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<document ID-CLB-Dataset="287679" ID-DOI="http://dx.doi.org/10.3897/evolsyst.8.114527" ID-GBIF-Dataset="e13392ec-8851-40da-9403-4f2574646f10" ID-Pensoft-Pub="2535-0730-1-15" ID-Pensoft-UUID="9667B588C08053D1BC52E6807E89A599" ID-ZooBank="42D6D571379D4EB0BC8DB3134A4E0912" ModsDocID="2535-0730-8-1-15" checkinTime="1707483718867" checkinUser="pensoft" docAuthor="Arteaga, Alejandro, Pyron, R. Alexander, Batista, Abel, Vieira, Jose, Meneses Pelayo, Elson, Smith, Eric N., Barrio Amoros, Cesar L., Koch, Claudia, Agne, Stefanie, Valencia, Jorge H., Bustamante, Lucas &amp; Harris, Kyle J." docDate="2024" docId="CF4FD9AC10AE55F1A8BCC918FD4DFB83" docLanguage="en" docName="EvolutSyst 8(1): 15-64" docOrigin="Evolutionary Systematics 8 (1)" docPubDate="2024-02-08" docSource="http://dx.doi.org/10.3897/evolsyst.8.114527" docTitle="Bothriechis klebbai Arteaga &amp; Pyron &amp; Batista &amp; Vieira &amp; Meneses Pelayo &amp; Smith &amp; Barrio Amorós &amp; Koch &amp; Agne &amp; Valencia &amp; Bustamante &amp; Harris 2024, sp. nov." docType="treatment" docUuid="20308073-EA2E-474F-951C-C2ABBFB7562F" docUuidSource="ZooBank" docVersion="5" id="9667B588C08053D1BC52E6807E89A599" lastPageNumber="15" masterDocId="9667B588C08053D1BC52E6807E89A599" masterDocTitle="Systematic revision of the Eyelash Palm-Pitviper Bothriechis schlegelii (Serpentes, Viperidae), with the description of five new species and revalidation of three" masterLastPageNumber="64" masterPageNumber="15" pageNumber="15" updateTime="1732668639335" updateUser="ExternalLinkService">
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<mods:title id="8F9FBF9DA4AD24A995F2FDB07B3442D0">Systematic revision of the Eyelash Palm-Pitviper Bothriechis schlegelii (Serpentes, Viperidae), with the description of five new species and revalidation of three</mods:title>
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<mods:namePart id="8557F2D4E0A61B00991760A95B5F1625">Arteaga, Alejandro</mods:namePart>
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<mods:namePart id="185A8A26FDEB35C2B219C9A2BA7AD390">Pyron, R. Alexander</mods:namePart>
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<mods:affiliation id="B7CBE1648B8AD459BB461B76770E2992">Department of Biological Sciences, The George Washington University, Washington DC, USA &amp; Division of Amphibians &amp; Reptiles, Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington DC, USA</mods:affiliation>
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<mods:affiliation id="8997EEF4558F348F591D2ACCC8CD02FF">Universidad Autonoma de Chiriqui (UNACHI), Instituto Interdisciplinario de Investigacion e Innovacion, David, Panama &amp; Fundacion Los Naturalistas, Boquete, Chiriqui, Panama &amp; Sistema Nacional de Investigacion (SNI), SENACYT, Clayton Panama, Panama</mods:affiliation>
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<mods:namePart id="057C8F4A2409D83FE948075FAB151D16">Vieira, Jose</mods:namePart>
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<mods:namePart id="005C07E4C23A2CACB440188B560C69E1">Meneses Pelayo, Elson</mods:namePart>
<mods:affiliation id="DA8429E54256FA77F9EB0335902853D5">Grupo de Estudios en Anfibios y Reptiles de Santander, Universidad Industrial de Santander, Bucaramanga, Colombia</mods:affiliation>
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<mods:namePart id="DB601F0E923572AB6D68F3B18F365F9D">Koch, Claudia</mods:namePart>
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<mods:affiliation id="3DC603FDBFF8C6E5A7330DC3A4155A1E">CRWild, Bahia Ballena, Uvita, Costa Rica</mods:affiliation>
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<mods:namePart id="B8198198C0DEA3C69E2338354245269E">Agne, Stefanie</mods:namePart>
<mods:affiliation id="98DB130C75E3D6FEE5303ECA6394F30A">Leibniz Institute for the Analysis of Biodiversity Change (LIB), Museum Koenig, Bonn, Germany</mods:affiliation>
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<mods:namePart id="DF790CFC7C0B8A8708248A4269EA79B2">Valencia, Jorge H.</mods:namePart>
<mods:affiliation id="483A49A16F02E8381396EFE3FD219BD9">Evolutionary Adaptive Genomics, Department of Mathematics and Natural Sciences, University of Potsdam, Potsdam, Germany &amp; Red de Estudios Moleculares Avanzados, Instituto de Ecologia, Xalapa, Veracruz, Mexico</mods:affiliation>
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<mods:namePart id="DD6C96ACF8094DD00715F4EBE023EB6A">Bustamante, Lucas</mods:namePart>
<mods:affiliation id="1C24C35EF366DBFD4C1F18CB1B898258">Tropical Herping S. A., Quito, Ecuador &amp; Red de Biogeografia y Ecologia Espacial (BioGeo 2), Universidad Regional Amazonica Ikiam, Tena, Ecuador</mods:affiliation>
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<mods:namePart id="4A724B3C3F865C291C185B674C6419EB">Harris, Kyle J.</mods:namePart>
<mods:affiliation id="61F579F073E860AECF32E39F8D0429D3">Savia Fund, Quito, Ecuador</mods:affiliation>
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<mods:date id="7EDF6BC56E83143D0907F0A2103B5628">2024</mods:date>
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<treatment id="CF4FD9AC10AE55F1A8BCC918FD4DFB83" ID-GBIF-Taxon="221225634" LSID="urn:lsid:zoobank.org:act:20308073-EA2E-474F-951C-C2ABBFB7562F" httpUri="http://treatment.plazi.org/id/CF4FD9AC10AE55F1A8BCC918FD4DFB83" lastPageNumber="15" pageId="0" pageNumber="15" scope_family="Viperidae">
<subSubSection id="62004EDD5CE38B22A55690390D208E87" pageId="0" pageNumber="15" type="nomenclature">
<paragraph id="39BF7B3D2517DC0625A5E1E3F4A99772" pageId="0" pageNumber="15">
<taxonomicName id="FCAE4AEDB50436143AF97A61A60DA1B5" LSID="https://zoobank.org/20308073-EA2E-474F-951C-C2ABBFB7562F" authority="Arteaga &amp; Pyron &amp; Batista &amp; Vieira &amp; Meneses Pelayo &amp; Smith &amp; Barrio Amorós &amp; Koch &amp; Agne &amp; Valencia &amp; Bustamante &amp; Harris, 2024" authorityName="Arteaga &amp; Pyron &amp; Batista &amp; Vieira &amp; Meneses Pelayo &amp; Smith &amp; Barrio Amorós &amp; Koch &amp; Agne &amp; Valencia &amp; Bustamante &amp; Harris" authorityYear="2024" family="Viperidae" genus="Bothriechis" higherTaxonomySource="treatment-meta" kingdom="Animalia" lsidName="Bothriechis klebbai" order="Squamata" pageId="0" pageNumber="15" rank="species" species="klebbai" status="sp. nov.">Bothriechis klebbai</taxonomicName>
<taxonomicNameLabel id="9740117459A615A6558BABC6AAA8EC5F" pageId="0" pageNumber="15">sp. nov.</taxonomicNameLabel>
</paragraph>
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<subSubSection id="EFDB10D86DBD8F2A14FD3124BBC6E972" pageId="0" pageNumber="15" type="description">
<paragraph id="DE54C5E148A6613E2B15E6A3AC990100" pageId="0" pageNumber="15">
<figureCitation id="802F94E6212E9ACBF58ACCB5AC0C75A3" captionStart="Figure 19" captionStartId="F19" captionText="Figure 19. Adult male holotype of Bothriechis klebbai sp. nov. ICN 2786 in a. Dorsal and b. Ventral view. Photos by Duvan Zambrano." httpUri="https://binary.pensoft.net/fig/981705" pageId="0" pageNumber="15">Figs 19</figureCitation>
<figureCitation id="6C50AB432E3BD67325BA9C2331FC2A77" captionStart="Figure 20" captionStartId="F20" captionText="Figure 20. Photographs of some specimens of Bothriechis klebbai sp. nov. in life from Finca la Arrinconada, vereda Esparta, minicipio de Santa Barbara, Santander department, Colombia. a. UIS-R 4139 adult male; b. INSV-SR- 00098 adult of undetermined sex; and c. UIS-R- 4296 juvenile male. Photos by Elson Meneses Pelayo and Jose Vieira." httpUri="https://binary.pensoft.net/fig/981706" pageId="0" pageNumber="15">, 20</figureCitation>
<figureCitation id="E08560298EAA461B8FD37E6209480C88" captionStart="Figure 21" captionStartId="F21" captionText="Figure 21. Hemipenial architecture of Bothriechis klebbai sp. nov. in sulcate, lateral, and asulcate views. a. ICN 2786 and b. ICN 2855 from Virolin, municipio de Charala, Santander department, Colombia; c. UIS-R 4803 from vereda Naranjal, municipio de Coromoro, Santander department, Colombia. Photos by Duvan Zambrano and Elson Meneses Pelayo." httpUri="https://binary.pensoft.net/fig/981707" pageId="0" pageNumber="15">, 21</figureCitation>
</paragraph>
</subSubSection>
<subSubSection id="1D2E9A562A786DFA13D0BA3064DC696F" pageId="0" pageNumber="15" type="type material">
<paragraph id="4D1D0ACEC486BAB8258B6237E5C8576E" pageId="0" pageNumber="15">Type material.</paragraph>
<paragraph id="9C863B4AA01F2EF86354152E2CFE9AED" pageId="0" pageNumber="15">
<emphasis id="04A98770225410F75B3C4DD42A0D6D15" bold="true" italics="true" pageId="0" pageNumber="15">Holotype</emphasis>
: ICN 2786 (Fig.
<figureCitation id="9F557987F62CDDDAFF0D4295E087949F" captionStart="Figure 19" captionStartId="F19" captionText="Figure 19. Adult male holotype of Bothriechis klebbai sp. nov. ICN 2786 in a. Dorsal and b. Ventral view. Photos by Duvan Zambrano." httpUri="https://binary.pensoft.net/fig/981705" pageId="0" pageNumber="15">19</figureCitation>
), adult male collected by Pedro M. Ruiz-Carranza on September 21, 1979 at Vereda
<normalizedToken id="239B956CFBA526EC70F2301081E9270C" originalValue="Virolín">Virolin</normalizedToken>
, municipio de
<normalizedToken id="332222185736C02DDCC4EBB9F8493112" originalValue="Charalá">Charala</normalizedToken>
, Santander department, Colombia (
<geoCoordinate id="B7B5650B4A671B68F2B1253099BD34DB" degrees="6.10720" direction="north" orientation="latitude" precision="1" value="6.1072">6.10720</geoCoordinate>
,
<geoCoordinate id="3CBB6A4BA2E4DD63922A2C4C1798E061" degrees="73.19781" direction="west" orientation="longitude" precision="1" value="-73.19781">-73.19781</geoCoordinate>
; 1819 m).
</paragraph>
<paragraph id="3D18CF0187505483D3E59227BBF14AC6" pageId="0" pageNumber="15">
<emphasis id="86428A36CF845F02BBCA3A96AF8E81E8" bold="true" italics="true" pageId="0" pageNumber="15">Paratypes</emphasis>
: All labeled
<taxonomicName id="FD106C959511089FD0F3D948C6D24D7D" authorityName="Arteaga &amp; Pyron &amp; Batista &amp; Vieira &amp; Meneses Pelayo &amp; Smith &amp; Barrio Amorós &amp; Koch &amp; Agne &amp; Valencia &amp; Bustamante &amp; Harris" authorityYear="2024" family="Viperidae" genus="Bothriechis" kingdom="Animalia" lsidName="Bothriechis klebbai" order="Squamata" pageId="0" pageNumber="15" rank="species" species="klebbai">
<emphasis id="58B8D2BAF942E980DC904E2CFEAAC0F3" italics="true" pageId="0" pageNumber="15">Bothriechis klebbai</emphasis>
</taxonomicName>
sp. nov. in Suppl. material 1.
</paragraph>
</subSubSection>
<subSubSection id="499A5D0374AAE7B504B4E9A663E2D1D3" pageId="0" pageNumber="15" type="proposed standard english name">
<paragraph id="CA1F7F406928DAB48330DE030FE8B8C9" pageId="0" pageNumber="15">Proposed standard English name.</paragraph>
<paragraph id="1CE44A2EDB439CA21B7E903AD062A6F6" pageId="0" pageNumber="15">
<normalizedToken id="85D5FF3A5D70D58DFD5C22AEBBD35E74" originalValue="Klebbas">Klebba's</normalizedToken>
Eyelash--Pitviper.
</paragraph>
</subSubSection>
<subSubSection id="97B711E1C4F604A0736754576E09D127" pageId="0" pageNumber="15" type="proposed standard spanish name">
<paragraph id="4F2098EEEE1DECB6D450B79FD929CDF9" pageId="0" pageNumber="15">Proposed standard Spanish name.</paragraph>
<paragraph id="7DACBFC6599406990CACA3FD9F36A972" pageId="0" pageNumber="15">
<normalizedToken id="E7584C1F6BD73C1024EB2189812B47FE" originalValue="Víbora">Vibora</normalizedToken>
de
<normalizedToken id="5B005F67025F440B7814275DEBFE63E1" originalValue="pestañas">pestanas</normalizedToken>
de Klebba.
</paragraph>
</subSubSection>
<subSubSection id="14A053CCD8B82F0E5D025DD03CEE76D5" pageId="0" pageNumber="15" type="diagnosis">
<paragraph id="A2DD58CBB4C06D13674D865563CEF690" pageId="0" pageNumber="15">Diagnosis.</paragraph>
<paragraph id="D4E82095FDDB852951E89A2CADB3153B" pageId="0" pageNumber="15">
<taxonomicName id="CBCB69614E54F2D216E09E19036209E5" authorityName="Arteaga &amp; Pyron &amp; Batista &amp; Vieira &amp; Meneses Pelayo &amp; Smith &amp; Barrio Amorós &amp; Koch &amp; Agne &amp; Valencia &amp; Bustamante &amp; Harris" authorityYear="2024" family="Viperidae" genus="Bothriechis" kingdom="Animalia" lsidName="Bothriechis klebbai" order="Squamata" pageId="0" pageNumber="15" rank="species" species="klebbai">
<emphasis id="6F7465FB7814EC7E916218B1E5F8DFFE" italics="true" pageId="0" pageNumber="15">Bothriechis klebbai</emphasis>
</taxonomicName>
sp. nov. is diagnosed based on the following combination of characters: (1) supraciliary scales low and granular or two raised, but not sharp; (2) anterior dorsal head scales smooth; (3) gular scales similar in size to chinshields; (4) 7-11 interoculolabials; (5) 2-4 canthals which may be flat or slightly raised forming a ridge along the canthus; (6) loreal in contact with preocular in ~1/3 of specimens; (7) yellow morph absent; (8) dorsal bands black; (9) opposing kidney shaped dorsal marks absent; (10) black speckles on dorsal scales present; (11) black speckling on ventral surfaces present, prominent; (12) ventral surfaces never entirely white; (13) iris rich dark reddish brown; (14) 21-23 dorsal scale rows at mid-body; (15) 144-151 ventrals in males, 141-151 in females; (16) maximum total length in males 671 mm, in females 874 mm.
</paragraph>
</subSubSection>
<subSubSection id="090DC31B2677E3D940D9B961061D9840" pageId="0" pageNumber="15" type="comparisons">
<paragraph id="02FB4B46B3C7DE3E8A616942848E17EE" pageId="0" pageNumber="15">Comparisons.</paragraph>
<paragraph id="20B55AB95CBA197D7FFA4C397CF48978" pageId="0" pageNumber="15">
<taxonomicName id="426911223BF3CB4BD8C5BE1BBAAFEDE0" authorityName="Arteaga &amp; Pyron &amp; Batista &amp; Vieira &amp; Meneses Pelayo &amp; Smith &amp; Barrio Amorós &amp; Koch &amp; Agne &amp; Valencia &amp; Bustamante &amp; Harris" authorityYear="2024" family="Viperidae" genus="Bothriechis" kingdom="Animalia" lsidName="Bothriechis klebbai" order="Squamata" pageId="0" pageNumber="15" rank="species" species="klebbai">
<emphasis id="A29803D7024A78C0A774D147FD41279B" italics="true" pageId="0" pageNumber="15">Bothriechis klebbai</emphasis>
</taxonomicName>
sp. nov. is compared to other species of the genus previously subsumed under
<taxonomicName id="8758F7293B72D59020AA49AB614F4DE3" lsidName="B. schlegelii" pageId="0" pageNumber="15" rank="species" species="schlegelii">
<emphasis id="C066E34AD8C28DEF64EEC419601A927E" italics="true" pageId="0" pageNumber="15">B. schlegelii</emphasis>
</taxonomicName>
<emphasis id="D1C6A15E0633CAFBD734927B1D46D995" italics="true" pageId="0" pageNumber="15">sensu lato</emphasis>
(differences summarized in Table
<tableCitation id="860742BC1AE71363DE7D1E99A49F7539" captionStart="Table 2" captionStartId="T2" captionText="Table 2. Differences in coloration, scale counts, hemipenial architecture, size, and median lethal dose (LD 50), between members of the Bothriechis schlegelii species complex. The range of each continuous variable is from our own sample (Suppl. material 1), Kuch and Freire (1995), and Solorzano et al. (1998)." httpUri="http://table.plazi.org/id/4C8B465DA02061B16CBCD59564B9E214" pageId="0" pageNumber="15" tableUuid="4C8B465DA02061B16CBCD59564B9E214">2</tableCitation>
). It differs from most of them by having supraciliaries low and granular or two raised but not sharp, a low number of canthals (2-4), gular scales similar in size to chinshields, and dorsal and ventral surfaces heavily speckled with black pigment.
<taxonomicName id="65CDC53D22C1F43483DF0965C6BF404B" authorityName="Arteaga &amp; Pyron &amp; Batista &amp; Vieira &amp; Meneses Pelayo &amp; Smith &amp; Barrio Amorós &amp; Koch &amp; Agne &amp; Valencia &amp; Bustamante &amp; Harris" authorityYear="2024" family="Viperidae" genus="Bothriechis" kingdom="Animalia" lsidName="Bothriechis klebbai" order="Squamata" pageId="0" pageNumber="15" rank="species" species="klebbai">
<emphasis id="AB78330BEA1D72E062FB01A52E90516B" italics="true" pageId="0" pageNumber="15">Bothriechis klebbai</emphasis>
</taxonomicName>
sp. nov. resembles
<taxonomicName id="ED36772D8F57FC46494FC8CF8C2C75A4" lsidName="B. rasikusumorum" pageId="0" pageNumber="15" rank="species" species="rasikusumorum">
<emphasis id="58D02932F3E316463EF65083E205E2FC" italics="true" pageId="0" pageNumber="15">B. rasikusumorum</emphasis>
</taxonomicName>
sp. nov., but this other species occurs on the southern portion of
<normalizedToken id="5A8F1DC5CE3C520190A69E1FA8956D68" originalValue="Colombias">Colombia's</normalizedToken>
Cordillera Oriental and is characterized by having more than 151 ventrals in females, loreal not in contact with preocular, and by lacking paired oblong blotches on each side of the dorsum (a characteristic variably present in
<taxonomicName id="E761F0EB15F6EF01A3812C02A0A6A879" lsidName="B. klebbai" pageId="0" pageNumber="15" rank="species" species="klebbai">
<emphasis id="D8A3B68FB7FF8E80D2CB16351010D731" italics="true" pageId="0" pageNumber="15">B. klebbai</emphasis>
</taxonomicName>
sp. nov.). Furthermore,
<taxonomicName id="33FA9677214133E96764A4F4D2101C97" authorityName="Arteaga &amp; Pyron &amp; Batista &amp; Vieira &amp; Meneses Pelayo &amp; Smith &amp; Barrio Amorós &amp; Koch &amp; Agne &amp; Valencia &amp; Bustamante &amp; Harris" authorityYear="2024" family="Viperidae" genus="Bothriechis" kingdom="Animalia" lsidName="Bothriechis klebbai" order="Squamata" pageId="0" pageNumber="15" rank="species" species="klebbai">
<emphasis id="6D3EE2C2F6F2CE756876ED1DA74B69A6" italics="true" pageId="0" pageNumber="15">Bothriechis klebbai</emphasis>
</taxonomicName>
sp. nov. lacks enlarged basal hooks on the hemipenial body (present in
<taxonomicName id="01B62C385A356802F691099CCAB59802" lsidName="B. rasikusumorum" pageId="0" pageNumber="15" rank="species" species="rasikusumorum">
<emphasis id="675C91E63C0A90B9F4FD8F48A2E1EFB2" italics="true" pageId="0" pageNumber="15">B. rasikusumorum</emphasis>
</taxonomicName>
sp. nov.).
<taxonomicName id="8E5263DC7CFBA737BD6FD468213498FD" authorityName="Arteaga &amp; Pyron &amp; Batista &amp; Vieira &amp; Meneses Pelayo &amp; Smith &amp; Barrio Amorós &amp; Koch &amp; Agne &amp; Valencia &amp; Bustamante &amp; Harris" authorityYear="2024" family="Viperidae" genus="Bothriechis" kingdom="Animalia" lsidName="Bothriechis klebbai" order="Squamata" pageId="0" pageNumber="15" rank="species" species="klebbai">
<emphasis id="2FEDF82E71F3C5B5BC51A9A61641BF32" italics="true" pageId="0" pageNumber="15">Bothriechis klebbai</emphasis>
</taxonomicName>
sp. nov. also resembles
<taxonomicName id="0F9CDE2FFEB9D2C76A5229BE49775915" lsidName="B. schlegelii" pageId="0" pageNumber="15" rank="species" species="schlegelii">
<emphasis id="5AD1E27A284A5ED58E32D444F9CC4250" italics="true" pageId="0" pageNumber="15">B. schlegelii</emphasis>
</taxonomicName>
, but this other species does not occur on the Cordillera Oriental and is characterized by having two triangular and moderately raised supraciliary scales, a different iris color (pale orange, light green, or golden yellow usually without speckles vs rich dark reddish brown), and by lacking black speckles on the dorsal and ventral scales in most populations (Figs
<figureCitation id="42A253F80A2CF6AB1C102F87388CB853" captionStart="Figure 19" captionStartId="F19" captionText="Figure 19. Adult male holotype of Bothriechis klebbai sp. nov. ICN 2786 in a. Dorsal and b. Ventral view. Photos by Duvan Zambrano." httpUri="https://binary.pensoft.net/fig/981705" pageId="0" pageNumber="15">19</figureCitation>
,
<figureCitation id="456D21F6046254AD5D9F226EE6A31D2B" captionStart="Figure 20" captionStartId="F20" captionText="Figure 20. Photographs of some specimens of Bothriechis klebbai sp. nov. in life from Finca la Arrinconada, vereda Esparta, minicipio de Santa Barbara, Santander department, Colombia. a. UIS-R 4139 adult male; b. INSV-SR- 00098 adult of undetermined sex; and c. UIS-R- 4296 juvenile male. Photos by Elson Meneses Pelayo and Jose Vieira." httpUri="https://binary.pensoft.net/fig/981706" pageId="0" pageNumber="15">20</figureCitation>
).
<taxonomicName id="859A87B8C0E2D81D154397149092E8F5" authorityName="Arteaga &amp; Pyron &amp; Batista &amp; Vieira &amp; Meneses Pelayo &amp; Smith &amp; Barrio Amorós &amp; Koch &amp; Agne &amp; Valencia &amp; Bustamante &amp; Harris" authorityYear="2024" family="Viperidae" genus="Bothriechis" kingdom="Animalia" lsidName="Bothriechis klebbai" order="Squamata" pageId="0" pageNumber="15" rank="species" species="klebbai">
<emphasis id="2A15F3C5507DFBF87A3E8C48124593BD" italics="true" pageId="0" pageNumber="15">Bothriechis klebbai</emphasis>
</taxonomicName>
sp. nov. occurs in sympatry with
<taxonomicName id="9E1AA901BE5B4481981437F671F05140" lsidName="B. khwargi" pageId="0" pageNumber="15" rank="species" species="khwargi">
<emphasis id="835F8692A78A30BBCAE34AF5155D3382" italics="true" pageId="0" pageNumber="15">B. khwargi</emphasis>
</taxonomicName>
sp. nov. along the upper slopes of the
<normalizedToken id="DEF094A1FCB9073836767388FDA54E14" originalValue="Serranía">Serrania</normalizedToken>
De Los
<normalizedToken id="D3C99733598A2B2CD94060BAC10061E3" originalValue="Yariguíes">Yariguies</normalizedToken>
, but it differs from this other species by being larger in body size, having a dark green belly, smooth anterior dorsal head scales, gulars similar in size to or large than chinshields, and by having prominent black speckling on dorsal and ventral surfaces.
</paragraph>
<caption id="BB63CAFD3375A23F250B0A7E4B419ADA" httpUri="https://binary.pensoft.net/fig/981705" pageId="0" pageNumber="15" start="Figure 19" startId="F19">
<paragraph id="054C92C37C117C027D31A3EB96C16864" pageId="0" pageNumber="15">
<emphasis id="591F5B4616F5A12D1CD897FFB9CA7D7A" bold="true" pageId="0" pageNumber="15">Figure 19.</emphasis>
Adult male holotype of
<taxonomicName id="B178CCD265A6927020C538199C0CD267" authorityName="Arteaga &amp; Pyron &amp; Batista &amp; Vieira &amp; Meneses Pelayo &amp; Smith &amp; Barrio Amorós &amp; Koch &amp; Agne &amp; Valencia &amp; Bustamante &amp; Harris" authorityYear="2024" family="Viperidae" genus="Bothriechis" kingdom="Animalia" lsidName="Bothriechis klebbai" order="Squamata" pageId="0" pageNumber="15" rank="species" species="klebbai">
<emphasis id="2D896F1D06D0BAFA7FC3D33FF45FD8EA" italics="true" pageId="0" pageNumber="15">Bothriechis klebbai</emphasis>
</taxonomicName>
sp. nov. ICN 2786 in
<emphasis id="6B5B6B97C099784806A5B0CCE15609B1" bold="true" pageId="0" pageNumber="15">a.</emphasis>
Dorsal and
<emphasis id="D47F089C33533D6F06C4762C67814F17" bold="true" pageId="0" pageNumber="15">b.</emphasis>
Ventral view. Photos by
<normalizedToken id="2CD495F3DC79C25530AEBC1F7E239FD8" originalValue="Duván">Duvan</normalizedToken>
Zambrano.
</paragraph>
</caption>
<caption id="9F8B0A57799E4D0FA4F943B655EAFA9C" httpUri="https://binary.pensoft.net/fig/981706" pageId="0" pageNumber="15" start="Figure 20" startId="F20">
<paragraph id="41F7DCF79F27594E83E45DFD62EE4C77" pageId="0" pageNumber="15">
<emphasis id="60130AB9E8EC86620F8C0CE8F17D4026" bold="true" pageId="0" pageNumber="15">Figure 20.</emphasis>
Photographs of some specimens of
<taxonomicName id="C1C394F6B58FB4BBD4BB754C943E9C32" authorityName="Arteaga &amp; Pyron &amp; Batista &amp; Vieira &amp; Meneses Pelayo &amp; Smith &amp; Barrio Amorós &amp; Koch &amp; Agne &amp; Valencia &amp; Bustamante &amp; Harris" authorityYear="2024" family="Viperidae" genus="Bothriechis" kingdom="Animalia" lsidName="Bothriechis klebbai" order="Squamata" pageId="0" pageNumber="15" rank="species" species="klebbai">
<emphasis id="833DDA74DFA772D9113FF3455B3CA1CF" italics="true" pageId="0" pageNumber="15">Bothriechis klebbai</emphasis>
</taxonomicName>
sp. nov. in life from Finca la Arrinconada, vereda Esparta, minicipio de Santa
<normalizedToken id="F24F496D328521D829EDA639C9D386FE" originalValue="Bárbara">Barbara</normalizedToken>
, Santander department, Colombia.
<emphasis id="E110CD42F5C789A5DB93A8B55D660908" bold="true" pageId="0" pageNumber="15">a.</emphasis>
UIS-R 4139 adult male;
<emphasis id="A35BE75F5CDC088ED9E95BE5F34A78C2" bold="true" pageId="0" pageNumber="15">b.</emphasis>
INSV-SR-00098 adult of undetermined sex; and
<emphasis id="925902D689779E9B21FBC3C768BD6D86" bold="true" pageId="0" pageNumber="15">c.</emphasis>
UIS-R-4296 juvenile male. Photos by Elson Meneses Pelayo and Jose Vieira.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="2BB7ACCDBCDF1A0284B8E83041C12C5C" pageId="0" pageNumber="15" type="hemipenial morphology">
<paragraph id="AF1DAD98E7840B8FBBCE96BA3D63409A" pageId="0" pageNumber="15">Hemipenial morphology.</paragraph>
<paragraph id="2D768CD1DEE8154E55FA4D713B66266B" pageId="0" pageNumber="15">
(n = 3; Fig.
<figureCitation id="E441B3432D7DB2EF5B6D400F0C1900BA" captionStart="Figure 21" captionStartId="F21" captionText="Figure 21. Hemipenial architecture of Bothriechis klebbai sp. nov. in sulcate, lateral, and asulcate views. a. ICN 2786 and b. ICN 2855 from Virolin, municipio de Charala, Santander department, Colombia; c. UIS-R 4803 from vereda Naranjal, municipio de Coromoro, Santander department, Colombia. Photos by Duvan Zambrano and Elson Meneses Pelayo." httpUri="https://binary.pensoft.net/fig/981707" pageId="0" pageNumber="15">21</figureCitation>
) Everted and inflated, the organ is deeply bilobed, calyculate and noncapitate; hemipenial lobes cylindrical or spindle shaped; in sulcate and asulcate views, lobe crotch ornamented with densely packed small spines that become larger distally; sulcus spermaticus centrolineal, bifurcate and with walls weakly defined (strongly in ICN 2786), bifurcation occurs below bilobation point and proximal to the base of the hemipenial body; sulcus spermaticus branch runs to lobe tips; distal half of each hemipenal lobe densely ornamented with small calyces with strongly defined spinulate edges. In sulcate view, hemipenial body surface spinulate medially; calcified spines at the base of the hemipenial body only slightly larger than adjacent rows of smaller obliquely arranged mesial spines that become gradually smaller towards each
<normalizedToken id="3FF812EB7EB35B211466FF5A3231AF7A" originalValue="lobes">lobe's</normalizedToken>
capitulum (=no obvious basal hooks); each hemipenial lobe ornamented with mesial and lateral spines homogenous in size; the spines in each lobe are replaced distally by calyces with strongly spinulate edges. In lateral view, hemipenial body and lobes with rows of spines, but otherwise non-spiculate; distal half strongly calyculate. In asulcate view, the center of hemipenial body is nude to barely spiculate; hemipenial lobes ornamented with 4 rows of spines that decrease in size towards the lobe crotch.
</paragraph>
<caption id="A48E69D4D469A3C11D0E4C339A0B7110" httpUri="https://binary.pensoft.net/fig/981707" pageId="0" pageNumber="15" start="Figure 21" startId="F21">
<paragraph id="C5AF29574009ED0E954185ED551F744B" pageId="0" pageNumber="15">
<emphasis id="9948220675B780209C0F1CF6A40909FE" bold="true" pageId="0" pageNumber="15">Figure 21.</emphasis>
Hemipenial architecture of
<taxonomicName id="79A6EE72AF75FFC4999C8954C7182357" authorityName="Arteaga &amp; Pyron &amp; Batista &amp; Vieira &amp; Meneses Pelayo &amp; Smith &amp; Barrio Amorós &amp; Koch &amp; Agne &amp; Valencia &amp; Bustamante &amp; Harris" authorityYear="2024" family="Viperidae" genus="Bothriechis" kingdom="Animalia" lsidName="Bothriechis klebbai" order="Squamata" pageId="0" pageNumber="15" rank="species" species="klebbai">
<emphasis id="DB6F01B4823227AD6B6DE943F80A7AED" italics="true" pageId="0" pageNumber="15">Bothriechis klebbai</emphasis>
</taxonomicName>
sp. nov. in sulcate, lateral, and asulcate views.
<emphasis id="4846784E5B2CC01643DB46A5DF5905C9" bold="true" pageId="0" pageNumber="15">a.</emphasis>
ICN 2786 and
<emphasis id="8676936B314DFB457A2463B58A1ECDCE" bold="true" pageId="0" pageNumber="15">b.</emphasis>
ICN 2855 from
<normalizedToken id="6062267707370D84D18D998DF7088675" originalValue="Virolín">Virolin</normalizedToken>
, municipio de
<normalizedToken id="E4A2AB16C3BD8752A5DA8711BA2E0AE6" originalValue="Charalá">Charala</normalizedToken>
, Santander department, Colombia;
<emphasis id="40F72BE643EBCDD105B4839A3E0994F1" bold="true" pageId="0" pageNumber="15">c.</emphasis>
UIS-R 4803 from vereda Naranjal, municipio de Coromoro, Santander department, Colombia. Photos by
<normalizedToken id="D690FF63C9DE23082B543B3AA44E5889" originalValue="Duván">Duvan</normalizedToken>
Zambrano and Elson Meneses Pelayo.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="4D6EAE187D5AC23F1066579EE7225B00" pageId="0" pageNumber="15" type="description">
<paragraph id="C531872161CB1C63DAFDB350C02082BF" pageId="0" pageNumber="15">Description of holotype.</paragraph>
<paragraph id="8D195AEC188DB0FC54B8781047465DC1" pageId="0" pageNumber="15">
An adult male, SVL 349 mm, tail length 75 mm (21.5% SVL); head length 27.8 mm (8.0% SVL) from tip of snout to angle of jaw; head width 20.1 mm (72% head length) taken at broadest point; rostral broader than high (2.7
<normalizedToken id="EA2439CC6522F88DD05370A40E626D84" originalValue="×">x</normalizedToken>
1.8 mm); nasal divided and not fused with first supralabial; loreal about 1/2 size of pit, contacting postnasal, 2 canthals, lacunolabial, supralacunal, and preocular; prefoveals 4/4; subfoveals 3/4; postfoveals 0; prelacunal fused with second supralabial; sublacunals 2/2 (one of which is small and granular); supralacunal elongated and in contact with orbit; preoculars 1/1 (2/2 if supralacunal is considered a preocular); suboculars 1/1; postoculars 1/1; loreal pit large, directed anteriorly, located slightly below line drawn from center of eye to naris; supralabials 8/8 (including lacunolabial); infralabials 11/11, first meet posteriorly; mental broader than long (3.3
<normalizedToken id="E0B1CD2540184D9B3F536CEE9CB6DE96" originalValue="×">x</normalizedToken>
3.0 mm); 1 pair of chinshields; 5 pairs of gulars between chinshields and preventrals; preventral 1; anterior internasals 2; canthals 3/3; supraciliary scales low and granular with the exception of one raised and triangular scale; supraoculars oblong, 1.8
<normalizedToken id="26FFF3546451ABCB07044783DC4B1CC3" originalValue="×">x</normalizedToken>
longer than wide; intersupraoculars 6; anterior dorsal head scales smooth; posterior head scales barely keeled; interrictals 27; dorsal scale rows 21/21/18; ventrals 148; cloacal plate entire; 56 undivided subcaudals; tail prehensile.
</paragraph>
</subSubSection>
<subSubSection id="3CB9BE4097EB572AD81F3808F5FB5871" pageId="0" pageNumber="15" type="natural history">
<paragraph id="D53A8CF992DEDA3DFF72D05CF93F2D94" pageId="0" pageNumber="15">Natural history.</paragraph>
<paragraph id="BE6B1DBDD7C7BDEDAD61B76DFD7467D1" pageId="0" pageNumber="15">
<taxonomicName id="38F718BB2EFB8E16E55A6B625F340E92" authorityName="Arteaga &amp; Pyron &amp; Batista &amp; Vieira &amp; Meneses Pelayo &amp; Smith &amp; Barrio Amorós &amp; Koch &amp; Agne &amp; Valencia &amp; Bustamante &amp; Harris" authorityYear="2024" family="Viperidae" genus="Bothriechis" kingdom="Animalia" lsidName="Bothriechis klebbai" order="Squamata" pageId="0" pageNumber="15" rank="species" species="klebbai">
<emphasis id="0EB3A5C9A7DABA0998E3A1863C39FF0D" italics="true" pageId="0" pageNumber="15">Bothriechis klebbai</emphasis>
</taxonomicName>
sp. nov. is an arboreal snake that inhabits montane cloud forests. Vipers of this species have been seen at night or during the day, either at ground level or on low understory vegetation. In captivity, this species is known to hybridize with
<taxonomicName id="61DE531BAA5DB7E37896B8A345A66B9C" lsidName="B. khwargi" pageId="0" pageNumber="15" rank="species" species="khwargi">
<emphasis id="138EDAE7A55313AADCF87EDCFA23D863" italics="true" pageId="0" pageNumber="15">B. khwargi</emphasis>
</taxonomicName>
sp. nov. INSV-SR-1015 is believed to be a hybrid between the two species. We (JV and EMP) have seen this species preying upon
<taxonomicName id="682A2088487C34475D6440C83EDE9D42" authorityName="Dumeril" authorityYear="1851" class="Reptilia" family="Iguanidae" genus="Anolis" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Anolis heterodermus" order="Squamata" pageId="0" pageNumber="15" phylum="Chordata" rank="species" species="heterodermus">
<emphasis id="7B1F440B3F5F38A7EC3702EF85A91157" italics="true" pageId="0" pageNumber="15">Anolis heterodermus</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="75085FBAC28537B98DF0CA3057DAA565" pageId="0" pageNumber="15" type="venom">
<paragraph id="586FC2A7C2FB3711A91075891739B4E9" pageId="0" pageNumber="15">Venom.</paragraph>
<paragraph id="788321D5D65CE3BF953C7E0DADE9959D" pageId="0" pageNumber="15">We know of no snakebites caused by this species in Colombia nor any studies on the venom that could be assigned to this species.</paragraph>
</subSubSection>
<subSubSection id="489A69D3A4284875FBE0273D0FD6A580" pageId="0" pageNumber="15" type="distribution">
<paragraph id="A3192813A03A46F785A2B51DC01F83FD" pageId="0" pageNumber="15">Distribution.</paragraph>
<paragraph id="3B5791B49DCA9ECBDD0BC911C013FEF7" pageId="0" pageNumber="15">
<taxonomicName id="4303F5BE471E89AE7083DFE4BDC25267" authorityName="Arteaga &amp; Pyron &amp; Batista &amp; Vieira &amp; Meneses Pelayo &amp; Smith &amp; Barrio Amorós &amp; Koch &amp; Agne &amp; Valencia &amp; Bustamante &amp; Harris" authorityYear="2024" family="Viperidae" genus="Bothriechis" kingdom="Animalia" lsidName="Bothriechis klebbai" order="Squamata" pageId="0" pageNumber="15" rank="species" species="klebbai">
<emphasis id="54C82EF8F61A441AF0F9BEDD87820B96" italics="true" pageId="0" pageNumber="15">Bothriechis klebbai</emphasis>
</taxonomicName>
sp. nov. is known from at least 23 localities (listed in Suppl. material 3) along the western slopes of
<normalizedToken id="15822142FF5E25DCDBEB4E27DFDC86D9" originalValue="Colombias">Colombia's</normalizedToken>
Cordillera Orien-tal. The species occurs over an estimated 20,061 km2 and has been recorded at elevations 1,418-2,380 m above sea level (Fig.
<figureCitation id="A6CFC8181298E8285FDBC3EB41AF3CC0" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Distribution of species of Bothriechis previously subsumed under B. schlegelii sensu lato. Each colored area is a geographic representation of the suitable environmental conditions for one of the clades recovered in the phylogeny of Fig. 2. Lime green shade in southeastern Panama represents overlap between between B. nigroadspersus and B. torvus." httpUri="https://binary.pensoft.net/fig/981689" pageId="0" pageNumber="15">3</figureCitation>
). Approximately 21% of the predicted area of distribution of
<taxonomicName id="C8E76D4C49B7129D01AE4E1BB2FA7D8E" lsidName="B. klebbai" pageId="0" pageNumber="15" rank="species" species="klebbai">
<emphasis id="996F8C3311263F44C715CDE2009ACD9A" italics="true" pageId="0" pageNumber="15">B. klebbai</emphasis>
</taxonomicName>
sp. nov. overlaps with that of
<taxonomicName id="AF3E2C0D5C7704BB192E45B8CBAE9186" lsidName="B. khwargi" pageId="0" pageNumber="15" rank="species" species="khwargi">
<emphasis id="41F24A2ACA6B0B1890D69B2321E3872D" italics="true" pageId="0" pageNumber="15">B. khwargi</emphasis>
</taxonomicName>
sp. nov.
</paragraph>
</subSubSection>
<subSubSection id="867E8CEACC01A580785B1C71C1B1BE16" pageId="0" pageNumber="15" type="etymology">
<paragraph id="89F45BEC9C5604483C340C0F01933D19" pageId="0" pageNumber="15">Etymology.</paragraph>
<paragraph id="682A3436231A63715EA7DAEDE181040E" pageId="0" pageNumber="15">
Named after Casey Klebba, whose dedication to the preservation of tropical biodiversity is exemplary. As a co-founder of Minifund.org, alongside Carly Jones, he has been one of the most invaluable supporters of
<normalizedToken id="2039505CCB152136D74A7F3A0DEEC056" originalValue="AAs">AA's</normalizedToken>
field expeditions to remote areas of Ecuador and Colombia, resulting in the discovery of many new species to science.
</paragraph>
</subSubSection>
<subSubSection id="6776F07F9B2C323E1FCF2CC4D1117C49" pageId="0" pageNumber="15" type="conservation status">
<paragraph id="5DA408ABFB7C266A2477F7B3B95BF1F7" pageId="0" pageNumber="15">Conservation status.</paragraph>
<paragraph id="4EE106C8ACC3C65EE831520AD4089B11" pageId="0" pageNumber="15">
We consider
<taxonomicName id="900161D16CF62BEEE2E5846B6AED5E98" authorityName="Arteaga &amp; Pyron &amp; Batista &amp; Vieira &amp; Meneses Pelayo &amp; Smith &amp; Barrio Amorós &amp; Koch &amp; Agne &amp; Valencia &amp; Bustamante &amp; Harris" authorityYear="2024" family="Viperidae" genus="Bothriechis" kingdom="Animalia" lsidName="Bothriechis klebbai" order="Squamata" pageId="0" pageNumber="15" rank="species" species="klebbai">
<emphasis id="C162B32D7FBD1EBDAEFF4BA36FD9590D" italics="true" pageId="0" pageNumber="15">Bothriechis klebbai</emphasis>
</taxonomicName>
sp. nov. to be included in the Near Threatened category following IUCN Red List criteria (
<bibRefCitation id="EB89668FD66EB3232268B86018C13FE1" author="Hurtado-Gomez, JP" journalOrPublisher="BSc thesis, Universidad de Antioquia" pageId="0" pageNumber="15" refId="B48" refString="IUCN (2012) IUCN Red List categories and criteria: Version 3.1. Second edition.IUCN Species Survival Commission, Gland and Cambridge, 32 pp." year="2009">IUCN 2012</bibRefCitation>
) primarily because the
<normalizedToken id="B1EA5B4FAFB8EBC0291A56F64D9EAE6A" originalValue="species">species'</normalizedToken>
extent of occurrence is estimated to be greater than the 20,000 km2 needed to meet B1 criteria for the Vulnerable category. However, although the species occurs in at least two national parks (
<normalizedToken id="64CE04C3B6CE8178CD51DA1C04290B31" originalValue="Serranía">Serrania</normalizedToken>
de los
<normalizedToken id="C8DC8048ADBC287FF6B9DDCCE14DB668" originalValue="Yariguíes">Yariguies</normalizedToken>
and
<normalizedToken id="EE487EB8E773F57B0C1F09804D0ACA26" originalValue="Guanentá-alto">Guanenta-alto</normalizedToken>
<normalizedToken id="78E6D0B864CCDD744692F56465680160" originalValue="Río">Rio</normalizedToken>
Fonce), the majority of the
<normalizedToken id="177FAB2B4B51B5A739FE148DDF2F2D4B" originalValue="species">species'</normalizedToken>
montane forest habitat has been destroyed. Based on the distribution model presented in Fig.
<figureCitation id="19C712A10F741E0855D5B76FBE18FC27" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Distribution of species of Bothriechis previously subsumed under B. schlegelii sensu lato. Each colored area is a geographic representation of the suitable environmental conditions for one of the clades recovered in the phylogeny of Fig. 2. Lime green shade in southeastern Panama represents overlap between between B. nigroadspersus and B. torvus." httpUri="https://binary.pensoft.net/fig/981689" pageId="0" pageNumber="15">3</figureCitation>
in combination with maps of vegetation cover of Colombia (
<bibRefCitation id="50B2893BF5405B94D3235179CB08783D" author="Hurtado-Gomez, JP" journalOrPublisher="BSc thesis, Universidad de Antioquia" pageId="0" pageNumber="15" refId="B47" refString="IDEAM (2014) Mapa de cobertura de la tierra adaptada para Colombia." year="2009">IDEAM 2014</bibRefCitation>
), we estimate that only ~14% of the
<normalizedToken id="D85A7AB3FA9D64E7D361CFB9C8DB4DDE" originalValue="species">species'</normalizedToken>
forest habitat is still standing.
</paragraph>
</subSubSection>
</treatment>
</document>