treatments-xml/data/3B/7F/86/3B7F86708DC858229A4A8510B21F23FA.xml

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<mods:title id="CE043F800217ED7356428DA81BE2543D">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:affiliation id="DB25F916E6025A4B1A4FD332B5A6D51E">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="545D9DF57EFE8834173C6A37D448E918">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:date id="9594801EF7A434CB03E490BAD94A0B32">2024</mods:date>
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<treatment id="3B7F86708DC858229A4A8510B21F23FA" ID-GBIF-Taxon="221225637" LSID="urn:lsid:plazi:treatment:3B7F86708DC858229A4A8510B21F23FA" httpUri="http://treatment.plazi.org/id/3B7F86708DC858229A4A8510B21F23FA" lastPageNumber="15" pageId="0" pageNumber="15" scope_family="Viperidae">
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<paragraph id="85939B9DF681806FE380C19445E00D04" pageId="0" pageNumber="15">
<taxonomicName id="36329AEEB82A5958E30F4440C0E79124" ID-CoL="563YW" LSID="3B7F8670-8DC8-5822-9A4A-8510B21F23FA" authority="(Posada Arango, 1889 a)" baseAuthorityName="Posada Arango" baseAuthorityYear="1889" family="Viperidae" genus="Bothriechis" higherTaxonomySource="treatment-meta" kingdom="Animalia" lsidName="Bothriechis torvus" order="Squamata" pageId="0" pageNumber="15" rank="species" species="torvus">Bothriechis torvus (Posada Arango, 1889a)</taxonomicName>
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<paragraph id="51FD57BD05903D5C6B778742600BBD44" pageId="0" pageNumber="15">
<figureCitation id="1674322D9120320521CB76F2E13C93E8" captionStart="Figure 15" captionStartId="F15" captionText="Figure 15. Photographs of some specimens of Bothriechis torvus in life. a. Adult and b. Juvenile (same individual) from Morromico Reserve, Choco department, Colombia; c. TH 145 adult female from Morromico Reserve, Choco department, Colombia; d. Adult and e. Juvenile from Uraba, Antioquia department, Colombia. Photos by Jose Vieira." httpUri="https://binary.pensoft.net/fig/981701" pageId="0" pageNumber="15">Figs 15</figureCitation>
<figureCitation id="C421199CD9B13BCA9607C0BACB2DCCDD" captionStart="Figure 16" captionStartId="F16" captionText="Figure 16. Hemipenial architecture of Bothriechis torvus in sulcate, lateral, and asulcate views. ICN 0421 from Valle del Cauca department, Colombia. Photos by Duvan Zambrano." httpUri="https://binary.pensoft.net/fig/981702" pageId="0" pageNumber="15">, 16</figureCitation>
</paragraph>
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<paragraph id="9479409A8D81BAD240A880C71A2F1400" pageId="0" pageNumber="15">
<taxonomicName id="8634C401EB1202EDB9FC1C9E53B55444" authorityName="Posada Arango" authorityYear="1889" family="Viperidae" genus="Thanatophis" higherTaxonomySource="treatment-meta" kingdom="Animalia" lsidName="Thanatophis torvus" order="Squamata" pageId="0" pageNumber="15" rank="species" species="torvus">Thanatophis torvus</taxonomicName>
Posada Arango, 1889a: 48. Type(s): not designated. Type locality: &quot;lugares 
<normalizedToken id="488FEBBBC7B07E4FE8388D29B04F4714" originalValue="cálidos">calidos</normalizedToken>
de Antioquia.&quot;
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<taxonomicName id="AB410C9AAB8B2AC4B070F2033B40B839" authorityName="Posada Arango" authorityYear="1889" family="Viperidae" genus="Thanatophis" higherTaxonomySource="treatment-meta" kingdom="Animalia" lsidName="Thanatophis torvus" order="Squamata" pageId="0" pageNumber="15" rank="species" species="torvus">Thanatophis torvus</taxonomicName>
Posada Arango, 1889b: 345. Type(s): not designated. Type locality: &quot;regions chaudes 
<normalizedToken id="DC606C145A31291385DA7E0E9AED124F" originalValue="d’Antioquia.”">d'Antioquia.&quot;</normalizedToken>
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<taxonomicName id="CC01B8ADC9C59FAA8FFE9ACF3F602CF3" family="Viperidae" genus="Teleuraspis" higherTaxonomySource="treatment-meta" kingdom="Animalia" lsidName="Teleuraspis birri" order="Squamata" pageId="0" pageNumber="15" rank="species" species="birri">Teleuraspis birri</taxonomicName>
Posada Arango, 1909: 231.
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<subSubSection id="926513AF88A3E0DB09CE35FD4191467E" pageId="0" pageNumber="15" type="referred specimens">
<paragraph id="EE4E7B44B59BCDD50DAC87D28D1A2463" pageId="0" pageNumber="15">Referred specimens.</paragraph>
<paragraph id="4112494C115C8EA794F943524A3E099D" pageId="0" pageNumber="15">
All labeled 
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<emphasis id="55F59000F69EF8A77A511625A69D47CA" italics="true" pageId="0" pageNumber="15">Bothriechis torvus</emphasis>
</taxonomicName>
in Suppl. material 1.
</paragraph>
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<subSubSection id="1F54CE2383D887ECA3F285DABD0F2744" pageId="0" pageNumber="15" type="proposed standard english name">
<paragraph id="516E2E46F9CDFF96F52747AA8838A3A6" pageId="0" pageNumber="15">Proposed standard English name.</paragraph>
<paragraph id="C5B5F872A30BB53FD0A129CD22225EBE" pageId="0" pageNumber="15">Birri Eyelash--Pitviper.</paragraph>
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<paragraph id="B82F8B3C2CAA6C418B4F2FD42E87684A" pageId="0" pageNumber="15">Spanish names.</paragraph>
<paragraph id="D386C4F1B49BB2B02AFDBFBA91FD45DF" pageId="0" pageNumber="15">
<normalizedToken id="B465397F8AB599C7105EC2730B7AA5BE" originalValue="Víbora">Vibora</normalizedToken>
<normalizedToken id="90D7B37FC73AB5A127D0E1C311F6242F" originalValue="birrí">birri</normalizedToken>
, serpiente guinda, 
<normalizedToken id="8FBA954DBAF73B8E8529414CEE642220" originalValue="pestañona">pestanona</normalizedToken>
.
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<subSubSection id="B98C0F366695AAD313BE5521F64B5081" pageId="0" pageNumber="15" type="diagnosis">
<paragraph id="75585D9521DAD1EA8D3EC9AA48E55764" pageId="0" pageNumber="15">Diagnosis.</paragraph>
<paragraph id="B23445115B87BC65E38C0B79D0DEBBA5" pageId="0" pageNumber="15">
<taxonomicName id="3EED6F0B3BBEF9AC276E1AD84C21EB3B" baseAuthorityName="Posada Arango" baseAuthorityYear="1889" family="Viperidae" genus="Bothriechis" kingdom="Animalia" lsidName="Bothriechis torvus" order="Squamata" pageId="0" pageNumber="15" rank="species" species="torvus">
<emphasis id="6CECBA49E086DA570310027ECA70D862" italics="true" pageId="0" pageNumber="15">Bothriechis torvus</emphasis>
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is diagnosed based on the following combination of characters: (1) 3 raised supraciliary scales, triangular, but rounded and not spinelike; (2) anterior dorsal head scales keeled; (3) gular scales much smaller than chinshields; (4) 6-18 interoculolabials; (5) 3-8 canthals, most of them with raised triangular projections; (6) loreal in contact with preocular in some populations; (7) yellow morph absent; (8) dorsal bands pink or red; (9) no opposing kidney shaped dorsal marks; (10) black speckles on dorsal scales absent; (11) black speckling on ventral surfaces absent or faint; (12) ventral surfaces entirely white in some individuals; (13) iris pale bluish-green or straw yellow with fine black speckling; (14) 21-23 dorsal scale rows at mid-body; (15) 148-159 ventrals in males, 141-153 in females; (16) maximum total length in males 378 mm, in females 657 mm.
</paragraph>
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<paragraph id="C6482FB0B508E7E5F8CFBCD9DEA6C58F" pageId="0" pageNumber="15">Comparisons.</paragraph>
<paragraph id="62D2D57AF14B1DF0475C474007AEA3FE" pageId="0" pageNumber="15">
<taxonomicName id="599CFEA7267A94D8336AB745D1D0382C" baseAuthorityName="Posada Arango" baseAuthorityYear="1889" family="Viperidae" genus="Bothriechis" kingdom="Animalia" lsidName="Bothriechis torvus" order="Squamata" pageId="0" pageNumber="15" rank="species" species="torvus">
<emphasis id="C963292D6110383E2DA76FBB9148B3CF" italics="true" pageId="0" pageNumber="15">Bothriechis torvus</emphasis>
</taxonomicName>
is compared to other species of the genus previously subsumed under 
<taxonomicName id="A16B07C99A3F8DA9F9B748A03E522C29" lsidName="B. schlegelii" pageId="0" pageNumber="15" rank="species" species="schlegelii">
<emphasis id="AD99AC7D7FB76498116E92B371674957" italics="true" pageId="0" pageNumber="15">B. schlegelii</emphasis>
</taxonomicName>
<emphasis id="72974BCBEB85D0604FA4A8820BDCF63D" italics="true" pageId="0" pageNumber="15">sensu lato</emphasis>
(differences summarized in Table 
<tableCitation id="07944388AA3A0FCBB88F0A57E290DFEF" 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 all of them by having three raised and triangular supraciliary scales, anterior dorsal head scales keeled, a high number of canthals (3-8) and interoculolabials (6-18), thin pink dorsal bands, and entirely pale white belly. 
<taxonomicName id="8758D2541F520BED15F8E052B69345A6" baseAuthorityName="Posada Arango" baseAuthorityYear="1889" family="Viperidae" genus="Bothriechis" kingdom="Animalia" lsidName="Bothriechis torvus" order="Squamata" pageId="0" pageNumber="15" rank="species" species="torvus">
<emphasis id="43053ABF00C074EE05A458706BB575CD" italics="true" pageId="0" pageNumber="15">Bothriechis torvus</emphasis>
</taxonomicName>
differs from 
<taxonomicName id="B6D99D7AA8700244FC9AD938B41B5A48" lsidName="B. schlegelii" pageId="0" pageNumber="15" rank="species" species="schlegelii">
<emphasis id="5BDE90070BF9B69B83A007F98B0B7BD0" italics="true" pageId="0" pageNumber="15">B. schlegelii</emphasis>
</taxonomicName>
<emphasis id="57EB960F35DEB589EA9A534E7D3582F5" italics="true" pageId="0" pageNumber="15">sensu stricto</emphasis>
by having anterior dorsal head scales keeled, 3-8 canthals (rather than 1-3), bands pink or red (rather than black or dark brown; Fig. 
<figureCitation id="9FE5CA8365253AAC1B4A49A3A741B44F" captionStart="Figure 15" captionStartId="F15" captionText="Figure 15. Photographs of some specimens of Bothriechis torvus in life. a. Adult and b. Juvenile (same individual) from Morromico Reserve, Choco department, Colombia; c. TH 145 adult female from Morromico Reserve, Choco department, Colombia; d. Adult and e. Juvenile from Uraba, Antioquia department, Colombia. Photos by Jose Vieira." httpUri="https://binary.pensoft.net/fig/981701" pageId="0" pageNumber="15">15</figureCitation>
), ventral surfaces entirely white, a higher number of ventrals in both males and females, smaller body size, and smaller basal hooks on the hemipenial body (Fig. 
<figureCitation id="6AB8F626B20A16EDE19D6BB77B9E3B4A" captionStart="Figure 16" captionStartId="F16" captionText="Figure 16. Hemipenial architecture of Bothriechis torvus in sulcate, lateral, and asulcate views. ICN 0421 from Valle del Cauca department, Colombia. Photos by Duvan Zambrano." httpUri="https://binary.pensoft.net/fig/981702" pageId="0" pageNumber="15">16</figureCitation>
). 
<taxonomicName id="C8ECC949E8BDFDB7ADC106B88DBBB7BD" baseAuthorityName="Posada Arango" baseAuthorityYear="1889" family="Viperidae" genus="Bothriechis" kingdom="Animalia" lsidName="Bothriechis torvus" order="Squamata" pageId="0" pageNumber="15" rank="species" species="torvus">
<emphasis id="6F9DB8FD968D5EB9A0325C397C69F52E" italics="true" pageId="0" pageNumber="15">Bothriechis torvus</emphasis>
</taxonomicName>
is sympatric with 
<taxonomicName id="B32E7DB36C5B91333706685B67FE52E7" lsidName="B. nigroadspersus" pageId="0" pageNumber="15" rank="species" species="nigroadspersus">
<emphasis id="51DA1910CB350A41A0859A050BD071B0" italics="true" pageId="0" pageNumber="15">B. nigroadspersus</emphasis>
</taxonomicName>
along the 
<normalizedToken id="3F3AB37F167F32F0B46DF2E910A9F32E" originalValue="Serranía">Serrania</normalizedToken>
de Pirre, 
<normalizedToken id="3EDF94A4A55C437EBC1E77D0685DD629" originalValue="Darién">Darien</normalizedToken>
province, Panama (lime green shade in Fig. 
<figureCitation id="C148AA3FF3D4B4DD3838C879B6A4D18D" 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>
). The two species differ in coloration (Figs 
<figureCitation id="AC46437517852F41F0010C718FECE937" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Photographs of some specimens of Bothriechis nigroadspersus in life. a. Adult female from a private collection in Costa Rica; b. MHCH 3269 adult from Guabito, Bocas del Toro province, Panama; c. Subadult from Parque Nacional Gandoca Manzanillo, Limon province, Costa Rica; d. Adult from a private collection in Costa Rica; e, f. Juveniles from Parque Nacional Gandoca Manzanillo, Limon province, Costa Rica; g. MHCH 3268 adult female from Porton, Chiriqui province, Panama; h. MHCH 3266 adult female from Chucanti Reserve, Darien province, Panama; i. Subadult from Parque Nacional Gandoca Manzanillo, Limon province, Costa Rica; j. Adult female from a private collection in Costa Rica; k. FP 001 from Cerro Gaital, Cocle province, Panama; l. MHCH 3267 juvenile male from Chucanti Reserve, Darien province, Panama. Photos by Jose Vieira." httpUri="https://binary.pensoft.net/fig/981691" pageId="0" pageNumber="15">5</figureCitation>
, 
<figureCitation id="33146549C5D79DF3E0FE42D73E322170" captionStart="Figure 15" captionStartId="F15" captionText="Figure 15. Photographs of some specimens of Bothriechis torvus in life. a. Adult and b. Juvenile (same individual) from Morromico Reserve, Choco department, Colombia; c. TH 145 adult female from Morromico Reserve, Choco department, Colombia; d. Adult and e. Juvenile from Uraba, Antioquia department, Colombia. Photos by Jose Vieira." httpUri="https://binary.pensoft.net/fig/981701" pageId="0" pageNumber="15">15</figureCitation>
). 
<taxonomicName id="594155667A3033526AADEA562239EFEF" baseAuthorityName="Posada Arango" baseAuthorityYear="1889" family="Viperidae" genus="Bothriechis" kingdom="Animalia" lsidName="Bothriechis torvus" order="Squamata" pageId="0" pageNumber="15" rank="species" species="torvus">
<emphasis id="75C730F5EDDCB150D48E5737E5C21ACE" italics="true" pageId="0" pageNumber="15">Bothriechis torvus</emphasis>
</taxonomicName>
has dorsal bands (vs opposing kidney shaped marks), three broad and triangular enlarged supraciliary scales (vs two thin and spinelike), a lower number of ventral scales, and lateral surface of the hemipenial body and lobes more heavily packed with larger spines (Fig. 
<figureCitation id="E8E3BFA82A7E758888BDB0058820F828" captionStart="Figure 16" captionStartId="F16" captionText="Figure 16. Hemipenial architecture of Bothriechis torvus in sulcate, lateral, and asulcate views. ICN 0421 from Valle del Cauca department, Colombia. Photos by Duvan Zambrano." httpUri="https://binary.pensoft.net/fig/981702" pageId="0" pageNumber="15">16</figureCitation>
). 
<taxonomicName id="EF639F028B4A703DFA07EFA62E714043" baseAuthorityName="Posada Arango" baseAuthorityYear="1889" family="Viperidae" genus="Bothriechis" kingdom="Animalia" lsidName="Bothriechis torvus" order="Squamata" pageId="0" pageNumber="15" rank="species" species="torvus">
<emphasis id="9AE62D1E50BC60EEABA391E1091320EF" italics="true" pageId="0" pageNumber="15">Bothriechis torvus</emphasis>
</taxonomicName>
resembles 
<taxonomicName id="CDE3B14DB8CBDA47CA7159ACB3E53ACB" lsidName="B. rahimi" pageId="0" pageNumber="15" rank="species" species="rahimi">
<emphasis id="F8A1201BBEF2CA74E0D01EDC3A546125" italics="true" pageId="0" pageNumber="15">B. rahimi</emphasis>
</taxonomicName>
sp. nov., but this other species occurs south of the known distribution of 
<taxonomicName id="60B66A2611CBF6720A19076ADD267EA2" lsidName="B. torvus" pageId="0" pageNumber="15" rank="species" species="torvus">
<emphasis id="CAA7E07DBE9B1603DC8B817F3A2C122B" italics="true" pageId="0" pageNumber="15">B. torvus</emphasis>
</taxonomicName>
and is characterized by having a lower number of ventrals, enlarged spinelike supraciliary scales, faint dorsal banding, and presence of the yellow morph. 
<taxonomicName id="1C4E8A8252CF1891911191EBDE50408B" baseAuthorityName="Posada Arango" baseAuthorityYear="1889" family="Viperidae" genus="Bothriechis" kingdom="Animalia" lsidName="Bothriechis torvus" order="Squamata" pageId="0" pageNumber="15" rank="species" species="torvus">
<emphasis id="AC7C80356A2BA803D16A02730350C42A" italics="true" pageId="0" pageNumber="15">Bothriechis torvus</emphasis>
</taxonomicName>
differs from 
<taxonomicName id="7BF2E41BF5D29DACF962E2A96D37A10A" lsidName="B. khwargi" pageId="0" pageNumber="15" rank="species" species="khwargi">
<emphasis id="C3724257FC0A050D3EE38FCC69678C65" italics="true" pageId="0" pageNumber="15">B. khwargi</emphasis>
</taxonomicName>
sp. nov. by having pink bands (bands absent or dark vertebral blotches in 
<taxonomicName id="5F8691093AD4BE2AD6E60ACC811B3730" lsidName="B. khwargi" pageId="0" pageNumber="15" rank="species" species="khwargi">
<emphasis id="6E2E2176DEEC70D6D6F30F029C8168C1" italics="true" pageId="0" pageNumber="15">B. khwargi</emphasis>
</taxonomicName>
sp. nov.).
</paragraph>
<caption id="7247005AE16FBCFA882E3F61D359C867" httpUri="https://binary.pensoft.net/fig/981701" pageId="0" pageNumber="15" start="Figure 15" startId="F15">
<paragraph id="DD258AFF4F214A19DCB833EF4D823052" pageId="0" pageNumber="15">
<emphasis id="7464E0652D9DB5D7BD0FAAA302D79DF7" bold="true" pageId="0" pageNumber="15">Figure 15.</emphasis>
Photographs of some specimens of 
<taxonomicName id="1425CF940CE6A37F7C1C5CC2F39E90E1" baseAuthorityName="Posada Arango" baseAuthorityYear="1889" family="Viperidae" genus="Bothriechis" kingdom="Animalia" lsidName="Bothriechis torvus" order="Squamata" pageId="0" pageNumber="15" rank="species" species="torvus">
<emphasis id="761943D0F842FF3252B3EEB61E20C9B7" italics="true" pageId="0" pageNumber="15">Bothriechis torvus</emphasis>
</taxonomicName>
in life. 
<emphasis id="8C2B3F51EED238B700FB70B23E9BEF09" bold="true" pageId="0" pageNumber="15">a.</emphasis>
Adult and 
<emphasis id="D4D3C8629FB8998E9A70F2FEB2BF9978" bold="true" pageId="0" pageNumber="15">b.</emphasis>
Juvenile (same individual) from Morromico Reserve, 
<normalizedToken id="F07680445F463E8805E9C037B4DBF12B" originalValue="Chocó">Choco</normalizedToken>
department, Colombia; 
<emphasis id="72A4A9DF003B3BEDEA2E41E82E0F631F" bold="true" pageId="0" pageNumber="15">c.</emphasis>
TH 145 adult female from Morromico Reserve, 
<normalizedToken id="9A46549887495291402623E17B02FB1A" originalValue="Chocó">Choco</normalizedToken>
department, Colombia; 
<emphasis id="49CC820B1EB0D446370575637EC035C8" bold="true" pageId="0" pageNumber="15">d.</emphasis>
Adult and 
<emphasis id="0848C0CCAAA96A284530696493AC795D" bold="true" pageId="0" pageNumber="15">e.</emphasis>
Juvenile from 
<normalizedToken id="4A42648C2368D55B8224D4B7C24CD77B" originalValue="Urabá">Uraba</normalizedToken>
, Antioquia department, Colombia. Photos by Jose Vieira.
</paragraph>
</caption>
<caption id="CA5A8273A9BC3510038711778812CB41" httpUri="https://binary.pensoft.net/fig/981702" pageId="0" pageNumber="15" start="Figure 16" startId="F16">
<paragraph id="F22A3182AAA8E58CA013D173630251EC" pageId="0" pageNumber="15">
<emphasis id="56900E62EE8F34434DAE8FE970F521DD" bold="true" pageId="0" pageNumber="15">Figure 16.</emphasis>
Hemipenial architecture of 
<taxonomicName id="8D053949D1A82D6FBB0E2F393923DF68" baseAuthorityName="Posada Arango" baseAuthorityYear="1889" family="Viperidae" genus="Bothriechis" kingdom="Animalia" lsidName="Bothriechis torvus" order="Squamata" pageId="0" pageNumber="15" rank="species" species="torvus">
<emphasis id="539866212DC4746BDC46C2F598FCC855" italics="true" pageId="0" pageNumber="15">Bothriechis torvus</emphasis>
</taxonomicName>
in sulcate, lateral, and asulcate views. ICN 0421 from Valle del Cauca department, Colombia. Photos by 
<normalizedToken id="404778F8178F2BDF974B79FC7F457898" originalValue="Duván">Duvan</normalizedToken>
Zambrano.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="C867CEBF532ADE0A673E4C566BD479D9" pageId="0" pageNumber="15" type="hemipenial morphology">
<paragraph id="B999C4ABAFE390C51ABC8FA88FEFB5F6" pageId="0" pageNumber="15">Hemipenial morphology.</paragraph>
<paragraph id="8DE0E9A177F04657140790E42AEC5D35" pageId="0" pageNumber="15">
(n = 1; Fig. 
<figureCitation id="91A044E84950D6D783E8F74375736166" captionStart="Figure 16" captionStartId="F16" captionText="Figure 16. Hemipenial architecture of Bothriechis torvus in sulcate, lateral, and asulcate views. ICN 0421 from Valle del Cauca department, Colombia. Photos by Duvan Zambrano." httpUri="https://binary.pensoft.net/fig/981702" pageId="0" pageNumber="15">16</figureCitation>
) Everted and inflated, the organ is deeply bilobed, calyculate and noncapitate; hemipenial lobes cylindrical, but tapering towards the capitulum; in sulcate and asulcate views, lobe crotch ornamented with densely packed spines that become larger distally; sulcus spermaticus centrolineal, bifurcate and with walls strongly defined, 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 defined spinulate edges. In sulcate view, hemipenial body surface spinulate medially with enlarged and strongly calcified lateral basal hooks followed by rows of smaller obliquely arranged mesial spines that become gradually smaller towards each 
<normalizedToken id="A3A0D114F6ED1ACDF77E37BD9D7A1334" originalValue="lobe’s">lobe's</normalizedToken>
capitulum; each hemipenial lobe ornamented with mesial and lateral spines about 1/5 the size of the large basal hooks; the spines in each lobe are replaced distally by calyces with strongly spinulate edges. In lateral view, hemipenial body and lobes non-spiculate but with rows of spines; distal half of lobes strongly calyculate. In asulcate view, the center of hemipenial body is nude to barely spiculate; hemipenial lobes ornamented with 2 rows of smaller spines (about 1/3 the size of the lateral basal hooks) that decrease in size towards the lobe crotch.
</paragraph>
</subSubSection>
<subSubSection id="3A69EBBB45FEB1E1F58BB0FE614D495E" pageId="0" pageNumber="15" type="natural history">
<paragraph id="0DFCE8915F0AF29CA46B375152B85707" pageId="0" pageNumber="15">Natural history.</paragraph>
<paragraph id="B36779AA9AED091940EAD4DEF1C0E704" pageId="0" pageNumber="15">
<taxonomicName id="C7FA64EBE98DFAC26873D5A29D805C30" baseAuthorityName="Posada Arango" baseAuthorityYear="1889" family="Viperidae" genus="Bothriechis" kingdom="Animalia" lsidName="Bothriechis torvus" order="Squamata" pageId="0" pageNumber="15" rank="species" species="torvus">
<emphasis id="C7F76A7EFBAA3745577A11BA2F3DC0A2" italics="true" pageId="0" pageNumber="15">Bothriechis torvus</emphasis>
</taxonomicName>
is an arboreal snake that inhabits evergreen lowland/foothill forests. We have found vipers of this species active at night crawling on ferns, tree branches, and trunks from 1.8 to 5 m above the ground or during the day perched on arboreal vegetation up to 3.5 m above the ground. Specimens from Cerro Sapo and Pirre Mountain ranges, eastern Panama, were found in a cloud forest consisting predominantly of trees covered with moss and a large variety of understory and midstory bromeliads. In captivity, TH 145 (Fig. 
<figureCitation id="FE625044C8127CAC218C8A485A8863FC" captionStart="Figure 15" captionStartId="F15" captionText="Figure 15. Photographs of some specimens of Bothriechis torvus in life. a. Adult and b. Juvenile (same individual) from Morromico Reserve, Choco department, Colombia; c. TH 145 adult female from Morromico Reserve, Choco department, Colombia; d. Adult and e. Juvenile from Uraba, Antioquia department, Colombia. Photos by Jose Vieira." httpUri="https://binary.pensoft.net/fig/981701" pageId="0" pageNumber="15">15c</figureCitation>
) and another specimen from Morromico, 
<normalizedToken id="CE56A7912F02B5859D696395EF640E74" originalValue="Chocó">Choco</normalizedToken>
department, Colombia, consumed lizards (
<taxonomicName id="C5EE88753277F60CE37E95A281CD93C5" class="Reptilia" family="Iguanidae" genus="Anolis" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Anolis" order="Squamata" pageId="0" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="74629300B2685C262F7019EF2BBD7AC0" italics="true" pageId="0" pageNumber="15">Anolis</emphasis>
</taxonomicName>
and 
<taxonomicName id="0B158EA0706F47D0DD95FF4517B5B6EB" class="Squamata" family="Gekkonidae" genus="Lepidodactylus" kingdom="Animalia" lsidName="Lepidodactylus" pageId="0" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="0B2F78C407267D8E819CC827E9325C5B" italics="true" pageId="0" pageNumber="15">Lepidodactylus</emphasis>
</taxonomicName>
) and frogs (
<taxonomicName id="5C30D897F789FF03362F5C995E0EB14E" authorityName="Cope" authorityYear="1862" class="Amphibia" family="Brachycephalidae" genus="Craugastor" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Craugastor" order="Anura" pageId="0" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="A85067F9D279D2E54A4F7098F55A34D6" italics="true" pageId="0" pageNumber="15">Craugastor</emphasis>
</taxonomicName>
and 
<taxonomicName id="0210BCED16D67381303AB05A9DD38CAE" class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Pristimantis" order="Anura" pageId="0" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="09488DC7C149121670989F74D734A8E4" italics="true" pageId="0" pageNumber="15">Pristimantis</emphasis>
</taxonomicName>
).
</paragraph>
</subSubSection>
<subSubSection id="6098FE81278519835CE1065ED7D41F5C" pageId="0" pageNumber="15" type="venom">
<paragraph id="5710B916B17AB1090E168D439B03294E" pageId="0" pageNumber="15">Venom.</paragraph>
<paragraph id="D219F2F5C7D15FBA5A987B0176E51656" pageId="0" pageNumber="15">
<bibRefCitation id="2AC94F064CD9086789ED4A34D517A3CA" DOI="https://doi.org/10.1177/1535370216660214" author="Prezotto-Neto, JP" journalOrPublisher="Experimental Biology and Medicine" pageId="0" pageNumber="15" pagination="2075 - 2085" refId="B86" refString="Prezotto-Neto, JP, Kimura, LF, Alves, AF, Gutierrez, JM, Otero, R, Suarez, AM, Santoro, ML, Barbaro, KC, 2016. Biochemical and biological characterization of Bothriechis schlegelii snake venoms from Colombia and Costa Rica. Experimental Biology and Medicine 241: 2075 - 2085, DOI: https://doi.org/10.1177/1535370216660214" title="Biochemical and biological characterization of Bothriechis schlegelii snake venoms from Colombia and Costa Rica." url="https://doi.org/10.1177/1535370216660214" volume="241" year="2016">Prezotto-Neto et al. (2016)</bibRefCitation>
studied the composition of the venom of specimens of 
<taxonomicName id="D2EB8B3365D9DBAA33A24440B085AAB5" baseAuthorityName="Posada Arango" baseAuthorityYear="1889" family="Viperidae" genus="Bothriechis" kingdom="Animalia" lsidName="Bothriechis torvus" order="Squamata" pageId="0" pageNumber="15" rank="species" species="torvus">
<emphasis id="9A8311C1D18F6392249235E7B620AD13" italics="true" pageId="0" pageNumber="15">Bothriechis torvus</emphasis>
</taxonomicName>
from Vegachi, Antioquia, and found that its properties differ drastically from specimens of 
<taxonomicName id="AA441296CEC8E72B35DAC0F16F232782" lsidName="B. nigroadspersus" pageId="0" pageNumber="15" rank="species" species="nigroadspersus">
<emphasis id="7A73E1C1FDDEB642DA4232763FD962DB" italics="true" pageId="0" pageNumber="15">B. nigroadspersus</emphasis>
</taxonomicName>
from Costa Rica. When compared to the latter, the venom of 
<taxonomicName id="C472CC7DF9C1CF92C8601CE9B65A6FD7" lsidName="B. torvus" pageId="0" pageNumber="15" rank="species" species="torvus">
<emphasis id="009A08750EF926D22EB8B9CB1F9D6705" italics="true" pageId="0" pageNumber="15">B. torvus</emphasis>
</taxonomicName>
was found to be less edematous, hemorrhagic, and lethal (LD50 estimated as 9.24 mg/kg vs 5.60 mg/kg).
</paragraph>
</subSubSection>
<subSubSection id="45E5F3654421BBC54398DCB6AC9E0009" pageId="0" pageNumber="15" type="distribution">
<paragraph id="02407B4531A8A69E737A30BC812B5D19" pageId="0" pageNumber="15">Distribution.</paragraph>
<paragraph id="6902378D5A1E9EE96C04C5FD797A9AF2" pageId="0" pageNumber="15">
<taxonomicName id="5E3885C21A6CED942FD068CF0ACE35D3" baseAuthorityName="Posada Arango" baseAuthorityYear="1889" family="Viperidae" genus="Bothriechis" kingdom="Animalia" lsidName="Bothriechis torvus" order="Squamata" pageId="0" pageNumber="15" rank="species" species="torvus">
<emphasis id="BCD8BB864540E07E23678C8FBB70E738" italics="true" pageId="0" pageNumber="15">Bothriechis torvus</emphasis>
</taxonomicName>
is known from at least 37 localities (listed in Suppl. material 3) along the 
<normalizedToken id="870A52E887EE34B39D8D8CE5E6FB21AD" originalValue="Chocó">Choco</normalizedToken>
and 
<normalizedToken id="186190C015E338F81B384C08DDE2C12A" originalValue="Río">Rio</normalizedToken>
Magdalena valley regions of northern Colombia and extreme southeastern Panama. The species occurs over an area of approximately 61,466 km2 and has been recorded at elevations 61-1,413 m above sea level (Fig. 
<figureCitation id="621F099DC13DF84A7EF0E227D914FEA3" 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>
). The ENM suggests the distribution is not continuous across the valley of the 
<normalizedToken id="533EED7B9FB96144FAC28136E05E523B" originalValue="Río">Rio</normalizedToken>
Atrato (see Discussion).
</paragraph>
</subSubSection>
<subSubSection id="83B8CF839ECDC4565CF3B7304C17E00B" pageId="0" pageNumber="15" type="etymology">
<paragraph id="A72613A55FCB18CEEC7B8C4272F44A81" pageId="0" pageNumber="15">Etymology.</paragraph>
<paragraph id="747597016A4A32E574F3CBB26045E7AE" pageId="0" pageNumber="15">
The specific epithet 
<emphasis id="740B66547911A1FFAA527C5DE926C604" italics="true" pageId="0" pageNumber="15">torvus</emphasis>
is a Latin word meaning 
<normalizedToken id="2F5082552F04A7D57ABC491A628B9E0C" originalValue="“fierce.”">&quot;fierce.&quot;</normalizedToken>
It refers to the protruding supraciliary scales, which according to 
<normalizedToken id="EBB27597F637ABFDAFE7F2480387A215" originalValue="Andrés">Andres</normalizedToken>
Posada Arango, give this species an &quot; 
<emphasis id="0B07597F312C12663D9D6030128CA161" italics="true" pageId="0" pageNumber="15">
air 
<normalizedToken id="A9A9CA26F09EE67D6A897C89759AA776" originalValue="féroce.”">feroce.&quot;</normalizedToken>
</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="A89C4ABF2FC48056B2D138041692B19A" pageId="0" pageNumber="15" type="conservation status">
<paragraph id="7BD1140988923A4389D486C4ED1BAA6F" pageId="0" pageNumber="15">Conservation status.</paragraph>
<paragraph id="B781122BA5894C901A5BB3197F92BFB8" pageId="0" pageNumber="15">
We consider 
<taxonomicName id="6070B17277C5F8223CDBCAB244FB4D98" baseAuthorityName="Posada Arango" baseAuthorityYear="1889" family="Viperidae" genus="Bothriechis" kingdom="Animalia" lsidName="Bothriechis torvus" order="Squamata" pageId="0" pageNumber="15" rank="species" species="torvus">
<emphasis id="A088868438784C14DCD7541B33984FD2" italics="true" pageId="0" pageNumber="15">Bothriechis torvus</emphasis>
</taxonomicName>
to be included in the Least Concern category following IUCN Red List criteria (
<bibRefCitation id="4EA1897447B57C48C674D8EEDC3149F8" 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>
) because the species is distributed over a region that holds large areas of continuous unspoiled forest. Based on the species distribution model presented in Fig. 
<figureCitation id="A240B1FFED93D551299004A1A980F246" 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="15CD522D947BC5FCBC664A2504622B9E" 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>
) and Panama (
<bibRefCitation id="54B07AFE1B8E411D0F8A5831091300EB" author="Castoe, TA" journalOrPublisher="Journal of Biogeography" pageId="0" pageNumber="15" refId="B23" refString="CATHALAC (2011) Mapa Centroamericano de cobertura y uso de la tierra, cambios de cobertura y uso de la tierra 1980-1990-2000-2010.Programa Regional para la Reduccion de la Vulnerabilidad y Degradacion Ambiental, Ciudad de Panama, 166 pp." year="2009">CATHALAC 2011</bibRefCitation>
), we estimate that more than half (~59%) of the 
<normalizedToken id="D1542AFBA3587AEBFDDC486C78041EEA" originalValue="species’">species'</normalizedToken>
forest habitat is still standing. Unfortunately, vast areas of the 
<normalizedToken id="FEEE07C7A45BCA4F7F445D943E60C0ED" originalValue="Chocó">Choco</normalizedToken>
rainforest in northern Colombia and towards central Panama have already been converted to pastures (
<bibRefCitation id="CCFD705E3412D631C879ED10B60D3952" DOI="https://doi.org/10.1038/35002501" author="Myers, N" journalOrPublisher="Nature" pageId="0" pageNumber="15" pagination="835 - 838" refId="B69" refString="Myers, N, Mittermeier, RA, Mittermeier, CG, da Fonseca, GAB, Kent, J, 2000. Biodiversity hotspots for conservation priorities. Nature 403: 835 - 838, DOI: https://doi.org/10.1038/35002501" title="Biodiversity hotspots for conservation priorities." url="https://doi.org/10.1038/35002501" volume="403" year="2000">Myers et al. 2000</bibRefCitation>
). However, 
<taxonomicName id="7BD39D6CDC3E125E3AB7073179C9FEDB" lsidName="B. torvus" pageId="0" pageNumber="15" rank="species" species="torvus">
<emphasis id="552BDAE2AB481D3D1B539A0634A3C2EB" italics="true" pageId="0" pageNumber="15">B. torvus</emphasis>
</taxonomicName>
occurs over an area greater than 50,000 km2 and is presumably not declining fast enough to qualify for a threatened category.
</paragraph>
</subSubSection>
</treatment>
</document>