Research Article |
Corresponding author: Pier Cacciali ( pier_cacciali@yahoo.com ) Academic editor: Johannes Penner
© 2018 Pier Cacciali, Mariana Morando, Luciano Javier Avila, Gunther Koehler.
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:
Cacciali P, Morando M, Avila LJ, Koehler G (2018) Description of a new species of Homonota (Reptilia, Squamata, Phyllodactylidae) from the central region of northern Paraguay. Zoosystematics and Evolution 94(1): 147-161. https://doi.org/10.3897/zse.94.21754
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Homonota is a gecko distributed in central and southern South America with 12 species allocated in three groups. In this work, we performed molecular and morphological analyses of samples of Homonota from the central region of northern Paraguay, comparing the data with those of related species of the group: H. horrida and H. septentrionalis. We found strong molecular evidence (based on 16S, Cyt-b, and PRLR gene sequences) to distinguish this lineage as a new species. Morphological statistical analysis showed that females of the three species are different in metric characters (SVL and TL as the most contributing variables), whereas males are less differentiated. No robust differences were found in meristic characters. The most remarkable trait for the diagnosis of the new species is the presence of well-developed keeled tubercles on the sides of the neck, and lack of a white band (crescent-shaped) in the occipital area, which is present in H. horrida and H. septentrionalis. Nevertheless, in our sample, we found three specimens (one juvenile and two young adults) that exhibit the white occipital band. Thus, this character seems only reliable in adults of the new species. The new species is parapatric to H. septentrionalis, both inhabiting the Dry Chaco of Paraguay.
Dry Chaco, Gekkota , phylogeny, South America, taxonomy
Homonota is a gecko, inhabiting mainly xeric and rocky areas in central and southern South America (
Four species of Homonota are recorded in Paraguay: H. borellii, H. rupicola, H. horrida, and H. septentrionalis. The most commonly known species was Homonota horrida recorded for the “Chaco” (
After the description of H. septentrionalis, the same authors continued to study and analyze the taxonomy of Paraguayan samples of Homonota from the Chaco, within the framework of a barcoding initiative of the herpetofauna from Paraguay. We performed genetic and morphological analyses among different populations of Homonota from the central region of northern Paraguay. Based on genetic and morphological differences, and applying a species delimitation algorithm, we found enough differences to consider these new samples as a different taxonomic unit from those previously recorded for Paraguay. We present here a detailed analysis along with the description of this new species.
We extracted DNA from three samples of Homonota from the central area of northern Paraguay (Occidental Region), which were compared with available sequences of the remaining members of the genus (except H. fasciata) to assess its taxonomic relationships in the gene tree. We sequenced fragments of mitochondrial genes rRNA 16S and Cytochrome b (Cytb) and the nuclear gene prolactine receptor (PRLR). Samples used and GenBank accession numbers are specified in Table
Specimens used for genetic analyses and GenBank accession numbers for every gene. Asterisks (*) indicate tissue samples without voucher. Numbers in bold are samples generated for this work.
Species | Voucher | 16S | Cytb | PRLR | GenSeq Nomenclature |
Homonota andicola |
|
MD | KJ484188 | KJ484274 | genseq-3 |
|
MD | KJ484189 | KJ484275 | genseq-3 | |
Homonota borellii |
|
MD | KJ484205 | KJ484276 | genseq-4 |
|
MD | KM677796 | MD | genseq-4 | |
|
MD | KJ484206 | KJ484277 | genseq-4 | |
Homonota darwinii |
|
MD | KJ484191 | MD | genseq-3 |
|
MD | MD | KJ484278 | genseq-3 | |
|
MD | KJ484190 | MD | genseq-3 | |
Homonota horrida | BYU 47941 | MF278828 | KJ484192 | MG950402 | genseq-3 |
|
MD | KM677795 | MD | genseq-3 | |
|
MF278829 | MD | MG950403 | genseq-3 | |
|
MF278830 | MD | MG950404 | genseq-3 | |
|
MD | KJ484208 | MD | genseq-3 | |
Homonota rupicola |
|
MD | KJ484193 | KJ484281 | genseq-3 |
|
MD | KJ484194 | KJ484282 | genseq-3 | |
Homonota septentrionalis |
|
MD | MF278843 | MF278849 | genseq-2 |
|
MD | MF278844 | MF278850 | genseq-2 | |
|
MF278831 | MD | MG950405 | genseq-3 | |
|
MF278832 | MD | MD | genseq-1 | |
|
MF278833 | MD | MG950406 | genseq-2 | |
Homonota taragui |
|
MD | KJ484195 | KJ484283 | genseq-3 |
|
MD | KJ484196 | KJ484284 | genseq-3 | |
Homonota underwoodi |
|
MD | KJ484197 | KJ484286 | genseq-4 |
|
MD | KJ484198 | KJ484297 | genseq-4 | |
Homonota uruguayensis | UFRGS 2139 | MD | MD | KJ484296 | genseq-4 |
UFRGS 5769 | MD | KM677689 | MD | genseq-4 | |
UFRGS 5770 | MD | KM677690 | MD | genseq-4 | |
UFRGS 5771 | MD | KM677691 | MD | genseq-4 | |
Homonota whitii |
|
MD | KJ484199 | MD | genseq-4 |
|
MD | KJ484200 | MD | genseq-4 | |
Homonota williamsii |
|
MD | KJ484201 | KJ484287 | genseq-3 |
|
MD | KJ484202 | KJ484288 | genseq-2 | |
Homonota sp. n. |
|
MD | MG950409 | MG950407 | genseq-2 |
|
MG947388 | MG950410 | MG950408 | genseq-2 | |
|
MD | MG950411 | MD | genseq-2 | |
Outgroups | |||||
Garthia gaudichaudii | E61214 | MD | FJ985045 | MD | |
IBE_G1(1) | MD | MD | KJ484289 | ||
Phyllopezus przewalskii | LG1093 | JN935567 | JQ826890 | JQ825640 | |
|
MD | KJ484203 | MF278849 |
Tissue samples were first washed for 15 h with 50 ml Phosphate-buffered saline (PBS) (diluted of 1:9 PBS: H2O). The DNA extraction was carried out with the DNeasey kit of Qiagen. We used 25 μl of reaction mix for every sample for the PCR (except for PRLR where we used 20 μl). Reagents and concentrations for the PCR mix for the amplification of every gene, are provided in Suppl. material 1: Appendix S1. Primers (produced by Eurofins MWG Operon) used for amplification and sequencing, along with PCR conditions for each gene are detailed in Suppl. material 1: Appendix S2.
We used SeqTrace 0.9.0 (
We performed two phylogenetic analyses, first using a Maximum Likelihood (ML) approach, and then a Bayesian inference (BI) to compare the trees topologies. These analyses were made for each gene individually and for a concatenated dataset of the three genes together. For the ML analysis we used IQ-Tree (
We assessed the degree of intraspecific divergence within the alignment (removing the outgroups) with the species delimitation test ABGD (
Additionally, we generated morphological data for 13 specimens (7 males and 6 females) of the new species and taxa with similar pattern (related taxa of the horrida group) looking for potential diagnostic characters. Thus, we used for comparison H. horrida (7 males and 5 females) and H. septentrionalis (10 males and 12 females), using standard variables (continuous data expressed in mm) already used by
SVL snout–vent length, from tip of snout to vent.
TrL trunk length, distance from axilla to groin from posterior edge of forelimb insertion to anterior edge of hind limb insertion.
FL foot length, from the tip of the claw of the 4th straightened toe to the back of the heel.
TL tibial length, measured between the level of the knee and the level of the heel, as shown by
AL arm length, from tip of claws of the 3rd finger to elbow.
HL head length, distance between anterior edge of auditory meatus and snout tip.
HW head width, taken at the level of the temporal region, corresponding to the widest part of the head.
HH head height, maximum height of head, at level of parietal area.
END eye–nostril distance, from the anterior edge of the eye to the posterior edge of the nostril.
ESD eye–snout distance, from the anterior edge of the eye to the tip of the snout.
EMD eye–meatus distance, from the posterior edge of the eye to the anterior border of the ear opening.
ID interorbital distance, shortest distance between orbits.
IND internostril distance, shortest distance between nares.
DT number of keeled dorsal tubercles from occipital area to cloaca level.
TVS number of transversal rows of ventral scales, counted longitudinally at midline from the chest (shoulder level) to inguinal level.
LVS number of longitudinal rows of ventral scales, counted transversally at midbody.
SL number of supralabial scales.
IL number of infralabial scales.
4TL number of lamellae under the fourth toe.
3FL number of lamellae under the third finger.
Measurements were taken with digital calipers (precision 0.01), but only the first decimal considered to limit discrepancies. For the morphological analyses only specimens of ~40 mm or larger were included. When paired structures exist, data are presented in left/right orientation, and only the left side was used for statistical analyses. In the color descriptions, the capitalized colors and the color codes (in parentheses) are those of
We compared the morphological variation among species through a discriminant function analysis (DFA), testing the normality of the variables with a Shapiro-Wilk (W) test (
The final alignments of 16S, Cytb, and PRLR consisted of 539, 793, and 457 bp, respectively. Alignments and trees are available at TreeBASE (ID: 22305). The best substitution model for 16S was GTR+G, for Cytb TVM+I(1stpos)|TIM+G(2ndpos)|SYM+G(3rdpos), and for PRLRK81(1stpos)|GTR+G(2nd+3rdpos). The complete table with scores is provided in Suppl. material 1: Appendix S4. The topology of the ML (Suppl. material 1: Fig. S1) and BI (Suppl. material 1: Fig. S2) trees using 16S coincide in recognizing three clusters, but the ML tree shows the new species as a sister clade to H. septentrionalis, whereas BI shows a trichotomy including H. horrida, H. septentrionalis, and the new species. Trees of ML and BI based on Cytb have the same topology (Suppl. material 1: Figs S3–S4), with strong support values. In these trees H. andicola and H. whitii are sister clades, as are H. darwinii and H. williamsii, and they are sister to the remaining Homonota species. The borellii group shows H. uruguayensis as sister to H. borellii + H. rupicola + H. taragui. Finally, within the horrida group, H. underwoodi appears as the sister to the remaining Homonota with banded coloration pattern. In this part of the tree H. horrida is rendered as sister to the clade H. septentrionalis plus the new taxon. The topologies of the ML and BI trees using PRLR are also the same (Suppl. material 1: Figs S5–S6). Species in the whitii group are clustered together, and the borellii group also shows monophyly but with a unresolved polytomy. In the horrida group H. underwoodi is also suggested as sister to the remaining species, with the new species and H. septentrionalis showing the most recent divergence. The trees using the concatenated dataset (with both ML and BI) show similar branch arrangement previously observed in trees of individual genes (Suppl. material 1: Figs S7–S8). Only two samples (UFRGS 2139 of H. uruguayensis and
The analysis of intraspecific genetic divergence with ABGD results in 12 groups (Suppl. material 1: Appendix S5), which represent nearly all described species (except H. fasciata) and the new species, providing evidence for its recognition as a distinct taxon. This is highly congruent with the clusters shown by the gene trees. The species tree shows consensus in the clusters of the three groups of the genus Homonota, with slight differences in the branch arrangements. The lower value (density of green lines in Figure
All continuous morphological variables had normal distributions (Table
Normality Shapiro-Wilk (W) values for metric (above) and meristic (below) characters showing the p value. Values shaded in gray did not reach normality. See Methods section for reference to the acronyms.
Continuous | |||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|
SVL | TrL | FL | TL | AL | HL | HW | HH | END | ESD | EMD | ID | IND | |
W | 0.978 | 0.979 | 0.957 | 0.987 | 0.982 | 0.979 | 0.976 | 0.983 | 0.979 | 0.967 | 0.969 | 0.975 | 0.952 |
p | 0.503 | 0.506 | 0.087 | 0.849 | 0.696 | 0.575 | 0.401 | 0.758 | 0.555 | 0.199 | 0.284 | 0.400 | 0.050 |
Discrete | |||||||||||||
DT | TVS | LVS | SL | IL | 4TL | 3FL | |||||||
W | 0.962 | 0.971 | 0.965 | 0.779 | 0.788 | 0.913 | 0.948 | ||||||
p | 0.109 | 0.291 | 0.164 | 3.05E-7 | 4.65E-7 | 0.008 | 0.023 |
Most contributing continuous (Cont.) and discrete (Disc.) variables (highlighted in bold) for Axis 1–3 of the DFA.
Variables | Axis 1 | Axis 2 | Axis 3 | |
---|---|---|---|---|
Cont. | SVL | 0,417 | 1,202 | –3,447 |
TrL | –0,187 | 0,690 | –1,798 | |
FL | 0,132 | 0,401 | –0,479 | |
TL | 0,228 | 0,193 | –0,417 | |
AL | 0,201 | 0,213 | –0,553 | |
HL | –0,017 | 0,357 | –0,605 | |
HW | –0,128 | 0,218 | –0,525 | |
HH | –0,199 | 0,187 | –0,258 | |
END | –0,021 | 0,099 | –0,193 | |
ESD | –0,052 | 0,111 | –0,392 | |
EMD | 0,144 | 0,134 | –0,221 | |
ID | –0,028 | –0,013 | –0,302 | |
IND | 0,002 | 0,127 | –0,050 | |
Disc. | DT | 1.424 | –1.232 | –1.338 |
TVS | 2.166 | 1.825 | 0.921 | |
LVS | <0.001 | 1.482 | –1.255 |
There is a strong molecular congruence in the recognition of 12 taxa within the genus Homonota (three of them with a banded coloration pattern), which added to the significant differences among the three species with banded pattern based on the DFA, and the additional morphological distinctions discussed below are used to identify the new taxon described here.
Occidental Region of Paraguay, indicating the political division, showing the known records for Homonota septentrionalis (white circles) and the analyzed records of Homonota marthae (black circles), and its type locality (star). Circle with a white cross, indicates origin of the genetic samples. High resolution elevation base map (30 seconds resolution) taken from Consortium for Spatial Information (CGIAR-CSI) available on http://www.diva-gis.org/gdata (
Paraguay: Boquerón Department: Comunidad Ayoreo Jesudi (
A species of Homonota assigned to the horrida group given its relationship (based on molecular evidence) with H. horrida, and by the color pattern composed of a vertebral and five to seven transversal clear lines appearing as a banded Homonota smilar to H. horrida and H. septentrionalis. Homonota marthae has a robust body, and prominently keeled tubercles disposed in four to eight longitudinal rows on the dorsum.
Homonota marthae can be differentiated from all species in the genus, except H. fasciata, H. horrida, H. darwinii, and H. septentrionalis by the color pattern of transversal bands on the dorsum (reticulated pattern in the remaining species). Homonota marthae is further differentiated from H. andicola, H. whitii, H. darwinii, and H. underwoodi by the keeled scales along the whole dorsum (vs. smooth dorsal scales in H. andicola, H. whitii, and H. underwoodi, and keeled scales restricted to the posterior part of the dorsum in H. darwinii). It differs from H. fasciata by having a serrated edge of the auditory meatus (vs. smooth anterior margin in H. fasciata); presence of one or two enlarged tubercles on the upper edge of the auditory meatus (vs. no enlarged tubercles in H. fasciata); and a smaller size of the postmental scales (vs. postmentals of the size of the first infralabials in H. fasciata). Homonota marthae differs from H. horrida by the higher position of the ear opening in relation to the level of the mouth (vs. lower positioned in H. horrida); from H. septentrionalis by more developed keeled tubercles on the sides of the neck (Fig.
Adult female, SVL 56 mm (4.1 times the HL), TrL 26 mm, tail length 70 mm, FL 11.0 mm, TL 9.6 mm, AL 13.3 mm, HL 13.6 mm, HW 10.8 mm, HH 8.3 mm, END 4.2 mm, ESD 5.9 mm, EMD 4.7 mm, ID 4.6 mm, IND 2.0 mm; rostral wider (2.7 mm) than high (1.5 mm) with a median groove covering the upper two thirds of the scale; nares surrounded by rostral, supranasal, and postnasal; SL 8/8; one elongated tubercular scale on the mouth commissure; muzzle slightly convex, covered by large homogeneous juxtaposed scales; head covered with big homogeneous juxtaposed scales on the dorsal area, intermixed with small granules; superciliary scales imbricated, associated to spiny-like scales on the posterior half of the orbit; scales on lateral surface of the head heterogeneously covered with strongly conical tubercles intermixed with small granules; auditory meatus oblique and with serrated edge, and one large elongated scale on the upper border; IL 6/6, the last less than half the size of the others; mental bell-shaped; two postmentals less than twice the size of the following posterior scales, contacting the mental, the first IL, and four posterior scales; scales under the head gradually reducing in size posteriorly; dorsal and lateral parts of the neck with granular juxtaposed scales mixed with tubercles; ventral side of the head covered by imbricate cycloid scales; body dorsally covered with 14–16 rows of strongly keeled scales, separated by one to two small granules in the pleural areas, and three to four granules in the vertebral area; ventral scales cycloid and imbricate arranged in 16 longitudinal rows at midbody; suprascapular, axillary, inguinal regions, and cloacal opening surrounded by small imbricate granules; anterior and dorsal surfaces of limbs covered by large imbricate scales, keeled on the dorsal surface; posterior region of limbs covered by small juxtaposed granules; ventral surface of forelimbs with juxtaposed granules, and ventral surface of hind limbs with large imbricate scales; subdigital lamellae of hands starting from pollex were recorded as follows: 8/8 – 10/12 – 15/13 – 16/16 – 11/11; subdigital lamellae of feet starting from hallux were recorded as follow: 15/13 – 19/17 – 15/16 – 12/12 – 9/9; tail with large imbricated and mucronate scales, 10–12 per caudal whorl.
After five years in preservative, the coloration was recorded as follows: Head Mikado Brown (42) with Warm Sepia (40) speckling on the dorsal surface; Warm Sepia (40) on the sides, with a Light Buff (2) line from nares to orbit, and continuing behind the orbit above the temporal region; supralabials and infralabials Medium Neutral Gray (298) with suffusions of Smoky White (261); and Fawn Color (258) ventrally. Dorsal background color of the body Beige (254) with Vandyke Brown (282) splotches, and poorly defined Chamois (84) transversal lines; Drab (19) laterally, with Dusky Brown (285) and Pale Buff (1) splotches; and Ground Cinnamon (270) ventrally, with Smoky White (261) suffusions. Tail with Grayish Horn Color (268), Sepia (286), and Cream White (52) transversal bands dorsally; Drab (19) laterally; and Smoky White (261) ventrally. Limbs dorsally covered with a reticulation of Drab (19), Chamois (84), and Dusky Brown (285), ventrally grading to Fawn Color (258) in forelimbs, and Ground Cinnamon (270) with suffusions of Smoky White (261) in hind limbs.
Coloration in life of a young male (
Coloration in life of a juvenile female (
One juvenile (
SVL 36–59 mm; TrL 16–27 mm (43.8–48.2% of SVL in females, 40.7–46.7% in males); FL 9–11 mm (x̅ 10±0.36) in females, 7–11 mm (x̅ 8.7±0.52) in males; TL 8.7–10.1 mm (x̅ 9.5±0.2) in females, 8–10.2 mm (x̅ 9.1±0.31) in males; AL 9.3–13.7 mm (x̅ 12.8±0.28) in females, 11.2–14 mm (x̅ 12.4±0.38) in males; HL 9.3–13.8 mm (x̅ 13.2±0.19) in females, 11.1–13.5 mm (x̅ 12.2±0.31) in males; HW 7.1–11.2 mm (79.4–88% of HL in females, 78.9–85.9% in males); HH 5.5–8.3 mm (52.6–61% of HL in females, 52.6–60.5% in males); END 2.8–5.1 mm (30.8–35.1% of HL in females, 31.4–38.9% in males); ESD 3.9–6.1 mm (40.3–45% of HL in females, 40.5–46.5% in males); EMD 3.1–5 mm (31.3–34.7% of HL in females, 33.8–37% in males); ID 3.8–5.8 mm (33.8–40% of HL in females, 37.1–44.7% in males); IND 1.4–2.1 mm (12.2–16% of HL in females, 11.8–14% in males); SL 5–8; one or two elongated tubercular scales on the mouth commissure; auditory meatus with one large scale on the upper border; IL 5–7; 14–20 longitudinal rows of ventral scales at mid-body; 34–49 transversal rows of ventral scales.
This species is named in honor of our indefatigable colleague Martha Motte, who is not only dedicated to safekeeping the herpetological collection of the “Museo Nacional de Historia Natural del Paraguay”, but also does a great job in providing selfless support to scientists that are striving to improve the knowledge of the Paraguayan herpetofauna.
Homonota marthae is known from the central area of the Paraguayan Dry Chaco in the Department of Boquerón (Fig.
This species is a dry forest inhabitant, but it is also frequently found in human dwellings.
The diversity of species groups within the genus Homonota was explored in the last decade, and resulted in the description of H. williamsii (
No obvious external synapomorphy is known to diagnose the horrida group. Three of the four species (H. horrida, H. septentrionalis, and H. marthae) have a pattern characterized by transversal body bands and the presence of a vertebral line. This coloration is different from the remaining species of the genus. The fourth species of the horrida group, H. underwoodi, has homogeneous body scalation and a completely different pattern, and therefore H. horrida and H. underwoodi were considered not to be in the same group (
The most obvious external difference between H. marthae and its presumed closest relative, H. septentrionalis, is the lack of a white occipital band in the former taxon, although we found some specimens (mainly juveniles or hatchlings) of H. marthae that do have the occipital band. Given that this white occipital band is also present in H. horrida, it could be a plesiomorphic character, and therefore the lack of it could be interpreted as the derived state.
Both species seem to inhabit in parapatry the Dry Chaco in Paraguay, and although a major revision of the whole distribution of the group is needed in order to know their actual ranges, H. septentrionalis is distributed in the north-westernmost part of the Dry Chaco, whereas H. marthae occurs in the central and easternmost areas of the Dry Chaco. Due to the lack of evident geographic barriers between these two species and considering their relatively low morphological variation (especially in males), they remained recognized as a single taxonomic unit until now. Parapatric speciation or breaks to gene flow without evident geographic barriers were observed and discussed by
The degree of genetic differentiation between these two species is evident, and larger than the degree of morphological differences. Small morphological differentiation or even complete crypsis is common for many organisms, especially when they use the same ecological niche. Specifically for geckos, a recent study showed that it is difficult to find morphological diagnostic characters that match those observed by genetic evidence, as it is the case of the genera Garthia and Homonota, which are very similar morphologically (
Finally, Homonota marthae is a common species that resists human perturbation and can be found in rural environments, and although its actual distribution limits are not yet known, and more revisions are needed to target this issue, probably the records of “Homonota fasciata” from Defensores del Chaco National Park referred by
Information to generate the key was based on
1 | Coloration based on irregular or reticulated pattern | 2 |
– | Coloration pattern composed of transversal bands | 10 |
2 | Dorsal scales homogeneously smooth | 3 |
– | Dorsal scales smooth and granular mixed with series of enlarged keeled scales | 5 |
3 | Ventral surface of the body immaculate due to lack of chromatophores | H. underwoodi |
– | Ventral surface of the body pigmented with chromatophores | 4 |
4 | 43–49 scales around midbody | H. andicola |
– | 55–59 scales around midbody | H. whitii |
5 | Series of keeled scales restricted to the posterior half of the dorsum | H. darwinii |
– | Series of keeled scales uniformly extended along the whole dorsum | 6 |
6 | Dorsal surface of thighs with keeled scales | 7 |
– | Dorsal surface of thighs with smooth scales | 8 |
7 | Dorsal surface of arms with keeled scales; temporal region with enlarged keeled scales | H. uruguayensis |
– | Dorsal surface of arms with smooth cycloid scales; temporal region homogeneously covered by granular scales | H. taragui |
8 | 146–161 dorsal scales from occipital area to the level of the cloaca; oblique ear opening | H. williamsii |
– | 94–139 dorsal scales from occipital area to the level of the cloaca; round ear opening | 9 |
9 | 45–50 scales around midbody; dorsal surface of the tail with a pattern of thin speckling | H. borellii |
– | 54–63 scales around midbody; dorsal surface of the tail with a pattern of black blotches | H. rupicola |
10 | Edge of ear opening smooth, without enlarged tubercular scales around; postmentals about five times larger than the scales behind it | H. fasciata |
– | Edge of ear opening serrated, with one or two tubercular scales above; postmentals twice larger than the scales behind it | 11 |
11 | Tubercles on the dorsal and lateral sides of the neck poorly developed; occipital area with a wide whitish crescent-shaped mark | H. septentrionalis |
– | Tubercles on the dorsal and lateral sides of the neck well developed; occipital coloration variable | 12 |
12 | Ear opening above the mouth level; occipital area with homogeneous coloration or with a faint reticulation in adults | H. marthae |
– | Ear opening at the level of the mouth; occipital area with a wide whitish crescent-shaped mark | H. horrida |
Collecting (N° 04/11) and export (N° 02/12) permits were provided by Secretaría del Ambiente. GK thanks Dulcy Vázquez and Thomas and Sabine Vinke for help during fieldwork. We are grateful to Martha Motte (Museo Nacional de Historia Natural del Paragua -
Examined specimens
Homonota horrida
ARGENTINA: La Pampa: Ruta Provincial 1, 23.6 km W from intersection with Ruta Nacional 151 (
Homonota septentrionalis
PARAGUAY: Boquerón: Cruce San Miguel (
Acronyms
Supplementary information
Data type: Adobe PDF file
Explanation note: Description of a new species of Homonota (Reptilia, Squamata, Phyllodactylidae) from the central region of northern Paraguay