Research Article |
Corresponding author: Alejandro Arteaga ( af.arteaga.navarro@gmail.com ) Academic editor: Robert Jadin
© 2018 Alejandro Arteaga, David Salazar-Valenzuela, Konrad Mebert, Nicolás Peñafiel, Gabriela Aguiar, Juan C. Sánchez-Nivicela, R. Alexander Pyron, Timothy J. Colston, Diego F. Cisneros-Heredia, Mario H. Yánez-Muñoz, Pablo J. Venegas, Juan M. Guayasamin, Omar Torres-Carvajal.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Arteaga A, Salazar-Valenzuela D, Mebert K, Peñafiel N, Aguiar G, Sánchez-Nivicela JC, Pyron RA, Colston TJ, Cisneros-Heredia DF, Yánez-Muñoz MH, Venegas PJ, Guayasamin JM, Torres-Carvajal O (2018) Systematics of South American snail-eating snakes (Serpentes, Dipsadini), with the description of five new species from Ecuador and Peru. ZooKeys 766: 79-147. https://doi.org/10.3897/zookeys.766.24523
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A molecular phylogeny of the Neotropical snail-eating snakes (tribe Dipsadini) is presented including 43 (24 for the first time) of the 77 species, sampled for both nuclear and mitochondrial genes. Morphological and phylogenetic support was found for four new species of Dipsas and one of Sibon, which are described here based on their unique combination of molecular, meristic, and color pattern characteristics. Sibynomorphus is designated as a junior subjective synonym of Dipsas. Dipsas latifrontalis and D. palmeri are resurrected from the synonymy of D. peruana. Dipsas latifasciata is transferred from the synonymy of D. peruana to the synonymy of D. palmeri. A new name, D. jamespetersi, is erected for the taxon currently known as Sibynomorphus petersi. Re-descriptions of D. latifrontalis and D. peruana are presented, as well as the first photographic voucher of an adult specimen of D. latifrontalis, along with photographs of all known Ecuadorian Dipsadini species. The first country record of D. variegata in Ecuador is provided and D. oligozonata removed from the list of Peruvian herpetofauna. With these changes, the number of Dipsadini reported in Ecuador increases to 22, 18 species of Dipsas and four of Sibon.
Dipsadini , Dipsas , Ecuador, new species, Peru, phylogeny, Sibon , Sibynomorphus , snail-eating snakes, systematics
With 70 currently recognized species (Table
One of the first modern attempts to clarify the taxonomy and summarize knowledge on the tribe Dipsadini was published by
After Peters, several authors continued to address the systematics of the group (
Here, we combine morphological analysis and molecular phylogenetics to revise generic and species limits within Dipsadini. We combine all available molecular sampling with new samples from Ecuador, Peru, Brazil and Costa Rica, and find support for five new species, as well as a number of changes to the geographic distribution of several Andean species.
Genus | Group | Species | Authority | Reference |
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Dipsas | D. articulata | D. articulata | Cope, 1868 |
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D. bicolor | Günther, 1895 |
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D. brevifacies | Cope, 1866 |
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D. gaigeae | Oliver, 1837 |
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D. gracilis | Boulenger, 1902 |
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D. maxillaris | Werner, 1910 |
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D. tenuissima | Taylor, 1954 |
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D. viguieri | Bocourt, 1884 |
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D. catesbyi | D. catesbyi | Sentzen, 1796 |
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D. copei | Günther, 1872 |
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D. pavonina | Schlegel, 1837 |
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D. elegans | D. elegans | Boulenger, 1896 |
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D. ellipsifera | Boulenger, 1898 |
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D. oreas | Cope, 1868 |
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D. incerta | D. alternans | Fischer, 1885 |
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D. incerta | Jan, 1863 |
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D. praeornata | Werner, 1909 |
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D. sazimai | Fernandes et al., 2010 |
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D. indica | D. bucephala | Shaw, 1802 |
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D. cisticeps | Boettger, 1885 |
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D. indica | Laurenti, 1768 |
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D. pratti | D. baliomelas | Harvey, 2008 |
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D. chaparensis | Reynolds & Foster, 1992 |
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D. peruana | Boettger, 1898 |
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D. pratti | Boulenger, 1897 |
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D. sanctijoannis | Boulenger, 1911 |
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D. schunkii | Boulenger, 1908 |
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D. temporalis | D. pakaraima | MacCulloch & Lathrop, 2004 |
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D. temporalis | Werner, 1909 |
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D. vermiculata | Peters, 1960 |
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D. variegata | D. albifrons | Sauvage, 1884 |
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D. andiana | Boulenger, 1896 |
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D. nicholsi | Dunn, 1933 |
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D. trinitatis | Parker, 1926 |
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D. variegata | Duméril et al., 1854 |
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Plesiodipsas | Unassigned | P. perijanensis | Aleman, 1953 | – |
Sibon | S. annulatus | S. annulatus | Günther, 1872 |
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S. anthracops | Cope, 1868 |
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S. dimidiatus | Günther, 1872 |
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S. lamari | Solórzano, 2001 |
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S. linearis | Pérez-Higareda et al., 2002 |
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S. manzanaresi | McCranie, 2007 |
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S. merendonensis | Rovito et al., 2012 |
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S. miskitus | McCranie, 2006 |
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S. sanniolus | Cope, 1866 |
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Sibon | S. argus | S. argus | Cope, 1875 |
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S. longifrenis | Stejneger, 1909 |
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S. nebulatus | S. carri | Shreve, 1951 |
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S. dunni | Peters, 1957 |
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S. nebulatus | Linnaeus, 1758 |
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Unassigned | S. noalamina | Lotzkat et al., 2012 | – | |
S. perissostichon | Köhler et al., 2010 | – | ||
Sibynomorphus | Unassigned | S. lavillai | Scrocchi et al., 1993 | – |
S. mikanii | Schlegel, 1837 | – | ||
S. neuwiedi | Ihering, 1911 | – | ||
S. oligozonatus | Orcés & Almendáriz, 1989 | – | ||
S. oneilli | Rossman & Thomas, 1979 | – | ||
S. petersi | Orcés & Almendáriz, 1989 | – | ||
S. turgidus | Cope, 1868 | – | ||
S. vagrans | Dunn, 1923 | – | ||
S. vagus | Jan, 1863 | – | ||
S. ventrimaculatus | Boulenger, 1885 | – | ||
S. williamsi | Carillo de Espinoza, 1974 | – | ||
Tropidodipsas | T. fasciata | T. fasciata | Günther, 1858 |
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T. philippii | Jan, 1863 |
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T. sartorii | T. annulifera | Boulenger, 1894 |
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T. sartorii | Cope, 1863 |
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T. zweifeli | Liner & Wilson, 1970 |
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Unassigned | T. fischeri | Boulenger, 1894 | – | |
T. repleta | Smith et al., 2005 | – |
This study was carried out in strict accordance with the guidelines for use of live amphibians and reptiles in field research (
Criteria for common name designation are as proposed by
Tissue samples from 85 individuals representing 28 species (including five new species described here) were sampled from Ecuador, Peru, Guatemala, Costa Rica, Nicaragua, Brazil, and Mexico. All specimens included in the genetic analyses were morphologically identified according to
Photographs of some species of Dipsas in life: a D. andianaMZUTI 5413 from Bilsa, province of Esmeraldas, Ecuador b D. andiana from Mindo, province of Pichincha, Ecuador c D. bobridgelyiMZUTI 5414 from Buenaventura, Province of El Oro, Ecuador d D. catesbyi from Gareno, province of Napo, Ecuador e D. catesbyi from Gareno, province of Napo, Ecuador f D. elegans from Calacalí–Mindo, province of Pichincha, Ecuador g D. ellipsifera from Pimampiro, province of Imbabura, Ecuador h D. gracilis from Canandé, province of Esmeraldas, Ecuador i D. gracilis from Mashpi, province of Pichincha, Ecuador j D. indica from Gareno, province of Napo, Ecuador k D. jamespetersiAMARU 1123 from province of Azuay, Ecuador l D. klebbai from El Chaco, province of Napo, Ecuador m D. klebbai from El Chaco, province of Napo, Ecuador n D. latifrontalis from San Isidro, state of Mérida, Venezuela o D. oligozonata from Poetate, province of Azuay, Ecuador p D. oreasMZUTI 5414 from Buenaventura, province of El Oro, Ecuador q D. oreas from Poetate–Corraleja, province of Azuay, Ecuador r D. palmeri from Agoyán, province of Tungurahua, Ecuador s D. palmeriMZUTI 4975 from Reserva San Francisco, province of Zamora, Ecuador t D. pavonina from Maycu, province of Zamora, Ecuador u D. temporalis from Colombia v D. variegata from Gareno, province of Napo, Ecuador w D. vermiculata from Miazi, province of Zamora, Ecuador, and x D. vermiculata from Narupa, province of Napo, Ecuador.
Photographs of some species of Sibon in life: a S. annulatus from Verdecanandé, province of Esmeraldas, Ecuador b Sibon bevridgelyi
Genomic DNA was extracted from 96% ethanol-preserved tissue samples (liver, muscle tissue or scales) using either a guanidinium isothiocyanate extraction protocol, or a modified salt precipitation method based on the Puregene DNA purification kit (Gentra Systems). We amplified the 16S gene using primer pairs 16Sar-L / 16Sbr-H-R from
A total of 298 DNA sequences were used to build a phylogenetic tree of the tribe Dipsadini, of which 222 were generated during this work and 76 were downloaded from GenBank. Among the new sequences, 103 are 201–520 bp long fragments of the 16S gene, 91 are 586–1,090 bp long fragments of the Cytb gene, 45 are 443–583 bp long fragments of the c-mos gene, 31 are 242–473 bp long fragments of the 12S gene, and 28 are 593–699 bp long fragments of the ND4 gene. New sequences were edited and assembled using the program Geneious ProTM 5.4.7 (
Phylogenetic relationships were assessed under both a Bayesian inference (BI) and a maximum likelihood (ML) approach in MrBayes 3.2.0 (
Terminology for Dipsadini cephalic shields follows proposals by
We consider strong support to be bootstrap values of >70% and posterior probability values >95% following
Phylogenetic relationships within Dipsadini derived from analysis of 3,375 bp of DNA (gene fragments 12S, 16S, Cytb, ND4 and c-mos). Support values on intraspecic branches are not shown for clarity. Voucher numbers for sequences are indicated for each terminal when available. a Maximum likelihood analysis. Black dots indicate clades with bootstrap values from 90–100%. Grey dots indicate values from 70–89%. White dots indicate values from 50–69% (values <50% not shown) b Bayesian inference analysis. Black dots indicate clades with posterior probability values from 95–100%. Grey dots indicate values from 70–94%. White dots indicate values from 50–69% (values <50% not shown).
Sibon longifrenis is recovered as the sister taxon to all other included species of Sibon. Deep intraspecific divergence is found between samples of S. annulatus from Central America (
Eight Sibynomorphus species were included in the molecular analyses. These are S. mikanii, S. neuwiedi, S. oligozonatus, S. petersi, S. turgidus, S. vagus, S. ventrimaculatus, and S. williamsi. In the ML analysis, all of them are nested within different Dipsas subclades, whereas in the BI analysis, the clade containing S. mikanii and S. turgidus is not nested within Dipsas. Crucially, Dipsas mikanii Schlegel, 1837 is the type species of Sibynomorphus (Fitzinger, 1843). Thus, we synonymize Sibynomorphus with Dipsas primarily based on the ML analysis, which mirrors the results of
Based on our transfer of the genus Sibynomorphus Fitzinger to the synonymy of Dipsas, we propose the following binomial nomenclature for the eleven species traditionally included in the genus Sibynomorphus: Dipsas lavillai comb. n., D. mikanii, D. neuwiedi comb. n., D. oligozonata comb. nov., D. oneilli comb. n., D. turgida comb. nov., D. vagrans comb. n., D. vaga comb. n., D. ventrimaculata comb. n., and D. williamsi comb. n. However, we refrain from applying D. “petersi” for Sibynomorphus petersi here, because the name Dipsas “indica” petersi (Hoge & Romano, 1975), another taxon and putative species from southeastern Brazil, is often already named as Dipsas petersi (e.g.,
There are several clades within Dipsas peruana sensu lato. One is D. peruana, the other is a new species from northern Ecuador, which we describe below, and the third is the lineage corresponding to the population distributed along the Amazonian slopes of the Andes between central Ecuador and northern Peru. Below, we resurrect the name D. palmeri (Boulenger, 1912) for this lineage, as the type locality of D. palmeri (El Topo, province of Tungurahua, Ecuador) is located within the geographic range of the included samples (Fig.
Dipsas oligozonata is the strongly supported sister lineage of a clade that includes three species: D. williamsi and two new species from western Ecuador and northern Peru, which we describe below. Dipsas indica is paraphyletic with respect to D. bucephala. Dipsas jamespetersi is paraphyletic with respect to a sample of D. vaga (
Based on the species included in the phylogenetic analysis, the Dipsas articulata and D. indica groups, sensu
One individual (Fig.
We seek here to name or provide re-descriptions only for species that are monophyletic in our molecular phylogeny and share diagnostic features of their coloration pattern and lepidosis. Based on these species delimitation criteria, which follow the general species concept of
Bev Ridgely’s Snail-Eater
Caracolera de Bev Ridgely
MZUTI 5416 (Figs
AMNH 22092, adult male collected by George H. Tate on December 01, 1921 at Bucay, province of Guayas, Ecuador (S2.19788, W79.12909; 433 m).
Sibon bevridgelyi is placed in the genus Sibon based on phylogenetic evidence (Fig.
Sibon bevridgelyi is most similar to S. nebulatus, from which it differs on the basis of its distinctive coloration (Figs
Adult male, SVL 602 mm, tail length 186 mm (31% SVL); head length 20.9 mm (3% SVL) from tip of snout to commissure of mouth; head width 12.4 mm (59% head length) taken at broadest point; snout-orbit distance 21 mm; head distinct from neck; snout short, blunt in dorsal and lateral outline; rostral 3.5 mm wide, broader than high; internasals 1.9 mm wide, broader than long; prefrontals 4.4 mm wide, longer than broad, entering orbit; supraocular 4.4 mm long, longer than broad; frontal 4.1 mm long, pentagonal and rounded, in contact with prefrontals, supraoculars, and parietals; parietals 7.7 mm long, longer than broad; nasal weakly divided, in contact with first three supralabials, loreal, prefrontal, internasal, and rostral; loreal 3.7 mm long, longer than high, entering the orbit; eye diameter 3.9 mm; pupil semi-elliptical; no preocular; two postoculars; temporals 1+3 on the right side, 2+3 on the left side; eight supralabials with 5th and 6th contacting orbit on the right side, seven supralabials with 4th and 5th contacting orbit on the left side; symphysial separated from chinshields by the first pair of infralabials; nine infralabials, 1–7 contacting chinshields; anterior pair of chinshields broader than long, posterior pair longer than broad; dorsal scales in 15/15/15 rows, smooth, without apical pits; 184 ventrals; 80 divided subcaudals; cloacal plate single.
Specimens of Sibon bevridgelyi have been found active at night (20h56–03h56) on arboreal vegetation 30–500 cm above the ground in secondary and primary semideciduous foothill forest, pastures, and cacao plantations, usually close to streams.
Northwestern Peru in the department of Piura, and southwestern Ecuador in the provinces of Azuay, Chimborazo, El Oro, Guayas, Los Ríos and Manabí at elevations between 5 and 1206 m (Fig.
The specific epithet honors the late Prof. Beverly S. Ridgely, life-long birder and conservationist, and father of Robert S. Ridgely, well known in Ecuadorian ornithological circles and co-author of The Birds of Ecuador. Though he never got to visit Buenaventura, from afar Bev continued to delight in the conservation successes of Fundación Jocotoco, which now owns and manages one of the few protected areas where the Vulnerable Sibon bevridgelyi is known to occur.
We consider Sibon bevridgelyi to be Vulnerable following B2a,b(i,iii) IUCN criteria (IUCN 2001) because its area of occupancy is estimated to be less than 2,000 km2, it is known only from 15 patches of forest lacking connectivity between them, and its habitat is severely fragmented and declining in extent and quality due to deforestation. Furthermore, only three of the localities (Parque Nacional Machalilla, Reserva Buenaventura, and Reserva Ayampe) where S. bevridgelyi occurs are currently protected.
Bob Ridgely’s Snail-Eater
Caracolera de Bob Ridgely
MZUTI 5417 (Figs
DHMECN 11527, adult female collected by Juan Carlos Sánchez-Nivicela, Karem López, Verónica Urgilés, Bruno Timbe, Elvis Celi and Valentina Posse at Remolino, province of El Oro, Ecuador (S3.56551, W79.91948; 229 m). MZUTI 3266, adult female collected by Lucas Bustamante on October 06, 2013. MZUTI 5414, adult male collected by Matthijs Hollanders and Paulina Romero on June 08, 2017.
Dipsas bobridgelyi is placed in the genus Dipsas based on phylogenetic evidence (Fig.
Dipsas bobridgelyi is most similar to D. gracilis, from which it differs in coloration. In D. gracilis (Figs
Adult male, SVL 372 mm, tail length 158 mm (43% SVL); head length 15.1 mm (4% SVL) from tip of snout to commissure of mouth; head width 8.1 mm (54% head length) taken at broadest point; snout-orbit distance 4.3 mm; head distinct from neck; snout short, blunt in dorsal and lateral outline; rostral 2.4 mm wide, broader than high; internasals 2.3 mm wide, broader than long; prefrontals 2.5 mm wide, longer than broad and contacting orbit; supraocular 3.2 mm long, longer than broad; frontal 3.9 mm long, hexagonal, in contact with prefrontals, supraoculars, and parietals; parietals 4.7 mm long, longer than broad; nasal divided, in contact with first three supralabials, loreal, prefrontal, internasal, and rostral; loreal 1.8 mm long, slightly higher than long, entering the orbit; eye diameter 2.7 mm; pupil semi-elliptical; no preocular; two postoculars; temporals 2+3; nine supralabials, 4th and 5th contacting orbit; symphysial separated from chinshields by the first pair of infralabials; 13 infralabials, 1–7 contacting chinshields; anterior pair of chinshields longer than broad, posterior pair broader than long; dorsal scales in 15/15/15 rows, smooth, without apical pits; 182 ventrals; 101 divided subcaudals; cloacal plate single.
Individuals of Dipsas bobridgelyi have been found active at night (19h00–23h26) on arboreal vegetation 100–250 cm above the ground in secondary semi-deciduous foothill forest. MZUTI 5414 was found feeding on a snail.
Foothills of the southwestern Ecuadorian Andes in the provinces of Azuay and El Oro, and northwestern Peruvian Andes in the department of Tumbes, at elevations between 39 and 572 m (Fig.
This species is named in honor of Dr. Robert “Bob” S. Ridgely, a leading ornithologist and distinguished conservationist who has dedicated almost 50 years of his life to the study and conservation of birds and biodiversity across Latin America. Bob is the President of Rainforest Trust and for the past twenty years has been a major driver of conservation in Ecuador through Fundación Jocotoco, which he helped establish twenty years ago. In 1980, Bob visited the type locality of Dipsas bobridgelyi (Buenaventura, meaning "good fortune"), now known to be a key area for the conservation of biodiversity. Bob embarked on conservation and worked diligently to raise funds through Rainforest Trust for the past 18 years to purchase private properties and establish what is now the Reserva Buenaventura of Fundación Jocotoco.
We consider Dipsas bobridgelyi to be Endangered following the IUCN criteria B1a,b(i,iii) (IUCN 2001) because its extent of occurrence is estimated to be less than 5,000 km2, it is known only from 4 patches of forest lacking connectivity between them, and its habitat is severely fragmented and declining in extent and quality due to deforestation. Furthermore, only two of the localities (Buenaventura reserve and Reserva Nacional de Tumbes) where D. bobridgelyi occurs are currently protected.
George Jett’s Snail-Eater
Caracolera de George Jett
MZUTI 5411 (Figs
DHMECN 11639, adult male collected by Jacinto Bravo in 2014 at Montecristi, province of Manabí, Ecuador (S1.04694, W80.65766; 136 m). DHMECN 11646, adult male collected by Félix Almeida in 2014 at Rocafuerte, province of Manabí, Ecuador (S0.92371, W80.45212; 19 m).
Dipsas georgejetti is placed in the genus Dipsas based on phylogenetic evidence (Fig.
Dipsas georgejetti is most similar to D. oswaldobaezi, D. williamsi, D. oligozonata, and D. vagrans, in that order, all of which were previously included in the genus Sibynomorphus. From D. oswaldobaezi (Figs
Adult male, SVL 315 mm, TL 87 mm (28% SVL); head length 13.6 mm (4% SVL) from tip of snout to commissure of mouth; head width 8.4 mm (62% head length) taken at broadest point; snout-orbit distance 3.5 mm; head distinct from neck; snout short, blunt in dorsal and lateral outline; rostral 2.0 mm wide, broader than high; internasals 1.7 mm wide, broader than long; prefrontals 2.5 mm wide, longer than broad and contacting orbit; supraocular 3.4 mm long, longer than broad; frontal 3.3 mm long, pentagonal, in contact with prefrontals, supraoculars, and parietals; parietals 5.5 mm long, longer than broad; nasal divided, in contact with first two supralabials, loreal, prefrontal, internasal, and rostral; loreal 1.7 mm long, slightly higher than long, entering orbit; eye diameter 2.8 mm; pupil semi-elliptical; no preocular; two postoculars; temporals 2+2; seven supralabials, 4th and 5th contacting orbit; symphysial in contact with first pair of chinshields; nine infralabials, 1–6 contacting chinshields; anterior pair of chinshields longer than broad, posterior pair broader than long; dorsal scales in 15/15/15 rows, smooth, without apical pits; 178 ventrals; 69 divided subcaudals; cloacal plate single.
The holotype was active during a dry night after a sunny day. It was perched on tangled vegetation 130 cm above the ground in dry shrubland besides recently cleared pasture.
Deciduous and semideciduous forests along the central Pacific coast in Ecuador in the provinces of Manabí and Guayas, at elevations between 5 and 317 m (Fig.
The specific name georgejetti honors George Jett, who has been a long-time donor to Rainforest Trust and has supported the reserves of Fundación Jocotoco in Ecuador. He is an international traveler with a passion for reptiles, amphibians, and birds.
We consider Dipsas georgejetti to be Vulnerable following the IUCN criteria A1c,B1a,b(iii, iv) (IUCN 2001) because its extent of occurrence is estimated to be 10,193 km2, it is known only from 9 localities effectively corresponding to 4 patches of forest lacking connectivity between them, and its habitat is severely fragmented and declining in extent and quality due to deforestation. At the type locality, D. georgejetti was found in a patch of deciduous forest of 13 km2 that was being cleared to accommodate cattle pastures. One of the localities, 15 km N of Guayaquil, where D. georgejetti was collected in 1959, is now completely deforested, which suggests that this arboreal species is no longer present there.
Sibynomorphus oligozonatus Cadle, 2007: 195 (part).
Oswaldo Báez’ Snail-Eater
Caracolera de Oswaldo Báez
BMNH1935.11.3.108, adult female collected by Clodoveo Carrión in the valley of Catamayo, province of Loja, Ecuador (S3.98064, W79.35928; 1289 m). MUSM 2192, adult male collected by Otavio Ruíz in Piura (department or city not specified), Peru.
Dipsas oswaldobaezi is placed in the genus Dipsas based on phylogenetic evidence (Fig.
Dipsas oswaldobaezi is most similar to D. williamsi, D. georgejetti, D. oligozonata, and D. vagrans, in that order, all of which were previously included in the genus Sibynomorphus. From D. williamsi, it differs in having 7–9 infralabials (vs. 10 in D. williamsi), first supralabial not in contact with prefrontal (vs. in broad contact in D. williamsi), and dorsal blotches that are lighter in the middle (vs. dark solid blotches). From D. georgejetti (Figs
Adult female, SVL 277 mm, tail length 85 mm (31% SVL); head length 9.5 mm (3.4% SVL) from tip of snout to commissure of mouth; head width 7.3 mm (76% head length) taken at broadest point; snout-orbit distance 3.3 mm; head distinct from neck; snout short, blunt in dorsal and lateral outline; rostral 2.1 mm wide, broader than high; internasals 1.2 mm wide, broader than long; prefrontals 2.2 mm wide, slightly broader than long and contacting orbit; supraocular 2.6 mm long, longer than broad; frontal 2.9 mm long, pentagonal, in contact with prefrontals, supraoculars, and parietals; parietals 4.2 mm long, longer than broad; nasal not divided, in contact with first supralabial, loreal, prefrontal, internasal, and rostral; loreal 1.3 mm long, longer than high, entering orbit; eye diameter 2.2 mm; pupil semi-elliptical; no preocular; two postoculars; temporals 2+2; 6 supralabials, 3rd and 4th contacting orbit; symphysial separated from chinshields by the first pair of infralabials; 9/8 (right/left) infralabials, 1–6/1–5 contacting chinshields; both pairs of chinshields longer than broad; dorsal scales in 15/15/15 rows, smooth, without apical pits; 179 ventrals; 70 divided subcaudals; cloacal plate single.
Individuals of Dipsas oswaldobaezi have been found active by night on vegetation or at ground level in forested environments, pastures, or rural gardens. One individual (
Deciduous and semideciduous lowland to lower montane forests and dry lowland shrublands in southwestern Ecuador (provinces of Loja and El Oro) and northwestern Peru (department of Tumbes), at elevation between 39 and 1289 m (Fig.
The specific name oswaldobaezi honors Dr. Oswaldo Báez, a renowned Ecuadorian biologist and researcher who has dedicated his life to the teaching of science, scientific thinking, and the conservation of nature. Oswaldo Báez has played a major role in science education in Ecuador through many popular science articles and books.
We consider Dipsas oswaldobaezi to be Vulnerable following the IUCN criteria B1a,b(iii, iv) (IUCN 2001) because its extent of occurrence is estimated to be 8,605 km2; it is known only from eight localities effectively corresponding to four patches of forest lacking connectivity between them, and its habitat is severely fragmented and declining in extent and quality due to deforestation.
In his revision of Dipsas oligozonata,
Locality data for specimens examined in this study. Coordinates represent actual GPS readings taken at the locality of collection or georeferencing attempts from gazetteers under standard guidelines, though some variation from the exact collecting locality will be present. Similarly, elevations are taken from Google Earth, and may not exactly match the elevations as originally reported. Specimens listed here but not under Appendix
Species | Voucher | Country | Province | Locality | Latitude | Longitude | Elev. (m) |
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D. andiana |
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Ecuador | Cañar | Ocaña | -2.48807, -79.18758 | 923 | |
D. andiana |
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Ecuador | Cotopaxi | Las Pampas | -0.43021, -78.96663 | 1590 | |
D. andiana | MZUTI 5413 | Ecuador | El Oro | Reserva Buenaventura | -3.65477, -79.76830 | 497 | |
D. andiana | MZUTI 3501 | Ecuador | Pichincha | Mashpi lodge | 0.16567, -78.88656 | 860 | |
D. andiana | MZUTI 3505 | Ecuador | Pichincha | Valle Hermoso–Los Bancos | -0.01371, -79.09462 | 571 | |
D. andiana |
|
Ecuador | Pichincha | Tandayapa | 0.00205, -78.67880 | 1734 | |
D. andiana |
|
Ecuador | Pichincha | Hacienda La Joya | 0.08291, -78.98311 | 763 | |
D. andiana |
|
Ecuador | Manabí | 5km W Puerto López | -1.59045, -80.84087 | 7 | |
D. bobridgelyi |
|
Ecuador | Azuay | Ponce Enríquez | -3.06547, -79.74358 | 39 | |
D. bobridgelyi | DHMECN 11527 | Ecuador | El Oro | Remolino | -3.56551, -79.91948 | 229 | |
D. bobridgelyi | MZUTI 3266 | Ecuador | El Oro | Reserva Buenaventura | -3.65467, -79.76794 | 524 | |
D. bobridgelyi | MZUTI 5414 | Ecuador | El Oro | Reserva Buenaventura | -3.65310, -79.76336 | 572 | |
D. bobridgelyi | MZUTI 5417 | Ecuador | El Oro | Reserva Buenaventura | -3.65467, -79.76794 | 524 | |
D. catesbyi |
|
Ecuador | Morona Santiago | Macas | -2.31670, -78.11670 | 972 | |
D. catesbyi |
|
Ecuador | Morona Santiago | San Pablo de Kantesiya | -0.25001, -76.41849 | 250 | |
D. catesbyi |
|
Ecuador | Morona Santiago | Sucúa | -2.45663, -78.16784 | 829 | |
D. catesbyi | DHMECN 11555 | Ecuador | Napo | El Reventador | -0.04669, -77.52898 | 1428 | |
D. catesbyi |
|
Ecuador | Napo | Hollín–Loreto | -0.74087, -77.51945 | 1020 | |
D. catesbyi |
|
Ecuador | Napo | San Rafael | -0.10354, -77.58337 | 1246 | |
D. catesbyi |
|
Ecuador | Napo | San Rafael | -0.09669, -77.58995 | 1464 | |
D. catesbyi |
|
Ecuador | Orellana | Hacienda Primavera | -0.48689, -76.63581 | 267 | |
D. catesbyi |
|
Ecuador | Pastaza | Puyo | -1.46678, -77.98335 | 953 | |
D. catesbyi |
|
Ecuador | Pastaza | Villano B | -1.49961, -77.48234 | 341 | |
D. catesbyi |
|
Ecuador | Sucumbíos | El Reventador | -0.04480, -77.52858 | 1476 | |
D. catesbyi |
|
Ecuador | Sucumbíos | El Reventador | -0.04669, -77.52898 | 1428 | |
D. catesbyi |
|
Ecuador | – | – | – | – | – |
D. catesbyi |
|
Ecuador | – | – | – | – | – |
D. catesbyi | MZUTI 4736 | Ecuador | – | – | – | – | – |
D. catesbyi | MZUTI 4999 | Ecuador | – | – | – | – | – |
D. elegans |
|
Ecuador | Cotopaxi | Cutzualo | -0.54497, -78.91891 | 1952 | |
D. elegans |
|
Ecuador | Cotopaxi | Galápagos | -0.40583, -78.96667 | 1781 | |
D. elegans | DHMECN 1693 | Ecuador | Cotopaxi | Hacienda “La Mariela” | -1.14757, -79.09126 | 1256 | |
D. elegans |
|
Ecuador | Cotopaxi | Las Damas | -0.38402, -78.96741 | 1678 | |
D. elegans |
|
Ecuador | Cotopaxi | Las Pampas | -0.43021, -78.96663 | 1590 | |
D. elegans |
|
Ecuador | Cotopaxi | Palo Quemado | -0.61962, -78.99066 | 2402 | |
D. elegans |
|
Ecuador | Pichincha | 2.9 km SW of Tandayapa | 0.00578, -78.67867 | 1844 | |
D. elegans |
|
Ecuador | Pichincha | Ilaló | -0.26166, -78.44444 | 2579 | |
D. elegans | MZUTI 3695 | Ecuador | Pichincha | Tambotanda | -0.02011, -78.65101 | 1875 | |
D. elegans | MZUTI 3317 | Ecuador | Pichincha | Tandapi | -0.42278, -78.79611 | 1550 | |
D. elegans |
|
Ecuador | Santo Domingo | Chiriboga | -0.22841, -78.76725 | 1813 | |
D. elegans |
|
Ecuador | Santo Domingo | Hacienda Las Palmeras | -0.24520, -78.84806 | 1876 | |
D. elegans |
|
Ecuador | – | – | – | – | – |
D. elegans | MZUTI 3316 | Ecuador | – | – | – | – | – |
D. ellipsifera | MZUTI 4931 | Ecuador | Carchi | Chilma Bajo | 0.86274, -78.05080 | 2071 | |
D. ellipsifera |
|
Ecuador | Carchi | Quebrada Golondrinas | 0.83210, -78.12324 | 1737 | |
D. ellipsifera |
|
Ecuador | Carchi | Río Pailón | 0.95643, -78.23448 | 1669 | |
D. ellipsifera |
|
Ecuador | Imbabura | Cotacachi | 0.29395, -78.26682 | 2446 | |
D. gracilis |
|
Ecuador | Cañar | Manta Real | -2.55367, -79.36425 | 257 | |
D. gracilis |
|
Ecuador | Esmeraldas | Angostura | 1.02164, -78.86295 | 31 | |
D. gracilis |
|
Ecuador | Esmeraldas | Caimito | 0.69546, -80.08990 | 118 | |
D. gracilis |
|
Ecuador | Esmeraldas | Estero Gasparito | 0.91296, -78.84066 | 80 | |
D. gracilis |
|
Ecuador | Esmeraldas | Fauna Granja Tropical | 0.66152, -79.53875 | 29 | |
D. gracilis |
|
Ecuador | Esmeraldas | La Mayronga | 1.04361, -79.27786 | 14 | |
D. gracilis |
|
Ecuador | Esmeraldas | Tundaloma | 1.18166, -78.74945 | 74 | |
D. gracilis |
|
Ecuador | Guayas | Naranjal | -2.72302, -79.63172 | 58 | |
D. gracilis |
|
Ecuador | Guayas | Naranjal | -2.72302, -79.63172 | 58 | |
D. gracilis |
|
Ecuador | Guayas | Río Patul | -2.55548, -79.37180 | 266 | |
D. gracilis |
|
Ecuador | Los Ríos | Buena Fe | -0.89306, -79.48957 | 104 | |
D. gracilis |
|
Ecuador | Los Ríos | Río Palenque | -0.58333, -79.36667 | 173 | |
D. gracilis |
|
Ecuador | Los Ríos | Río Palenque | -0.58333, -79.36667 | 173 | |
D. gracilis |
|
Ecuador | Los Ríos | Río Palenque | -0.58333, -79.36667 | 173 | |
D. gracilis |
|
Ecuador | Los Ríos | Río Palenque | -0.58333, -79.36667 | 173 | |
D. gracilis |
|
Ecuador | Los Ríos | Río Palenque | -0.58333, -79.36667 | 173 | |
D. gracilis |
|
Ecuador | Los Ríos | Río Palenque | -0.58333, -79.36667 | 173 | |
D. gracilis | DHMECN 2902 | Ecuador | Manabí | El Aguacate | 0.65348, -80.05190 | 43 | |
D. gracilis |
|
Ecuador | Manabí | Jama Coaque | -0.11455, -80.12337 | 321 | |
D. gracilis |
|
Ecuador | Manabí | Lalo Loor | -0.08337, -80.15004 | 75 | |
D. gracilis |
|
Ecuador | Manabí | Maicito | -0.27265, -79.57179 | 173 | |
D. gracilis |
|
Ecuador | Manabí | Maicito | -0.27265, -79.57179 | 173 | |
D. gracilis |
|
Ecuador | Manabí | Maicito | -0.27265, -79.57179 | 173 | |
D. gracilis |
|
Ecuador | Manabí | Maicito | -0.27265, -79.57179 | 173 | |
D. gracilis |
|
Ecuador | Manabí | Reserva Jama Coaque | -0.11556, -80.12472 | 299 | |
D. gracilis |
|
Ecuador | Manabí | Zapallo Grande | 0.78165, -78.98345 | 100 | |
D. gracilis |
|
Ecuador | Pichincha | Cachaco–Lita | 0.78886, -78.36794 | 1108 | |
D. gracilis | MZUTI 1386 | Ecuador | Pichincha | El Abrazo del Árbol | -0.00913, -78.81321 | 1064 | |
D. gracilis |
|
Ecuador | Pichincha | El Monte | -0.06912, -78.76195 | 1316 | |
D. gracilis |
|
Ecuador | Pichincha | Finca Ecológica Orongo | 0.15304, -78.66737 | 1173 | |
D. gracilis | MZUTI 3503 | Ecuador | Pichincha | Mashpi lodge | 0.16681, -78.88111 | 905 | |
D. gracilis |
|
Ecuador | Pichincha | Rainforest Monterreal | 0.01557, -78.88407 | 860 | |
D. gracilis |
|
Ecuador | Pichincha | Road to Mindo | -0.03116, -78.75617 | 1638 | |
D. gracilis |
|
Ecuador | Santo Domingo | 8.5 km NW Santo Domingo | -0.17700, -79.21099 | 454 | |
D. gracilis |
|
Ecuador | Santo Domingo | 8.5 km NW Santo Domingo | -0.17700, -79.21099 | 454 | |
D. gracilis |
|
Ecuador | Santo Domingo | Finca de Germán Cortez | -0.00027, -79.41194 | 194 | |
D. gracilis |
|
Ecuador | Santo Domingo | La Perla | 0.13417, -79.49432 | 132 | |
D. gracilis | DHMECN 129 | Ecuador | – | – | – | – | – |
D. gracilis | MZUTI 4199 | Ecuador | – | – | – | – | – |
D. indica |
|
Ecuador | Morona Santiago | Rosa de Oro | – | – | – |
D. indica |
|
Ecuador | Orellana | Coca | -0.46167, -76.99310 | 253 | |
D. indica |
|
Ecuador | Orellana | Hacienda Primavera | -0.48689, -76.63581 | 267 | |
D. indica | MZUTI 4735 | Ecuador | Pastaza | Tzarentza | -1.35696, -78.05814 | 1355 | |
D. jamespetersi |
|
Ecuador | Azuay | La Paz | -3.31481, -79.15166 | 3148 | |
D. jamespetersi | MZUTI 5307 | Ecuador | Azuay | Poetate | -3.41645, -79.26964 | 2269 | |
D. jamespetersi |
|
Ecuador | Loja | 0.5 km E of Loja | -3.99277, -79.18327 | 2263 | |
D. jamespetersi |
|
Ecuador | Loja | 24 km S Loja | -4.22083, -79.24164 | 1562 | |
D. jamespetersi |
|
Ecuador | Loja | 24 km S Loja | -4.22083, -79.24164 | 1562 | |
D. jamespetersi |
|
Ecuador | Loja | 5 km E Loja | -3.98899, -79.16576 | 2610 | |
D. jamespetersi |
|
Ecuador | Loja | 5 km E Loja | -3.98899, -79.16576 | 2610 | |
D. jamespetersi |
|
Ecuador | Loja | 5 km E Loja | -3.98899, -79.16576 | 2610 | |
D. jamespetersi |
|
Ecuador | Loja | 5 km E Loja | -3.98899, -79.16576 | 2610 | |
D. jamespetersi |
|
Ecuador | Loja | 5 km E Loja | -3.98899, -79.16576 | 2610 | |
D. jamespetersi |
|
Ecuador | Loja | Guachanamá | -4.04081, -79.88290 | 2787 | |
D. jamespetersi |
|
Ecuador | Loja | Loja | -4.00789, -79.21128 | 2166 | |
D. latifrontalis | BMNH1946.1.20 | Venezuela | Mérida | Aricagua | 8.16162, -71.15776 | 1078 | |
D. klebbai |
|
Ecuador | Napo | 2 km E Borja | -0.41543, -77.83032 | 1608 | |
D. klebbai | DHMECN 568 | Ecuador | Napo | Borja | -0.42624, -77.84277 | 1698 | |
D. klebbai |
|
Ecuador | Napo | El Chaco | -0.33763, -77.80957 | 1595 | |
D. klebbai |
|
Ecuador | Napo | El Chaco | -0.33763, -77.80957 | 1595 | |
D. klebbai |
|
Ecuador | Napo | El Chaco | -0.33763, -77.80957 | 1595 | |
D. klebbai |
|
Ecuador | Napo | El Chaco | -0.33763, -77.80957 | 1595 | |
D. klebbai | MZUTI 5412 | Ecuador | Napo | Pacto Sumaco | -0.66377, -77.59895 | 1556 | |
D. klebbai |
|
Ecuador | Napo | Río Azuela | -0.14869, -77.65463 | 1402 | |
D. klebbai |
|
Ecuador | Napo | Río Azuela | -0.14869, -77.65463 | 1402 | |
D. klebbai |
|
Ecuador | Napo | Río Azuela | -0.14869, -77.65463 | 1402 | |
D. klebbai |
|
Ecuador | Napo | San Rafael | -0.09669, -77.58995 | 1464 | |
D. klebbai |
|
Ecuador | Napo | San Rafael | -0.09669, -77.58995 | 1464 | |
D. klebbai | MZUTI 63 | Ecuador | Napo | Yanayacu | -0.60042, -77.89053 | 2110 | |
D. klebbai |
|
Ecuador | Sucumbíos | El Reventador | -0.04480, -77.52858 | 1476 | |
D. klebbai |
|
Ecuador | Sucumbíos | El Reventador | -0.04480, -77.52858 | 1476 | |
D. klebbai |
|
Ecuador | Sucumbíos | El Reventador | -0.04669, -77.52898 | 1428 | |
D. klebbai |
|
Ecuador | Sucumbíos | La Bonita | 0.47209, -77.54661 | 1953 | |
D. klebbai |
|
Ecuador | – | – | – | – | – |
D. klebbai |
|
Ecuador | Napo | Cascada de San Rafael | -0.10007, -77.58034 | 1182 | |
D. georgejetti |
|
Ecuador | Guayas | 10 mi N of Guayaquil | -1.96418, -79.87988 | 5 | |
D. georgejetti |
|
Ecuador | Guayas | Cerro Blanco | -2.17465, -80.02135 | 147 | |
D. georgejetti | ENS 12817 | Ecuador | Manabí | 17 km NW Portoviejo | -1.00209, -80.31334 | 187 | |
D. georgejetti | MZUTI 5411 | Ecuador | Manabí | Cabuyal | -0.19698, -80.29059 | 15 | |
D. georgejetti |
|
Ecuador | Manabí | El Aromo | -1.04665, -80.83276 | 295 | |
D. georgejetti | DHMECN 11639 | Ecuador | Manabí | Montecristi | -1.04694, -80.65766 | 136 | |
D. georgejetti |
|
Ecuador | Manabí | El Aromo | -1.04665, -80.83276 | 295 | |
D. georgejetti |
|
Ecuador | Manabí | El Aromo | -1.04665, -80.83276 | 295 | |
D. georgejetti | DHMECN 11646 | Ecuador | Manabí | Rocafuerte | -0.92371, -80.45212 | 19 | |
D. georgejetti |
|
Ecuador | Manabí | Cerro La Mocora, foothill | -1.59817, -80.65431 | 308 | |
D. oligozonata |
|
Ecuador | Azuay | Girón | -3.15891, -79.14755 | 2102 | |
D. oligozonata |
|
Ecuador | Azuay | Granja Orgánica Susudel | -3.38885, -79.17847 | 2802 | |
D. oligozonata |
|
Ecuador | Azuay | Susudel | -3.40543, -79.18378 | 2376 | |
D. oligozonata |
|
Ecuador | Azuay | Via a Shaglli | -3.19178, -79.39623 | 2891 | |
D. oligozonata |
|
Ecuador | – | – | – | – | – |
D. oligozonata |
|
Ecuador | – | – | – | – | – |
D. oreas |
|
Ecuador | Azuay | Luz María | -2.68548, -79.40992 | 1661 | |
D. oreas | DHMECN 3478 | Ecuador | Azuay | Naranjo Lanto | -2.92628, -79.39963 | 1847 | |
D. oreas | DHMECN 7647 | Ecuador | Azuay | Reserva Biológica Yunguilla | -3.22684, -79.27520 | 1748 | |
D. oreas | DHMECN 7666 | Ecuador | Azuay | Reserva Biológica Yunguilla | -3.22684, -79.27520 | 1748 | |
D. oreas |
|
Ecuador | Azuay | San Rafael de Sharug | -3.27311, -79.54543 | 1593 | |
D. oreas |
|
Ecuador | Azuay | San Rafael de Sharug | -3.27311, -79.54543 | 1593 | |
D. oreas |
|
Ecuador | Azuay | Vía La Paz–Cuenca | -3.09021, -79.00800 | 2726 | |
D. oreas |
|
Ecuador | Chimborazo | Pallatanga–Guayaquil | -2.07459, -78.98123 | 1404 | |
D. oreas |
|
Ecuador | Chimborazo | Pallatanga–Guayaquil | -2.07459, -78.98123 | 1404 | |
D. oreas |
|
Ecuador | Chimborazo | Pallatanga–Guayaquil | -2.07459, -78.98123 | 1404 | |
D. oreas | DHMECN 10785 | Ecuador | El Oro | Playa Limón | -3.50096, -79.74701 | 816 | |
D. oreas | DHMECN 2572 | Ecuador | El Oro | Reserva Buenaventura | -3.65467, -79.76794 | 524 | |
D. oreas | MZUTI 3351 | Ecuador | El Oro | Reserva Buenaventura | -3.64882, -79.75640 | 898 | |
D. oreas | MZUTI 5415 | Ecuador | El Oro | Reserva Buenaventura | -3.63432, -79.74985 | 1048 | |
D. oreas | MZUTI 5418 | Ecuador | El Oro | Reserva Buenaventura | -3.63370, -79.75040 | 1068 | |
D. oreas |
|
Ecuador | Loja | 33 km E San Pedro | -3.97222, -79.25983 | 2493 | |
D. oreas |
|
Ecuador | Loja | 6 km S Loja | -4.03770, -79.19975 | 2144 | |
D. oreas |
|
Ecuador | Loja | Cazerío Balzones | -4.01502, -80.01635 | 1346 | |
D. oreas |
|
Ecuador | Loja | Jimbura | -4.66668, -79.45322 | 2513 | |
D. oreas |
|
Ecuador | Loja | Vía al Cerro Toledo | -4.38444, -79.15992 | 2214 | |
D. oreas |
|
Ecuador | Loja | Vilcabamba | -4.25792, -79.21962 | 1546 | |
D. oreas |
|
Ecuador | Loja | Yangana–Vilcabamba | -4.32455, -79.20041 | 1742 | |
D. palmeri |
|
Ecuador | Morona Santiago | 9 de Octubre–Macas | -2.21820, -78.29920 | 1767 | |
D. palmeri |
|
Ecuador | Morona Santiago | Chiguinda | -3.28125, -78.69829 | 2223 | |
D. palmeri | DHMECN 11197 | Ecuador | Morona Santiago | Concesión ECSA | -3.57524, -78.43609 | 1211 | |
D. palmeri |
|
Ecuador | Morona Santiago | Laguna Chimerella | -2.07956, -78.20338 | 1795 | |
D. palmeri |
|
Ecuador | Morona Santiago | Laguna Cormorán | -2.07153, -78.21590 | 1747 | |
D. palmeri |
|
Ecuador | Pastaza | Tzarentza | -1.35696, -78.05814 | 1355 | |
D. palmeri |
|
Ecuador | Tungurahua | 3 km E Río Verde | -1.40249, -78.28369 | 1474 | |
D. palmeri | AMNH 24126 | Ecuador | Tungurahua | Abitagua | -1.41667, -78.16667 | 1353 | |
D. palmeri | MZUTI 4804 | Ecuador | Tungurahua | Agoyán | -1.39795, -78.38415 | 1661 | |
D. palmeri |
|
Ecuador | Tungurahua | Baños | -1.39650, -78.42945 | 1847 | |
D. palmeri |
|
Ecuador | Tungurahua | Baños | -1.39650, -78.42945 | 1847 | |
D. palmeri |
|
Ecuador | Tungurahua | Baños | -1.39650, -78.42945 | 1847 | |
D. palmeri |
|
Ecuador | Tungurahua | Caserío Machay | -1.40062, -78.28085 | 1531 | |
D. palmeri | DHMECN 9229 | Ecuador | Tungurahua | Chamanapamba | -1.40114, -78.39975 | 1808 | |
D. palmeri | DHMECN 9230 | Ecuador | Tungurahua | Chamanapamba | -1.40114, -78.39975 | 1808 | |
D. palmeri | MZUTI 3956 | Ecuador | Tungurahua | La Candelaria | -1.43051, -78.31246 | 1920 | |
D. palmeri | AMNH 37939 | Ecuador | Tungurahua | Palmera | -1.41613, -78.19663 | 1225 | |
D. palmeri | DHMECN 9232 | Ecuador | Tungurahua | Parque Juan Montalvo | -1.40005, -78.42070 | 1803 | |
D. palmeri |
|
Ecuador | Tungurahua | Río Verde | -1.39406, -78.30405 | 1603 | |
D. palmeri |
|
Ecuador | Tungurahua | Río Verde | -1.39406, -78.30405 | 1603 | |
D. palmeri | DHMECN 12841 | Ecuador | Tungurahua | Ulba | -1.39622, -78.39418 | 1702 | |
D. palmeri | DHMECN 9219 | Ecuador | Tungurahua | Vizcaya | -1.34789, -78.40518 | 2282 | |
D. palmeri |
|
Ecuador | Zamora Chinchipe | 18.2 km W Zamora | -3.97643, -79.02075 | 1609 | |
D. palmeri |
|
Ecuador | Zamora Chinchipe | 182 km Zamora–Loja | -3.95600, -79.02599 | 1665 | |
D. palmeri |
|
Ecuador | Zamora Chinchipe | Estación San Francisco | -3.96128, -79.05556 | 1775 | |
D. palmeri |
|
Ecuador | Zamora Chinchipe | Reserva Numbami | -4.17233, -78.95928 | 1615 | |
D. palmeri | MZUTI 4971 | Ecuador | Zamora Chinchipe | Reserva San Francisco | -3.97051, -79.07814 | 1850 | |
D. palmeri | MZUTI 4975 | Ecuador | Zamora Chinchipe | Reserva San Francisco | -3.97140, -79.07909 | 1730 | |
D. palmeri |
|
Ecuador | Zamora Chinchipe | Reserva San Francisco | -3.97051, -79.07814 | 1850 | |
D. palmeri | MZUTI 5419* | Ecuador | Zamora Chinchipe | Romerillos Alto | -4.23230, -78.94222 | 1547 | |
D. palmeri |
|
Ecuador | Zamora Chinchipe | Zumba | -4.86517, -79.13384 | 1230 | |
D. palmeri |
|
Ecuador | – | – | – | – | – |
D. palmeri |
|
Peru | Cajamarca | Jaén | -5.72978, -78.84836 | 1438 | |
D. palmeri |
|
Peru | Cajamarca | Tabaconas | -5.31429, -79.29622 | 1892 | |
D. pavonina |
|
Ecuador | Morona Santiago | Kushapuk | -3.04373, -78.03648 | 326 | |
D. pavonina |
|
Ecuador | Morona Santiago | Tiink | -3.34389, -78.46805 | 730 | |
D. pavonina |
|
Ecuador | Napo | Archidona | -0.90856, -77.80814 | 571 | |
D. pavonina |
|
Ecuador | Napo | Tena | -0.98330, -77.81670 | 522 | |
D. pavonina | MZUTI 4972 | Ecuador | Zamora Chinchipe | Maycu | -4.38030, -78.74584 | 981 | |
D. peruana |
|
Peru | Amazonas | 28 km SE Ingenio | -6.05753, -77.98919 | 2235 | |
D. peruana |
|
Peru | Amazonas | Pomacochas | -5.82155, -77.91692 | 2150 | |
D. peruana |
|
Peru | Cuzco | Amaibamba | -13.27703, -73.28636 | 1858 | |
D. peruana |
|
Peru | Cuzco | Bosque Aputinye | -12.92300, -72.67455 | 1502 | |
D. peruana |
|
Peru | Cuzco | Machu Picchu | -13.17104, -72.50585 | 2400 | |
D. peruana | AMNH 147037 | Peru | Cuzco | Paucartambo Mirador | -13.06972, -71.55527 | 1818 | |
D. peruana | AMNH 147037 | Peru | Cuzco | Paucartambo Mirador | -13.06972, -71.55527 | 1810 | |
D. peruana |
|
Peru | Cuzco | Pucyura | -13.07450, -72.93437 | 2666 | |
D. peruana |
|
Peru | Cuzco | Rocotal | -13.10627, -71.57064 | 2004 | |
D. peruana | SMF 20801 | Peru | Cuzco | Santa Ana | -12.86755, -72.71670 | 1639 | |
D. peruana |
|
Peru | Huánuco | Playa Pampa | -9.95160, -75.69605 | 2091 | |
D. peruana |
|
Peru | Pasco | Huancabamba | -10.42265, -75.51718 | 1775 | |
D. peruana |
|
Peru | Puno | 10 km NNE Ollachea | -13.78330, -70.46730 | 2598 | |
D. peruana |
|
Peru | Puno | 11 km NNE Ollachea | -13.78661, -70.47248 | 2601 | |
D. peruana |
|
Peru | Puno | 12 km NNE Ollachea | -13.78330, -70.46730 | 2598 | |
D. peruana | AMNH 52444 | Peru | San Martín | Cumbre Ushpayacu-Mishquiyacu | -6.99468, -76.03371 | 1279 | |
D. temporalis | MZUTI 3331 | Ecuador | Esmeraldas | Tundaloma Lodge | 1.18317, -78.75245 | 74 | |
D. temporalis |
|
Ecuador | Imbabura | 16 km W Lita | 0.90235, -78.54504 | 799 | |
D. vagrans | AMNH 63373 | Peru | San Martín | Bellavista | -7.05346, -76.58928 | 316 | |
D. vermiculata |
|
Ecuador | Morona Santiago | 69 km S Vilcabamba | -4.84920, -79.12731 | 1310 | |
D. vermiculata | DHMECN 11197 | Ecuador | Morona Santiago | Concesión ECSA | -3.57245, -78.46982 | 790 | |
D. vermiculata |
|
Ecuador | Napo | El Reventador | -0.04480, -77.52858 | 1476 | |
D. vermiculata | MZUTI 5080 | Ecuador | Pastaza | Kallana | -1.469629, -77.27838 | 325 | |
D. vermiculata |
|
Ecuador | Pastaza | Sendero Higuerones | -4.11464, -78.96702 | 981 | |
D. vermiculata | MZUTI 4738 | Ecuador | Pastaza | Tzarentza | -1.35696, -78.05814 | 1355 | |
D. vermiculata | MZUTI 3663 | Ecuador | Zamora Chinchipe | Maycu | -4.20719, -78.63987 | 869 | |
D. vermiculata |
|
Ecuador | Zamora Chinchipe | Nangaritza | -4.43169, -78.63869 | 1011 | |
D. oswaldobaezi |
|
Ecuador | El Oro | Arenillas | -3.62110, -80.17513 | 41 | |
D. oswaldobaezi |
|
Ecuador | El Oro | Guabillo | -3.60346, -80.18139 | 44 | |
D. oswaldobaezi |
|
Ecuador | El Oro | Huaquillas | -3.54115, -80.08646 | 39 | |
D. oswaldobaezi |
|
Ecuador | Loja | Quebrada El Faique | -4.17889, -80.04226 | 1004 | |
D. oswaldobaezi |
|
Ecuador | Loja | Reserva La Ceiba-Pilares | -4.27502, -80.32805 | 534 | |
D. oswaldobaezi | BMNH1935.11.3.108 | Ecuador | Loja | Catamayo | -3.98064, -79.35928 | 1289 | |
D. oswaldobaezi | MUSM 2192 | Peru | Piura | Piura | -5.17882, -80.62231 | 32 | |
S. annulatus | MZUTI 3034 | Ecuador | Esmeraldas | Reserva Itapoa | 0.51307, -79.13401 | 321 | |
S. bevridgelyi |
|
Ecuador | Azuay | 2 km N Palmales Nuevo | -3.65158, -80.09625 | 129 | |
S. bevridgelyi |
|
Ecuador | Azuay | 30 KM E Pasaje | -3.31439, -79.57970 | 561 | |
S. bevridgelyi |
|
Ecuador | Azuay | Ponce Enríquez–El Coca | -3.03197, -79.64615 | 1206 | |
S. bevridgelyi |
|
Ecuador | Azuay | Proyecto Minas San Francisco | -3.30829, -79.47079 | 862 | |
S. bevridgelyi |
|
Ecuador | Azuay | Sarayunga | -3.31431, -79.58069 | 552 | |
S. bevridgelyi |
|
Ecuador | Chimborazo | Valle del Chanchán | -2.27383, -79.08735 | 697 | |
S. bevridgelyi | DHMECN 11526 | Ecuador | El Oro | Remolino | -3.56551, -79.91948 | 229 | |
S. bevridgelyi | DHMECN 9483 | Ecuador | El Oro | Reserva Buenaventura | -3.65467, -79.76794 | 524 | |
S. bevridgelyi | MZUTI 3269 | Ecuador | El Oro | Reserva Buenaventura | -3.65343, -79.76722 | 473 | |
S. bevridgelyi | MZUTI 5416 | Ecuador | El Oro | Reserva Buenaventura | -3.65467, -79.76794 | 524 | |
S. bevridgelyi | AMNH 22092 | Ecuador | Guayas | Reserva Ayampe | -1.65417, -80.81833 | 43 | |
S. bevridgelyi |
|
Ecuador | Guayas | Río Daule | -1.87009, -80.00539 | 5 | |
S. bevridgelyi |
|
Ecuador | Los Ríos | Jauneche | -1.33333, -79.58333 | 41 | |
S. bevridgelyi | DHMECN 8976 | Ecuador | Manabí | San Sebastián | -1.60002, -80.69974 | 602 | |
S. bevridgelyi | DHMECN 10061 | Ecuador | Manabí | Puerto López | -1.55598, -80.81200 | 3 | |
S. bevridgelyi |
|
Ecuador | Manabí | Cerro La Mocora, tophill | -1.60379, -80.70191 | 818 | |
S. bevridgelyi |
|
Peru | Tumbes | El Caucho | -3.81438, -80.27101 | 379 | |
S. bevridgelyi |
|
Peru | Tumbes | El Caucho | -3.81438, -80.27101 | 379 | |
S. bevridgelyi |
|
Peru | Tumbes | El Caucho | -3.81844, -80.26856 | 478 | |
S. bevridgelyi |
|
Peru | Tumbes | El Caucho | -3.81244, -80.26716 | 481 | |
S. nebulatus | MZUTI 4810 | Ecuador | Cotopaxi | El Jardín de los Sueños | -0.83142, -79.21337 | 349 | |
S. nebulatus | DHMECN 9585 | Ecuador | Esmeraldas | Canandé | 0.52580, -79.20880 | 310 | |
S. nebulatus | DHMECN 5645 | Ecuador | Esmeraldas | Lita–San Lorenzo | 1.18236, -78.79528 | 42 | |
S. nebulatus | MZUTI 3911 | Ecuador | Esmeraldas | Reserva Itapoa | 0.51307, -79.13401 | 321 | |
S. nebulatus | DHMECN 5647 | Ecuador | Esmeraldas | Tundaloma | 1.18236, -78.75250 | 74 | |
S. nebulatus | DHMECN 10312 | Ecuador | Imbabura | Selva Alegre | 0.26667, -78.58333 | 1299 | |
S. nebulatus |
|
Ecuador | Los Ríos | Hacienda Cerro Chico | -0.62444, -79.42940 | 170 | |
S. nebulatus |
|
Ecuador | Los Ríos | Macul | -1.12980, -79.65730 | 65 | |
S. nebulatus |
|
Ecuador | Los Ríos | Río Palenque | -0.58333, -79.36667 | 173 | |
S. nebulatus |
|
Ecuador | Los Ríos | Río Palenque | -0.58333, -79.36667 | 173 | |
S. nebulatus |
|
Ecuador | Los Ríos | Río Palenque | -0.58333, -79.36667 | 173 | |
S. nebulatus | DHMECN 2882 | Ecuador | Manabí | Aguacate | 0.65348, -80.05190 | 43 | |
S. nebulatus | MZUTI 5342 | Ecuador | Manabí | Jama Coaque | -0.11556, -80.12472 | 299 | |
S. nebulatus | DHMECN 1704 | Ecuador | Pichincha | Curipogio | 0.13112, -78.67632 | 1171 | |
S. nebulatus |
|
Ecuador | Santo Domingo | Rancho Santa Teresita | -0.25277, -79.37946 | 288 |
Based on differences in coloration and the topology of the molecular phylogeny obtained here (Fig.
Dipsas peruana Harvey & Embert, 2008: 79 (part).
Klebba’s Snail-Eater
Caracolera de Klebba
MZUTI 5412 (Figs
DHMECN 568, adult female collected by Thomas Begher on 1980 at Borja, province of Napo, Ecuador (S0.42054, W77.84104; 1717 m).
Dipsas klebbai is placed in the genus Dipsas based on phylogenetic evidence (Fig.
Dipsas klebbai is compared to species previously subsumed under D. peruana: D. latifrontalis, D. palmeri, and D. peruana. From D. latifrontalis (Fig.
Adult male, SVL 608 mm, tail length 262 mm (43% SVL); head length 20.3 mm (3% SVL) from tip of snout to commissure of mouth; head width 12.7 mm (62% head length) taken at broadest point; snout-orbit distance 5.4 mm; head distinct from neck; snout short, blunt in dorsal and lateral outline; rostral 4.0 mm wide, broader than high; internasals 2.6 mm wide, as broad as long; prefrontals 3.9 mm wide, broader than long, excluded from entering orbit by preocular; supraocular 4.3 mm long, broader than long; frontal 4.5 mm long, hexagonal, in contact with prefrontals, supraoculars, and parietals; parietals 6.6 mm long, longer than broad; nasal divided, in contact with first two supralabials, loreal, prefrontal, internasal, and rostral; loreal 2.6 mm long, slightly longer than high, entering orbit; eye diameter 4.5 mm; pupil semi-elliptical; one preocular; two postoculars; temporals 2+2; ten supralabials, 5th and 6th contacting orbit; symphysial separated from chinshields by the first pair of infralabials; 14 infralabials, 2–7 contacting chinshields; anterior pair of chinshields longer than broad, posterior pair broader than long; dorsal scales in 15/15/15 rows, smooth, without apical pits; 188 ventrals; 116 divided subcaudals; cloacal plate single.
At night (21h53–02h13), specimens of Dipsas klebbai have been found active during or after light rain on arboreal vegetation 50–500 cm above the ground in a variety of environments ranging from primary montane cloud forests and evergreen montane forests to silvopastures and forest borders, occasionally close to rivers. By day, individuals have been found hidden underground in pastures or among shrubs in rural gardens, or coiled on leaves at 300 cm above the ground. At dusk, after warm days, individuals of Dipsas klebbai have been seen crossing roads.
Endemic to the eastern slopes of the Ecuadorian Andes in the provinces of Napo and Sucumbíos at elevations between 1246 and 2120 m (Fig.
Named after Casey Klebba, in recognition of his appreciation of and passion for Andean wildlife, and his invaluable support of AA’s field expeditions to remote areas of Ecuador. After a visit to Peru in 2011, Casey became an active supporter of conservation and scientific projects in Ecuador.
All known localities of occurrence for Dipsas klebbai fall within the limits or within the buffer zone of the following protected areas: Parque Nacional Cayambe Coca, Parque Nacional Sumaco Napo Galeras, Reserva Ecológica Antisana, and Reserva Ecológica Cofán Bermejo. Furthermore, the species is common in degraded environments, which suggests a degree of tolerance for habitat modification. For these reasons, and because it does not meet the criteria (IUCN 2001) for qualifying in a threatened category, we here list it as Least Concern following IUCN guidelines.
In their revision of Dipsas peruana,
Leptognathus
palmeri
Boulenger, 1912: 422. Holotype
Leptognathus
latifasciatus
Boulenger, 1913: 72. Holotype
Dipsas peruana Harvey & Embert, 2008: 79 (part).
Palmer’s Snail-Eater
Caracolera de Palmer
Dipsas palmeri differs from all described species of Dipsas based on the following combination of characters: (1) 15/15/15 smooth dorsals with enlarged vertebral row; (2) one loreal and one preocular in contact with orbit; (3) 8–10 supralabials with (usually) 4th to 6th contacting orbit; (4) one pair of infralabials in contact behind symphysial; (5) 172–202 ventrals in males, 181–200 in females; (6) 91–118 divided subcaudals in males, 86–102 in females; (7) dorsal and ventral ground color light brown with various degrees of fine black speckling and with 32–41 brown to blackish, white-edged circular blotches that are longer than interspaces in the first half of the body, but shorter in the second half (Figs
Dipsas palmeri is compared to species previously subsumed under D. peruana: D. latifrontalis, D. klebbai (Fig.
Eastern slopes of the Ecuadorian and Peruvian Andes south of the Jatunyacu–Napo river valley in Ecuador and north of the Huancabamba depression at elevations between 1211 and 2282 m (Fig.
An estimated 31 out of the 42 known localities of occurrence for Dipsas palmeri are located within the limits or the buffer area of the following protected areas: Bosque Protector del Alto Nangaritza, Parque Nacional Llanganates, Parque Nacional Podocarpus and Parque Nacional Sangay. Furthermore, the presence of the species in degraded environments suggests a degree of tolerance for habitat modification. For these reasons, and because it does not meet the criteria for qualifying in a threatened category, we here list it as Least Concern following IUCN guidelines.
Neither
Two other junior synonyms of Dipsas peruana are D. latifasciata and D. polylepis, both of which occur in Peru (Fig.
Leptognathus peruana Boettger, 1898: 128. Holotype SMF 20801, a female from Santa Ana, department of Cuzco, Peru.
Leptognathus boettgeri Werner, 1901: 11. Holotype MTKD D 1671 M, a female from Chanchamayo, department of Junín, Peru.
Leptognathus boliviana Werner, 1909: 240. Holotype ZMH, a female from department of Beni, Bolivia.
Leptognathus
polylepis
Boulenger, 1912: 422. Holotype
Peruvian Snail-Eater
Caracolera Peruana
Dipsas peruana differs from all described species of Dipsas based on the following combination of characters: (1) 15/15/15 smooth dorsals with moderately enlarged vertebral row; (2) one loreal and one preocular in contact with orbit; (3) 8–9 supralabials with 4–6 or 3–5 contacting orbit; (4) one pair of infralabials in contact behind symphysial; (5) 177–200 ventrals in males, 180–203 in females; (6) 75–127 divided subcaudals in males, 79–105 in females; (7) dorsal and ventral ground color brown to dark brown (light brown in juveniles) with 33–43 blackish brown to complete black, white to cream edged circular to vertically elliptical blotches that are longer than interspaces; head dark brown with dingy cream reticulations and different degrees of whitish edging on the labial scales, and a thin (1–3 scales long) white to light grayish brown irregular nuchal collar; dorsal blotches extending marginally onto ventrals and rarely fusing midventrally; (8) 199 mm SVL in males, 610–725 mm in females; (9) 85 mm TL in males, 155–241 mm in females.
Dipsas peruanasensu stricto is compared to species previously subsumed under D. peruanasensu lato: D. latifrontalis, D. palmeri, and D. klebbai. From D. latifrontalis and D. palmeri, it differs in having dorsal blotches along the entire body similar in length or longer than interspaces (shorter than interspaces in D. latifrontalis and D. palmeri), and in having melanized interspaces in some adult individuals. With the exception of
Eastern slopes of the Peruvian and Bolivian Andes south of the Huancabamba depression at elevations between 1279 and 2671 m (Fig.
Leptognathus
latifrontalis
Boulenger, 1905: 561. Holotype
Dipsas peruana Harvey & Embert, 2008: 79 (part).
Broad-fronted Snail-Eater
Caracolera frentona
Dipsas latifrontalis differs from all described species of Dipsas based on the following combination of characters: (1) 15/15/15 smooth dorsals with moderately enlarged vertebral row; (2) one loreal and one preocular in contact with orbit; (3) 8–10 supralabials with 3rd to 6th contacting orbit; (4) one pair of infralabials in contact behind symphysial; (5) 192 ventrals in one male (CVULA 7883), 194 in the female holotype; (6) 109 divided subcaudals in the single male, 95 in the female holotype; (7) dorsal and ventral ground color bronze (light brown in juveniles) with 32–36 dark reddish brown to black, circular to vertically elliptical blotches that are longer than interspaces and white to cream edged on first half of body; head grayish brown to black with different degrees of whitish edging on the labial scales, and with or without a thin (1–2 scales long) dingy white irregular nuchal collar; dorsal blotches extending marginally onto ventrals and occasionally fusing on the anterior part of the body; (8) 800 mm SVL in the holotype female; (9) 220 mm TL in the holotype female.
Dipsas latifrontalis is compared to species previously subsumed under D. peruana: D. palmeri, D. peruana, and the herein described D. klebbai. From D. palmeri, it differs in having the first 9–10 dorsal blotches edged with white or cream, vs. the first 19–35 in D. palmeri. The only known adult of D. latifrontalis photographed in life has bronze interspaces (Fig.
Known only from two localities in the Venezuelan Andes and one in the Northern Colombian Andes at elevations between 1000 and 1400 m (Fig.
Neither
All Dipsas latifrontalis depicted in
Higher-level relationships within Dipsadini are still far from being resolved. The monotypic Plesiodipsas perijanensis was not included in our analysis or other recent molecular phylogenies. The species of Dipsas+Sibynomorphus and Sibon included here form monophyletic groups, but this is not the case for the genus Tropidodipsas, for which T. sartorii and T. fasciata + T. fischeri are the successive sister lineages of Dipsas+Sibynomorphus and Sibon (Fig.
Decades ago,
Additionally, many traditional infrageneric groups are either non-monophyletic, or poorly supported and weakly placed. We recognize that this may reflect inadequate sampling of taxa (only 43 of 77 species are included) or characters (only four mtDNA and one nuclear locus were used). From the eight Dipsas species groups recognized by
Dipsas bobridgelyi is most similar in coloration to D. gracilis (Fig.
Although we did not examine MUSM 17589 from Tumbes department, Peru, the description of the coloration and head scales of this specimen provided by
Sibon bevridgelyi and S. nebulatus leucomelas were not recovered as sister taxa in our phylogenetic analyses (Fig.
Unlike the previous examples, the pattern of cladogenesis recovered in our phylogeny for the species of the Dipsas peruana complex (Fig.
A different scenario of speciation can be interpreted from the current distribution (Fig.
We suspect that there are numerous additional species to be described across all genera of Dipsadini. Our results and the results of other recent researchers such as
Conceived and designed the work: AA JMG DSV OTC. Performed the analyses: AA RAP NP. Gathered morphological data: AA KM GA JCSN TJC RAP DSV DFCH PJV MYM OTC. Contributed reagents/materials/analysis tools: RAP JMG DSV NP GRC PJV TJC DFCH. Wrote the paper: AA DSV KM NP GA JCSN RAP DFCH PJV MYM JMG OTC.
This article was greatly improved by comments of Robert Jadin, Sebastian Lotzkat, and one anonymous reviewer. For granting access to their protected forests, we are grateful to Martin Schaefer and David Agro of Fundación Jocotoco, Ana Cristina de la Torre of Pacoche Lodge, Andrés Chiriboga of Tundaloma Lodge, and Renzo Paladines of Naturaleza y Cultura Internacional. Special thanks to Lucas Bustamante, Jose Vieira, Gabriela Morales, Melissa Costales, Frank Pichardo, Sebastián Di Doménico, Jorge Castillo, James Muchmore, Matthijs Hollanders, Paulina Romero, Aaron Pomerantz, Phil Torres, Ernesto Arbeláez, Fausto Siavichay, Diego Piñán, Carlos Morochz, Hannah Som, Carlos Gómez, Carlos Londoño, Valentina Rubio, Darwin Núñez, and Abel Batista for their assistance and companionship in the field. For providing ecological information about Sibon bevridgelyi, we are grateful to Jose Manuel Falcón. For providing live images of Dipsas and Sibon, we are grateful to Jose Vieira, Sebastián Di Doménico, Frank Pichardo, Matthijs Hollanders, Lucas Bustamante, Daniel Quihua, Luis Vera, Alessandro Catennazi, and Juan Carlos Chaparro. For providing images of preserved specimens, we are grateful to Gustavo Pazmiño, Diego Quirola, César Barrio, Micaela Stacey, Luke Welton, Jackson Roberts, and Joseph Martinez. For granting access to specimens under their care, we are grateful to Andreas Schmitz (
GenBank accession numbers for loci and terminals of taxa and outgroups sampled in this study. Novel sequence data produced in this study are marked with an asterisk (*).
Species | Voucher | Country | 12S | 16S | CYTB | ND4 | c-mos |
---|---|---|---|---|---|---|---|
A. iridescens | MZUTI 4178 | Ecuador | - | KT944040 | KY610080 | - | KT944066 |
D. albifrons | MZUSP 13993 | Brazil | JQ598803 | JQ598866 | JQ598925 | - | - |
D. andiana | MZUTI 3501 | Ecuador | - | MH341009* | MH375032* | - | - |
D. andiana | MZUTI 3505 | Ecuador | - | MH341010* | MH374974* | - | - |
D. andiana | MZUTI 5413 | Ecuador | - | MH341011* | MH374978* | - | - |
D. andiana |
|
Ecuador | - | MH341014* | MH375012* | - | - |
D. andiana |
|
Ecuador | - | MH341015* | MH375018* | - | - |
D. andiana |
|
Ecuador | - | MH341012* | MH375005* | - | - |
D. andiana |
|
Ecuador | - | MH341013* | MH375011* | - | - |
D. articulata |
|
Panama | JQ598804 | JQ598867 | - | - | - |
D. bobridgelyi | MZUTI 5414 | Ecuador | - | MH341016* | MH374984* | - | - |
D. bobridgelyi | MZUTI 5417 | Ecuador | - | MH341017* | MH374985* | - | - |
D. bucephala | GRCOLLI 25659 | Brazil | MH341087* | MH341018* | MH375026* | MH375052* | MH374932* |
D. bucephala | IBSP72899 | Brazil | GQ457789 | GQ457730 | - | - | GQ457850 |
D. catesbyi |
|
Peru | - | - | EF078537 | EF078585 | - |
D. catesbyi | LSUMNS 13989 | Brazil | - | KX660267 | KX660536 | - | - |
D. catesbyi | MZUSP 14664 | Brazil | JQ598805 | KX694637 | KX694856 | - | JQ598977 |
D. catesbyi |
|
Ecuador | MH341088* | MH341019* | MH374975* | MH375042* | MH374933* |
D. elegans | DHMECN 10311 | Ecuador | - | MH341020* | MH374979* | - | - |
D. elegans | MZUTI 3317 | Ecuador | - | MH341021* | MH375033* | - | - |
D. elegans | MZUTI 3695 | Ecuador | - | MH341022* | MH375031* | - | - |
D. elegans |
|
Ecuador | - | - | MH374994* | - | - |
D. elegans |
|
Ecuador | - | MH341023* | MH374992* | - | - |
D. ellipsifera | MZUTI 4931 | Ecuador | - | MH341024* | MH375030* | - | MH374934* |
D. ellipsifera | TH | Ecuador | - | - | MH374966* | - | MH374935* |
D. georgejetti |
|
Ecuador | - | MH341025* | MH375024* | - | MH374936* |
D. georgejetti |
|
Ecuador | - | MH341026* | MH375025* | - | MH374937* |
D. georgejetti |
|
Ecuador | - | MH341027* | - | - | - |
D. gracilis | JMG 070 | Ecuador | - | MH341028* | MH374980* | - | MH374938* |
D. gracilis | MZUTI 1386 | Ecuador | - | MH341029* | MH374970* | - | - |
D. gracilis | MZUTI 3331 | Ecuador | - | MH341030* | MH374995* | - | - |
D. gracilis | MZUTI 3503 | Ecuador | - | MH341031* | MH375023* | - | - |
D. gracilis |
|
Ecuador | - | MH341033* | MH375000* | - | - |
D. gracilis |
|
Ecuador | - | MH341034* | MH375001* | - | - |
D. gracilis |
|
Ecuador | - | MH341035* | MH375002* | - | - |
D. gracilis |
|
Ecuador | - | MH341036* | MH375013* | - | - |
D. gracilis |
|
Ecuador | - | - | MH374998* | - | - |
D. gracilis |
|
Ecuador | - | MH341032* | MH374999* | - | - |
D. indica | - | French Guiana | NN | AF158488 | - | - | - |
D. indica ecuadoriensis |
|
Ecuador | MH341089* | MH341037* | MH375006* | MH375043* | MH374939* |
D. indica ecuadoriensis |
|
Ecuador | MH341090* | MH341038* | MH375007* | MH375044* | MH374940* |
D. indica ecuadoriensis |
|
Ecuador | MH341091* | MH341039* | MH375008* | MH375045* | MH374941* |
D. jamespetersi | AMARU 1123 | Ecuador | - | MH341040* | - | - | MH374943* |
D. jamespetersi | AMARU 383 | Ecuador | - | - | - | - | MH374942* |
D. jamespetersi | CAMPO 488 | Ecuador | - | MH341041* | MH375028* | - | MH374944* |
D. jamespetersi |
|
Ecuador | - | MH341042* | MH375014* | - | - |
D. klebbai | JMG 050 | Ecuador | - | MH341043* | MH375022* | - | MH374945* |
D. klebbai | MZUTI 5412 | Ecuador | - | MH341045* | MH374977* | - | - |
D. klebbai | MZUTI 63 | Ecuador | - | MH341044* | MH374986* | - | - |
D. klebbai |
|
Ecuador | - | MH341046* | MH375019* | - | - |
D. klebbai |
|
Ecuador | - | MH341047* | MH374996* | - | - |
D. klebbai |
|
Ecuador | - | MH341048* | - | - | - |
D. klebbai |
|
Ecuador | - | MH341049* | - | - | - |
D. mikanii | MZUSP 14658 | Brazil | GQ457832 | GQ457771 | KX694855 | - | GQ457892 |
D. neuwiedi | MCP13291 | Brazil | GQ457831 | GQ457770 | - | - | GQ457891 |
D. neuwiedi | MZUSP 13972 | Brazil | JQ598838 | JQ598898 | - | - | - |
D. oligozonata |
|
Ecuador | - | MH341050* | MH375029* | - | - |
D. oreas | DHMECN 7647 | Ecuador | - | MH341051* | MH374971* | - | - |
D. oreas | DHMECN 7648 | Ecuador | - | MH341052* | MH374967* | - | - |
D. oreas |
|
Ecuador | - | MH341053* | MH374987* | - | - |
D. oreas | MZUTI 3351 | Ecuador | - | MH341054* | - | MH375038* | - |
D. oreas | MZUTI 5415 | Ecuador | - | MH341055* | - | - | - |
D. oreas | MZUTI 5418 | Ecuador | - | MH341056* | MH374981* | - | - |
D. oreas |
|
Ecuador | - | MH341057* | MH375015* | - | - |
D. oreas |
|
Ecuador | - | MH341058* | MH375016* | - | - |
D. oreas |
|
Ecuador | - | MH341059* | MH375017* | - | - |
D. oswaldobaezi |
|
Ecuador | - | MH341060* | MH374997* | - | - |
D. palmeri | JMG 069 | Ecuador | - | MH341061* | MH374976* | - | MH374946* |
D. palmeri | MZUTI 4804 | Ecuador | - | MH341062* | MH374982* | - | MH374947* |
D. palmeri | MZUTI 4975 | Ecuador | - | MH341063* | - | - | - |
D. palmeri | MZUTI 5419 | Ecuador | - | MH341064* | MH374988* | - | MH374948* |
D. palmeri |
|
Ecuador | MH341092* | MH341065* | MH375009* | MH375046* | MH374949* |
D. palmeri |
|
Ecuador | MH341093* | MH341066* | MH375004* | MH375047* | MH374950* |
D. palmeri |
|
Ecuador | MH341094* | MH341067* | MH375010* | MH375048* | MH374951* |
D. pavonina | LSUMNS 14372 | Brazil | - | KX660268 | KX660537 | - | - |
D. pavonina | MZUTI 4972 | Ecuador | - | MH341068* | MH374983* | - | MH374952* |
D. peruana | LSUMNS 1532 | Peru | - | - | KX660538 | - | KX660406 |
D. pratti | MHUA 14278 | Colombia | - | - | GQ334482 | GQ334583 | - |
D. temporalis |
|
Ecuador | - | MH341069* | MH375003* | - | - |
D. turgida | FML 14969 | Argentina | JQ598839 | JQ598899 | KX660547 | - | - |
D. turgida | LSUMNS 6459 | - | - | KX660279 | - | KX660659 | KX660418 |
D. vaga |
|
Peru | - | KX660252 | - | - | KX660393 |
D. variegata | MZUSP 14665 | Brazil | - | GQ457731 | - | - | GQ457851 |
D. variegata | - | - | AF158406 | AF158476 | - | - | - |
D. ventrimaculata | MCP4870 | Brazil | JQ598840 | JQ598900 | - | - | JQ598997 |
D. vermiculata | MZUTI 3663 | Ecuador | - | MH341070* | MH374989* | - | - |
D. vermiculata |
|
Ecuador | MH341095* | MH341071* | MH374972* | MH375049* | MH374953* |
D. vermiculata |
|
Ecuador | MH341096* | MH341072* | - | MH375040* | MH374954* |
D. vermiculata |
|
Ecuador | - | MH341073* | MH374973* | MH375050* | MH374955* |
D. vermiculata | SBI 171139 | Peru | Z46459 | Z46496 | - | - | - |
D. williamsi |
|
Peru | - | - | MH374968* | MH375041* | - |
D. williamsi |
|
Peru | - | - | MH374969* | MH375039* | - |
G. godmani | - | - | JQ598814 | JQ598877 | JQ598932 | - | - |
S. annulatus | ADM 0007 | Costa Rica | - | KX660170 | KX660444 | KX660573 | KX660309 |
S. annulatus | ADM 242 | Costa Rica | - | KX660169 | KX660443 | KX660572 | KX660308 |
S. annulatus |
|
Nicaragua | MH341097* | MH341074* | MH375034* | MH375053* | MH374956* |
S. annulatus | MZUTI 3034 | Ecuador | - | MH341075* | MH375021* | - | - |
S. anthracops |
|
Costa Rica | MH341098* | MH341076* | MH375035* | MH375054* | MH374957* |
S. bevridgelyi |
|
Ecuador | - | - | MH374990* | - | - |
S. bevridgelyi | MZUTI 3269 | Ecuador | - | MH341077* | MH374962* | - | - |
S. bevridgelyi | MZUTI 5416 | Ecuador | - | MH341078* | MH374963* | - | - |
S. dimidiatus | LSUMNS 6689 | - | - | KX660278 | - | - | KX660417 |
S. dunni | CAMPO 533 | Ecuador | - | MH341079* | MH374991* | - | - |
S. longifrenis |
|
Costa Rica | MH341099* | MH341080* | MH375036* | MH375055* | MH374958* |
S. merendonensis |
|
Guatemala | MH341100* | MH341081* | MH375037* | MH375056* | MH374959* |
S. nebulatus hartwegi | MHUA14511 | Colombia | - | - | GQ334556 | GQ334662 | - |
S. nebulatus leucomelas | DHMECN 9585 | Ecuador | - | MH341082* | - | - | - |
S. nebulatus leucomelas | MZUTI 3911 | Ecuador | - | MH341083* | MH374964* | - | - |
S. nebulatus leucomelas | MZUTI 4810 | Ecuador | - | MH341084* | MH374965* | - | MH374960* |
S. nebulatus nebulatus | Belize | Belize | AF544777 | AF544806 | - | - | AF544736 |
S. nebulatus nebulatus |
|
Costa Rica | EU728583 | EU728583 | EU728583 | EU728583 | - |
T. fasciata | TJC 666 | Mexico | MH341101* | MH341085* | MH375027* | MH375057* | MH374961* |
T. fischeri |
|
Guatemala | MH341102* | MH341086* | MH374993* | MH375051* | - |
T. sartorii |
|
El Salvador | - | - | EF078540 | EF078588 | - |
List of PCR and sequencing primers and their respective PCR conditions (denaturation, annealing, extension and number of corresponding cycles) used in this study. All PCR protocols included an initial 3-min step at 94 °C and a final extension of 10 min at 72 °C.
Locus | Primer name | Sequence (5’-3’) | Reference | PCR profile: |
---|---|---|---|---|
16S | 16Sar-L | CGCCTGTTTATCAAAAACAT |
|
30 cycles of 94 °C (45 sec), 53 °C (45 sec), 72 °C (1 min) |
16Sbr-H-R | CCGGTCTGAACTCAGATCACGT | |||
Cytb | GLUDG-L | TGACTTGAARAACCAYCGTTG |
|
35–42 cycles of 95°C (30 sec) , 50 or 56 °C (45 sec), 72 °C (45 sec) |
ATRCB3 | TGAGAAGTTTTCYGGGTCRTT |
|
||
ND4 | ND4 | CACCTATGACTACCAAAAGCTCATGTAGAAGC |
|
94 °C (25 sec), 56 or 60 °C (1 min), 72 °C (2 min) [x25–30] |
Leu | CATTACTTTTACTTGGATTTGCACCA | |||
c-mos | S77 | CATGGACTGGGATCAGTTATG |
|
1 cycle of 94 °C (3 min), 56 °C (45 sec), 72 °C (1 min), followed by 34 cycles of 94 °C (45 sec), 56 °C (45 sec), 72 °C (1 min) |
S78 | CCTTGGGTGTGATTTTCTCACCT |
Morphological data and sex for specimens of Dipsadini species examined. Codes: V = ventrals; SC = subcaudals; D1–3 = dorsal scale rows at neck, midbody, and vent; PO = postoculars; SL = supralabials; IL = infralabials; SVL = snout-vent length (mm); TL = tail length (mm); M = Male, F = Female.
Species | Voucher | V | SC | D1 | D2 | D3 | PO | SL | IL | SVL | TL | Sex |
---|---|---|---|---|---|---|---|---|---|---|---|---|
D. andiana |
|
187 | 96 | 15 | 15 | 15 | 3 | 9 | 11 | 744 | 196 | M |
D. andiana |
|
194 | 85 | 15 | 15 | 15 | 2 | 9 | 12 | 292 | 71 | F |
D. andiana | MZUTI 5413 | 190 | 101 | 14 | 15 | 15 | 2 | 10 | 11 | 471 | 165 | M |
D. andiana | MZUTI 3501 | 187 | 98 | 15 | 15 | 15 | 2 | 9 | 12 | 398 | 137 | M |
D. andiana | MZUTI 3505 | 192 | – | 15 | 15 | 15 | 2 | 8 | 10 | 674 | 167 | F |
D. andiana |
|
189 | 84 | 15 | 15 | 15 | 2 | 10 | 10 | 680 | 150 | F |
D. andiana |
|
186 | 90 | 15 | 15 | 15 | 2 | 10 | 10 | 453 | 149 | M |
D. andiana |
|
189 | 101 | 15 | 15 | 15 | 2 | 9 | 9 | 405 | 139 | M |
D. bobridgelyi |
|
201 | 117 | 15 | 15 | 15 | 2 | 9 | 12 | 445 | 212 | M |
D. bobridgelyi | DHMECN 11527 | 178 | 98 | 15 | 15 | 15 | 2 | 9 | 12 | 404 | 158 | F |
D. bobridgelyi | MZUTI 3266 | 184 | 96 | 15 | 15 | 15 | 2 | 9 | 11 | 286 | 117 | F |
D. bobridgelyi | MZUTI 5414 | 180 | 95 | 15 | 15 | 15 | 2 | 9 | 13 | 478 | 195 | M |
D. bobridgelyi | MZUTI 5417 | 182 | 101 | 15 | 15 | 15 | 2 | 9 | 13 | 372 | 158 | M |
D. catesbyi |
|
180 | 98 | 13 | 13 | 13 | 1 | 8 | 9 | 366 | 147 | F |
D. catesbyi |
|
176 | 94 | 13 | 13 | 13 | 2 | 9 | 10 | 420 | 155 | F |
D. catesbyi |
|
168 | 81 | 13 | 13 | 13 | 1 | 7 | 8 | 276 | 98 | F |
D. catesbyi | DHMECN 11555 | 164 | 97 | – | – | – | 2 | 7 | – | 222 | 80 | – |
D. catesbyi |
|
172 | 93 | 13 | 13 | 13 | 2 | 9 | 10 | 470 | 169 | F |
D. catesbyi |
|
175 | 83 | 13 | 13 | 13 | 1 | 8 | 10 | 505 | 180 | F |
D. catesbyi |
|
199 | 97 | 13 | 13 | 13 | 2 | 8 | 9 | 441 | 165 | F |
D. catesbyi |
|
183 | 108 | 13 | 13 | 13 | 1 | 8 | 9 | 480 | 202 | M |
D. catesbyi |
|
184 | 98 | 13 | 13 | 13 | 1 | 8 | 10 | 308 | 117 | F |