Research Article |
Corresponding author: Andrés R. Acosta-Galvis ( aacosta@humboldt.org.co ) Corresponding author: Mauricio Torres ( mauriciotorresmejia@gmail.com ) Academic editor: Anthony Herrel
© 2019 Andrés R. Acosta-Galvis, Mauricio Torres, Paola Pulido-Santacruz.
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:
Acosta-Galvis AR, Torres M, Pulido-Santacruz P (2019) A new species of Caecilia (Gymnophiona: Caeciliidae) from the Magdalena valley region of Colombia, with comments on the genus Parvicaecilia. ZooKeys 884: 135-157. https://doi.org/10.3897/zookeys.884.35776
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A new species of the genus Caecilia (Caeciliidae) from the western foothills of the Serranía de los Yariguíes in Colombia is described. Caecilia pulchraserrana sp. nov. is similar to C. degenerata and C. corpulenta but differs from these species in having fewer primary annular grooves and a shorter body length. With this new species, the currently recognized species in the genus are increased to 35. Mitochondrial DNA sequences, including newly sequenced terminals representing two additional, previously unanalyzed species, corroborate the phylogenetic position of the new species within Caecilia and the monophyly of the genus. This analysis also included newly sequenced terminals of Epicrionops aff. parkeri (Rhinatrematidae) and trans-Andean Microcaecilia nicefori (Siphonopidae). Evidence was found for the non-monophyly of the family Siphonopidae and the siphonopid genera Microcaecilia and Siphonops. The implications of these results for caecilian systematics are discussed and the status of the trans-Andean populations of Caecilia degenerata is commented upon.
Amphibia, Caecilia degenerata, Epicrionops, Microcaecilia, paraphyly, phylogeny, Siphonopidae, South America, taxonomy, tropical humid forest
The Neotropical caecilian amphibian genus Caecilia Linnaeus, 1758 (Gymnophiona: Caeciliidae) currently comprises 34 nominal species (
A second group includes three species that have secondary annular grooves: C. guntheri Dunn, 1942, with a wide distribution from northern Ecuador to Colombia, where the records are discontinuous and include the sub-Andean forests of the Cordillera Occidental and the region of Muzo at Quípama Municipality, Boyacá Department, western slope of the Cordillera Oriental, 1000 m a.s.l.; C. subnigricans Dunn, 1942, from northern Venezuela and lowlands of the Caribbean and Magdalena Valley regions of Colombia, with a record from Mariquita Municipality, Tolima Department; and C. thompsoni Boulenger, 1902b, endemic to the middle Magdalena valley in Colombia, 240–1571 m a.s.l. (
During a recent herpetological survey in wet tropical forests of the Serranía de los Yariguíes, in the Department of Santander, Colombia (Fig.
A Map of Colombia showing the known localities of the species of Caecilia that occur in the Magdalena valley region. Key: C. caribea (blue triangle), C. corpulenta (black dot), C. degenerata (black cross), C. guntheri (violet asterisk), C. subnigricans (yellow triangle), C. subdermalis (green star), C. thompsoni (black star), Caecilia pulchraserrana sp. nov. (red triangle) B Type locality of Caecilia pulchraserrana sp. nov. (red triangle) at Serranía de los Yariguíes, Santander Department, Colombia.
The new species was collected during fieldwork carried out in the Serranía de los Yariguíes, vereda La Belleza, municipality of El Carmen de Chucurí, Santander Department, Colombia (06°34'N, 73°34'W, 731–789 m a.s.l.; Fig.
Previous fieldwork conducted between 1998–1999 by John Lynch in collaboration with the first author, successfully allowed the detection of microhabitats and several specimens of Microcaecilia nicefori (
The collecting technique, which was used to obtain specimens of the new species, consists of first asking local people about the locations where they have spotted caecilians using the common names of “blind snakes”, or “captain worms” (“lombrices capitanas”), or “motolas” (this common name is specific for the Department of Santander). Subsequently, the reported sites are visited and inspected to select sites under the shade of vegetation, and where the soil is not compact and very humid (usually associated with water springs that form a mosaic of marshy and dry areas). Collecting efforts are focused in the selected damp microhabitats, digging with a hoe to a depth of approximately 20 cm (approximate sampling effort of 2-person-hour to collect five specimens). Coordinates and elevations were obtained with a Garmin GPSMAP 64SC (map datum WGS 84). Collected specimens were euthanized using 20% benzocaine (
To test the generic assignment of the new species and to explore the relationships of other endemic caecilians from Colombia, available mitochondrial DNA sequences of the genes 16S and CO1 from members of all Neotropical caecilian families (Caeciliidae, Typhlonectidae, Siphonopidae, Dermophiidae, and Rhinatrematidae) were analyzed (Table
List of species examined and GenBank or Barcode of life Data Systems (BOLD) accession numbers for each gene analyzed in this study. See Appendix
Species | Family | Tissue code | 16S GenBank; BOLD number | CO1 GenBank; BOLD number | Source |
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Caecilia gracilis | Caeciliidae | KX757086 | NC_023508 |
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Caecilia isthmica | Caeciliidae | IAvH-CT-22982 | MN555719; SABIO393-19 | MN555727; SABIO393-19 | This study |
Caecilia pulchraserrana sp. nov | Caeciliidae | IAvH-CT-227334 | MN555715; SABIO005-18 | MN555723; SABIO005-18 | This study |
Caeciliidae | IAvH-CT-22733 | MN555718; SABIO002-18 | MN555726; SABIO002-18 | This study | |
Caecilia tentaculata | Caeciliidae | NC_023507 | NC_023507 |
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Caecilia thompsoni | Caeciliidae | IAvH-CT-22986 | MN555717; SABIO392-19 | MN555725; SABIO392-19 | This study |
Caecilia volcani | Caeciliidae | FJ784371 | NC_020137 |
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Oscaecilia ochrocephala | Caeciliidae | GQ244474 | GQ244474 |
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Dermophis mexicanus | Dermophiidae | – | NC_020138 |
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Epicrionops cf. marmoratus | Rhinatrematidae | KF540151 | KF540151 |
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Rhinatrema nigrum | Rhinatrematidae | GQ244468 | GQ244468 |
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Epicrionops aff. parkeri | Rhinatrematidae | IAvH-CT-21477 | MN555716; CBIHA031-17 | MN555724; CBIHA031-17 | This study |
Microcaecilia dermatophaga | Siphonopidae | NC_023514 | NC_023514 |
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Microcaecilia sp. | Siphonopidae | GQ244473 | GQ244473 |
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Microcaecilia unicolor | Siphonopidae | NC_023515 | NC_023515 |
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Microcaecilia nicefori | Siphonopidae | IAvH-CT-22985 | MN555722; CAECI002-19 | MN555729; CAECI002-19 | This study |
Siphonops annulatus | Siphonopidae | KU495581 | KU495581 |
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Siphonops hardyii | Siphonopidae | KU495582 | KU494789 |
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Siphonops insulanus | Siphonopidae | KU495583 | KU494790 |
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Siphonops paulensis | Siphonopidae | KU495584 | KU494791 |
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Potomotyphlus kaupii | Typhlonectidae | NC_023516 | NC_023516 |
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Typhlonectes compressicauda | Typhlonectidae | KU495605 | KU494812 |
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Typhlonectes natans | Typhlonectidae | AF154051 | AF154051 |
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Typhlonectidae | IAvH-CT-22983 | MN555720; SABIO394-19 | MN555728; SABIO394-19 | This study | |
Typhlonectidae | IAvH-CT-22984 | MN555721; CAECI001-19 | – | This study |
Criteria and terminology for morphological descriptions, diagnostic characters, and data for other species of Caecilia follow
ADD anal disc diameter;
AM anteromedial limit of the mouth on the upper jaw;
BH body height at midbody;
C1 first collar length;
C2 second collar length;
CM corner of the mouth;
CMB circumference at midbody;
D diameter at midbody;
ED eye diameter;
END distance between eye and naris;
HH head height at level with CM;
HL head length;
HW head width at CM;
HWNG1 head width at NG1;
IND distance between nares;
IOD interorbital distance;
TL total length;
TL/D TL divided by diameter at midbody (ratio of length/diameter);
LPOD distance between eye and lip;
ND naris diameter;
NG1 first nuchal groove;
NG2 second nuchal groove;
NG3 third nuchal groove;
PA primary annulus;
PAG primary annular groove;
PM premaxillary-maxillary tooth;
ST snout tip;
STD distance between snout tip and anterior margin of mouth;
STND distance between ST and naris;
STLPD distance between ST and lip;
STOD distance between ST and eye;
TA tentacular aperture;
INTA distance between TAs;
TAOD distance between TA and eye;
TALPD distance between TA and lip;
TANRD distance between TA and naris;
TASTD distance between TA and ST;
VP vomeropalatine tooth;
WC2 width at second collar;
WCH width of choanae;
WBV width of body at vent level;
WMB width at midbody;
TL/HL TL divided by HL;
TL/WMB TL divided by WMB;
TL/HW TL divided by HW;
HL/HW HL divided by HW.
Dermal scale pockets and subdermal scales were searched using the criteria proposed by
The final concatenated molecular dataset consisted of a matrix of 1273 bp, 567 sites were parsimony-informative, 111 were singletons, and 595 were constant sites. The best fitting substitution model for both CO1 and 16S was TIM2+F+I+G4 after testing the large selection of models in IQ-TREE. The ML tree is shown in Fig.
Maximum Likelihood tree inferred from the analysis of a concatenated dataset comprising partial sequences of two mitochondrial genes. Numbers above branches indicate bootstrap support values (percent) (* = 100% bootstrap). Scale bar indicates nucleotide substitutions per site. The phylogenetic position of Caecilia pulchraserrana sp. nov. is shown in bold.
Generic assignment. The new species is assignable to the genus Caecilia because its eyes are not covered by bone and it has tentacles below the nostrils (Type D sensu Lynch, 1999, Fig.
IAvH-Am-15487 (field number ARA 7872; Figs
(Fig.
Four specimens (Fig.
UIS-MHN-A-6576–7 (field numbers ARA 7692–3, respectively), juveniles, 06°34'41.1"N, 73°34'28.9"W, 731 m a.s.l., collected 19 February 2018 by A. R. Acosta-Galvis and Miguel Torres. Tissues for molecular analysis (IAvH-CT-22733–4) were extracted from these specimens.
Caecilia pulchraserrana sp. nov. differs from its congeners by the combination of having 100–104 dorsally incomplete primary annular grooves, a small size (195–232 mm), lips and ventral margin of upper jaw with a pink-orange (salmon) color (Fig.
Regarding the species of the genus Caecilia, the absence of secondary annular grooves distinguishes C. pulchraserrana sp. nov. from C. abitaguae Dunn, 1942, C. albiventris Daudin, 1803, C. armata Dunn, 1942, C. antioquiaensis Taylor, 1968, C. bokermanni Taylor, 1968, C. dunni Hershkovitz, 1938, C. flavopunctata Roze & Solano, 1963, C. gracilis Shaw, 1802, C. guntheri Dunn, 1942, C. isthmica Cope, 1878, C. leucocephala Taylor, 1968, C. marcusi Wake, 1985, C. mertensi Taylor, 1973, C. museugoeldi Maciel & Hoogmoed, 2018, C. nigricans Boulenger, 1902, C. occidentalis Taylor, 1968, C. pressula Taylor, 1968, C. perdita Taylor, 1968, C. subnigricans Dunn, 1942, C. subterminalis Taylor, 1968, C. tentaculata Linnaeus, 1758, C. tenuissima (Taylor, 1973), C. thompsoni Boulenger, 1902, and C. volcani Taylor, 1969.
Caecilia pulchraserrana sp. nov. shares with C. attenuata Taylor, 1968, C. caribea Dunn, 1942, C. corpulenta Taylor, 1968, C. crassisquama Taylor, 1968, C. degenerata Dunn, 1942, C. inca Taylor, 1973, C. orientalis Taylor, 1968, C. pachynema Günther, 1859, and C. subdermalis Taylor, 1968 the absence of secondary annular grooves and the presence of incomplete primary annular grooves. However, the new species can be distinguished from these nine species by having a lower number of primary annular grooves (100–104 vs. 114–199). Caecilia pulchraserrana sp. nov. most closely resembles C. degenerata, which also lacks subdermal scales, but differs from it in having fewer primary annuli.
An adult female (Fig.
(Fig.
(ethanol 70%; Fig.
(Tables
Morphological data of the Colombian species of Caecilia that lack secondary annular grooves and possess incomplete primary annular grooves. Abbreviations are given in Material and methods.
Species | PAG | TL (mm) | TL/D | Dermal scale pockets | Sample size | Source |
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C. caribea | 142–152 | 390–585 | 53–55 | Absent | 4 |
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C. corpulenta | 129–132 | 152–441 | 19–35 | Absent | 6 |
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C. degenerata | 123–137 | 390–1050 | 38–58 | Absent | 9 |
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C. orientalis | 114–124 | 231–673 | 29–55 | Present | 8 |
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C. subdermalis | 116–138 | 131–680 | 28–54 | Present | 32 |
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C. pulchraserrana sp. nov. | 100–104 | 195–232 | 9–12 | Absent | 7 | This study |
Morphometric (in mm) and meristic data of the type series of Caecilia pulchraserrana sp. nov. Abbreviations are given in Materials and methods.
IAvH-Am-15487 Holotype | IAvH-Am-15490 Paratype | IAvH-Am-15489 Paratype | IAvH-Am-15488 Paratype | UIS-MHN-A-6575 Paratype | |
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Sex | F | M | M | F | F |
PAG | 104 | 100 | 101 | 103 | 100 |
TL | 206 | 214 | 200 | 232 | 195 |
HW | 5.4 | 5.3 | 5.0 | 4.8 | 4.3 |
HWNG1 | 5.2 | 4.2 | 4.4 | 4.9 | 4.3 |
WC2 | 5.8 | 4.6 | 4.0 | 5.2 | 4.8 |
WMB | 8.5 | 6.2 | 5.5 | 8.1 | 6.2 |
CMB | 22 | 18 | 17 | 23 | 18 |
WBV | 5.2 | 3.7 | 4.0 | 4.4 | 3.5 |
HL | 7.4 | 5.8 | 6.4 | 6.0 | 5.1 |
HH | 5.1 | 4.8 | 4.0 | 4.4 | 3.8 |
IND | 1.7 | 1.6 | 1.5 | 2.0 | 1.2 |
IOD | 2.9 | 2.6 | 2.3 | 2.8 | 2.5 |
ED | 0.3 | 0.4 | 0.4 | 0.2 | 0.4 |
ND | 0.18 | 0.18 | 0.16 | 0.16 | 0.15 |
END | 2.3 | 2.3 | 1.6 | 2.5 | 2.1 |
STD | 6.9 | 5.6 | 5.7 | 6.0 | 5.2 |
STND | 0.8 | 0.6 | 0.4 | 0.7 | 0.7 |
STLPD | 2.2 | 2.1 | 2.1 | 2.0 | 1.8 |
STOD | 3.3 | 2.7 | 2.5 | 3.4 | 2.9 |
TA | 0.27 | 0.19 | 0.30 | 0.26 | 0.33 |
INTA | 2.3 | 2.2 | 1.8 | 2.3 | 1.9 |
TAOD | 2.5 | 2.1 | 1.9 | 2.6 | 2.1 |
TALPD | 1.0 | 1.3 | 0.6 | 1.4 | 0.99 |
TANRD | 0.99 | 0.67 | 0.69 | 0.75 | 0.7 |
TASTD | 0.6 | 0.7 | 0.7 | 0.2 | 0.7 |
LPOD | 1.0 | 1.2 | 0.9 | 1.0 | 0.7 |
WCH | 0.16 | 0.11 | 0.09 | 0.11 | 0.14 |
C1 | 1.6 | 1.2 | 1.6 | 1.1 | 0.9 |
C2 | 1.7 | 1.5 | 2.4 | 1.5 | 1.1 |
BH | 7.0 | 4.4 | 4.1 | 6.5 | 5.1 |
ADD | 2.9 | 2.6 | 2.9 | 2.7 | 2.6 |
VP | 11 | 9 | 10 | 9 | 11 |
Premaxillary-maxillary teeth | 13 | 11 | 14 | 14 | 12 |
Dentary teeth | 12 | 13 | 10 | 11 | 12 |
Ratios and percentages of measurements of the type series of Caecilia pulchraserrana sp. nov. Abbreviations are given in Materials and methods.
IAvH-Am-15490 Paratype | IAvH-Am-15489 Paratype | IAvH-Am-15488 Paratype | IAvH-Am-15487 Holotype | UIS-MHN-A-6575 Paratype | |
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Sex | M | M | F | F | F |
C1/C2 | 75.9 | 66.1 | 70.5 | 96.4 | 82.7 |
TL/D | 11.8 | 11.7 | 10.0 | 9.3 | 10.8 |
TL/HL | 39.9 | 40.0 | 48.2 | 38.1 | 44.5 |
TL/ WMB | 34.1 | 35.9 | 28.3 | 24.1 | 31.5 |
L/HW | 36.7 | 30.8 | 38.2 | 27.7 | 37.8 |
HL/HW | 92.0 | 77.0 | 79.2 | 72.9 | 85.0 |
Caecilia pulchraserrana sp. nov. is currently known from two adjacent, relictual tropical wet forest localities on the western slope of the Cordillera Oriental of Colombia (Serranía de los Yariquíes; Fig.
Habitat of Caecilia pulchraserrana sp. nov. in the Serranía de los Yariguíes in Santander Department, El Carmen de Chucurí Municipality, vereda La Belleza, Cascajales River, 06°34'8.9"N, 73°34'20.2"W, 789 m a.s.l.. A View showing standing water in marshy area B Transitional change of wetter (right) to drier (left) microhabitat.
Caecilia pulchraserrana sp. nov. was obtained during the initial 10 minutes of removal with a hoe.We extracted the first specimen in intermediate substrates between marshy and dry areas; after 40 minutes of excavation in these selected areas, we obtained four additional specimens. Using these same criteria, when moving two kilometers above the original point, an area with similar characteristics was located and within 20 minutes we collected two additional specimens. Caecilia pulchraserrana sp. nov. was collected on black sandy soils with high organic matter content. These caecilians move quickly under the substrate, so once the first specimen is detected it is important to quickly create channels to surround and block them from escaping.
The specific epithet is formed from the Latin pulchra (nominative feminine singular of pulcher), meaning beauty, and the Spanish adjective serrana (feminine singular of serrano), from the sierra or serranía. This specific name refers to the type locality of the species: vereda La Belleza (beauty in English) in the western foothills of the Serranía de Los Yariguíes. The specific name was chosen using a citizen science approach. First, scientists and inhabitants of the El Carmen de Chucurí municipality gathered a list of possible names for the new species. Then, the list of potential names and their meanings was shared with the local people, who voted to choose their preferred name.
Our description of Caecilia pulchraserrana sp. nov. brings the number of known species of Caecilia to 35 (
Our phylogenetic analysis only included two mitochondrial loci and a small number of species and should not be considered as a robust resolution of caecilian relationships. Nevertheless, our results highlight several potential cases of non-monophyletic taxa and suggest that a taxonomic revision, including a major generic rearrangement, is warranted. Our study includes, for the first time, the Colombian endemics Epicrionops aff. parkeri (Rhinatrematidae) and Microcaecilia nicefori in molecular phylogenetic analyses. On one hand, recent contributions (
On the other hand, Microcaecilia nicefori was recovered as the sister taxon to a clade formed by the dermophiids Gymnophis multiplicata + Dermophis mexicanus and the remaining siphonopids, including Microcaecilia, Brasilotyphlus guarantanus, Siphonops, and Luetkenotyphlus. In addition, Microcaecilia and Siphonops were recovered as paraphyletic with respect to Brasilotyphlus guarantanus and Luetkenotyphlus brasiliensis, respectively (Fig.
Consistent with previous findings (i.e.,
Caecilia pulchraserrana sp. nov. is described as an endemic species from the Serranía de los Yariguies. The species is similar to C. degenerata, from which it can be distinguished using morphological characters. According to their morphology, we hypothesize there is a group of closely related species that comprises C. caribea, C. corpulenta, C. degenerata, C. orientalis, and C. subdermalis. The trans-Andean Microcaecilia nicefori is an endemic and poorly known species from Colombia. We provide here the first analysis of molecular data that tests its phylogenetic position. Our results address the need to evaluate with more evidence the status of the genus Parvicaecilia Taylor, 1968 (currently under the synonymy of Microcaecilia), and the potential non-monophyly of the family Siphonopidae. Further analyses sampling additional taxa and molecular markers are required to establish a more robust classification for Gymnophiona.
This research was supported by Santander Bio, a project funded by the Sistema General de Regalías, administered by the Departamento Nacional de Planeación (BPIN 2017000100046), executed by the Gobernación de Santander, and operated by the Instituto de Investigación de Recursos Biológicos Alexander von Humboldt and the Universidad Industrial de Santander (Inter-administrative Agreement 2243, Gobernación de Santander). Specimens were collected under a permit issued by the Instituto de Investigación de Recursos Biológicos Alexander von Humboldt (Decree 1376 of 2013). We thank Javier Barriga for providing invaluable support during fieldwork. We specially thank Marjorie Pinzón Arias for interchanging knowledge with local people from El Carmen de Chucurí to come up with the name of the new species. Special thanks to the inhabitants of the Vereda La Belleza, El Carmen de Chucurí municipality, for participating in and allowing us to carry out the biodiversity inventories. We are also grateful to Miguel Torres and Daniela García for collaborating actively on discovering and collecting several specimens of the new species. In addition, we thank the collaboration of Yeison Tolosa in the process of obtaining material of Microcaecilia nicefori. Mark Wilkinson, David Gower, and Santiago J. Sánchez-Pacheco provided comments, suggestions and corrections that greatly improved the manuscript. Special thanks to Eduardo Tovar-Luque for his expert technical assistance with the molecular work.
Additional specimens examined in this study. Number of specimens examined of each species in parenthesis.
Caecilia guntheri (2): COLOMBIA: NARIÑO: La Planada Natural Reserve, 7 km South of Chucunes, 1780 m above sea level; IAvH-Am-1396; RISARALDA: Pueblo Rico Municipality, Vereda Montebello, Montezuma Reserve, 4°33'40.5"N, 74°21'4.9"W, 1650 m above sea level, IAvH-Am-8872.
Caecilia isthmica (1): COLOMBIA: SUCRE: San Benito Abad Municipality, Vereda La Caimanera, site La Caimanera, 9°2'33.7"N, 74°54'17.6"W, 26 m above sea level, IAvH-Am-8246 (tissue IAvH-CT-22982).
Caecilia subdermalis (10): COLOMBIA: CALDAS: Norcasia Municipality, Hidromiel camp, 5°34'16.4"N, 74°53'24.8"W, 850 m above sea level. IAvH-Am-9663; HUILA, Acevedo Municipality, Cueva de los Guácharos National Natural Park, 1820 m above sea level. IAvH-Am-0687, IAvH-Am-3541, IAvH-Am-3549, IAvH-Am-4316-7, IAvH-Am-4322-23, IAvH-Am-4708, IAvH-Am-5388.
Caecilia thompsoni (1): COLOMBIA: CUNDINAMARCA: La Mesa Municipality, site Payacal, La Gran Via, Tacarcuna Farm, 04°39'6,77"N, 74°25'1.0"W; 1100 m above sea level, MUJ 3713 (tissue IAvH-CT-22986).
Epicrionops aff. parkeri (2): COLOMBIA: ANTIOQUIA: municipality of El Carmen de Viboral, vereda El Porvenir, creek afferent to the Melcocho River, 5°54'7.9"N, 75°10'25.6"W, 898 m above sea level, IAvH-Am-14608, IAvH-Am-14609 (tissue IAvH-CT-21477).
Microcaecilia nicefori (1): COLOMBIA: TOLIMA: municipality of Coello, El Neme farm (outside of town), 4°7'12.50"N, 74°55'21.10"W, 327 m above sea level, IAvH-Am-14879 (tissue IAvH-CT-22985).
Typhlonectes natans (2): COLOMBIA: SUCRE: San Benito Abad Municipality, Vereda La Caimanera, site La Caimanera, 9°27'1"N, 74°54'26.7"W, 25 m above sea level, IAvH-Am-8275 (tissue IAvH-CT-22983). NORTE DE SANTANDER: San José de Cúcuta Municipality, Aguasal Creek, Footbridge about 1.2 km northeast of the community of Aguasal, 08°13'05"N, 072°32'31.2"W, 62 m above sea level, IAvH-Am-14559 (tissue IAvH-CT-22984).