Research Article |
Corresponding author: Alejandro Arteaga ( af.arteaga.navarro@gmail.com ) Academic editor: Robert Jadin
© 2023 Alejandro Arteaga, Abel Batista.
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, Batista A (2023) A consolidated phylogeny of snail-eating snakes (Serpentes, Dipsadini), with the description of five new species from Colombia, Ecuador, and Panama. ZooKeys 1143: 1-49. https://doi.org/10.3897/zookeys.1143.93601
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A molecular phylogeny of the Neotropical snail-eating snakes (tribe Dipsadini Bonaparte, 1838) is presented that includes 60 of the 133 species currently recognized. There is morphological and phylogenetic support for four new species of Sibon Fitzinger, 1826 and one of Dipsas Laurenti, 1768, which are described here based on their unique combination of molecular, meristic, and color pattern characteristics. Plesiodipsas
Caenophidia, Colubroidea, Dipsas, Plesiodipsas, Sibon, Squamata, systematics, taxonomy
The snail-eating snake tribe Dipsadini is one of the most diverse yet taxonomically complex group of snakes in the Neotropics. Many authors (
In his unpublished PhD thesis,
First,
Here, we combine the datasets of
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
Our terminology for Dipsadini cephalic shields follows proposals by
Tissue samples from 19 individuals representing eight species (including the five new species described here) were obtained in Colombia, Ecuador, and Panama. All specimens included in the genetic analyses were morphologically identified according to
Genomic DNA was extracted from 96% ethanol-preserved tissue samples (liver, muscle tissue, or scales) using either a guanidinium isothiocyanate extraction protocol (
A total of 343 DNA sequences was used to build a phylogenetic tree of the tribe Dipsadini, of which 35 were generated during this work and 308 were downloaded from GenBank, most of which were produced by
Phylogenetic relationships were assessed under a Bayesian inference (BI) approach in MrBayes 3.2.0 (
We present ranges of occurrence for eleven species of Dipsadini, including five new species described here. Presence localities are derived from museum vouchers (Suppl. materal 1), photographic records (iNaturalist), and the literature (all summarized in Suppl. materal 2). For each species, a binary environmental niche model (ENM) accompanies the dot maps. These models estimate potential areas of distribution on the basis of observed presences and a set of environmental predictors (
For the first explorative exercise, we used the 19 climate layers from the WorldClim project and assessed which variables were the most important for the model, according to the Jackknife test calculated in MaxEnt (
Selected partitions and models of evolution are presented in Table
Phylogenetic relationships within Dipsadini inferred using a Bayesian inference and derived from analysis of DNA gene fragments 12S, 16S, COI, CYTB, ND4, DNAH3, and NT3. Support values on intra-specific branches are not shown for clarity. Voucher numbers for sequences are indicated for each terminal. 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). Colored clades correspond to the species’ distribution presented in the maps. New or redefined species are indicated in bold type.
Partition scheme and models of evolution used in phylogenetic analyses. Numbers in parentheses indicate codon position.
Partitition | Best model | Gene regions | Number of aligned sites |
---|---|---|---|
1 | GTR+I+G | 12S, 16S, COI(1), CYTB(3), ND4(1) | 1742 |
2 | HKY+I+G | CYTB(1), ND4(2) | 614 |
3 | GTR+I+G | COI(3), CYTB(2), ND4(3) | 833 |
4 | K80+I | DNAH3(1), DNAH3(2) | 450 |
5 | K80+G | DNAH3(3), NT3(1) | 381 |
6 | K80+G | NT3(2), NT3(3) | 313 |
7 | F81 | COI(2) | 219 |
Tropidodipsas fischeri Boulenger, 1894 is recovered as sister to all other sampled Dipsadini, with the exception of Geophis sanniolus, a species that did not form a group with the remaining Geophis. Neither of these or any other higher relationships within Dipsadini are strongly supported in our analysis, but all other sampled species of Dipsadini are included in their corresponding genera sensu
With the exception of Geophis sanniolus, relationships within Geophis are identical to those presented in
Dipsas gaigeae is recovered as the sister species of all other Dipsas, albeit with low support, but not as sister to (see
Relationships within Sibon are most similar to those presented in
Finally, we excluded Sibon noalamina
We name and provide 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
Holotype
: MHCH 3143 (Figs
Photographs of some species of Sibon in life a S. irmelindicaprioae sp. nov. MHCH 3269 from Chucantí Reserve, Darién province, Panama b S. irmelindicaprioae sp. nov. from Morromico Reserve, Chocó department, Colombia c S. canopy sp. nov. from Cerro Gaital, Coclé province, Panama d, e S. annulatus from Centro Manu, Limón province, Costa Rica f S. ayerbeorum
Photographs of some species of Sibon and Dipsas in life a S. canopy sp. nov. from El Valle de Antón, Coclé province, Panama b S. irmelindicaprioae sp. nov. holotype MHCH 3143 from Puerto Indio, Darién province, Panama c S. marleyae sp. nov. from Verdecanandé, Esmeraldas Province, Ecuador d S. vieirai sp. nov. from Mashpi Amagusa Reserve, Pichincha province, Ecuador e Dipsas sp. from Cerro Gaital, Coclé province, Panama f D. welborni sp. nov.
Differences in head morphology between species of Sibon previously subsumed under S. annulatus a S. annulatus MHCH 1982 from Bonyic, Bocas del Toro province, Panama b S. canopy sp. nov. holotype MHCH 3110 from Cerro Gaital, Coclé province, Panama c S. irmelindicaprioae sp. nov. MHCH 3120 from Pirré, Darién province, Panama d S. marleyae sp. nov. holotype
Paratypes : MHCH 3145, adult female collected by Abel Batista and Milan Vesely on 26 September 2012 at Ambroya, Panama province, Panama (8.91680, -78.61779; 484 m a.s.l.). MHCH 3146, adult male collected by Abel Batista on 16 November 2012 at Cerro Garra Garra, Pavarandó, Comarca Emberá-Wounaan, Panama (7.76400, -78.10063; 655 m a.s.l.). MHCH 3111, adult male collected by Abel Batista, Madian Miranda, Orlando Garcés, Rogemif Fuentes on 15 October 2016 at Chucantí, Darién province, Panama (8.79773, -78.46225; 1295 m a.s.l.). MHCH 3120, adult male collected by Abel Batista, Madian Miranda, Michelle Quiroz, Marcos Ponce on 18 June 2015 at Pirré, Darién province, Panama (7.99695, -77.71040; 550 m a.s.l.). COLZOOCH-H 0792, adult male collected by Jhon Tailor Rengifo Mosquera on 6 March 2005 at El Afirmado, Chocó department, Colombia (5.64190, -77.07550; 216 m a.s.l.).
Sibon irmelindicaprioae sp. nov. is placed in the genus Sibon based on phylogenetic evidence (Fig.
Sibon irmelindicaprioae sp. nov. is compared to other species of Sibon previously subsumed under S. annulatus sensu lato (differences summarized in Table
Differences in coloration, scale counts, and size between Sibon annulatus, S. canopy sp. nov., S. irmelindicaprioae sp. nov., and S. marleyae sp. nov. The range of each continuous variable is from our own sample,
Variable | Sibon annulatus | Sibon canopy sp. nov. | Sibon irmelindicaprioae sp. nov. | Sibon marleyae sp. nov. | ||||
---|---|---|---|---|---|---|---|---|
Dorsum pattern | Reddish bands distinct and extending over the entire dorsal and lateral surfaces | Reddish bands distinct and broken along vertebral line in about half of individuals | Reddish bands faint and broken along vertebral line | Reddish bands distinct and usually broken along vertebral line | ||||
Reddish vertebral spots in interspaces | No | Yes | No | No | ||||
Head pattern | Symmetrical broad blotches | Irregular broad blotches | Finely variegated | Irregular/symmetrical broad blotches | ||||
Supralabials | 7–8 | 6–8 | 7–9 | 7–8 | ||||
Infralabials | 7–9 | 6–8 | 8–10 | 8–9 | ||||
Postmentals | 2 | 1 | 2 | 2 (1 in |
||||
Sex | Males (n = 10) | Females (n = 15) | Males (n = 12) | Females (n = 8) | Males (n = 7) | Females (n = 1) | Males (n = 6) | Females (n = 5) |
Maximum TOL | 707 mm | 611 mm | 648 mm | 536 mm | 580 mm | 606 mm | 657 mm | 551 mm |
Ventrals | 170–192 | 161–186 | 180–189 | 170–185 | 187–196 | 174 | 186–204 | 176–193 |
Subcaudals | 108–135 | 113–126 | 113–130 | 107–124 | 110–128 | 117 | 130–143 | 109–128 |
Adult male, SVL 387 mm, tail length 193 mm (49% SVL); head length 14.3 mm (3.7% SVL) from tip of snout to angle of jaw; head width 9.0 mm (88% head length) taken at broadest point; snout-orbit distance 2.3 mm; head distinct from neck; snout short, blunt in dorsal outline and rounded in profile; rostral 1.8 mm wide, higher than broad; internasals 1.8 mm wide, broader than long; prefrontals 2.3 mm wide, longer than broad, entering orbit; supraocular 3.6 mm long, longer than broad; frontal 3.7 mm long, pentagonal and with an inward-bent anterior border, in contact with prefrontals, supraoculars, and parietals; parietals 5.8 mm long, longer than broad; nasal divided, in contact with first two supralabials, loreal, prefrontal, internasal, and rostral; loreal 1.4 mm long, longer than high, entering the orbit; eye diameter 3.7 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, eight supralabials with 5th and 6th contacting orbit on the left side; symphysial precluded from contacting chinshields by a pair of postmentals; ten infralabials, 3rd–7th contacting chinshields; two pair of chinshields longer than wide; dorsal scales in 15/15/15 rows, smooth, without apical pits; 193 ventrals; 128 paired subcaudals; cloacal plate single.
Specimens of Sibon irmelindicaprioae sp. nov. have been found at night foraging on shrubs, trees, and palm fronds 200–300 cm above the ground in old-growth to moderately disturbed evergreen lowland/foothill forests. Snakes of this species are docile and never attempt to bite. When threatened, individuals may hide the head among body coils and produce a musky and distasteful odor.
Sibon irmelindicaprioae sp. nov. is known from 16 localities (listed in Suppl. material
The specific epithet irmelindicaprioae is a patronym honoring Irmelin DiCaprio (1945–present), mother of Leonardo DiCaprio, long-time advocate and supporter of biodiversity conservation around the world.
We consider Sibon irmelindicaprioae sp. nov. to be included in the Near Threatened category following IUCN Red List criteria (
Holotype
: MHCH 3110 (Figs
Paratypes
: MHCH 1067,
Sibon canopy sp. nov. is placed in the genus Sibon based on phylogenetic evidence (Fig.
Sibon canopy sp. nov. is compared to other species of Sibon previously subsumed under S. annulatus sensu lato (differences summarized in Table
Adult female, SVL 321 mm, tail length 157 mm (48% SVL); head length 15.4 mm (4.7% SVL) from tip of snout to commissure of mouth; head width 8.0 mm (76% head length) taken at broadest point; snout-orbit distance 3.3 mm; head distinct from neck; snout short, blunt in dorsal outline and rounded in profile; rostral 2.1 mm wide, higher than broad; internasals 1.6 mm wide, broader than long; prefrontals 1.9 mm wide, longer than broad, entering orbit; supraocular 3.7 mm long, longer than broad; frontal 3.2 mm long, pentagonal and with a straight anterior border, in contact with prefrontals, supraoculars, and parietals; parietals 5.2 mm long, longer than broad; nasal divided, in contact with first three supralabials, loreal, prefrontal, internasal, and rostral; loreal 1.7 mm long, longer than high, entering the orbit; eye diameter 3.0 mm; pupil semi-elliptical; no preocular; two postoculars; temporals 1+2; 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 in contact with chinshields; nine infralabials with 2nd–5th contacting chinshields; two pair of chinshields longer than wide; dorsal scales in 15/15/15 rows, smooth, without apical pits; 172 ventrals; 93+ divided subcaudals; cloacal plate entire.
Sibon canopy sp. nov. is known from 25 localities (listed in Suppl. material
The specific epithet canopy is used as a noun in apposition and honors the Canopy Family system of reserves, particularly its Canopy Lodge in Valle de Antón, Coclé province, Panama, where the new species occurs. Though best known for its world-class eco-tourism focused on birds, the Canopy Family also protects habitat that is critical for dozens of poorly studied Panamanian snakes such as S. canopy sp. nov. and S. irmelindicaprioae sp. nov. The project was founded in 1994 by Raúl Arias de Para and Denise Barakat de Arias, two champions of Panamanian conservation who are deeply intertwined with the Political history of the country. In 2019, the Canopy Family invited us to explore their system of reserves in order to discover their herpetofauna. As a result of this invitation, both S. canopy sp. nov. and a new species of Dipsas were discovered.
We consider Sibon canopy sp. nov. to be included in the Near Threatened category following IUCN Red List criteria (
Holotype
:
Paratypes
: MZUTI 3034, adult male collected by Jaime Culebras on 22 July 2013 at Reserva Itapoa, Esmeraldas Province, Ecuador (0.51307, -79.13401; 321 m a.s.l.). ICN 10834, adult male collected at San José del Palmar, Chocó department, Colombia (4.96841, -76.22751; 1338 m a.s.l.).
Sibon marleyae sp. nov. is placed in the genus Sibon based on phylogenetic evidence (Fig.
Sibon marleyae sp. nov. is compared to other species of Sibon previously subsumed under S. annulatus sensu lato (differences summarized in Table
Adult male, SVL 335 mm, tail length 167 mm (49.8% SVL); head length 12.8 mm (3.8% SVL) from tip of snout to angle of jaw; head width 7.6 mm (59% head length) taken at broadest point; snout-orbit distance 3.1 mm; head distinct from neck; snout short, blunt in dorsal outline and rounded in profile; rostral 2.1 mm wide, higher than broad; internasals 1.3 mm wide, broader than long; prefrontals 1.6 mm wide, longer than broad, entering orbit; supraocular 2.8 mm long, longer than broad; frontal 2.9 mm long, pentagonal and with a straight anterior border, in contact with prefrontals, supraoculars, and parietals; parietals 4.3 mm long, longer than broad; nasal divided, in contact with two supralabials, loreal, prefrontal, internasal, and rostral; loreal 1.2 mm long, longer than high, entering the orbit; eye diameter 2.9 mm; pupil semi-elliptical; no preocular; two postoculars; temporals 1+2; seven supralabials with 4th and 5th contacting orbit; symphysial precluded from contacting chinshields by the presence of two small postmentals; eight infralabials with 2nd–6th contacting chinshields on the right side, nine infralabials with 2nd–7th contacting chinshields on the left side; two pairs of chinshields longer than wide; dorsal scales in 15/15/15 rows, smooth, without apical pits; 204 ventrals; 132 divided subcaudals; cloacal plate entire.
Specimens of Sibon marleyae sp. nov. have been found at night foraging on shrubs and trees 1–6 m above the ground in old-growth evergreen lowland/foothill forests, particularly along streams and small rivers. Snakes of this species are docile and never attempt to bite. When threatened, individuals may hide the head among body coils and produce a musky and distasteful odor. One female (Fig.
Sibon marleyae sp. nov. is known from 17 localities (listed in Suppl. material
The specific epithet marleyae is a patronym honoring a young nature lover, Marley Sheth, the 11-year old daughter of Brian and Adria Sheth, both long-time supporters of biodiversity conservation around the world.
We consider Sibon marleyae sp. nov. to be included in the Least Concern category following IUCN Red List criteria (
Holotype
:
Paratypes
:
Sibon vieirai sp. nov. is placed in the genus Sibon based on phylogenetic evidence (Fig.
Photographs of species of the Sibon nebulatus leucomelas complex in life a S. leucomelas from Morromico Reserve, Chocó department, Colombia b S. vieirai sp. nov.
Differences in throat color pattern between species of the Sibon nebulatus leucomelas complex a S. vieirai sp. nov.
Sibon vieirai sp. nov. is most similar to S. leucomelas, from which it differs primarily on the basis of coloration (differences summarized under Table
Differences in coloration, scale counts, and size between Sibon leucomelas and S. vieirai sp. nov. The range of each continuous variable is from our own sample,
Variable | Sibon leucomelas | Sibon vieirai sp. nov. | ||
---|---|---|---|---|
White dorsal bands | Distinct, 1–2 scales wide | Incomplete, broken into dots | ||
Complete black bands extending over the entire dorsal and lateral surfaces | Present, distinct | Usually absent; if present, indistinct and broken | ||
Color of pale dorsal markings | Rosy white | White | ||
Throat pattern | Entirely black with fine white speckling | Checkerboard, with large black and white markings | ||
Sex | Males (n = 5) | Females (n = 7) | Males (n = 8) | Females (n = 5) |
Maximum TOL | 809 mm | 700 mm | 732 mm | 714 mm |
Ventral scales | 190–198 | 187–194 | 183–195 | 178–189 |
Subcaudal scales | 86–101 | 84–100 | 95–105 | 78–92 |
Adult male, SVL 515 mm, tail length 199 mm (38.6% SVL); head length 20.7 mm (4.0% SVL) from tip of snout to angle of jaw; head width 11.6 mm (55% head length) taken at broadest point; snout-orbit distance 4.9 mm; head distinct from neck; snout short, blunt in dorsal outline and rounded in profile; rostral 3.8 mm wide, higher than broad; internasals 2.1 mm wide, broader than long; prefrontals 3.4 mm wide, slightly broader than long, entering orbit; supraocular 3.6 mm long, longer than broad; frontal 4.3 mm long, with a rounded triangular shape, in contact with prefrontals, supraoculars, and parietals; parietals 6.4 mm long, longer than broad; nasal divided, in contact with two supralabials, loreal, prefrontal, internasal, and rostral; loreal 2.3 mm long, longer than high, entering the orbit; eye diameter 3.9 mm; pupil semi-elliptical; no preocular; two postoculars; temporals 1+2; seven supralabials with 4th and 5th contacting orbit; symphysial precluded from contacting chinshields by first pair of infralabials; nine infralabials with 1st–6th contacting chinshields; two pairs of chinshields longer than wide; dorsal scales in 15/15/15 rows, smooth, without apical pits; 195 ventrals; 105 divided subcaudals; cloacal plate entire.
Specimens of Sibon vieirai sp. nov. have been found in old growth to heavily disturbed evergreen lowland/foothill forests as well as in rural gardens and plantations. Active snakes have been seen at night foraging at ground level or on vegetation up to 3 m above the ground. One snake was spotted as it emerged from under a pile of logs at sunset. Based on our own field experience, individuals appear to be more active when it is raining or drizzling. In the field in Ecuador, specimens of S. vieirai sp. nov. have been observed feeding on slugs and snails. A female from Hostería Selva Virgen, Pichincha province, Ecuador laid a clutch of four eggs.
Sibon vieirai sp. nov. is known from at least 95 localities (listed in Suppl. material
The specific epithet vieirai is a patronym honoring Jose Vieira, a Venezuelan biologist and wildlife photographer who created the Ex-Situ project, a free-access photo bank depicting Latin American fauna on a white background. Jose Vieira’s photos have been crucial in illustrating field guides about herpetofauna, educational posters, and research publications. Most of the images in this work were created by Jose Vieira. Additionally, after nearly six years of active collaboration with one of us (AA), it has become evident that Jose is one of the most tireless and focused young field biologists ever to sample the jungles of the tropics, a work ethic that has resulted in the generation of photo and museum vouchers for hundreds of poorly studied species of herpetofauna, including the holotype of this new Sibon.
We consider Sibon vieirai sp. nov. to be included in the Least Concern category following IUCN Red List criteria (
Holotype
: MZUTI 3663 (Figs
Paratypes
:
Dipsas welborni sp. nov. is placed in the genus Dipsas based on phylogenetic evidence (Fig.
Dipsas welborni sp. nov. differs from the majority of its congeners by having dorsal scales arranged in 13/13/13 rows, loreal entering the orbit, and dorsum of head strongly vermiculated. The new species is most similar to D. vermiculata, from which it differs on the basis of the following characters of coloration and lepidosis (Fig.
Photographs of species of Dipsas previously subsumed under D. vermiculata a D. welborni sp. nov. paratype
Differences in coloration, scale counts, and size between Dipsas vermiculata and D. welborni sp. nov. The range of each continuous variable is from our own sample,
Variable | Dipsas vermiculata | Dipsas welborni sp. nov. | ||
---|---|---|---|---|
Background color of ventral surfaces | Always white | Usually yellow, occasionally pale yellowish white | ||
Prefrontals fused | No (partially fused in |
Yes | ||
Rows of spines on hemipenis body (asulcate surface) | 1–2 rows of curved spines | 2 rows of straight spines followed by 2–3 rows of curved spines | ||
Sex | Males (n = 8) | Females (n = 3) | Males (n = 7) | Females (n = 3) |
Maximum SVL | 515 mm | 501 mm | 542 mm | 595 mm |
Maximum TOL | 735 mm | 701 mm | 689 mm | 876 mm |
Ventral scales | 181–192 | 173–174 | 181–193 | 177–179 |
Subcaudal scales | 103–113 | 99–103 | 107–116 | 105–106 |
Adult male, SVL 542 mm, tail length 195 mm (incomplete); head length 16.4 mm (3.0% SVL) from tip of snout to angle of jaw; head width 10.2 mm (62% head length) taken at broadest point; snout-orbit distance 3.8 mm; head distinct from neck; snout short, blunt in dorsal outline and rounded in profile; rostral 2.4 mm wide, higher than broad; internasals 1.8 mm wide, broader than long; prefrontals 2.7 mm wide, longer than broad, not entering orbit; supraocular 3.8 mm long, longer than broad; frontal 4.1 mm long, hexagonal and with angled anterior border, in contact with prefrontals, supraoculars, and parietals; parietals 5.5 mm long, longer than broad; nasal divided, in contact with two supralabials, loreal, prefrontal, internasal, and rostral; loreal 2.1 mm long, longer than high, entering the orbit; eye diameter 3.4 mm; pupil semi-elliptical; one small preocular above loreal; two postoculars; temporals 2+2; seven supralabials with 4th–5th contacting orbit; symphysial precluded from contacting chinshields by first pair of infralabials; nine infralabials with 1st to 5th contacting chinshields; three pairs of chinshields, first longer than wide; dorsal scales in 13/13/13 rows, smooth, without apical pits; 185 ventrals; 80+ divided subcaudals; cloacal plate entire.
Specimens of Dipsas welborni sp. nov. have been found foraging on vegetation 20–350 cm above the ground in old-growth to moderately disturbed evergreen montane forests. Snakes of this species are docile and never attempt to bite. When threatened, individuals may flatten their body and expand their head to simulate a triangular shape as well as produce a musky and distasteful odor.
Dipsas welborni sp. nov. is known from 26 localities (listed in Suppl. material
The specific epithet welborni is a patronym honoring David Welborn, a lifelong champion of ecosystem and species conservation who supports and serves on several nonprofit boards dedicated to the environment. David retired from the board of Nature and Culture International in 2021 after 18 years of service, including four as board chairman. Nature and Culture International, a non-profit organization, has conserved more than 9 million hectares of tropical Latin American ecosystems, including key habitat in the Maycu Reserve of southeastern Ecuador, where Dipsas welborni sp. nov. was discovered.
We consider Dipsas welborni sp. nov. to be in the Near Threatened category following IUCN Red List criteria (
We expand the distribution of Sibon ayerbeorum, a species previously known only from departments Cauca (
This article was greatly improved by comments of Sebastian Lotzkat, Julie M. Ray, and Robert Jadin. For granting access to the protected forests under their care, we are grateful to Daniel Arias and Raúl Arias of the Canopy Family lodges, to Martin Schaefer and David Agro of Fundación Jocotoco, and to Matthew Clark, Renzo Paladines, and Felipe Serrano of Nature and Culture International. For providing DNA sequence data of Sibon and the enigmatic Dipsas perijanensis, we are grateful to Elson Meneses-Pelayo (UIS). For granting access to specimens under their care, we are grateful to David Salazar-Valenzuela (MZUTI), Gustavo Adolfo Londoño Guerrero (CZI), Wilmar Bolívar (
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 | 12S | 16S | COI | CYTB | ND4 | DNAH3 | NT3 |
---|---|---|---|---|---|---|---|---|
Atractus ukupacha |
|
– | MH790540 | – | MN887689 | MN887714 | – | – |
Dipsas albifrons | MZUSP 13993 | JQ598803 | JQ598866 | – | JQ598925 | – | – | – |
Dipsas andiana | MZUTI 3505 | – | MH341010 | – | MH374974 | – | – | – |
Dipsas articulata | USNM 348490 | JQ598804 | MH140680 | MH140122 | – | – | – | – |
Dipsas bobridgelyi | MZUTI 5414 | – | MH341016 | – | MH374984 | – | – | – |
Dipsas bucephala | GRCOLLI 25659 | MH341087 | MH341018 | – | MH375026 | MH375052 | – | – |
Dipsas catesbyi |
|
MH341088 | MH341019 | – | MH374975 | MH375042 | – | – |
Dipsas elegans | MZUTI 3317 | – | MH341021 | – | MH375033 | – | – | – |
Dipsas ellipsifera | MZUTI 4931 | – | MH341024 | – | MH375030 | – | – | – |
Dipsas gaigeae | JAC 28587 | – | – | – | JX398613 | JX398462 | JX293850 | JX398735 |
Dipsas georgejetti |
|
– | MH341025 | – | MH375024 | – | – | – |
Dipsas gracilis | MZUTI 3331 | – | MH341030 | – | MH374995 | – | – | – |
Dipsas indica |
|
MH341089 | MH341037 | – | MH375006 | MH375043 | – | – |
Dipsas jamespetersi |
|
– | MH341042 | – | MH375014 | – | – | – |
Dipsas klebbai | MZUTI 5412 | – | MH341045 | – | MH374977 | – | – | – |
Dipsas sp. | CH 8479 | – | MH140683 | – | – | – | – | – |
JM 663 | – | MH140698 | – | JX398625 | JX398475 | – | – | |
JM 664 | – | MH140697 | – | JX398626 | JX398476 | – | – | |
JM 758 | – | – | – | JX398627 | JX398477 | – | JX398752 | |
JM 795 | – | MH140692 | – | JX398628 | JX398478 | – | JX398753 | |
|
– | OP879850* | – | – | – | – | – | |
Dipsas mikanii | MZUSP 14658 | GQ457832 | GQ457771 | – | KX694855 | – | – | – |
Dipsas neuwiedi | MZUSP 13972 | JQ598838 | JQ598898 | – | – | – | – | – |
Dipsas nicholsi | JM 812 | – | – | – | JX398619 | JX398469 | – | – |
Dipsas oligozonata |
|
– | MH341050 | – | MH375029 | – | – | – |
Dipsas oreas | MZUTI 5418 | – | MH341054 | – | MH374981 | – | – | – |
Dipsas pakaraima | USNM 561837 | – | – | MH273777 | – | – | – | – |
Dipsas palmeri |
|
MH341092 | MH341065 | – | MH375009 | MH375046 | – | – |
Dipsas pavonina | MZUTI 4972 | – | MH341068 | – | MH374983 | – | – | – |
Dipsas perijanensis | UIS R-4180 | – | – | – | – | OP897299* | – | – |
Dipsas peruana | LSUMZ 1532 | – | – | – | JX398622 | JX398472 | JX293856 | JX398750 |
Dipsas pratti | MBUCV 6837 | – | – | – | JX398624 | JX398473 | – | JX398751 |
Dipsas temporalis | MHCH 2878 | – | OP879851* | OP873116* | – | – | – | – |
MHUA 14278 | – | – | – | GQ334482 | GQ334583 | – | GQ334667 | |
|
– | MH341069 | – | MH375003 | – | – | – | |
|
– | – | OP873117* | – | – | – | – | |
Dipsas trinitatis | UWIZM 2011.20.25 | – | – | – | JX398629 | JX398479 | – | – |
Dipsas turgida | LSUMZ 6458 | JQ598839 | KX660279 | – | JX398696 | JX398556 | JX293899 | JX398819 |
Dipsas vagus | KU 219121 | – | KX660252 | – | – | – | – | – |
Dipsas variegata | UTA R-15772 | – | – | – | JX398601 | JX398482 | JX293858 | JX398736 |
Dipsas ventrimaculata | MCP 4870 | JQ598840 | JQ598900 | – | – | – | – | – |
Dipsas vermiculata | MZUTI 4738 | OP839489* | OP879846* | – | – | OP897291* | – | – |
|
MH341095 | MH341071 | – | MH374972 | MH375049 | – | – | |
|
MH341096 | MH341072 | – | – | MH375040 | – | – | |
|
OP839490* | OP879847* | – | OP897289* | OP897293* | – | – | |
|
OP839491* | OP879848* | – | OP897290* | OP897292* | – | – | |
Dipsas viguieri | MHCH 2875 | – | OP879852* | OP873118* | – | – | – | – |
Dipsas welborni sp. nov. | MZUTI 3663 | – | MH341070 | – | MH374989 | OP897294* | – | – |
|
– | MH341073 | – | MH374973 | MH375050 | – | – | |
UTA R-55939 | – | – | – | JX398632 | JX398483 | JX293859 | JX398754 | |
Dipsas williamsi | CORBIDI 12695 | – | – | – | MH374968 | MH375041 | – | – |
Geophis annuliferus | JAC 27792 | – | – | – | JX398699 | JX398559 | JX293914 | – |
Geophis bicolor | MX29-53 | – | – | – | JX398637 | JX398487 | JX293862 | JX398759 |
Geophis nigrocinctus | JAC 30704 | – | – | – | JX398638 | JX398488 | – | – |
Geophis omiltemanus | ENS 11496 | – | – | – | JX398639 | – | – | JX398760 |
Geophis sanniolus | MX21-36 | – | – | – | JX398692 | JX398553 | JX293895 | JX398815 |
Geophis sartorii | USNM 564144 | – | – | – | JX398717 | JX398585 | JX293912 | JX398831 |
Geophis tarascae | MX28-19 | – | – | – | JX398640 | JX398489 | JX293870 | JX398761 |
Sibon aff. hartwegi | ICN 11463 | – | – | – | – | JX398532 | – | – |
ICN 11510 | – | – | – | – | JX398533 | – | JX398803 | |
Sibon aff. nebulatus | CH 6614 | – | – | MH140390 | – | – | – | – |
UTA R-42429 | – | – | – | – | JX398534 | JX293887 | – | |
JAC 28055 | – | – | – | JX398678 | JX398535 | JX293889 | – | |
JAC 28140 | – | – | – | – | JX398536 | JX293893 | – | |
JAC 28589 | – | – | – | – | JX398537 | JX293894 | – | |
JAC 30102 | – | – | – | – | JX398538 | – | – | |
MVZ 233298 | EU728583 | EU728583 | EU728583 | EU728583 | EU728583 | FJ455221 | FJ455189 | |
N068 | – | – | – | JX398682 | JX398542 | – | JX398807 | |
USNM 564142 | – | – | – | – | JX398547 | – | JX398810 | |
USNM 564143 | – | – | – | – | JX398548 | – | JX398811 | |
UTA R-42431 | – | – | – | JX398690 | JX398549 | JX293891 | JX398812 | |
Sibon annulatus | ADM 242 | – | KX660169 | – | KX660443 | KX660572 | – | KX651996 |
ADM0007 | – | KX660170 | – | KX660444 | KX660573 | – | KX651997 | |
B45-57 | – | – | – | – | JX398499 | – | JX398770 | |
D167 | – | – | – | JX398652 | JX398501 | JX293869 | JX398772 | |
MVZ 269290 | MH341097 | MH341074 | – | MH375034 | MH375053 | – | – | |
N740 | – | – | – | – | JX398505 | – | JX398777 | |
Sibon anthracops | MVZ 215680 | MH341098 | MH341076 | – | MH375035 | MH375054 | – | – |
Sibon argus | USNM 579852 | – | MH140960 | MH140380 | JX398662 | JX398511 | – | JX398783 |
Sibon ayerbeorum |
|
OP839492* | OP879849* | – | – | OP897298* | – | – |
Sibon bevridgelyi |
|
– | – | – | MH374990 | – | – | – |
MZUTI 3269 | – | MH341077 | – | MH374962 | – | – | – | |
MZUTI 5416 | – | MH341078 | – | MH374963 | – | – | – | |
RSCDSP 0391 | – | – | – | JX398683 | JX398543 | JX293890 | JX398808 | |
Sibon canopy sp. nov. | JM 705 | – | MH140951 | – | JX398654 | JX398503 | – | JX398774 |
JM 759 | – | MH140949 | – | JX398655 | – | – | JX398775 | |
USNM 579846 | – | – | – | JX398653 | JX398502 | – | JX398773 | |
USNM 579849 | – | MH140947 | MH140366 | JX398656 | JX398504 | – | JX398776 | |
Sibon carri | UTA R-45493 | – | – | – | JX398665 | JX398514 | JX293876 | JX398786 |
Sibon irmelindicaprioae sp. nov. | MHCH 3111 | – | OP879853* | – | – | – | – | – |
MHCH 3145 | – | OP879854* | OP873119* | – | – | – | – | |
MHCH 3146 | – | OP879855* | OP873121* | – | – | – | – | |
|
– | OP879856* | OP873120* | – | – | – | – | |
|
– | OP879857* | – | – | – | – | – | |
UIS R-3515 | – | – | – | – | OP897296* | – | – | |
UIS R-3701 | – | – | – | – | OP897297* | – | – | |
Sibon dimidiatus | USNM 565824 | – | – | – | JX398668 | JX398517 | – | JX398789 |
Sibon dunni | CAMPO 533 | – | MH341079 | – | MH374991 | – | – | – |
Sibon hartwegi | MHUA 14511 | – | – | – | GQ334556 | GQ334662 | GQ334579 | GQ334685 |
SN 0001 | – | – | – | JX398684 | JX398544 | JX293892 | JX398809 | |
Sibon lamari | ASL 362 | – | – | – | JX398670 | JX398519 | – | – |
No voucher | – | – | – | JX398671 | JX398520 | JX293879 | JX398791 | |
Sibon leucomelas | CH 5296 | – | MH140972 | MH140392 | – | – | – | – |
CH 9135 | – | MH140971 | MH140391 | – | – | – | – | |
JM 703 | – | – | – | JX398679 | JX398539 | – | JX398804 | |
JM 722 | – | MH140967 | – | JX398680 | JX398540 | – | JX398805 | |
MHCH 3149 | – | OP879860* | – | – | – | – | – | |
Sibon leucomelas | USNM 579854 | – | – | MH140393 | JX398681 | JX398541 | – | JX398806 |
Sibon longifrenis | MVZ 215681 | MH341099 | – | – | MH375036 | MH375055 | – | – |
Sibon marleyae sp. nov. |
|
– | OP879861* | – | – | OP897295* | – | – |
MZUTI 3034 | – | MH341075 | – | MH375021 | – | – | – | |
Sibon nebulatus | SN02 | – | – | – | – | JX398545 | – | – |
UWIZM 2011.20.26 | – | – | – | JX398687 | JX398551 | – | – | |
Sibon perissostichon |
|
– | – | – | JX398688 | JX398552 | JX293888 | JX398814 |
Sibon vieirai sp. nov. | DHMECN 9585 | – | MH341082 | – | – | – | – | – |
ENS 12500 | – | – | – | – | JX398531 | – | JX398802 | |
ENS 12459 | – | – | – | – | JX398530 | – | JX398801 | |
MZUTI 3911 | – | MH341083 | – | MH374964 | – | – | – | |
MZUTI 4810 | – | MH341084 | – | MH374965 | – | – | – | |
Tropidodipsas fasciata | MX14 | – | – | – | JX398703 | – | JX293901 | JX398821 |
Tropidodipsas fischeri | UTA R-38932 | – | – | – | JX398707 | JX398566 | JX293903 | JX398823 |
Tropidodipsas guerreroensis | INIRENA 2781 | – | – | – | MZ287381 | MZ287395 | MZ287403 | MZ287420 |
Tropidodipsas papavericola | INIRENA 2805 | – | – | – | MZ287392 | MZ287392 | MZ287400 | MZ287418 |
Tropidodipsas philipii | JAC 24811 | – | – | – | JX398710 | JX398570 | JX293909 | JX398826 |
Tropidodipsas tricolor | CIG 1837 | – | – | – | MZ287386 | MZ287394 | MZ287404 | MZ287415 |
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 | Sequence (5’-3’) | Reference | PCR profile: |
---|---|---|---|---|
12S | H1557mod | GTACRCTTACCWTGTTACGACTT |
|
93 °C (1 min), 54 °C (1 min), 72 °C (2–5 min) [x25–40] |
L1091mod | CAAACTAGGATTAGATACCCTACTAT | |||
16S | 16Sar-L | CGCCTGTTTATCAAAAACAT |
|
94 °C (45 sec), 53 °C (45 sec), 72 °C (1 min) [x30] |
16Sbr-H-R | CCGGTCTGAACTCAGATCACGT | |||
COI | RepCOI-F | TNTTMTCAACNAACCACAAAGA |
|
94 °C (3 min), 48.5 °C (30 sec), 72 °C (1 min) [x40] |
RepCOI-R | ACTTCTGGRTGKCCAAARAATCA | |||
Cytb | L14910 | GACCTGTGATMTGAAAACCAYCGTTGT |
|
94 °C (1 min), 58 °C (1 min), 72 °C (2 min) [x30–36] |
H16064 | CTTTGGTTTACAAGAACAATGCTTTA | |||
ND4 | ND4 | CACCTATGACTACCAAAAGCTCATGTAGAAGC |
|
94 °C (25 sec), 56 or 60 °C (1 min), 72 °C (2 min) [x25–30] |
Leu | CATTACTTTTACTTGGATTTGCACCA |
Morphological and locality data for specimens of Dipsadini species examined, either directly, indirectly through digital photographs, or both. Codes: SVL = snout-vent length (mm); TL = tail length (mm); M = Male, F = Female.
Data type: Morphological
Locality data used to create distribution maps. Type localities indicated in bold type.
Data type: Occurrences