Genoways &amp; P. K. Tucker" pageId="23" pageNumber="24" pagination="289 - 303" refId="ref18385" refString="Engstrom, M. D., H. H. Genoways, and P. K. Tucker. 1987. Morphological variation, karyology, and systematic relationships of Heteromys gaumeri (Rodentia: Heteromyidae). In B. D. Patterson and R. M. Timm (editors), Studies in Neotropical mammalogy: essays in honor of Philip Hershkovitz. Fieldiana: Zoology (new series) 39: 289 - 303." type="journal article" year="1987">Engstrom et al., 1987</bibRefCitation>
; Rogers, 1989); however, the species with a diploid number of 2n 5 60 vary widely in fundamental number (FN). At least nine karyotypic forms exist within the
<taxonomicName id="4C734D28FF9B2560FDF3FF1D0292FEBD" authorityName="Gray" authorityYear="1868" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="desmarestianus">
<emphasis id="B907EAB9FF9B2560FDF3FF1D0292FEBD" italics="true" pageId="24" pageNumber="25">H. desmarestianus</emphasis>
</taxonomicName>
species complex (including
<taxonomicName id="4C734D28FF9B2560FDD6FEE302FCFEDE" authorityName="Merriam" authorityYear="1902" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="goldmani">
<emphasis id="B907EAB9FF9B2560FDD6FEE302FCFEDE" italics="true" pageId="24" pageNumber="25">H. goldmani</emphasis>
</taxonomicName>
and a sample of
<taxonomicName id="4C734D28FF9B2560FE65FEC00074FEDE" box="[408,574,300,324]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B907EAB9FF9B2560FE65FEC00058FEDE" box="[408,530,300,324]" italics="true" pageId="24" pageNumber="25">Heteromys</emphasis>
sp.
</taxonomicName>
from near Ciudad Nielly in southwestern
<collectingCountry id="F364763BFF9B2560FDB6FEA602FDFE1A" name="Costa Rica" pageId="24" pageNumber="25">Costa Rica</collectingCountry>
;
<bibRefCitation id="EFE24B5AFF9B2560FF35FE84006FFE1A" author="Mascarello, J. T. &amp; D. S. Rogers" box="[200,549,360,384]" pageId="24" pageNumber="25" pagination="126 - 130" refId="ref20059" refString="Mascarello, J. T., and D. S. Rogers. 1988. Banded chromosomes of Liomys salvini, Heteromys oresterus, and H. desmarestianus. Journal of Mammalogy 69: 126 - 130." type="journal article" year="1988">Mascarello and Rogers, 1988</bibRefCitation>
; Rogers, 1989; see also
<bibRefCitation id="EFE24B5AFF9B2560FEDCFE69005FFE07" author="Genoways, H. H." box="[289,533,389,413]" pageId="24" pageNumber="25" pagination="1 - 368" refId="ref18726" refString="Genoways, H. H. 1973. Systematics and evolutionary relationships of spiny pocket mice, genus Liomys. Special Publications of the Museum, Texas Tech University 5: 1 - 368." type="journal article" year="1973">Genoways, 1973: 298</bibRefCitation>
;
<bibRefCitation id="EFE24B5AFF9B2560FDDCFE6902BBFE21" author="Burton, D. W. &amp; J. W. Bickham &amp; H. H. Genoways &amp; T. J. McCarthy" pageId="24" pageNumber="25" pagination="103 - 112" refId="ref17913" refString="Burton, D. W., J. W. Bickham, H. H. Genoways, and T. J. McCarthy. 1987. Karyotypic analysis of five rodents and a marsupial from Belize, Central America. Annals of Carnegie Museum 56: 103 - 112." type="journal article" year="1987">Burton et al., 1987</bibRefCitation>
;
<bibRefCitation id="EFE24B5AFF9B2560FEF1FE4F0078FE21" author="Engstrom, M. D. &amp; H. H. Genoways &amp; P. K. Tucker" box="[268,562,419,443]" pageId="24" pageNumber="25" pagination="289 - 303" refId="ref18385" refString="Engstrom, M. D., H. H. Genoways, and P. K. Tucker. 1987. Morphological variation, karyology, and systematic relationships of Heteromys gaumeri (Rodentia: Heteromyidae). In B. D. Patterson and R. M. Timm (editors), Studies in Neotropical mammalogy: essays in honor of Philip Hershkovitz. Fieldiana: Zoology (new series) 39: 289 - 303." type="journal article" year="1987">Engstrom et al., 1987</bibRefCitation>
). Four standard karyotypes are known from populations of
<taxonomicName id="4C734D28FF9B2560FF10FE3303C1FE6D" authority="in" authorityName="DESMARESTIANUS" authorityYear="1817" box="[237,395,479,503]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="24" pageNumber="23" phylum="Chordata" rank="genus">
<emphasis id="B907EAB9FF9B2560FF10FE33032DFE6D" box="[237,359,479,503]" italics="true" pageId="24" pageNumber="25">Heteromys</emphasis>
in
</taxonomicName>
<collectingCountry id="F364763BFF9B2560FE67FE33006BFE6D" box="[410,545,479,503]" name="Costa Rica" pageId="24" pageNumber="25">Costa Rica</collectingCountry>
(fig. 10; appendices 1, 2). The cytotype F of Rogers (1989) has a FN 5 86 and was found for individuals from the
<collectingRegion id="49B7F849FF9B2560FE92FDD400C1FDCA" box="[367,651,568,592]" country="Costa Rica" name="Puntarenas" pageId="24" pageNumber="25">Provincia de Puntarenas</collectingRegion>
: Monteverde, John Campbells woods (appendix 1) that represent the species now known as
<taxonomicName id="4C734D28FF9B2560FF7DFD7D0351FD33" baseAuthorityName="Rogers" baseAuthorityYear="1989" box="[128,283,657,681]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="nubicolens">
<emphasis id="B907EAB9FF9B2560FF7DFD7D0351FD33" box="[128,283,657,681]" italics="true" pageId="24" pageNumber="25">H. nubicolens</emphasis>
</taxonomicName>
. Cytotype F also was encountered in populations of the
<taxonomicName id="4C734D28FF9B2560FE42FD4300C7FD5D" authorityName="Gray" authorityYear="1868" box="[447,653,687,711]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="desmarestianus">
<emphasis id="B907EAB9FF9B2560FE42FD4300C7FD5D" box="[447,653,687,711]" italics="true" pageId="24" pageNumber="25">H. desmarestianus</emphasis>
</taxonomicName>
species complex from the
<collectingRegion id="49B7F849FF9B2560FE10FD210343FC98" country="Costa Rica" name="Guanacaste" pageId="24" pageNumber="25">Provincia de Guanacaste</collectingRegion>
: 4.1
<quantity id="4C8B9B4EFF9B2560FEBAFD0603CBFC98" box="[327,385,746,770]" metricMagnitude="3" metricUnit="m" metricValue="5.0" pageId="24" pageNumber="25" unit="km" value="5.0">5 km</quantity>
NE (by road) Tilarán,
<quantity id="4C8B9B4EFF9B2560FF7DFCE4028CFCBA" box="[128,198,776,800]" metricMagnitude="2" metricUnit="m" metricValue="6.5" pageId="24" pageNumber="25" unit="m" value="650.0">650 m</quantity>
(appendix 1) in northwestern
<collectingCountry id="F364763BFF9B2560FDB6FCE402FFFCA4" name="Costa Rica" pageId="24" pageNumber="25">Costa Rica</collectingCountry>
and from the
<collectingRegion id="49B7F849FF9B2560FE76FCCA00C0FCA4" box="[395,650,806,830]" country="Costa Rica" name="Limon" pageId="24" pageNumber="25">Provincia de Limón</collectingRegion>
: 4.6 km W (by road)
<collectingRegion id="49B7F849FF9B2560FE78FCA8039EFCC6" box="[389,468,836,860]" country="Costa Rica" name="Limon" pageId="24" pageNumber="25">Limón</collectingRegion>
,
<quantity id="4C8B9B4EFF9B2560FE1AFCA8006AFCC6" box="[487,544,836,860]" metricMagnitude="1" metricUnit="m" metricValue="2.5" pageId="24" pageNumber="25" unit="m" value="25.0">25 m</quantity>
[9
<emphasis id="B907EAB9FF9B2560FDBEFCA80007FCC6" bold="true" box="[579,589,836,860]" pageId="24" pageNumber="25">°</emphasis>
599N,
<geoCoordinate id="EE47506CFF9B2560FF7DFC8D02AFFCE3" box="[128,229,865,889]" degrees="83" direction="west" minutes="049" orientation="longitude" pageId="24" pageNumber="25" precision="925" value="-83.816666">
83
<emphasis id="B907EAB9FF9B2560FF67FC8D02EEFCE3" bold="true" box="[154,164,865,889]" pageId="24" pageNumber="25">°</emphasis>
049W
</geoCoordinate>
,
<bibRefCitation id="EFE24B5AFF9B2560FF0AFC8D033DFCE3" author="IGN" box="[247,375,865,889]" pageId="24" pageNumber="25" refId="ref19631" refString="IGN. 1978. Hoja Rio Banano, escala 1: 50,000, 2 nd ed. San Jose', Costa Rica: Instituto Geografico Nacional." type="book" year="1978">IGN, 1978</bibRefCitation>
] in southeastern
<collectingCountry id="F364763BFF9B2560FDB6FC8D02FFFC0D" name="Costa Rica" pageId="24" pageNumber="25">Costa Rica</collectingCountry>
(Rogers, 1989). The second variant, cytotype G of Rogers (1989), corresponds to samples of the
<taxonomicName id="4C734D28FF9B2560FEAFFC570061FC49" authorityName="Gray" authorityYear="1868" box="[338,555,955,979]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="desmarestianus">
<emphasis id="B907EAB9FF9B2560FEAFFC570061FC49" box="[338,555,955,979]" italics="true" pageId="24" pageNumber="25">H. desmarestianus</emphasis>
</taxonomicName>
species complex from the
<collectingRegion id="49B7F849FF9B2560FEA4FC34001CFC6A" box="[345,598,984,1008]" country="Costa Rica" name="Cartago" pageId="24" pageNumber="25">Provincia de Cartago</collectingRegion>
: Río Reventazón, 5.6 km SE (by road) Turrialba,
<quantity id="4C8B9B4EFF9B2560FF7DFBF8028CFBB6" box="[128,198,1044,1068]" metricMagnitude="2" metricUnit="m" metricValue="4.5" pageId="24" pageNumber="25" unit="m" value="450.0">450 m</quantity>
[9
<emphasis id="B907EAB9FF9B2560FF0CFBF802B1FBB6" bold="true" box="[241,251,1044,1068]" pageId="24" pageNumber="25">°</emphasis>
539N, 83
<emphasis id="B907EAB9FF9B2560FE91FBF8033CFBB6" bold="true" box="[364,374,1044,1068]" pageId="24" pageNumber="25">°</emphasis>
399W,
<bibRefCitation id="EFE24B5AFF9B2560FE2FFBF80011FBB6" author="IGN" box="[466,603,1044,1068]" pageId="24" pageNumber="25" refId="ref19684" refString="IGN. 1981. Hoja Tucurrique, escala 1: 50,000, 2 nd ed. San Jose', Costa Rica: Instituto Geografico Nacional." type="book" year="1981">IGN, 1981</bibRefCitation>
] in southeastern
<collectingCountry id="F364763BFF9B2560FEEAFBDE03D3FBD0" box="[279,409,1074,1098]" name="Costa Rica" pageId="24" pageNumber="25">Costa Rica</collectingCountry>
, and it displays a FN 5 80. Individuals of the
<taxonomicName id="4C734D28FF9B2560FE46FBBC00C7FBF2" authorityName="Gray" authorityYear="1868" box="[443,653,1104,1128]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="desmarestianus">
<emphasis id="B907EAB9FF9B2560FE46FBBC00C7FBF2" box="[443,653,1104,1128]" italics="true" pageId="24" pageNumber="25">H. desmarestianus</emphasis>
</taxonomicName>
species complex (
<taxonomicName id="4C734D28FF9B2560FEA8FB8103B4FB1F" box="[341,510,1133,1157]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B907EAB9FF9B2560FEA8FB810385FB1F" box="[341,463,1133,1157]" italics="true" pageId="24" pageNumber="25">Heteromys</emphasis>
sp.
</taxonomicName>
of Rogers, 1989) from the
<collectingRegion id="49B7F849FF9B2560FEABFB6700C1FB39" box="[342,651,1163,1187]" country="Costa Rica" name="Puntarenas" pageId="24" pageNumber="25">Provincia de Puntarenas</collectingRegion>
: 1.1 km SE (by road) Ciudad Nielly,
<quantity id="4C8B9B4EFF9B2560FDA9FB4500C4FB5B" box="[596,654,1193,1217]" metricMagnitude="1" metricUnit="m" metricValue="2.5" pageId="24" pageNumber="25" unit="m" value="25.0">25 m</quantity>
[8
<emphasis id="B907EAB9FF9B2560FF69FB2A02D4FB44" bold="true" box="[148,158,1222,1246]" pageId="24" pageNumber="25">°</emphasis>
399N, 82
<emphasis id="B907EAB9FF9B2560FEFDFB2A0340FB44" bold="true" box="[256,266,1222,1246]" pageId="24" pageNumber="25">°</emphasis>
579W,
<bibRefCitation id="EFE24B5AFF9B2560FEA8FB2A0385FB44" author="IGN" box="[341,463,1222,1246]" pageId="24" pageNumber="25" refId="ref19658" refString="IGN. 1980. Hoja Canoas, escala 1: 50,000, 2 nd ed. San Jose', Costa Rica: Instituto Geografico Nacional." type="book" year="1980">IGN, 1980</bibRefCitation>
] in southwestern
<collectingCountry id="F364763BFF9B2560FF7DFB08034FFB66" box="[128,261,1252,1276]" name="Costa Rica" pageId="24" pageNumber="25">Costa Rica</collectingCountry>
constitute the third moiety and show a FN 5 90 (Rogers, 1989). By comparison, the karyotype of
<taxonomicName id="4C734D28FF9B2560FE6DFACC0056FAA2" box="[400,540,1312,1336]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="oresterus">
<emphasis id="B907EAB9FF9B2560FE6DFACC0056FAA2" box="[400,540,1312,1336]" italics="true" pageId="24" pageNumber="25">H. oresterus</emphasis>
</taxonomicName>
from the Provincia de San Jose´: 2.2 km E (by road) La
<collectingCountry id="F364763BFF9B2560FF7DFAB702AEFAE9" box="[128,228,1371,1395]" name="Trinidad and Tobago" pageId="24" pageNumber="25">Trinidad</collectingCountry>
de Dota,
<quantity id="4C8B9B4EFF9B2560FE8BFAB70380FAE9" box="[374,458,1371,1395]" metricMagnitude="3" metricUnit="m" metricValue="2.6" pageId="24" pageNumber="25" unit="m" value="2600.0">2600 m</quantity>
(see above) in central
<collectingCountry id="F364763BFF9B2560FF24FA950316FA0B" box="[217,348,1401,1425]" name="Costa Rica" pageId="24" pageNumber="25">Costa Rica</collectingCountry>
has a FN 5 78 (Rogers, 1989). In addition to these differences in fundamental number, individuals of the
<taxonomicName id="4C734D28FF9B2560FD8DFA59036DFA71" authorityName="Gray" authorityYear="1868" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="desmarestianus">
<emphasis id="B907EAB9FF9B2560FD8DFA59036DFA71" italics="true" pageId="24" pageNumber="25">H. desmarestianus</emphasis>
</taxonomicName>
species complex from near Ciudad Nielly and from near Tilarán (see above) also differed strongly in karyological banding patterns (
<bibRefCitation id="EFE24B5AFF9B2560FE9BF9C002F2F9FB" author="Mascarello, J. T. &amp; D. S. Rogers" pageId="24" pageNumber="25" pagination="126 - 130" refId="ref20059" refString="Mascarello, J. T., and D. S. Rogers. 1988. Banded chromosomes of Liomys salvini, Heteromys oresterus, and H. desmarestianus. Journal of Mammalogy 69: 126 - 130." type="journal article" year="1988">Mascarello and Rogers, 1988</bibRefCitation>
; Rogers, 1989). While karyological data alone cannot be used to elucidate species boundaries accurately (note similarity in standard karyotype between
<taxonomicName id="4C734D28FF9B2560FE16F94F00C7F921" baseAuthorityName="Rogers" baseAuthorityYear="1989" box="[491,653,1699,1723]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="nubicolens">
<emphasis id="B907EAB9FF9B2560FE16F94F00C7F921" box="[491,653,1699,1723]" italics="true" pageId="24" pageNumber="25">H. nubicolens</emphasis>
</taxonomicName>
and some populations of the
<emphasis id="B907EAB9FF9B2560FE26F92D00A8FF71" box="[475,738,211,1753]" italics="true" pageId="24" pageNumber="25">H. desmarestianus</emphasis>
species complex), these data reflect substantial cytogenetic discontinuities within the
<taxonomicName id="4C734D28FF9B2560FD40FEE201C0FEBC" authorityName="Gray" authorityYear="1868" box="[701,906,270,294]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="desmarestianus">
<emphasis id="B907EAB9FF9B2560FD40FEE201C0FEBC" box="[701,906,270,294]" italics="true" pageId="24" pageNumber="25">H. desmarestianus</emphasis>
</taxonomicName>
species complex.
</paragraph>
<paragraph id="8BCC36ABFF9B2560FD25FEC00653FA66" blockId="24.[701,1228,211,1532]" pageId="24" pageNumber="25">
Members of the
<taxonomicName id="4C734D28FF9B2560FC62FEC00681FEDE" authorityName="Gray" authorityYear="1868" box="[927,1227,300,324]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="desmarestianus">
<emphasis id="B907EAB9FF9B2560FC62FEC00681FEDE" box="[927,1227,300,324]" italics="true" pageId="24" pageNumber="25">Heteromys desmarestianus</emphasis>
</taxonomicName>
species complex also show extensive variation in protein allozymes (Rogers, 1990; based on many of the same individuals as the karyological studies cited above; fig. 10; appendices 1, 2). Two samples from southwestern
<collectingCountry id="F364763BFF9B2560FB74FE5300B8FE6F" name="Costa Rica" pageId="24" pageNumber="25">Costa Rica</collectingCountry>
referred to as
<taxonomicName id="4C734D28FF9B2560FC41FE31062CFE6F" box="[956,1126,477,501]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B907EAB9FF9B2560FC41FE31067CFE6F" box="[956,1078,477,501]" italics="true" pageId="24" pageNumber="25">Heteromys</emphasis>
sp.
</taxonomicName>
showed a strong divergence in allozymes from other samples of the
<taxonomicName id="4C734D28FF9B2560FC6DFDF40623FDAA" authorityName="Gray" authorityYear="1868" box="[912,1129,536,560]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="desmarestianus">
<emphasis id="B907EAB9FF9B2560FC6DFDF40623FDAA" box="[912,1129,536,560]" italics="true" pageId="24" pageNumber="25">H. desmarestianus</emphasis>
</taxonomicName>
species complex:
<collectingRegion id="49B7F849FF9B2560FCCBFDD9062CFDD7" box="[822,1126,565,589]" country="Costa Rica" name="Puntarenas" pageId="24" pageNumber="25">Provincia de Puntarenas</collectingRegion>
: 1.1 km SE (by road) Ciudad Nielly,
<quantity id="4C8B9B4EFF9B2560FBF6FDBF060FFDF1" box="[1035,1093,595,619]" metricMagnitude="1" metricUnit="m" metricValue="2.5" pageId="24" pageNumber="25" unit="m" value="25.0">25 m</quantity>
(see above; locality 16 of Rogers, 1990); Provincia de San Jose´: 16.3 km SE (by road) San Isidro,
<quantity id="4C8B9B4EFF9B2560FB78FD620686FD3C" box="[1157,1228,654,678]" metricMagnitude="2" metricUnit="m" metricValue="5.25" pageId="24" pageNumber="25" unit="m" value="525.0">525 m</quantity>
[9
<emphasis id="B907EAB9FF9B2560FD2FFD470096FD59" bold="true" box="[722,732,683,707]" pageId="24" pageNumber="25">°</emphasis>
159N, 83
<emphasis id="B907EAB9FF9B2560FCBEFD470107FD59" bold="true" box="[835,845,683,707]" pageId="24" pageNumber="25">°</emphasis>
389W,
<bibRefCitation id="EFE24B5AFF9B2560FC63FD470655FD59" author="IGN" box="[926,1055,683,707]" pageId="24" pageNumber="25" refId="ref19583" refString="IGN. 1970. Hoja Talamanca, escala 1: 200,000. San Jose', Costa Rica: Instituto Geografico Nacional." type="book" year="1970">IGN, 1970</bibRefCitation>
; locality 17 of Rogers, 1990]. These two samples were most similar to a sample of two individuals from eastern
<collectingCountry id="F364763BFF9B2560FCE4FCE8013CFC86" box="[793,886,772,796]" name="Panama" pageId="24" pageNumber="25">Panama</collectingCountry>
in the Provincia del
<collectingRegion id="49B7F849FF9B2560FB88FCE80682FC86" box="[1141,1224,772,796]" country="Panama" name="Darien" pageId="24" pageNumber="25">Darién</collectingRegion>
: ca.
<quantity id="4C8B9B4EFF9B2560FD10FCCD0162FCA3" box="[749,808,801,825]" metricMagnitude="3" metricUnit="m" metricValue="6.0" pageId="24" pageNumber="25" unit="km" value="6.0">6 km</quantity>
NW Cana, E. slope Cerro Pirre,
<quantity id="4C8B9B4EFF9B2560FD40FCD2015BFCCC" box="[701,785,830,854]" metricMagnitude="3" metricUnit="m" metricValue="1.4" pageId="24" pageNumber="25" unit="m" value="1400.0">1400 m</quantity>
[7
<emphasis id="B907EAB9FF9B2560FCC0FCD2010DFCCC" bold="true" box="[829,839,830,854]" pageId="24" pageNumber="25">°</emphasis>
519N, 77
<emphasis id="B907EAB9FF9B2560FC47FCD2018EFCCC" bold="true" box="[954,964,830,854]" pageId="24" pageNumber="25">°</emphasis>
449W,
<bibRefCitation id="EFE24B5AFF9B2560FBDCFCD20121FCEE" author="Fairchild, G. B. &amp; C. O. Handley, Jr." pageId="24" pageNumber="25" pagination="9 - 22" refId="ref18465" refString="Fairchild, G. B., and C. O. Handley, Jr. 1966. Gazetteer of collecting localities in Panama. In R. L. Wenzel and V. J. Tipton (editors), Ectoparasites of Panama: 9 - 22. Chicago: Field Museum of Natural History." type="book chapter" year="1966">Fairchild and Handley, 1966</bibRefCitation>
; locality 26 of Rogers, 1990; there considered
<taxonomicName id="4C734D28FF9B2560FC6DFC960651FC08" baseAuthorityName="Anderson" baseAuthorityYear="1999" box="[912,1051,890,914]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="australis">
<emphasis id="B907EAB9FF9B2560FC6DFC960651FC08" box="[912,1051,890,914]" italics="true" pageId="24" pageNumber="25">H. australis</emphasis>
</taxonomicName>
]. These specimens from
<collectingCountry id="F364763BFF9B2560FCC3FC7B01D1FC35" box="[830,923,919,943]" name="Panama" pageId="24" pageNumber="25">Panama</collectingCountry>
represent a member of the
<taxonomicName id="4C734D28FF9B2560FD40FC5901C4FC57" authorityName="Gray" authorityYear="1868" box="[701,910,949,973]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="desmarestianus">
<emphasis id="B907EAB9FF9B2560FD40FC5901C4FC57" box="[701,910,949,973]" italics="true" pageId="24" pageNumber="25">H. desmarestianus</emphasis>
</taxonomicName>
species complex and were considered
<emphasis id="B907EAB9FF9B2560FCBAFC3E0655FC70" box="[839,1055,978,1002]" italics="true" pageId="24" pageNumber="25">
H. d.
<taxonomicName id="4C734D28FF9B2560FC67FC3E0655FC70" box="[922,1055,978,1002]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="crassirostris">crassirostris</taxonomicName>
</emphasis>
by
<bibRefCitation id="EFE24B5AFF9B2560FBA1FC3E0142FB9D" author="Anderson, R. P." pageId="24" pageNumber="25" pagination="613 - 630" refId="ref17183" refString="Anderson, R. P. 1999 [2000]. Preliminary review of the systematics and biogeography of the spiny pocket mice (Heteromys) of Colombia. Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales 23 (suplemento especial): 613 - 630." type="book chapter" year="1999">Anderson (1999)</bibRefCitation>
. A single specimen from a slightly lower elevation (Provincia del
<collectingRegion id="49B7F849FF9B2560FC2FFBE1066FFBBF" box="[978,1061,1037,1061]" country="Panama" name="Darien" pageId="24" pageNumber="25">Darién</collectingRegion>
: ca.
<quantity id="4C8B9B4EFF9B2560FBA4FBE106DEFBBF" box="[1113,1172,1037,1061]" metricMagnitude="3" metricUnit="m" metricValue="6.0" pageId="24" pageNumber="25" unit="km" value="6.0">6 km</quantity>
NW Cana, E. slope Cerro Pirre,
<quantity id="4C8B9B4EFF9B2560FC08FBC60606FBD8" box="[1013,1100,1066,1090]" metricMagnitude="3" metricUnit="m" metricValue="1.2" pageId="24" pageNumber="25" unit="m" value="1200.0">1200 m</quantity>
; locality 27 of Rogers, 1990; there considered
<taxonomicName id="4C734D28FF9B2560FBA1FBA400A3FBE7" authorityName="Thompson" authorityYear="1815" class="Mammalia" family="Muridae" genus="Mus" kingdom="Animalia" order="Rodentia" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="anomalus">
<emphasis id="B907EAB9FF9B2560FBA1FBA400A3FBE7" italics="true" pageId="24" pageNumber="25">H. anomalus</emphasis>
</taxonomicName>
) represents true
<taxonomicName id="4C734D28FF9B2560FC45FB8900B5FB01" authority="(Anderson, 1999)" baseAuthorityName="Anderson" baseAuthorityYear="1999" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="australis">
<emphasis id="B907EAB9FF9B2560FC45FB890675FBE7" box="[952,1087,1125,1149]" italics="true" pageId="24" pageNumber="25">H. australis</emphasis>
(
<bibRefCitation id="EFE24B5AFF9B2560FBA9FB8900BDFB01" author="Anderson, R. P." pageId="24" pageNumber="25" pagination="613 - 630" refId="ref17183" refString="Anderson, R. P. 1999 [2000]. Preliminary review of the systematics and biogeography of the spiny pocket mice (Heteromys) of Colombia. Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales 23 (suplemento especial): 613 - 630." type="book chapter" year="1999">Anderson, 1999</bibRefCitation>
)
</taxonomicName>
. Several species-level taxa described based on material from
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and
<collectingCountry id="F364763BFF9B2560FD40FB520150FB4C" box="[701,794,1214,1238]" name="Panama" pageId="24" pageNumber="25">Panama</collectingCountry>
currently are considered synonyms of
<taxonomicName id="4C734D28FF9B2560FD0EFB37019CFB69" authorityName="Gray" authorityYear="1868" box="[755,982,1243,1267]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="desmarestianus">
<emphasis id="B907EAB9FF9B2560FD0EFB37019CFB69" box="[755,982,1243,1267]" italics="true" pageId="24" pageNumber="25">H. desmarestianus</emphasis>
</taxonomicName>
(
<bibRefCitation id="EFE24B5AFF9B2560FC00FB37068DFB69" author="Goodwin, G. G." box="[1021,1223,1243,1267]" pageId="24" pageNumber="25" pagination="271 - 473" refId="ref18791" refString="Goodwin, G. G. 1946. Mammals of Costa Rica. Bulletin of the American Museum of Natural History 87: 271 - 473." type="journal article" year="1946">Goodwin, 1946</bibRefCitation>
; Williams et al., 1993). Detailed specimenbased revisionary studies of alpha-level taxonomy are necessary to determine the true species limits in the
<taxonomicName id="4C734D28FF9B2560FC5AFABD063EFAF3" authorityName="Gray" authorityYear="1868" box="[935,1140,1361,1385]" class="Mammalia" family="Heteromyidae" genus="Heteromys" kingdom="Animalia" order="Rodentia" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="desmarestianus">
<emphasis id="B907EAB9FF9B2560FC5AFABD063EFAF3" box="[935,1140,1361,1385]" italics="true" pageId="24" pageNumber="25">H. desmarestianus</emphasis>
</taxonomicName>
species complex in central and eastern
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(and throughout
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), clarify the synonymy of named forms, and determine the nomenclatural status of the species found near Ciudad Nielly and San Isidro.
</paragraph>
</subSubSection>
</treatment>
</document>