<documentid="A85874AF593B24CD50DE4AE003D2CF49"ID-DOI="10.11646/zootaxa.5153.1.1"ID-ISSN="1175-5326"ID-Zenodo-Dep="6630973"ID-ZooBank="A658ADE4-F352-4D16-9DC7-2721BCBE1EEF"IM.materialsCitations_approvedBy="felipe"IM.metadata_approvedBy="felipe"IM.tables_requiresApprovalFor="existingObjects,plazi"IM.taxonomicNames_approvedBy="felipe"checkinTime="1654852256489"checkinUser="plazi"docAuthor="Albuquerque, Nelson Rufino De & Fernandes, Daniel S."docDate="2022"docId="039B220BFFE4D147FF6B97CCFDC8E99A"docLanguage="en"docName="zootaxa.5153.1.1.pdf"docOrigin="Zootaxa 5153 (1)"docSource="http://zoobank.org/a658ade4-f352-4d16-9dc7-2721bcbe1eef"docStyle="DocumentStyle:5EBBA59367AD13919D70D935FA04F6A3.14:Zootaxa.2013-.monograph"docStyleId="5EBBA59367AD13919D70D935FA04F6A3"docStyleName="Zootaxa.2013-.monograph"docStyleVersion="14"docTitle="Leptophis bolivianus Oliver 1942"docType="treatment"docVersion="4"lastPageNumber="20"masterDocId="FFA25A73FFF6D154FFFC9632FFE3EE47"masterDocTitle="Taxonomic revision of the parrot snake Leptophis ahaetulla (Serpentes, Colubridae)"masterLastPageNumber="69"masterPageNumber="1"pageNumber="19"updateTime="1720181101203"updateUser="ExternalLinkService">
<bibRefCitationid="EFA3EEECFFE4D146FE6697CCFDD2EC5F"author="Oliver, J. A."box="[410,561,510,536]"pageId="18"pageNumber="19"pagination="1 - 19"refId="ref48637"refString="Oliver, J. A. (1942) A check list of the snakes of the genus Leptophis, with descriptions of new forms. Occasional Papers of the Museum of Zoology, University of Michigan, 462, 1 - 19."type="journal article"year="1942">Oliver, 1942</bibRefCitation>
<figureCitationid="13098F98FFE4D146FF639411FF0CEC7A"box="[159,239,547,573]"captionStart="FIGURE 9"captionStartId="18.[151,250,1698,1722]"captionTargetBox="[157,1431,1238,1674]"captionTargetId="figure-421@18.[157,1431,1238,1674]"captionTargetPageId="18"captionText="FIGURE 9. Color patterns and color tones variability of Leptophis bolivianus: MNKR 4332 (SVL 920 mm), a female from Estancia Caparu, Provincia Chiquitos, Santa Cruz, Bolivia. Note the (A) general color pattern and the (B) postocular extending beyond seven scales onto the nuchal region (indicated by the arrow) in this specimen, respectively. Photos by L. Gonzales."figureDoi="http://doi.org/10.5281/zenodo.6630992"httpUri="https://zenodo.org/record/6630992/files/figure.png"pageId="18"pageNumber="19">Figs. 9</figureCitation>
,
<figureCitationid="13098F98FFE4D146FF079411FEFAEC7A"box="[251,281,547,573]"captionStart="FIGURE 10"captionStartId="20.[151,250,1719,1743]"captionTargetBox="[300,1287,181,1695]"captionTargetId="figure-32@20.[300,1287,181,1695]"captionTargetPageId="20"captionText="FIGURE 10. Color patterns and color tones variability of Leptophis bolivianus: (A) Paratype FMNH 35614, SVL 727 mm, (B) Paratype FMNH 35620, SVL 1118 mm, (C) Holotype UMMZ 67973, SVL 1079 mm and (D) Paratype UMMZ 67976b, SVL 755 mm—all with incomplete tail—showing the variation in the distribution, number and extent of black spots on the supraocular and parietal scales (see text for discussion); (E) AMNH 141443, SVL 367 mm, a female from San Antonio de Parapeti, Santa Cruz, Bolivia exhibiting the banded pattern typical of juveniles of most species of Leptophis, scale bar = 0.5 cm; (F) Female paratopotype that also bears the USNM number 67977, note that the dorsal scales of its tail are keeled only from the point of reduction from six to four rows (indicated by the arrow), scale bar = 1 cm; (G) Holotype UMMZ 67973, SVL 1079 mm showing the ventral scales not edged. Courtesy of The Field Museum, FMNH 35614 and FMNH 35620 (C), D. Kizirian (E) and N.R. Albuquerque (F–G)."figureDoi="http://doi.org/10.5281/zenodo.6630994"httpUri="https://zenodo.org/record/6630994/files/figure.png"pageId="18"pageNumber="19">10</figureCitation>
,
<figureCitationid="13098F98FFE4D146FED99411FE96EC7A"box="[293,373,547,573]"captionStart="FIGURE 11"captionStartId="21.[151,250,1031,1055]"captionTargetBox="[164,1423,181,1006]"captionTargetId="figure-18@21.[164,1423,181,1006]"captionTargetPageId="21"captionText="FIGURE 11. Sulcate (left) and asulcate (right) sides of the hemipenes of Leptophis bolivianus (AMNH 104564) from Estancia Yutiole, department of Beni, Bolivia (A–B); L. coeruleodorsus (TCWC 46262) from municipality of San Fernando de Apure, state of Apure, Venezuela (C–D); L. liocercus (CZGB 1149) from municipality of Ilhéus, state of Bahia, Brazil (E–F). Note that although the hemipenis of CZGB 1149 is slightly bilobed, its sulcus spermaticus remains unbifurcated until the apical portion of the organ; L. nigromarginatus (LPHA 1731) from municipality of Santarém, state of Pará, (G–H); and (TCWC 42177) from Centro Union, departament of Loreto, Peru (I–J). Although the first row of basal spines is small in the four species, the two spines adjacent to sulcus spermaticus are larger than those in other rows. Scale bars = 0.5 cm. Photos by D. Kizirian (A–B) and N.R. Albuquerque (C–J)."figureDoi="http://doi.org/10.5281/zenodo.6630996"httpUri="https://zenodo.org/record/6630996/files/figure.png"pageId="18"pageNumber="19">11A–B</figureCitation>
<treatmentCitationid="0A93B50CFFE4D146FE849455FDC9EC38"author="Griffin, L. E."box="[376,554,615,639]"page="184"pageId="18"pageNumber="19"year="1916">
<bibRefCitationid="EFA3EEECFFE4D146FE849455FDC9EC38"author="Griffin, L. E."box="[376,554,615,639]"pageId="18"pageNumber="19"pagination="163 - 228"refId="ref46335"refString="Griffin, L. E. (1916) A catalogue of the Ophidia from South America at present (June, 1916) contained in the Carnegie Museum with descriptions of some new species. Memoirs of the Carnegie Museum, 7, 163 - 228. https: // doi. org / 10.5962 / bhl. title. 43279"type="journal article"year="1916">Griffin 1916: 184</bibRefCitation>
<bibRefCitationid="EFA3EEECFFE4D146FB1294B4FA7BECD9"author="Oliver, J. A."box="[1262,1432,646,670]"pageId="18"pageNumber="19"pagination="1 - 19"refId="ref48637"refString="Oliver, J. A. (1942) A check list of the snakes of the genus Leptophis, with descriptions of new forms. Occasional Papers of the Museum of Zoology, University of Michigan, 462, 1 - 19."type="journal article"year="1942">Oliver (1942: 2)</bibRefCitation>
<bibRefCitationid="EFA3EEECFFE4D146FF3B9496FDBCECFA"author="Wallach, V. & Williams, K. L. & Boundy, J."box="[199,607,676,701]"pageId="18"pageNumber="19"refId="ref50952"refString="Wallach, V., Williams, K. L. & Boundy, J. (2014) Snakes of the World - A Catalogue of Living and Extinct Species. CRC press, Boca Raton, Florida, 1237 pp. https: // doi. org / 10.1201 / b 16901"type="book"year="2014">Wallach, Williams & Boundy 2014: 372</bibRefCitation>
<bibRefCitationid="EFA3EEECFFE4D146FE2894F1FD8FEC9C"author="Oliver, J. A."box="[468,620,707,731]"pageId="18"pageNumber="19"pagination="1 - 19"refId="ref48637"refString="Oliver, J. A. (1942) A check list of the snakes of the genus Leptophis, with descriptions of new forms. Occasional Papers of the Museum of Zoology, University of Michigan, 462, 1 - 19."type="journal article"year="1942">Oliver, 1942: 1</bibRefCitation>
<bibRefCitationid="EFA3EEECFFE4D146FCD794D0FB6FECBD"author="Peters, J. A. & Orejas-Miranda, B."box="[811,1164,738,762]"pageId="18"pageNumber="19"pagination="1 - 347"refId="ref49487"refString="Peters, J. A. & Orejas-Miranda, B. (1970) Catalogue of the Neotropical Squamata: Part I - Snakes. Bulletin of the United States National Museum, 297, 1 - 347. https: // doi. org / 10.5479 / si. 03629236.297.1"type="journal article"year="1970">Peters & Orejas-Miranda 1970: 63</bibRefCitation>
;
<bibRefCitationid="EFA3EEECFFE4D146FB6A94D0FAABECBD"author="Tipton, B. L."box="[1174,1352,738,762]"pageId="18"pageNumber="19"refId="ref50461"refString="Tipton, B. L. (2005) Snakes of the Americas. Checklist and Lexicon. Krieger Publishing Company, Malabar, 492 pp."type="book"year="2005">Tipton 2005: 161</bibRefCitation>
<bibRefCitationid="EFA3EEECFFE4D146FE0F9532FD41ED5F"author="Oliver, J. A."box="[499,674,768,793]"pageId="18"pageNumber="19"pagination="157 - 280"refId="ref48705"refString="Oliver, J. A. (1948) The relationships and zoogeography of the genus Thalerophis Oliver. Bulletin of the American Museum of Natural History, 92, 157 - 280."type="journal article"year="1948">Oliver 1948: 225</bibRefCitation>
<bibRefCitationid="EFA3EEECFFE4D146FE05952DFB7BED70"author="International Commission of Zoological Nomenclature"box="[505,1176,799,823]"pageId="18"pageNumber="19"pagination="263 - 276"refId="ref47024"refString="International Commission of Zoological Nomenclature. (1958) Opnion 524, Interpretation of the nominal species Coluber ahaetulla Linnaeus, 1758, and addition to the Official List of Generic Names in Zoology of the generic name Ahaetulla Link, 1807 with Ahaetulla mycterizans Link, 1807 as type species (Class Reptilia). Opinions and declarations rendered by the ICZN, 19, 263 - 276."type="journal article"year="1958">International Commission of Zoological Nomenclature 1958: 270</bibRefCitation>
<bibRefCitationid="EFA3EEECFFE4D146FE7E950CFD34ED11"author="Torres-Carvajal, O. & Teran, C."box="[386,727,830,854]"pageId="18"pageNumber="19"pagination="1 - 7"refId="ref50594"refString="Torres-Carvajal, O. & Teran, C. (2021) Molecular phylogeny of Neotropical Parrot Snakes (Serpentes: Colubrinae: Leptophis) supports underestimated species richness. Molecular Phylogenetics and Evolution, 164, 1 - 7. https: // doi. org / 10.1016 / j. ympev. 2021.107267"type="journal article"year="2021">Torres-Carvajal & Téran, 2021: 6</bibRefCitation>
head scales distinctly edged with black and usually with two or more poorly defined, diffuse black spots on each parietal; a distinct, elongated black spot (occasionally two small spots) on each supraocular; preocular stripe absent;
4332 (SVL 920 mm), a female from Estancia Caparu, Provincia Chiquitos, Santa Cruz, Bolivia. Note the (A) general color pattern and the (B) postocular extending beyond seven scales onto the nuchal region (indicated by the arrow) in this specimen, respectively. Photos by L. Gonzales.
complex by the combination of head scales distinctly edged with black and usually with two or more poorly defined, diffuse black spots on each parietal; a distinct, elongated black spot (occasionally two small spots) on each supraocular (
<figureCitationid="13098F98FFE4D146FB5D914AFACAE9D4"box="[1185,1321,1912,1939]"captionStart="FIGURE 10"captionStartId="20.[151,250,1719,1743]"captionTargetBox="[300,1287,181,1695]"captionTargetId="figure-32@20.[300,1287,181,1695]"captionTargetPageId="20"captionText="FIGURE 10. Color patterns and color tones variability of Leptophis bolivianus: (A) Paratype FMNH 35614, SVL 727 mm, (B) Paratype FMNH 35620, SVL 1118 mm, (C) Holotype UMMZ 67973, SVL 1079 mm and (D) Paratype UMMZ 67976b, SVL 755 mm—all with incomplete tail—showing the variation in the distribution, number and extent of black spots on the supraocular and parietal scales (see text for discussion); (E) AMNH 141443, SVL 367 mm, a female from San Antonio de Parapeti, Santa Cruz, Bolivia exhibiting the banded pattern typical of juveniles of most species of Leptophis, scale bar = 0.5 cm; (F) Female paratopotype that also bears the USNM number 67977, note that the dorsal scales of its tail are keeled only from the point of reduction from six to four rows (indicated by the arrow), scale bar = 1 cm; (G) Holotype UMMZ 67973, SVL 1079 mm showing the ventral scales not edged. Courtesy of The Field Museum, FMNH 35614 and FMNH 35620 (C), D. Kizirian (E) and N.R. Albuquerque (F–G)."figureDoi="http://doi.org/10.5281/zenodo.6630994"httpUri="https://zenodo.org/record/6630994/files/figure.png"pageId="18"pageNumber="19">Fig. 10A–D</figureCitation>
) (vs. head scales not edged with black, with or with no black spot on each parietal; a small black spot on each supraocular). It differs from parapatric
by having ventrals not edged with dark greenish blue or blue (
<figureCitationid="13098F98FFE4D147FA9391F2FF2AEEF6"captionStart="FIGURE 10"captionStartId="20.[151,250,1719,1743]"captionTargetBox="[300,1287,181,1695]"captionTargetId="figure-32@20.[300,1287,181,1695]"captionTargetPageId="20"captionText="FIGURE 10. Color patterns and color tones variability of Leptophis bolivianus: (A) Paratype FMNH 35614, SVL 727 mm, (B) Paratype FMNH 35620, SVL 1118 mm, (C) Holotype UMMZ 67973, SVL 1079 mm and (D) Paratype UMMZ 67976b, SVL 755 mm—all with incomplete tail—showing the variation in the distribution, number and extent of black spots on the supraocular and parietal scales (see text for discussion); (E) AMNH 141443, SVL 367 mm, a female from San Antonio de Parapeti, Santa Cruz, Bolivia exhibiting the banded pattern typical of juveniles of most species of Leptophis, scale bar = 0.5 cm; (F) Female paratopotype that also bears the USNM number 67977, note that the dorsal scales of its tail are keeled only from the point of reduction from six to four rows (indicated by the arrow), scale bar = 1 cm; (G) Holotype UMMZ 67973, SVL 1079 mm showing the ventral scales not edged. Courtesy of The Field Museum, FMNH 35614 and FMNH 35620 (C), D. Kizirian (E) and N.R. Albuquerque (F–G)."figureDoi="http://doi.org/10.5281/zenodo.6630994"httpUri="https://zenodo.org/record/6630994/files/figure.png"lastPageId="19"lastPageNumber="20"pageId="18"pageNumber="19">Fig. 10G</figureCitation>
) and distinct keels on the dorsal scales of tail (vs. ventrals edged with dark greenish blue or blue,
<figureCitationid="13098F98FFE5D147FACE96A5FA7BEEF6"box="[1330,1432,151,177]"captionStart="FIGURE 23"captionStartId="36.[151,250,1314,1338]"captionTargetBox="[161,1426,417,1291]"captionTargetId="figure-149@36.[161,1426,417,1291]"captionTargetPageId="36"captionText="FIGURE 23. Color patterns and color tones variability of Leptophis nigromarginatus: (A) dorsal view of the head and (B) ventral view of the body of the holotype (BMNH 1946.1.5.7) from “Upper Amazons”. Arrow indicating ventral scale edged anteriorly and laterally with greenish blue, a characteristic thought by Oliver (1948: 246) to be typical of L. a. ortoni. Note two black spots in center of each parietal scale, a larger black spot on right supraocular scale, and two smaller spots on left supraocular scale; (C) KU 112279 and (D) KU 155512, a female and a male, respectively, from Santa Cecília, Ecuador. Both show distinct differences in the prominence of supracephalic black borders and spots, being both more prominent in the female specimen. Scale bar = 0.5 cm. Photos by N.R. Albuquerque (A –B) and A. Motta (C–D)."figureDoi="http://doi.org/10.5281/zenodo.6631022"httpUri="https://zenodo.org/record/6631022/files/figure.png"pageId="19"pageNumber="20">Fig. 23B</figureCitation>
, and no keels on dorsals of the tail), by higher number of ventrals for males 95% CI = 156.2–161 and females 95% CI 162.4–166 (vs. 151.6–153 and 155.9–157, respectively).
<tableCitationid="C6B0A6A6FFE5D147FC479731FBF1EF5A"box="[955,1042,259,285]"captionStart="TABLE 2"captionStartId="46.[151,239,490,516]"captionText="TABLE 2. Quantitative variation of selected characters of 10 species of Leptophis that were formerly subspecies of L. ahaetulla. M=male, F=female. Sample sizes given for paired characters of pholidosis (e.g., number of teeth) refer to the number of sides examined for all specimens, whereas sample sizes given for other (nonpaired) characters (e.g., ventrals, subcaudals) refer to the number of specimens examined. 95% confidence limits (Cl) follow ranges, means ± standard deviations when appropriate. The ratio tail length (TL) snout-vent length (SVL) did not differ significantly between females and males in all the species examined, so the confidence interval of 95% (CI) encompasses the variation of both sexes. 1. L. ahaetulla, 2. L. bocourti, 3. L. bolivianus, 4. L. coeruleodorsus, 5. L. liocercus, 6. L. marginatus, 7. L. nigromarginatus, 8. L. occidentalis, 9. L. praestans, 10. L. urostictus."pageId="19"pageNumber="20">Table 2</tableCitation>
74), with fourth–fifth (94.5%, n = 70), or, rarely, fifth–sixth (5.5%, n = 4) bordering orbit; infralabials 9–12 (10.0 ± 0.5, n = 72), with first 5 (79.7%, n = 59), first 6 (17.6%, n = 13), or, rarely, first 4 (2.7%, n = 2) contacting first chin shields; preocular 1 (n = 36) and a single specimen with 2 on right side; postoculars 2 (n = 36) and a single specimen with 3 on right side; anterior temporal 1 (n = 37); posterior temporal 1–2 (1.7 ± 0.4, n
<specimenCodeid="DB943B66FFE5D147FECD9475FE2CEC26"box="[305,463,582,609]"collectionCode="FMNH"country="USA"httpUri="http://biocol.org/urn:lsid:biocol.org:col:34795"lsid="urn:lsid:biocol.org:col:34795"name="Field Museum of Natural History"pageId="19"pageNumber="20"type="Museum">FMNH 35614</specimenCode>
</materialsCitation>
possess two small black spots on each supraocular scale,
<specimenCodeid="DB943B66FFE5D147FB3A9475FA87EC26"box="[1222,1380,582,609]"collectionCode="FMNH"country="USA"httpUri="http://biocol.org/urn:lsid:biocol.org:col:34795"lsid="urn:lsid:biocol.org:col:34795"name="Field Museum of Natural History"pageId="19"pageNumber="20"type="Museum">FMNH 35620</specimenCode>
</materialsCitation>
possess two small black spots on left supraocular,
<specimenCodeid="DB943B66FFE5D147FCE49459FC5DECC2"box="[792,958,619,645]"collectionCode="UMMZ"country="USA"httpUri="http://grbio.org/cool/ra64-3eva"name="University of Michigan, Museum of Zoology"pageId="19"pageNumber="20"type="University or college">UMMZ 67973</specimenCode>
</materialsCitation>
possess one elongated black spot on each supraocular, and
<specimenCodeid="DB943B66FFE5D147FE3A94BDFD8CECEE"box="[454,623,655,681]"collectionCode="UMMZ"country="USA"httpUri="http://grbio.org/cool/ra64-3eva"name="University of Michigan, Museum of Zoology"pageId="19"pageNumber="20"type="University or college">UMMZ 67976</specimenCode>
</materialsCitation>
b possess two small black spots on left supraocular and one elongated black spot on right supraocular; parietal scales possess a poorly defined, diffuse black spot on each specimen though they are more elongated in the
<specimenCodeid="DB943B66FFE5D147FDA594E5FD1EECB6"box="[601,765,727,753]"collectionCode="UMMZ"country="USA"httpUri="http://grbio.org/cool/ra64-3eva"name="University of Michigan, Museum of Zoology"pageId="19"pageNumber="20"type="University or college">UMMZ 67973</specimenCode>
(
<figureCitationid="13098F98FFE5D147FCF094E5FC90ECB6"box="[780,883,727,753]"captionStart="FIGURE 10"captionStartId="20.[151,250,1719,1743]"captionTargetBox="[300,1287,181,1695]"captionTargetId="figure-32@20.[300,1287,181,1695]"captionTargetPageId="20"captionText="FIGURE 10. Color patterns and color tones variability of Leptophis bolivianus: (A) Paratype FMNH 35614, SVL 727 mm, (B) Paratype FMNH 35620, SVL 1118 mm, (C) Holotype UMMZ 67973, SVL 1079 mm and (D) Paratype UMMZ 67976b, SVL 755 mm—all with incomplete tail—showing the variation in the distribution, number and extent of black spots on the supraocular and parietal scales (see text for discussion); (E) AMNH 141443, SVL 367 mm, a female from San Antonio de Parapeti, Santa Cruz, Bolivia exhibiting the banded pattern typical of juveniles of most species of Leptophis, scale bar = 0.5 cm; (F) Female paratopotype that also bears the USNM number 67977, note that the dorsal scales of its tail are keeled only from the point of reduction from six to four rows (indicated by the arrow), scale bar = 1 cm; (G) Holotype UMMZ 67973, SVL 1079 mm showing the ventral scales not edged. Courtesy of The Field Museum, FMNH 35614 and FMNH 35620 (C), D. Kizirian (E) and N.R. Albuquerque (F–G)."figureDoi="http://doi.org/10.5281/zenodo.6630994"httpUri="https://zenodo.org/record/6630994/files/figure.png"pageId="19"pageNumber="20">Fig. 10C</figureCitation>
<specimenCodeid="DB943B66FFE5D147FC7894E5FBD7ECB6"box="[900,1076,727,753]"collectionCode="AMNH"country="USA"httpUri="http://biocol.org/urn:lsid:biocol.org:col:34925"lsid="urn:lsid:biocol.org:col:34925"name="American Museum of Natural History"pageId="19"pageNumber="20">AMNH 141443</specimenCode>
(
<figureCitationid="13098F98FFE5D147FBB894E5FB44ECB6"box="[1092,1191,727,753]"captionStart="FIGURE 10"captionStartId="20.[151,250,1719,1743]"captionTargetBox="[300,1287,181,1695]"captionTargetId="figure-32@20.[300,1287,181,1695]"captionTargetPageId="20"captionText="FIGURE 10. Color patterns and color tones variability of Leptophis bolivianus: (A) Paratype FMNH 35614, SVL 727 mm, (B) Paratype FMNH 35620, SVL 1118 mm, (C) Holotype UMMZ 67973, SVL 1079 mm and (D) Paratype UMMZ 67976b, SVL 755 mm—all with incomplete tail—showing the variation in the distribution, number and extent of black spots on the supraocular and parietal scales (see text for discussion); (E) AMNH 141443, SVL 367 mm, a female from San Antonio de Parapeti, Santa Cruz, Bolivia exhibiting the banded pattern typical of juveniles of most species of Leptophis, scale bar = 0.5 cm; (F) Female paratopotype that also bears the USNM number 67977, note that the dorsal scales of its tail are keeled only from the point of reduction from six to four rows (indicated by the arrow), scale bar = 1 cm; (G) Holotype UMMZ 67973, SVL 1079 mm showing the ventral scales not edged. Courtesy of The Field Museum, FMNH 35614 and FMNH 35620 (C), D. Kizirian (E) and N.R. Albuquerque (F–G)."figureDoi="http://doi.org/10.5281/zenodo.6630994"httpUri="https://zenodo.org/record/6630994/files/figure.png"pageId="19"pageNumber="20">Fig. 10E</figureCitation>
<specimenCodeid="DB943B66FFE5D147FE8994C9FDFEED52"box="[373,541,763,789]"collectionCode="UMMZ"country="USA"httpUri="http://grbio.org/cool/ra64-3eva"name="University of Michigan, Museum of Zoology"pageId="19"pageNumber="20"type="University or college">UMMZ 67977</specimenCode>
total length (but tail incomplete), are ornamented with bands in anterior and middle region of body, similar to those found in juveniles of other species of
<bibRefCitationid="EFA3EEECFFE5D147FAA9952DFF31ED1A"author="Oliver, J. A."pageId="19"pageNumber="20"pagination="157 - 280"refId="ref48705"refString="Oliver, J. A. (1948) The relationships and zoogeography of the genus Thalerophis Oliver. Bulletin of the American Museum of Natural History, 92, 157 - 280."type="journal article"year="1948">Oliver 1948</bibRefCitation>
; this study). Females have more ventrals than males (F
= 0.0735; P = 0.7856). Dorsal scales of tail can be keeled from the point of reduction from six to four rows (a female that also bears the
<collectionCodeid="ED230BD8FFE5D147FC9395B9FC23EDE2"box="[879,960,907,933]"country="USA"httpUri="http://biocol.org/urn:lsid:biocol.org:col:34871"lsid="urn:lsid:biocol.org:col:34871"name="Smithsonian Institution, National Museum of Natural History"pageId="19"pageNumber="20"type="Museum">USNM</collectionCode>
number 67977) (
<figureCitationid="13098F98FFE5D147FB7B95B9FB0AEDE2"box="[1159,1257,907,933]"captionStart="FIGURE 10"captionStartId="20.[151,250,1719,1743]"captionTargetBox="[300,1287,181,1695]"captionTargetId="figure-32@20.[300,1287,181,1695]"captionTargetPageId="20"captionText="FIGURE 10. Color patterns and color tones variability of Leptophis bolivianus: (A) Paratype FMNH 35614, SVL 727 mm, (B) Paratype FMNH 35620, SVL 1118 mm, (C) Holotype UMMZ 67973, SVL 1079 mm and (D) Paratype UMMZ 67976b, SVL 755 mm—all with incomplete tail—showing the variation in the distribution, number and extent of black spots on the supraocular and parietal scales (see text for discussion); (E) AMNH 141443, SVL 367 mm, a female from San Antonio de Parapeti, Santa Cruz, Bolivia exhibiting the banded pattern typical of juveniles of most species of Leptophis, scale bar = 0.5 cm; (F) Female paratopotype that also bears the USNM number 67977, note that the dorsal scales of its tail are keeled only from the point of reduction from six to four rows (indicated by the arrow), scale bar = 1 cm; (G) Holotype UMMZ 67973, SVL 1079 mm showing the ventral scales not edged. Courtesy of The Field Museum, FMNH 35614 and FMNH 35620 (C), D. Kizirian (E) and N.R. Albuquerque (F–G)."figureDoi="http://doi.org/10.5281/zenodo.6630994"httpUri="https://zenodo.org/record/6630994/files/figure.png"pageId="19"pageNumber="20">Fig. 10F</figureCitation>
) to the point of hundredth tenth subcaudal scales, corresponding to 36.4% TL (
<specimenCodeid="DB943B66FFE5D147FCA4959DFBE0ED8D"box="[856,1027,943,970]"collectionCode="UMMZ"country="USA"httpUri="http://grbio.org/cool/ra64-3eva"name="University of Michigan, Museum of Zoology"pageId="19"pageNumber="20"type="University or college">UMMZ 67974</specimenCode>
). On the other hand, all scales of upper surface of tail are keeled in
<specimenCodeid="DB943B66FFE5D147FE0095E1FD7CEDAA"box="[508,671,979,1005]"collectionCode="AMNH"country="USA"httpUri="http://biocol.org/urn:lsid:biocol.org:col:34925"lsid="urn:lsid:biocol.org:col:34925"name="American Museum of Natural History"pageId="19"pageNumber="20">AMNH 22446</specimenCode>
<specimenCodeid="DB943B66FFE5D147FD2995E1FC69EDAA"box="[725,906,978,1005]"collectionCode="AMNH"country="USA"httpUri="http://biocol.org/urn:lsid:biocol.org:col:34925"lsid="urn:lsid:biocol.org:col:34925"name="American Museum of Natural History"pageId="19"pageNumber="20">AMNH 104563</specimenCode>
</materialsCitation>
. The TL/SVL showed no significant difference between females and males (F
Single retracted organ examined extends up to seven-eight subcaudals. Everted hemipenis unilobed, noncapitate; sulcus spermaticus centrolineal, undivided, extending from base to tip of lobe; basal portion bears small spines distributed in 6 rows approximatelly encircling the organ; first row bears 7 hooked spines slightly larger than those in other rows; spinules widely scattered adjacent to sulcus spermaticus; small calyces ornamented with 10–12 fringing papillae concentrated above most distal row of basal spines whereas large-sized calyces have 5–8 fleshy papillae concentrated along distal portion of hemipenial body; papillae gradually decrease in length toward distal portion of hemipenis; distal portion of lobe completely calyculate; asulcate side similar to sulcate side (
<figureCitationid="13098F98FFE5D147FED49325FE52EB76"box="[296,433,1303,1329]"captionStart="FIGURE 11"captionStartId="21.[151,250,1031,1055]"captionTargetBox="[164,1423,181,1006]"captionTargetId="figure-18@21.[164,1423,181,1006]"captionTargetPageId="21"captionText="FIGURE 11. Sulcate (left) and asulcate (right) sides of the hemipenes of Leptophis bolivianus (AMNH 104564) from Estancia Yutiole, department of Beni, Bolivia (A–B); L. coeruleodorsus (TCWC 46262) from municipality of San Fernando de Apure, state of Apure, Venezuela (C–D); L. liocercus (CZGB 1149) from municipality of Ilhéus, state of Bahia, Brazil (E–F). Note that although the hemipenis of CZGB 1149 is slightly bilobed, its sulcus spermaticus remains unbifurcated until the apical portion of the organ; L. nigromarginatus (LPHA 1731) from municipality of Santarém, state of Pará, (G–H); and (TCWC 42177) from Centro Union, departament of Loreto, Peru (I–J). Although the first row of basal spines is small in the four species, the two spines adjacent to sulcus spermaticus are larger than those in other rows. Scale bars = 0.5 cm. Photos by D. Kizirian (A–B) and N.R. Albuquerque (C–J)."figureDoi="http://doi.org/10.5281/zenodo.6630996"httpUri="https://zenodo.org/record/6630996/files/figure.png"pageId="19"pageNumber="20">Fig. 11A–B</figureCitation>
<emphasisid="B9464F0FFFE5D147FF3B9308FE7EEB13"bold="true"box="[199,413,1338,1364]"pageId="19"pageNumber="20">Coloration in life.</emphasis>
Dorsum of the head Light Emerald Green (142), distinctly edged with black; usually two or more poorly defined, diffuse black spots on each parietal and distinct, 1–2 black spots (occasionally elongate) on each supraocular scale; rows II–IV (occasionally V) in anterior portion of the body (before body scale reduction from 15 to 11) Pale Greenish Yellow (86); rows IV–VII (occasionally V–VII) in anterior portion of the body Light Emerald Green (142); dorsal scales slightly edged with black; keels of dorsal scales slightly black; narrow Jet Black (300) postocular stripe covering upper edge of lower postocular, lower edge of upper postocular, lower margin to one-third of anterior temporal, half of lower posterior temporal and upper edges of last two supralabials; postocular stripe may extend beyond seven scales onto nuchal region; first five to seven supralabials, as well as infralabials, chin, throat, and venter white; last two supralabials below black upper edges of postocular stripe white.
<emphasisid="B9464F0FFFE5D147FF3B904CFDA8E8DF"bold="true"box="[199,587,1662,1689]"pageId="19"pageNumber="20">Distribution and natural history.</emphasis>
asl in the tropical and subtropical dry broadleaf forests ecoregion, as defined by
<bibRefCitationid="EFA3EEECFFE5D147FF6B90F5FE88E8A6"author="Olson, D. M. & Dinerstein, E. & Wikramanayake, E. D. & Burgess, N. D. & Powell, G. V. N. & Underwood, E. C. & D'Amico, J. A. & Itoua, I. & Strand, H. E. & Morrison, J. C. & Loucks, C. J. & Allnutt, T. F. & Ricketts, T. H. & Kura, Y. & Lamoreux, J. F. & Wetengel, W. W. & Hedao, P. & Kassem, K. R."box="[151,363,1735,1761]"pageId="19"pageNumber="20"pagination="933 - 938"refId="ref48823"refString="Olson, D. M., Dinerstein, E., Wikramanayake, E. D., Burgess, N. D., Powell, G. V. N., Underwood, E. C., D'Amico, J. A., Itoua, I., Strand, H. E., Morrison, J. C., Loucks, C. J., Allnutt, T. F., Ricketts, T. H.; Kura, Y., Lamoreux, J. F., Wetengel, W. W., Hedao, P. & Kassem, K. R. (2001) Terrestrial Ecoregions of the World: A New Map of Life on Earth. Bioscience, 51 (11), 933 - 938."type="journal article"year="2001">
<figureCitationid="13098F98FFE5D147FE8690F5FE2EE8A6"box="[378,461,1735,1761]"captionStart="FIGURE 18"captionStartId="30.[151,250,1971,1995]"captionTargetBox="[155,1428,1107,1946]"captionTargetId="figure-467@30.[151,1436,1100,1947]"captionTargetPageId="30"captionText="FIGURE 18. Distribution of Leptophis bolivianus, L. liocercus, L. marginatus, and L. nigromarginatus based upon the material examined."figureDoi="http://doi.org/10.5281/zenodo.6631010"httpUri="https://zenodo.org/record/6631010/files/figure.png"pageId="19"pageNumber="20">Fig. 18</figureCitation>
<bibRefCitationid="EFA3EEECFFE5D147FE2190D9FD56E941"author="Oliver, J. A."box="[477,693,1771,1798]"pageId="19"pageNumber="20"pagination="157 - 280"refId="ref48705"refString="Oliver, J. A. (1948) The relationships and zoogeography of the genus Thalerophis Oliver. Bulletin of the American Museum of Natural History, 92, 157 - 280."type="journal article"year="1948">Oliver (1948: 226)</bibRefCitation>
one of the most remarkable variations in the head color pattern of
consists of differences of the distribution and extent of spots on the dorsal surface of the head (
<figureCitationid="13098F98FFE5D147FF279101FE85E90A"box="[219,358,1843,1869]"captionStart="FIGURE 10"captionStartId="20.[151,250,1719,1743]"captionTargetBox="[300,1287,181,1695]"captionTargetId="figure-32@20.[300,1287,181,1695]"captionTargetPageId="20"captionText="FIGURE 10. Color patterns and color tones variability of Leptophis bolivianus: (A) Paratype FMNH 35614, SVL 727 mm, (B) Paratype FMNH 35620, SVL 1118 mm, (C) Holotype UMMZ 67973, SVL 1079 mm and (D) Paratype UMMZ 67976b, SVL 755 mm—all with incomplete tail—showing the variation in the distribution, number and extent of black spots on the supraocular and parietal scales (see text for discussion); (E) AMNH 141443, SVL 367 mm, a female from San Antonio de Parapeti, Santa Cruz, Bolivia exhibiting the banded pattern typical of juveniles of most species of Leptophis, scale bar = 0.5 cm; (F) Female paratopotype that also bears the USNM number 67977, note that the dorsal scales of its tail are keeled only from the point of reduction from six to four rows (indicated by the arrow), scale bar = 1 cm; (G) Holotype UMMZ 67973, SVL 1079 mm showing the ventral scales not edged. Courtesy of The Field Museum, FMNH 35614 and FMNH 35620 (C), D. Kizirian (E) and N.R. Albuquerque (F–G)."figureDoi="http://doi.org/10.5281/zenodo.6630994"httpUri="https://zenodo.org/record/6630994/files/figure.png"pageId="19"pageNumber="20">Fig. 10A–D</figureCitation>
).
<bibRefCitationid="EFA3EEECFFE5D147FE859101FDA9E909"author="Oliver, J. A."box="[377,586,1843,1870]"pageId="19"pageNumber="20"pagination="157 - 280"refId="ref48705"refString="Oliver, J. A. (1948) The relationships and zoogeography of the genus Thalerophis Oliver. Bulletin of the American Museum of Natural History, 92, 157 - 280."type="journal article"year="1948">Oliver (1948: 226)</bibRefCitation>
<bibRefCitationid="EFA3EEECFFE5D147FABD9149FF01E9FE"author="Oliver, J. A."pageId="19"pageNumber="20"pagination="157 - 280"refId="ref48705"refString="Oliver, J. A. (1948) The relationships and zoogeography of the genus Thalerophis Oliver. Bulletin of the American Museum of Natural History, 92, 157 - 280."type="journal article"year="1948">Oliver’s (1948)</bibRefCitation>
revision, however, reveals that the number of ventrals (153) of AMNH 22446 falls within the variation range for males of
<tableCitationid="C6B0A6A6FFE5D147FE3A91F1FDFDE99A"box="[454,542,1987,2013]"captionStart="TABLE 2"captionStartId="46.[151,239,490,516]"captionText="TABLE 2. Quantitative variation of selected characters of 10 species of Leptophis that were formerly subspecies of L. ahaetulla. M=male, F=female. Sample sizes given for paired characters of pholidosis (e.g., number of teeth) refer to the number of sides examined for all specimens, whereas sample sizes given for other (nonpaired) characters (e.g., ventrals, subcaudals) refer to the number of specimens examined. 95% confidence limits (Cl) follow ranges, means ± standard deviations when appropriate. The ratio tail length (TL) snout-vent length (SVL) did not differ significantly between females and males in all the species examined, so the confidence interval of 95% (CI) encompasses the variation of both sexes. 1. L. ahaetulla, 2. L. bocourti, 3. L. bolivianus, 4. L. coeruleodorsus, 5. L. liocercus, 6. L. marginatus, 7. L. nigromarginatus, 8. L. occidentalis, 9. L. praestans, 10. L. urostictus."pageId="19"pageNumber="20">Table 2</tableCitation>
