Corresponding author: Alejandro Arteaga (
Academic editor: R. Jadin
We present a molecular phylogeny of snake genus
Arteaga A, Mebert K, Valencia JH, Cisneros-Heredia DF, Peñafiel N, Reyes-Puig C, Vieira-Fernandes JL, Guayasamin JM (2017) Molecular phylogeny of
With 140 species,
One recent work by
To resolve these pending issues and to shed light on potentially unclear species boundaries, we report on new material of
This study was carried out in strict accordance with the guidelines for use of live amphibians and reptiles in field research compiled by the American Society of Ichthyologists and Herpetologists (
Tissue samples from 39 individuals representing 22 species (including three new species described here) were obtained throughout Ecuador. The majority of individuals were located by space-constrained visual examination of ground-level substrates (
Genomic DNA was extracted from 96% ethanol-preserved tissue samples (liver, muscle tissue or scales) using a modified salt precipitation method based on the Puregene DNA purification kit (Gentra Systems). We amplified the 16S gene using the primers 16Sar-L and 16Sbr-H-R from
A total of 126 mtDNA sequences were used to build a mitochondrial phylogenetic tree of the genus
Our terminology for
Locality data for specimens examined in this study. Coordinates represent 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.
Species | Voucher | Locality | Latitude | Longitude | Elev. |
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Loja, Utuana | -4.36642 | -79.72483 | 2517 | |
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Esmeraldas, Copa Quininde (in error) | 0.06181 | -78.72641 | 1688 | |
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Loja, Utuana | -4.36642 | -79.72483 | 2517 | |
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Loja, Utuana | -4.36642 | -79.72483 | 2517 | |
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Loja, Utuana | -4.36642 | -79.72483 | 2517 | |
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Napo, Chiriboga (in error) | - | - | - | |
|
Ecuador | - | - | - | |
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Napo, Yanayacu | -0.60071 | -77.88927 | 1924 | |
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Napo, Yanayacu | -0.59939 | -77.89050 | 2064 | |
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Carchi, Gualpi | 0.86439 | -78.22435 | 2104 | |
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Pichincha, Río Cambugán | 0.17697 | -78.50779 | 1828 | |
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Imbabura, Junín | 0.27009 | -78.64975 | 1688 | |
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Pichincha, Tambo Quinde | 0.00967 | -78.66906 | 1870 | |
|
Pichincha, Pahuma | 0.02757 | -78.63208 | 1914 | |
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Santo Domingo, Chiriboga | -0.22841 | -78.76725 | 1813 | |
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Santo Domingo, Chiriboga | -0.22841 | -78.76725 | 1813 | |
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Santo Domingo, La Favorita | -0.22833 | -78.76503 | 1810 | |
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Santo Domingo, La Favorita | -0.22833 | -78.76503 | 1810 | |
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Santo Domingo, La Favorita | -0.22833 | -78.76503 | 1810 | |
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Santo Domingo, La Favorita | -0.22833 | -78.76503 | 1810 | |
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Cotopaxi, Cutzualo | -0.54497 | -78.91891 | 1952 | |
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Santo Domingo, La Favorita | -0.22841 | -78.76725 | 1813 | |
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Cotopaxi, Otonga | -0.41549 | -79.00480 | 2095 | |
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Pichincha, Tandayapa–Bellavista | -0.00843 | -78.67619 | 1919 | |
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Pichincha, Tandayapa Lodge | 0.00268 | -78.68131 | 1757 | |
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Imbabura, Santa Rosa de Intag | 0.37616 | -78.46054 | 2077 | |
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Imbabura, Santa Rosa de Intag | 0.37616 | -78.46054 | 2077 | |
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Imbabura, Santa Rosa de Intag | 0.37616 | -78.46054 | 2077 | |
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Imbabura, Below of Siempre Verde | 0.37782 | -78.46901 | 1974 | |
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Imbabura, Toisán | 0.53297 | -78.52924 | 2286 | |
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Imbabura, Toisán | 0.53297 | -78.52924 | 2286 | |
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Santo Domingo, Guajalito | -0.22875 | -78.82248 | 1801 | |
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Tungurahua, Río Verde | -1.40344 | -78.30099 | 1507 | |
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Morona Santiago, Tundayme | -3.57244 | -78.46982 | 790 | |
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Morona Santiago, Macas | -2.31670 | -78.11670 | 972 | |
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Santo Domingo, Chiriboga | -0.22841 | -78.76725 | 1813 | |
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Cotopaxi, Otonga | -0.41549 | -79.00480 | 2095 | |
|
Pichincha, Las Gralarias | -0.00807 | -78.73238 | 1985 | |
|
Esmeraldas, Canande | 0.52993 | -79.03541 | 594 | |
|
Esmeraldas, Tundaloma | 1.18236 | -78.75250 | 74 | |
|
Esmeraldas, Canande | 0.52993 | -79.03541 | 594 | |
|
Carchi, Río Blanco | 1.18993 | -78.50413 | 223 | |
|
Esmeraldas, Tsejpi | 0.79930 | -78.84527 | 152 | |
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Esmeraldas, Tundaloma | 1.18166 | -78.74945 | 74 | |
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Esmeraldas, Tundaloma | 1.18166 | -78.74945 | 74 | |
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Pichincha, Puerto Quito | 0.11667 | -79.26661 | 143 | |
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Esmeraldas, Canande | 0.52993 | -79.03541 | 594 | |
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Esmeraldas, Tundaloma | 1.18166 | -78.74945 | 74 | |
|
Azuay, Reserva Yunguilla | -3.22684 | -79.27520 | 1748 | |
|
Azuay, Reserva Yunguilla | -3.22684 | -79.27520 | 1748 | |
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ANF 1545 | Orellana, Estación Científica Yasuní | -0.67781 | -76.39819 | 246 |
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Sucumbíos, Bloque 27 | 0.32273 | -76.19369 | 272 | |
|
Ecuador | - | - | - | |
|
Zamora Chinchipe, Maycu | -4.38030 | -78.74584 | 981 | |
|
El Oro, Buenaventura | -3.65467 | -79.76794 | 524 | |
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El Oro, Buenaventura | -3.65467 | -79.76794 | 524 | |
|
El Oro, Zambo Tambo | -3.67861 | -79.68001 | 1014 | |
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El Oro, El Progreso | -3.26998 | -79.73452 | 176 | |
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El Oro, El Progreso | -3.26998 | -79.73452 | 176 | |
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El Oro, El Progreso | -3.26998 | -79.73452 | 176 | |
|
El Oro, El Progreso | -3.26998 | -79.73452 | 176 | |
|
El Oro, El Progreso | -3.26998 | -79.73452 | 176 | |
|
Manabí, Jama Coaque | -0.11556 | -80.12472 | 299 | |
|
Los Ríos, Río Palenque | -0.59273 | -79.36369 | 163 | |
|
Loja, Olmedo | -3.94994 | -79.66667 | 1545 | |
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Los Ríos, Rio Palenque | -0.58333 | -79.36667 | 173 | |
|
Los Ríos, Rio Palenque | -0.58333 | -79.36667 | 173 | |
|
El Oro, Piñas | -3.68041 | -79.68253 | 1019 | |
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Carchi, Chical | 0.90327 | -78.16201 | 1437 | |
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Pichincha, Maquipucuna | 0.11757 | -78.67446 | 1490 | |
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Pichincha, Maquipucuna | 0.11757 | -78.67446 | 1490 | |
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Cotopaxi, Las Pampas | -0.44036 | -78.96663 | 1590 | |
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Pichincha, Gualea | 0.08536 | -78.74092 | 1557 | |
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Esmeraldas, Canandé | 0.52581 | -79.2088 | 310 | |
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Pichincha, Mindo | -0.04872 | -78.77520 | 1277 | |
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Santo Domingo, La Favorita | -0.22833 | -78.76503 | 1810 | |
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Cotopaxi, Las Pampas | -0.44036 | -78.96663 | 1590 | |
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Pichincha, Tandapi | -0.41522 | -78.79728 | 1455 | |
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Cotopaxi, Las Pampas | -0.44036 | -78.96663 | 1590 | |
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Pichincha, Tandapi | -0.41522 | -78.79728 | 1455 | |
|
Pichincha, Tandapi | -0.41522 | -78.79728 | 1455 | |
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Pichincha, Nanegalito | 0.06181 | -78.72641 | 1688 | |
|
Pichincha, Yellow House | -0.04492 | -78.75843 | 1504 | |
|
Pichincha, Yellow House | -0.05199 | -78.76923 | 1325 | |
|
Pichincha, Yellow House | -0.04371 | -78.75351 | 1520 | |
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Pichincha,Las Gralarias | -0.00615 | -78.73381 | 1985 | |
|
Pichincha, Pedro Vicente Maldonado | 0.05361 | -78.92109 | 938 | |
|
Santa Elena, Comuna Loma Alta | -1.83442 | -80.70291 | 72 | |
|
Santo Domingo, Finca La Esperanza | -0.27160 | -79.10568 | 616 | |
|
Santo Domingo, Finca La Esperanza | -0.27160 | -79.10568 | 616 | |
|
Santo Domingo, Finca La Esperanza | -0.27160 | -79.10568 | 616 | |
|
Santo Domingo, Finca La Esperanza | -0.27160 | -79.10568 | 616 | |
|
Pichincha, Puerto Quito | 0.11667 | -79.26661 | 143 | |
|
Santo Domingo, Santo Domingo | -0.25351 | -79.17297 | 554 | |
|
Pichincha, Río Cinto | -0.09070 | -78.80299 | 1409 | |
|
El Oro, Buenaventura | -3.65467 | -79.76794 | 524 | |
|
Pichincha, Río Cinto | -0.09070 | -78.80299 | 1409 | |
|
Tungurahua, Puntzan | -1.41359 | -78.40951 | 1962 | |
|
Chimborazo, Vicinity of Tixán | -2.16174 | -78.81227 | 2892 | |
|
Chimborazo, Vicinity of Tixán | -2.16174 | -78.81227 | 2892 | |
|
Azuay, Hierba Mala | -2.76439 | -79.43816 | 3029 | |
|
El Oro, Guanazán | -3.44139 | -79.49417 | 2596 | |
|
El Oro, Guanazán | -3.44139 | -79.49417 | 2596 | |
|
El Oro, Guanazán | -3.44139 | -79.49417 | 2596 | |
|
El Oro, Guanazán | -3.44139 | -79.49417 | 2596 | |
|
Carchi, Río la Plata | 0.82381 | -78.04584 | 2256 | |
|
Carchi, Chilma Bajo | 0.86495 | -78.04978 | 2058 | |
|
Morona Santiago, Gualaquiza | -3.39914 | -78.57859 | 835 | |
|
Morona Santiago, Gualaquiza | -3.39914 | -78.57859 | 835 | |
|
ANF 2390 | Pastaza, Tzarentza | -1.35696 | -78.05814 | 1355 |
|
Imbabura, Paramba (in error) | 0.81671 | -78.35002 | 698 | |
|
Imbabura, Paramba (in error) | 0.81671 | -78.35002 | 698 | |
|
Esmeraldas, Canandé | 0.52993 | -79.03541 | 594 | |
|
Esmeraldas, Gualpi | 0.78173 | -79.15993 | 63 | |
|
Esmeraldas, Gualpi | 0.78173 | -79.15993 | 63 | |
|
Esmeraldas, Itapoa | 0.51307 | -79.13400 | 321 |
The electronic edition of this article conforms to the requirements of the amended International Code of Zoological Nomenclature, and hence the new names contained herein are available under that Code from the electronic edition of this article. This published work and the nomenclatural acts it contains have been registered in ZooBank, the online registration system for the
The overall topology and support (Fig.
Bayesian consensus phylogeny depicting relationships within colubrid snakes of the genus
The resulting hypotheses of species relationships for our mitochondrial phylogenetic tree supports
We seek here to only name or redelimit
Photographs of some Ecuadorian species of
Figure
Figure
Figure
Morphometric data for members of the
Species |
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Males | Females | Males | Females | ||||||
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152–157 | – | 25–26 | – | 17 | 2 | 7 | 7 | 7 |
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125–136 | 138–150 | 26–39 | 19–26 | 17 | 2 | 6–7 | 6–8 | 5–7 |
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127 | – | 36 | – | 15 | 2 | 7 | 7 | 6 |
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149 | 156 | 41 | 30 | 17 | 2 | 7 | 7 | 5 |
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127–150 | 135–144 | 33–42 | 25–37 | 17 | 2 | 6–7 | 6–7 | 5–6 |
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133–150 | 144–163 | 32–40 | 24–29 | 17 | 1–2 | 7 | 6–7 | 5–7 |
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129–141 | 128–149 | 33–39 | 20–37 | 17 | 2 | 6–7 | 6–7 | 5–7 |
Distribution of Ecuadorian snakes of the
Morphometric data for members of the
Species |
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Males | Females | Males | Females | ||||||
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136–151 | 143–161 | 25–34 | 18–32 | 15 | 1 | 6 | 6 | 7–10 |
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141–144 | 148–153 | 25–29 | 20–21 | 15–17 | 1 | 5 | 6 | 8–11 |
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– | 143 | – | 16 | 15 | 1 | 6 | 5 | 8 |
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135–146 | 143–156 | 20–27 | 14–23 | 15 | 1 | 5–6 | 6–7 | 9–13 |
Distribution of Ecuadorian snakes of the
Cerberus Groundsnake
Tierrera cancerbera
Adult male holotype
The dorsal ground color is brown with five feebly visible dark-brown to black longitudinal lines that are not continuous throughout the length of the body but broken into spots along some sections (Fig.
Adult male,
The two known specimens of
Known only from the type locality, Pacoche, in the Ecuadorian province of Manabí at 280–324 m (Fig.
The specific epithet “
Although
Indistinct Groundsnake
Tierrera indistinta
Adult male holotype of
The dorsal ground color is dark brown with either six longitudinal black lines separated by lighter areas or a pattern of dark brown longitudinally arranged spots that correspond to the longitudinal lines. On each side, the line or series of dark spots along the 2nd and 3rd dorsal scale row is feebly visible, but the other lines or spots are conspicuous. The dorsal surface of the head is dark brown and there is a clearly marked dark postocular stripe running from behind the eye to the edge of the mouth (Fig.
Adult male,
The two known specimens of
Known only from the type locality, Caimito, in the Ecuadorian province of Esmeraldas at 102 m (Fig.
The specific epithet
We consider
Pyron’s Groundsnake
Tierrera de Pyron
Adult female holotype of
The dorsal ground color is blackish with a dark vertebral (mid-dorsal) scale row flanked by a dark yellow scale row on either side (the 7th dorsal scale row), irregularly adjoined by one to few additional yellow scales on the 6th dorsal scale row, rendering an appearance of an irregularly edged mid-dorsal striped pattern (Fig.
Adult female,
The only known specimen of
Known only from the type locality, between Balzapamba and Bilován, in the Ecuadorian province of Bolívar at 2026 m (Fig.
Named after R. Alexander Pyron, one of the most prolific contemporary herpetologists, in recognition of his invaluable contribution to systematics and evolution of the world’s reptiles.
We consider
Species relationships and taxonomy in the colubrid snake genus
From the five members of the
We also re-delimit the
To further clarify the landscape of
Finally, although
Our analysis of new
With these changes, the species number reported in Ecuador increases to 27:
We hope that the novel genetic and morphological data provided herein will promote future researchers to examine species boundaries in
Conceived and designed the work: AA. Performed the analyses: AA NP. Gathered morphological data: KB JHV DFCH CRP JLVF AA. Analyzed the data: AA KM DFCH JMG. Contributed reagents/materials/analysis tools: JMG NP. Wrote the paper: AA KM JHV DFCH NP CRP JLVF JMG.
This article was greatly improved by comments of three anonymous reviewers. For granting access to their protected forests, we are grateful to Francisco Sornoza and Martin Schaefer of Fundación Jocotoco, Ana Cristina de la Torre of Pacoche Lodge, and Andrés Chiriboga of Tundaloma Lodge. Special thanks to Lucas Bustamante, Alex Pyron, Gabriela Morales, Carlos Durán, Carlos Gómez, Gabriela Aguiar, James Muchmore, Ryan Lynch, Rita Hidalgo, Ángela León, Silvia Cevallos, Paulina Romero, Jaime Culebras, Carlos Londoño, Andy Proaño, and Daniel Romero for their assistance and companionship in the field. For providing the picture of
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 | 16S | CYTB | ND4 |
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ANF 2390 | |||
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MHUA 14368 | – |
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Outgroups | ||||
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MVZ 233298 |
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List of
Locus | Primer name | Sequence (5’-3’) | Reference | |
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16S | 16Sar-L | CGCCTGTTTATCAAAAACAT |
|
94 °C (45 sec), 53 or 56 °C (45 sec), 72 °C (1 min) [x25-30] |
16Sbr-H-R | CCGGTCTGAACTCAGATCACGT | |||
Cytb | L14910 | GACCTGTGATMTGAAAACCAYCGTTGT |
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94 °C (1 min), 58 °C (1 min), 72 °C (2 min) [x30-36] |
H16064 | CTTTGGTTTACAAGAACAATGCTTTA | |||
ND4 | ND4 | CACCTATGACTACCAAAAGCTCATGTAGAAGC |
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94 °C (25 sec), 58 or 60 °C (1 min), 72 °C (2 min) [x25-30] |
Leu | CATTACTTTTACTTGGATTTGCACCA |
Morphometric data and sex for specimens of
Species | Voucher |
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Sex |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|
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144 | 32 | 15 | 15 | 15 | 1 | 6 | 6 | 7 | 361 | 59 |
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157 | 23 | 15 | 15 | 15 | 1 | 6 | 6 | 8 | 333 | 39 |
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149 | 28 | 15 | 15 | 15 | 1 | 6 | 6 | 7 | 354 | 58 |
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144 | 31 | 15 | 15 | 15 | 1 | 6 | 6 | 8 | 371 | 53 |
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152 | 25 | 17 | 17 | 17 | 2 | 7 | 7 | 7 | 309 | 36 |
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157 | 26 | 17 | 17 | 17 | 2 | 7 | 7 | 7 | 212 | 23 |
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166 | 22 | 15 | 15 | 15 | 2 | 7 | 6 | 6 | 455 | 34 |
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164 | 17 | 15 | 15 | 15 | 2 | 8 | 7 | – | 131 | 11 |
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141 | 36 | 17 | 17 | 17 | 2 | 6 | 7 | 7 | 279 | 39 | – | |
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144 | 24 | 17 | 17 | 17 | 2 | 7 | 7 | 6 | 278 | 35 |
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141 | 24 | 17 | 17 | 17 | 2 | 6 | 6 | 6 | 352 | 48 |
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143 | 21 | 17 | 17 | 17 | 2 | 6 | 6 | – | 101 | 19 | – | |
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129 | 35 | 17 | 17 | 17 | 2 | 5 | 6 | 6 | 266 | 65 | – | |
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– | – | – | – | – | – | – | – | – | 355 | 46 |
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– | – | – | – | – | – | – | – | – | 295 | 63 |
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128 | 36 | 17 | 17 | 17 | 2 | 7 | 7 | 6 | 219 | 48 |
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143 | 26 | 17 | 17 | 17 | 2 | 7 | 7 | 5 | 278 | 33 |
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132 | 35 | 17 | 17 | 17 | 2 | 7 | 7 | 6 | 297 | 61 |
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125 | 35 | 17 | 17 | 17 | 2 | 7 | 7 | 6 | 231 | 52 |
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145 | 20 | 17 | 17 | 17 | 2 | 7 | 7 | 6 | 205 | 22 |
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129 | 34 | 17 | 17 | 17 | 2 | 7 | 6 | 5 | 197 | 39 |
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136 | 39 | 16 | 17 | 17 | 2 | 7 | 6 | 5 | 114 | 22 |
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134 | 29 | 17 | 17 | 17 | 2 | 7 | 7 | 6 | 189 | 28 |
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139 | 24 | 17 | 17 | 17 | 2 | 7 | 7 | 5 | 329 | 36 |
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149 | 21 | 17 | 17 | 17 | 2 | 7 | 7 | 6 | 152 | 17 | – | |
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130 | 37 | 17 | 17 | 17 | 2 | 7 | 7 | 6 | 126 | 19 | – | |
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140 | 25 | 17 | 17 | 17 | 2 | 7 | 7 | – | 118 | 15 |
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138 | 24 | 17 | 17 | 17 | 2 | 7 | 7 | – | 335 | 36 |
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136 | 34 | 17 | 18 | 17 | 2 | 7 | 7 | 6 | 242 | 53 |
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129 | 35 | 15 | 17 | 17 | 2 | 7 | 7 | 5 | 242 | 53 |
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149 | 41 | 17 | 17 | 17 | 2 | 7 | 7 | 5 | 232 | 53 |
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156 | 30 | 17 | 17 | 17 | 2 | 7 | 7 | 5 | 241 | 34 |
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136 | 34 | 17 | 17 | 17 | 2 | 7 | 7 | 5 | 129 | 23 |
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168 | 34 | 19 | 17 | 17 | 2 | 6 | 6 | 3 | 272 | 40 |
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177 | 31 | 17 | 17 | 17 | 2 | 6 | 6 | 5 | 1060 | 116 |
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138 | 28 | 17 | 17 | 17 | 2 | 6 | 7 | 6 | 252 | 36 |
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141 | 32 | 17 | 17 | 17 | 2 | 6 | 6 | 6 | 272 | 46 |
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129 | 42 | 16 | 17 | 17 | 2 | 6 | 6 | 6 | 219 | 62 |
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139 | 32 | 17 | 17 | 17 | 2 | 7 | 7 | 5 | 204 | 32 |
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131 | 34 | 17 | 17 | 17 | 2 | 7 | 7 | 6 | 200 | 44 |
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140 | 40 | 17 | 17 | 17 | 2 | 7 | 7 | 6 | 210 | 46 |
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148 | 17 | 17 | 17 | 2 | 7 | 5 | 211 | 37 |
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127 | 38 | 17 | 17 | 17 | 2 | 7 | 7 | 5 | 209 | 46 |
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144 | 29 | 15 | 15 | 15 | 1 | 5 | 6 | 11 | 300 | 35 |
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144 | 23 | 15 | 15 | 15 | 1 | 5 | 7 | 10 | 321 | 42 | – | |
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174 | 35 | 17 | 17 | 17 | 2 | 7 | 7 | 6 | 586 | 86 |
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144 | 39 | 17 | 17 | 17 | 1 | 7 | 6 | 6 | 239 | 45 |
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149 | 37 | 17 | 17 | 17 | 2 | 7 | 7 | 7 | 239 | 51 |
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133 | 34 | 18 | 17 | 17 | 2 | 7 | 6 | 5 | 269 | 55 |
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144 | 25 | 17 | 17 | 17 | 2 | 7 | 7 | 6 | 300 | – |
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147 | 26 | 17 | 17 | 17 | 2 | 7 | 6 | 5 | 225 | 28 |
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144 | 24 | 17 | 17 | 17 | 2 | 7 | 6 | 5 | 217 | 28 |
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137 | 36 | 17 | 17 | 17 | 2 | 7 | 6 | 5 | 239 | 53 |
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163 | 29 | 17 | 17 | 17 | 2 | 7 | 7 | 7 | 222 | 27 |
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147 | 40 | 17 | 17 | 17 | 2 | 7 | 7 | 7 | 178 | 37 |
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152 | 26 | 17 | 17 | 17 | 2 | 7 | 7 | – | 335 | 45 |
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163 | 28 | 17 | 17 | 17 | 2 | 7 | 7 | – | 212 | 21 |
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|
|
45 | 17 | 17 | 17 | 2 | 7 | 6 | – | 344 | 41 | – | ||
|
165 | 41 | 17 | 17 | 17 | 2 | 7 | 7 | 5 | 110 | 20 |
|
|
|
186 | 27 | 17 | 17 | 17 | 2 | 7 | 7 | 6 | 294 | 38 |
|
|
|
146 | 21 | 15 | 15 | 15 | 2 | 7 | 6 | 6 | 200 | 23 |
|
|
|
147 | 42 | 17 | 17 | 17 | 2 | 6 | 7 | 5 | 273 | 59 |
|
|
|
128 | 37 | 17 | 17 | 17 | 2 | 7 | 7 | 7 | 188 | 40 |
|
|
|
145 | 20 | 17 | 17 | 17 | 2 | 6 | 7 | 5 | 262 | 25 |
|
|
|
141 | 35 | 17 | 17 | 17 | 2 | 6 | 7 | 5 | 272 | 55 |
|
|
|
137 | 38 | 17 | 17 | 17 | 2 | 6 | 7 | 5 | 117 | 21 |
|
|
|
138 | 35 | 17 | 17 | 17 | 2 | 7 | 7 | 5 | 253 | 55 |
|
|
|
129 | 33 | 17 | 17 | 17 | 2 | 7 | 6 | 5 | 122 | 23 |
|
|
|
134 | 37 | 17 | 17 | 17 | 2 | 7 | 7 | – | 274 | 68 |
|
|
|
134 | 36 | 17 | 17 | 16 | 2 | 7 | 7 | 6 | 223 | 35 |
|
|
|
149 | 24 | 17 | 17 | 17 | 2 | 7 | 7 | – | 191 | 21 |
|
|
|
134 | 39 | 17 | 17 | 17 | 2 | 7 | 7 | 7 | 332 | 67 |
|
|
|
171 | 43 | 17 | 17 | 17 | 2 | 7 | 7 | 7 | 290 | 50 |
|
|
|
163 | 43 | 17 | 17 | 17 | 2 | – | 7 | 7 | 249 | 50 |
|
|
|
– | – | – | – | – | – | – | – | – | 246 | 53 |
|
|
|
– | – | – | – | – | – | – | – | – | 237 | 51 |
|
|
|
156 | 46 | 15 | 15 | 15 | 2 | 7 | 6 | 6 | 196 | 45 |
|
|
|
186 | 40 | 17 | 17 | 17 | 2 | 7 | 7 | – | 262 | 42 |
|
|
|
143 | 16 | 15 | 15 | 15 | 1 | 6 | 5 | 8 | 443 | 34 |
|
|
|
135 | 27 | 15 | 15 | 15 | 1 | 6 | 6 | 9 | 337 | 48 |
|
|
|
146 | 25 | 15 | 15 | 15 | 1 | 5 | 6 | 9 | 309 | 39 |
|
|
|
156 | 19 | 15 | 15 | 15 | 1 | 6 | 7 | 11 | 392 | 37 |
|
|
|
149 | 17 | 15 | 15 | 15 | 1 | 6 | 6 | 11 | 139 | 13 |
|
|
|
143 | 19 | 15 | 15 | 15 | 1 | 6 | 6 | 13 | 230 | 21 |
|
|
|
166 | 25 | 17 | 17 | 17 | 2 | 6 | 7 | 7 | 214 | 23 |
|
|
|
151 | 29 | 17 | 17 | 17 | 2 | 7 | 7 | 7 | 283 | 35 |
|
|
|
160 | 27 | 17 | 17 | 17 | 2 | 7 | 7 | 8 | 315 | 35 |
|
|
|
ANF 2390 | 176 | 31 | 17 | 17 | 17 | 2 | 7 | 7 | 7 | 652 | 71 |
|
|
141 | 19 | 15 | 15 | 15 | 2 | 7 | 7 | 8 | 179 | 19 |
|
|
|
132 | 21 | 15 | 15 | 15 | 2 | 7 | 7 | 7 | 182 | 20 |
|
|
|
153 | 47 | 15 | 15 | 15 | 2 | 7 | 6 | 7 | 187 | 31 |
|
|
|
166 | 41 | 15 | 15 | 15 | 2 | 7 | 7 | 6 | 370 | 68 |
|
|
|
158 | 48 | 16 | 16 | 16 | 2 | 7 | 7 | 7 | 349 | 87 |
|