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<mods:title id="07971BF471964B531A3D54C7D6CF0B0F">Cytotaxonomy and DNA taxonomy of lizards (Squamata, Sauria) from a tropical dry forest in the Chamela-Cuixmala Biosphere Reserve on the coast of Jalisco, Mexico</mods:title>
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<mods:namePart id="19451326AA93B1E9F2A2B686B6EEE8A8">Castiglia, Riccardo</mods:namePart>
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<mods:namePart id="251CD8D226D2F252FC5E1F6A638C6CF8">Bezerra, Alexandra M. R.</mods:namePart>
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<mods:namePart id="53C36E15F936C0D20A6CDB55DF96E399">García, Andrés</mods:namePart>
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<treatment id="03E2993FFFECF441FF48FF617BECFAC8" ID-DOI="http://doi.org/10.5281/zenodo.6210582" ID-GBIF-Taxon="119388425" ID-Zenodo-Dep="6210582" LSID="urn:lsid:plazi:treatment:03E2993FFFECF441FF48FF617BECFAC8" httpUri="http://treatment.plazi.org/id/03E2993FFFECF441FF48FF617BECFAC8" lastPageNumber="16" pageId="15" pageNumber="16">
<subSubSection id="C3517BA2FFECF441FF48FF617C15FEE3" pageId="15" pageNumber="16" type="nomenclature">
<paragraph id="8BF42829FFECF441FF48FF617C7AFF4D" blockId="15.[151,803,151,177]" box="[151,803,151,177]" pageId="15" pageNumber="16">
<heading id="D0BC9F45FFECF441FF48FF617C7AFF4D" bold="true" box="[151,803,151,177]" fontSize="11" level="1" pageId="15" pageNumber="16" reason="1">
<emphasis id="B93FF43BFFECF441FF48FF617C7AFF4D" bold="true" box="[151,803,151,177]" pageId="15" pageNumber="16">
<taxonomicName id="4C4B53AAFFECF441FF48FF617D4EFF4D" authority="Dumeril" authorityName="Dumeril" box="[151,535,151,177]" class="Reptilia" family="Phrynosomatidae" genus="Urosaurus" kingdom="Animalia" order="Squamata" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="bicarinatus">
<emphasis id="B93FF43BFFECF441FF48FF617EF1FF4D" bold="true" box="[151,424,151,177]" italics="true" pageId="15" pageNumber="16">Urosaurus bicarinatus</emphasis>
Duméril
</taxonomicName>
(Tropical tree lizard)
</emphasis>
</heading>
</paragraph>
<paragraph id="8BF42829FFECF441FF48FF297C15FEE3" blockId="15.[151,1437,223,1332]" pageId="15" pageNumber="16">
<emphasis id="B93FF43BFFECF441FF48FF297EC5FF05" bold="true" box="[151,412,223,249]" pageId="15" pageNumber="16">Specimens analysed:</emphasis>
one male from Chamela (CEAC19), one specimen from Rio Grande, Oaxaca (MZFC 12046), one specimen from Epatlan, Puebla (MZFC 6863).
</paragraph>
</subSubSection>
<subSubSection id="C3517BA2FFECF441FF1BFEDC7AC1FE03" pageId="15" pageNumber="16" type="distribution">
<paragraph id="8BF42829FFECF441FF1BFEDC7BC9FEB8" blockId="15.[151,1437,223,1332]" box="[196,1168,298,324]" pageId="15" pageNumber="16">
<emphasis id="B93FF43BFFECF441FF1BFEDC7E3EFEB8" bold="true" box="[196,359,298,324]" pageId="15" pageNumber="16">Distribution:</emphasis>
Mexican endemic. Pacific coast of
<collectingCountry id="F35C68B9FFECF441FCD6FEDC7C3FFEB8" box="[777,870,298,324]" name="Mexico" pageId="15" pageNumber="16">Mexico</collectingCountry>
, from Sonora to Chiapas.
</paragraph>
<paragraph id="8BF42829FFECF441FF1BFEB97AC1FE03" blockId="15.[151,1437,223,1332]" pageId="15" pageNumber="16">
<emphasis id="B93FF43BFFECF441FF1BFEB97E00FE95" bold="true" box="[196,345,335,361]" pageId="15" pageNumber="16">Subspecies:</emphasis>
<emphasis id="B93FF43BFFECF441FEBDFEA67D68FE95" box="[354,561,336,361]" italics="true" pageId="15" pageNumber="16">
U. b.
<taxonomicName id="4C4B53AAFFECF441FE78FEA67D68FE95" box="[423,561,336,361]" class="Reptilia" family="Phrynosomatidae" genus="Urosaurus" kingdom="Animalia" order="Squamata" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="bicarinatus">bicarinatus</taxonomicName>
</emphasis>
, distributed from Michoacán to central Guerrero, and in the Río Balsas basin up to Morelos and southern Puebla;
<emphasis id="B93FF43BFFECF441FD57FE837C20FE72" box="[648,889,373,398]" italics="true" pageId="15" pageNumber="16">U. b. anonymorphus</emphasis>
, found in east Guerrero, Oaxaca, and possibly western Chiapas;
<emphasis id="B93FF43BFFECF441FEADFE6C7D56FE4F" box="[370,527,410,435]" italics="true" pageId="15" pageNumber="16">U. b. nelsoni</emphasis>
, localized in northern Oaxaca;
<emphasis id="B93FF43BFFECF441FC4BFE6C7B2BFE4F" box="[916,1138,410,435]" italics="true" pageId="15" pageNumber="16">U. b. tuberculatus</emphasis>
, distributed in Southern Sonora southward to Jalisco and Colima with isolated populations in Sinaloa;
<emphasis id="B93FF43BFFECF441FB4BFE367A14FE25" box="[1172,1357,448,473]" italics="true" pageId="15" pageNumber="16">U. b. spinosus</emphasis>
, from southwestern Chiapas. However,
<bibRefCitation id="EFDA55D8FFECF441FDC0FE137DE7FE03" author="Wiens" box="[543,702,485,511]" pageId="15" pageNumber="29" refString="Wiens, J. J. (1993) Phylogenetic Systematics of the Tree Lizards (genus Urosaurus). Herpetologica, 49, 399 - 420." type="journal article" year="1993">Wiens (1993)</bibRefCitation>
did not find morphological differences among the subspecies.
</paragraph>
</subSubSection>
<subSubSection id="C3517BA2FFECF441FF1BFDFC7BECFAC8" pageId="15" pageNumber="16" type="description">
<paragraph id="8BF42829FFECF441FF1BFDFC7BC0FDD8" blockId="15.[151,1437,223,1332]" box="[196,1177,522,548]" pageId="15" pageNumber="16">
<emphasis id="B93FF43BFFECF441FF1BFDFC7E0AFDD8" bold="true" box="[196,339,522,548]" pageId="15" pageNumber="16">Karyotype:</emphasis>
Unfortunately, we did not obtained good metaphases from this species.
</paragraph>
<paragraph id="8BF42829FFECF441FF1BFDD97A48FBD3" blockId="15.[151,1437,223,1332]" pageId="15" pageNumber="16">
<emphasis id="B93FF43BFFECF441FF1BFDD97ECBFDB5" bold="true" box="[196,402,559,585]" pageId="15" pageNumber="16">DNA taxonomy:</emphasis>
There is no sequence deposited in GenBank for this species. The available rDNA 16S sequences in GenBank are for
<taxonomicName id="4C4B53AAFFECF441FE22FDA37D23FD92" box="[509,634,597,622]" class="Reptilia" family="Phrynosomatidae" genus="Urosaurus" kingdom="Animalia" order="Squamata" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="ornatus">
<emphasis id="B93FF43BFFECF441FE22FDA37D23FD92" box="[509,634,597,622]" italics="true" pageId="15" pageNumber="16">U. ornatus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C4B53AAFFECF441FD56FDA37C60FD92" box="[649,825,597,622]" class="Reptilia" family="Phrynosomatidae" genus="Urosaurus" kingdom="Animalia" order="Squamata" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="nigricaudus">
<emphasis id="B93FF43BFFECF441FD56FDA37C60FD92" box="[649,825,597,622]" italics="true" pageId="15" pageNumber="16">U. nigricaudus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C4B53AAFFECF441FC97FDA37B56FD92" box="[840,1039,597,622]" class="Reptilia" family="Phrynosomatidae" genus="Urosaurus" kingdom="Animalia" order="Squamata" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="microscutatus">
<emphasis id="B93FF43BFFECF441FC97FDA37B56FD92" box="[840,1039,597,622]" italics="true" pageId="15" pageNumber="16">U. microscutatus</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C4B53AAFFECF441FB90FDA37AC1FD93" authority="Reeder 1995" authorityName="Reeder" authorityYear="1995" box="[1103,1432,597,623]" class="Reptilia" family="Phrynosomatidae" genus="Urosaurus" kingdom="Animalia" order="Squamata" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="graciosus">
<emphasis id="B93FF43BFFECF441FB90FDA37BBCFD92" box="[1103,1253,597,622]" italics="true" pageId="15" pageNumber="16">U. graciosus</emphasis>
(
<bibRefCitation id="EFDA55D8FFECF441FB2BFDA37AD6FD93" author="Reeder" box="[1268,1423,597,623]" pageId="15" pageNumber="29" refString="Reeder, T. W. (1995) Phylogenetic relationships among phrynosomatid lizards as inferred from mitochondrial ribosomal DNA sequences: substitutional bias and information content of transitions relative to transversions. Molecular Phylogenetics and Evolution, 4, 203 - 222." type="journal article" year="1995">Reeder 1995</bibRefCitation>
)
</taxonomicName>
. We aligned these sequences with the sequence of
<taxonomicName id="4C4B53AAFFECF441FCFBFD8C7CBBFD6F" box="[804,994,634,659]" class="Reptilia" family="Phrynosomatidae" genus="Urosaurus" kingdom="Animalia" order="Squamata" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="bicarinatus">
<emphasis id="B93FF43BFFECF441FCFBFD8C7CBBFD6F" box="[804,994,634,659]" italics="true" pageId="15" pageNumber="16">U. bicarinatus</emphasis>
</taxonomicName>
from Chamela belonging to
<emphasis id="B93FF43BFFECF441FA83FD8C7E73FD45" italics="true" pageId="15" pageNumber="16">U. b. tuberculatus</emphasis>
and with sequences from two additional individuals (Rio Grande, Oaxaca and Epatlan, Puebla) possibly belonging to
<emphasis id="B93FF43BFFECF441FE41FD337D6EFD22" box="[414,567,709,734]" italics="true" pageId="15" pageNumber="16">U. b. nelsoni</emphasis>
and performed a phylogenetic analysis using
<taxonomicName id="4C4B53AAFFECF441FB86FD337A09FD22" box="[1113,1360,709,734]" class="Reptilia" family="Phrynosomatidae" genus="Sceloporus" kingdom="Animalia" order="Squamata" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="utiformis">
<emphasis id="B93FF43BFFECF441FB86FD337A09FD22" box="[1113,1360,709,734]" italics="true" pageId="15" pageNumber="16">Sceloporus utiformis</emphasis>
</taxonomicName>
as the outgroup. The obtained tree is shown in
<figureCitation id="137034ACFFECF441FDABFD1C7D80FCF8" box="[628,729,746,772]" captionStart="FIGURE 8" captionStartId="14.[151,257,1441,1465]" captionTargetBox="[226,1361,975,1406]" captionTargetId="figure@14.[174,1412,927,1420]" captionTargetPageId="14" captionText="FIGURE 8. Neighbour-Joining tree of 16 S rDNA haplotypes (455 bp) of Urosaurus species. Bootstrap values (%) obtained by the NJ, ML and MP are shown. The substitution model selected for ML was Tamura-Nei model (Tamura &amp; Nei 1993) with rate variation among sites (+ G), and a gamma distribution shape parameter of 0.0855." httpUri="https://zenodo.org/record/196014/files/figure.png" pageId="15" pageNumber="16">Figure 8</figureCitation>
. Interestingly, the phylogenetic relationships among species are different from those identified using morphological characters by
<bibRefCitation id="EFDA55D8FFECF441FC02FCE67BD9FCD6" author="Wiens" box="[989,1152,784,810]" pageId="15" pageNumber="29" refString="Wiens, J. J. (1993) Phylogenetic Systematics of the Tree Lizards (genus Urosaurus). Herpetologica, 49, 399 - 420." type="journal article" year="1993">Wiens (1993)</bibRefCitation>
and are congruent with Reeders (1995) results. Molecular analysis shows that
<taxonomicName id="4C4B53AAFFECF441FCFEFCC37C93FCB2" box="[801,970,821,846]" class="Reptilia" family="Phrynosomatidae" genus="Urosaurus" kingdom="Animalia" order="Squamata" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="bicarinatus">
<emphasis id="B93FF43BFFECF441FCFEFCC37C93FCB2" box="[801,970,821,846]" italics="true" pageId="15" pageNumber="16">U. bicarinatus</emphasis>
</taxonomicName>
has an external position with respect to the other species, which form a monophyletic group (supported only by NJ, 62%). Moreover in our tree
<taxonomicName id="4C4B53AAFFECF441FAA0FCAC7FAEFC65" class="Reptilia" family="Phrynosomatidae" genus="Urosaurus" kingdom="Animalia" order="Squamata" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="ornatus">
<emphasis id="B93FF43BFFECF441FAA0FCAC7FAEFC65" italics="true" pageId="15" pageNumber="16">U. ornatus</emphasis>
</taxonomicName>
is clearly the sister species of
<taxonomicName id="4C4B53AAFFECF441FDA3FC767C44FC65" box="[636,797,896,921]" class="Reptilia" family="Phrynosomatidae" genus="Urosaurus" kingdom="Animalia" order="Squamata" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="graciosus">
<emphasis id="B93FF43BFFECF441FDA3FC767C44FC65" box="[636,797,896,921]" italics="true" pageId="15" pageNumber="16">U. graciosus</emphasis>
</taxonomicName>
(supported by 8799%). Conversely, phylogenetic relationships based on morphological characters show that
<taxonomicName id="4C4B53AAFFECF441FCBFFC537CA3FC42" box="[864,1018,933,958]" class="Reptilia" family="Phrynosomatidae" genus="Urosaurus" kingdom="Animalia" order="Squamata" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="graciosus">
<emphasis id="B93FF43BFFECF441FCBFFC537CA3FC42" box="[864,1018,933,958]" italics="true" pageId="15" pageNumber="16">U. graciosus</emphasis>
</taxonomicName>
was external to
<taxonomicName id="4C4B53AAFFECF441FB1EFC537A36FC42" box="[1217,1391,933,958]" class="Reptilia" family="Phrynosomatidae" genus="Urosaurus" kingdom="Animalia" order="Squamata" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="bicarinatus">
<emphasis id="B93FF43BFFECF441FB1EFC537A36FC42" box="[1217,1391,933,958]" italics="true" pageId="15" pageNumber="16">U. bicarinatus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C4B53AAFFECF441FAA0FC537E7FFC1F" class="Reptilia" family="Phrynosomatidae" genus="Urosaurus" kingdom="Animalia" order="Squamata" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="nigricaudus">
<emphasis id="B93FF43BFFECF441FAA0FC537E7FFC1F" italics="true" pageId="15" pageNumber="16">U. nigricaudus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C4B53AAFFECF441FEEAFC3C7EEFFC1F" box="[309,438,970,995]" class="Reptilia" family="Phrynosomatidae" genus="Urosaurus" kingdom="Animalia" order="Squamata" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="ornatus">
<emphasis id="B93FF43BFFECF441FEEAFC3C7EEFFC1F" box="[309,438,970,995]" italics="true" pageId="15" pageNumber="16">U. ornatus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C4B53AAFFECF441FE2DFC3C7C30FC18" authority="Wiens 1993" authorityName="Wiens" authorityYear="1993" box="[498,873,970,996]" class="Reptilia" family="Phrynosomatidae" genus="Urosaurus" kingdom="Animalia" order="Squamata" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="microscutatus">
<emphasis id="B93FF43BFFECF441FE2DFC3C7DE7FC1F" box="[498,702,970,995]" italics="true" pageId="15" pageNumber="16">U. microscutatus</emphasis>
(
<bibRefCitation id="EFDA55D8FFECF441FD11FC3C7C39FC18" author="Wiens" box="[718,864,970,996]" pageId="15" pageNumber="29" refString="Wiens, J. J. (1993) Phylogenetic Systematics of the Tree Lizards (genus Urosaurus). Herpetologica, 49, 399 - 420." type="journal article" year="1993">Wiens 1993</bibRefCitation>
)
</taxonomicName>
. The topology obtained with molecular data is congruent with the distribution of the species.
<taxonomicName id="4C4B53AAFFECF441FD66FC067C3BFBF5" box="[697,866,1008,1033]" class="Reptilia" family="Phrynosomatidae" genus="Urosaurus" kingdom="Animalia" order="Squamata" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="bicarinatus">
<emphasis id="B93FF43BFFECF441FD66FC067C3BFBF5" box="[697,866,1008,1033]" italics="true" pageId="15" pageNumber="16">U. bicarinatus</emphasis>
</taxonomicName>
is nested in the southern part of the range of the genus while the other species, which cluster together in the tree, are localized in the northern part.
</paragraph>
<paragraph id="8BF42829FFECF441FF1BFBCC7BECFAC8" blockId="15.[151,1437,223,1332]" pageId="15" pageNumber="16">
The highest interspecific distance has been found between
<taxonomicName id="4C4B53AAFFECF441FC46FBCC7B13FBAF" box="[921,1098,1082,1107]" class="Reptilia" family="Phrynosomatidae" genus="Urosaurus" kingdom="Animalia" order="Squamata" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="bicarinatus">
<emphasis id="B93FF43BFFECF441FC46FBCC7B13FBAF" box="[921,1098,1082,1107]" italics="true" pageId="15" pageNumber="16">U. bicarinatus</emphasis>
</taxonomicName>
and the other species (8.4 9.4%), while lower values have been found between the other species (3.57.7%). A low divergence value (1.8%) was found between sequences of the Rio Grande (Oaxaca) and Epatlan (Puebla) populations of
<taxonomicName id="4C4B53AAFFECF441FAA0FB737E44FB38" class="Reptilia" family="Phrynosomatidae" genus="Urosaurus" kingdom="Animalia" order="Squamata" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="bicarinatus">
<emphasis id="B93FF43BFFECF441FAA0FB737E44FB38" italics="true" pageId="15" pageNumber="16">U. bicarinatus</emphasis>
</taxonomicName>
. Greater distance was found between these two localities and the sequences from Chamela (4.3 4.5%) belonging to a different subspecies. In the absence of additional data, it is very difficult to infer a conclusion regarding the taxonomic status of the Chamela population. These findings suggest that a complete intra and interspecific revision of the genus is needed using additional molecular markers.
</paragraph>
</subSubSection>
</treatment>
</document>