Echinosaura embera, Vásquez-Restrepo & Daza, 2025

Vásquez-Restrepo, Juan D. & Daza, Juan M., 2025, A new species of hedgehog-lizard of the genus Echinosaura (Squamata: Gymnophthalmidae) from Colombia and Panama with a reevaluation of the conservation status of the genus, Zootaxa 5583 (1), pp. 128-142 : 133-137

Echinosaura embera sp. nov.

LSID: urn:lsid:zoobank.org:act:BC88160A-51E8-4A2E-9FE3-E70BCE3B5B3A

Echinosaura horrida palmeri (in part)— Uzzell (1965)

Echinosaura palmeri (in part)— Fritts et al. (2002)

Echinosaura palmeri (in part)—Vásquez-Restrepo et al. (2020)

Holotype. MHUA-R 12591 : an adult male ( Fig. 3 ; Fig. 4A ) from Finca La Mejía , vereda Juradó , Chigorodó municipality, Antioquia department, Colombia (76° 35’ 38.328” W, 7° 31’ 30.323” N, WGS84 datum, at 94 m above sea level), collected on 2013 by Juan M. Daza ( JMD 1825 ).

Paratypes. A series encompassing 16 adult specimens (14 males + 2 females). MHUA-R 10702 : an adult male from La Playona , Acandí municipality, Chocó department, Colombia (77° 13’ 1.920” W, 8° 26’ 57.281” N, WGS84 datum, at 10 m above sea level), collected on 2002 by Juan M. Daza. MHUA-R 12306 : an adult male from Finca La Mejía , vereda Juradó , Chigorodó municipality, Antioquia department, Colombia (76° 33’ 36.719” W, 7° 31’ 15.959” N, WGS84 datum, at 56 m above sea level), collected on 2010 by Juan P. Hurtado ( JPH 342 ) . MHUA-R 12328 : an adult male from Camagüey , Mutatá municipality, Antioquia department, Colombia (76° 24’ 46.155” W, 7° 11’ 49.679” N, WGS84 datum, at 304 m above sea level), collected on 2010 by Claudia Molina (CMZ 384) . MHUA-R 13256 : an adult female from the border between the corregimientos of Campo Bonito (Quibdó municipality) and Chachajo (Alto Baudó municipality) , Chocó department, Colombia (76° 55’ 38.938” W, 5° 42’ 35.650” N, WGS84 datum, at 187 m above sea level), collected on February 18, 2013 by Guido F. Medina ( GFM 668 ) . CBUCES-D 921 : an adult male from Piedras Blancas , Chigorodó municipality, Antioquia department, Colombia (76° 33’ 47.870” W, 7° 43’ 7.910” N, WGS84 datum, at 123 m above sea level), collected on May 24, 2023 by Esteban Garzón-Franco ( EGF 002 ) . CBUCES-D 1867 : an adult male from Quebrada Chorro Frío , vereda Guaduas , Carmen de Atrato municipality, Chocó department, Colombia (76° 11’ 32.520” W, 5° 43’ 22.950” N, WGS84 datum, at 1295 m above sea level), collected on March 10, 2024 by Esteban Garzón-Franco (EGF 401) . CBUCES-D 1876 : an adult male from Finca La Guaduala , vereda Guaduas , Carmen de Atrato municipality, Chocó department, Colombia (76° 11’ 42.990” W, 5° 44’ 9.570” N, WGS84 datum, at 1000 m above sea level), collected on March 10, 2024 by Esteban Garzón-Franco ( EGF 410 ) . KUH 113622 : an adult male from SE slope Cerro Pirre , Darien province, Panama (approx. 77° 42’ 9.000” W, 7° 55’ 41.070” N, WGS84 datum), collected on 1965 by Charles W. Myers ( CWM 5003 ) . AMNH-R 49187 , 49199 : adult males from Chalichiman’s Creek , Darien province, Panama (approx. 77° 46’ 53.760” W, 8° 47’ 34.660” N, WGS84 datum, at ca. 390 m above sea level), collected on February 1924. AMNH-R 49195 : an adult male from Camp Townsend , Darien province, Panama (approx. 77° 41’ 30.930” W, 8° 9’ 22.680” N, WGS84 datum, at ca. 10 m above sea level), collected on February 1924. AMNH-R 49201 : an adult female from Three Falls Creek , Darien province, Panama (approx. 77° 41’ 30.930” W, 8° 9’ 22.680” N, WGS84 datum, at ca. 10 m above sea level), collected on February 1924. CH 9058 : an adult male from Cerro Sapo , Darien province, Panama (78° 21’ 26.570” W, 7° 58’ 50.450” N, WGS84 datum, at 694 m above sea level), collected on June 2010 by Andrew Crawford et al. CH 9109 : an adult male from Cabecera del río Cana , campamento de Pirre , Darien province, Panama (approx. 77° 41’ 24.870” W, 7° 45’ 15.300” N, WGS84 datum, at 604 m above sea level), collected on July 2010 by Kelly Zamudio. CH 9117 : an adult male from Sendero al campamento de Pirre , Darien province, Panama (approx. 77° 41’ 24.870” W, 7° 45’ 15.300” N, WGS84 datum, at 604 m above sea level), collected on July 2010 by Karen Lips. CH 9151 : an adult male from Campamento Serranía de Pirre, Darien province, Panama (77° 41’ 24.870” W, 7° 45’ 15.300” N, WGS84 datum, at 604 m above sea level), collected on July 2010 by Myra Hughey et al .

Etymology. The specific epithet embera derives from the name of the indigenous people inhabiting in Panama and Colombia, from the Darien region along the Pacific coast in the Chocó. We chose this name to reflect the new species’ distribution, which closely resembles part of the historical territory occupied by the Emberá people. In particular, among the Emberá in western Colombia, there is an ethnic subgroup known as Emberá-Dobidá, referred to as “river people”. This designation beautifully aligns with the behavioral characteristics of our newly described species, which is frequently observed wandering along small creeks and streams. Thus, the name not only reflects the geographical overlap but also draws a poetic parallel between the “river people” and this creek-dwelling lizard. Here, we used this word as a noun in apposition (Art. 11.9.1.2 of the ICZN).

Distribution. This species is distributed in the Pacific versant of Colombia and Panama, along the Atrato River basin in Colombia, primarily in the Chocó department and part of the western portion of Antioquia, as well as the Darien region in both Colombia and Panama ( Fig. 2A ). Its altitudinal range extends from near sea level to approximately 1300 meters, with most occurrences below 800 meters ( Fig. 2B ).

Diagnosis. Echinosaura embera sp. nov. can be distinguished from all congeners by the combination of the following characteristics (in the same order as in Vásquez-Restrepo et al. 2020): (1) snout pointed; (2) internasal divided longitudinally; (3) frontonasals paired; (4) frontal single (rarely divided); (5) frontoparietals paired; (6) supraoculars two or three; (7) supralabials 4–5 (usually five); (8) infralabials 4–5 (usually four); (9) postmental single (rarely divided); (10) one pair of enlarged chin shields (rarely two pairs); (11) two paravertebral ridges, separated from each other by usually five or more small irregular scales; (12) spine-like scales forming oblique lines on lateral surface of body; (13) ventral scales squared, usually smooth (rarely keeled); (14) subdigital lamellae on the fourth finger 12–17; (15) subdigital lamellae on the fourth toe 21–25; (16) femoral pores per hind limb in males 6–10; (17) dorsal surface of tail with two oblique ridges per autotomic segment disposed in a V-shaped pattern ( Fig. 5 ), formed by a series of increasing in size scales from the anterior margin of each caudal segment (present in 92% of examined specimens, barely to none visible in juveniles); in lateral view, usually with only more developed scales on the distal portion of each segment, forming a ring; (18) subcaudals per caudal segment three (anterior third of tail excluded); (19) in life, dorsum brown or dark brown, relatively uniform, sometimes with yellowish marks and/or two spots more or less distinguishable at the base of tail ( Fig. 4B ); ventral surface of head and neck mainly light brown, except on the anterior part where there is a darker ground color with some transverse cream blotches extending to labial scales; ventral surface of body (including limbs) mainly light brown spotted with dark brown or cream, while that of tail almost completely dark; (20) hemipenes (in males) lacking enlarged hook-like spines on sides and asulcate face. The in life and in preservative coloration is similar, sometimes slightly darker in some preserved specimens.

Comparisons with similar species. Echinosaura embera sp. nov. can be distinguished from the other species of Echinosaura (in parenthesis) by having: three transverse rows of subcaudal scales per autotomic segment (four in E. brachycephala and five to six in E. orcesi ); internasal scale divided (single in E. brachycephala , E. horrida , E. fischerorum , and E. orcesi ); paravertebral ridges separated from each other by small irregular scales (juxtaposed in E. fischerorum and E. horrida ); frontal scale single (transversely divided in E. panamensis and rarely in E. keyi ); scales in the nuchal region spine-like but blunt-tipped (conical and sharp-tipped in E. fischerorum ); tail with small scales obliquely disposed forming a V shape on each autotomic segment (more developed scales only to the distal portion of each segment in E. brachycephala , E. palmeri , and E. panamensis , well-developed scales forming parallel crest in E. centralis and E. horrida , and well-developed conical scales forming whorls in E. keyi ). For additional traits see the taxonomic key in Vásquez-Restrepo et al. (2020) and the species comparison table in Yánez-Muñoz et al. (2021). Regarding hemipenial morphology, E. embera differs from other species by the absence of enlarged hook-like spines laterally and dorsally [see Pattern 2 in Fig. 6 ] (one or two pairs of enlarged hook-like spines in E. centralis , E. palmeri , and E. panamensis [see Pattern 1 in Fig. 6 ], three to five flounces of spinules on each side of the sulcus in E. horrida and E. fischerorum , condition of E. brachycephala , E. keyi , and E. orcesi unknown). It is probable that most species in the genus present an apical disc. However, due to the technical difficulty of removing the thin sheath of tissue at the basal crotch of the lobes without decapitating the hemipenis, most preparations tend to exhibit a “wrinkled pocket” structure in the apical portion, likely resulting from an incomplete eversion ( Uzzell 1965; Sales-Nunes 2011; Yánez-Muñoz et al. 2021).

Description of the holotype. An adult male; snout-vent length = 55 mm; trunk length = 26 mm, snout length = 5.10 mm; head length = 13.45 mm; head width = 7.62 mm; tail length (non-regenerated) = 96 mm. Head heartshaped and pointed. Rostral scale single. Internasals two, longer than wide, grooved, in contact with rostral anteriorly, nasals laterally, and frontonasals posteriorly. Nasals subtriangular, in contact with internasals and first supralabials. Frontonasals about three times longer than wide. Frontal single, grooved, wider anteriorly and thinner posteriorly. Supraoculars three, the first one in contact with the frontal posteriorly, preceded by a series of small irregular scales, the first two supraoculars larger than the third. Lower eyelid with a palpebral disc divided into three enlarged unpigmented scales on the left side and four on the right side. Loreal in contact with preciliary, the frontonasal and nasal on each side. Suboculars in four small rows, in contact with a fifth row formed by four large scales which contact the third to fifth supralabials. Frenocular scale present, in contact with the first enlarged subocular, the second supralabial, and the loreal. Frontoparietals paired, small and well defined. Parietal region covered with small irregular scales. Five supralabials on both sides. Three infralabials on the left side and four on the right. Mental posteriorly in contact with a postmental. Postmental trapezoidal, in contact with the first infralabial and the anterior half of the second infralabial on the right side, while in the left touching only the first infralabial. Chin shields large, in one pair, about three times longer than wide, in contact with the second and third infralabials on the right side, while with the first and second on the left. Gular region with small, semicircular and tubercular scales, becoming spine-like scales toward the posterior part. Dorsally two lines of spine-like scales extending on paravertebral region, parallel and continuous in the first third of body, becoming discontinuous to the tail, and separated from each other by four to six irregular small scales. Two additional rows of spine-like scales extending on upper costal region but forming a discontinuous and sinuous line. On the sides, there is a series of spine-like scales forming oblique lines. The spine-like scales on dorsum separated from each other by small, irregular or tubercular scales. Limbs pentadactyl, digits clawed. Subdigital lamellae on the fourth finger 14/13, subdigital lamellae on the fourth toe 21/23, in both cases including ungual sheath. Femoral pores seven on the left side and six on the right. Scales on pectoral region slightly keeled, rounded in the anterior portion, becoming six rows of squared scales toward the belly. Tail with two parallel ridges dorsally, formed by a series of three to four increasing in size scales from the posterior margin of each caudal segment, oblique towards the anterior margin forming a kind of V-shaped pattern. Tail complete, with three subcaudal scales per caudal segment. Hemipenes extracted [their location is unknown]. Dorsum light brown, barely yellowish blotches on limbs, and two cream spots at the base of the tail. Scales of ventral surface of head dark-brown on the anterior part, most of them with a cream spot on the center. Mental and labials dark with cream coloration on the edges of the scales. Suture between postmental and chin shields with a white transverse stripe. Scales on belly with an irregular chessboard pattern of black and cream scales. Ventral surface of tail dark brown from the base, except by a few whitish scales on the anterior and posterior parts. The coloration in life and in preservative is quite similar, slightly darker in the latter case.

Variation. A few of the examined specimens presented some degree of variation on their morphological features departing from the most common states. For instance, MHUA-R 12306 and IAvH-R 1534 specimens present a divided postmental scale. MHUA-R 12305 specimen has an additional division in one of its chin shields, while MHUA-R 12590 exhibited both chin shields divided to form two pairs. Moreover, MHUA-R 13256 presents a divided frontal scale and additional small irregular scales on the head. Here, we consider these like individual variations rather than character-states.

Hemipenial morphology. Hemipenial body globular and bilobed; sulcus spermaticus divided by a fleshy triangular fold at the lobular fork; the sulcate face bare, lacking spine flounces bordering the sulcus; on the sides towards the basal portion, there is a series of three to four small spines arranged in a transversal row; the asulcate face features two longitudinal rows of spines, each consisting of four to five small spines (non-enlarged or hook-like), usually accompanied by an additional row formed by two to four smaller spines on each side; towards the distal portion at the base of each lobe, between the sides and the asulcate face, there are approximately eight rows of semicircular (U-shaped) spinulated folds; presence of two flat, unornamented discs at the apex of each lobe. See Pattern 2 in Fig. 6 .

Extinction risk. Based on the combined evidence of EOO, number of localities, and trend of forest loss as proxy of habitat quality, we assessed or reassessed all species in the genus Echinosaura , as presented in Table 2 . According to the IUCN, to be considered in a threatened category following the geographic B1 criteria (EOO ≤ 20000 km 2), a species must also satisfy at least two of the three subcriteria, in our case, number of locations ≤ 10 (B1a) and observed, estimated, inferred, or projected decline in habitat quality [B1b(iii)]. For most species, the mean forest cover in the EOO at present is over 95% of the available cover in the year 2000. For this reason, species that do not meet the B1a criteria were assessed as Least Concern (LC), is the case of E. embera sp. nov. and E. palmeri . Both E. centralis and E. horrida were also assigned to LC despite presenting current forest covers below the 95% threshold (but still over 80%) since the species may not be threatened in a near future given their relatively large distribution. On the other hand, species fulfilling the B1a criteria but with current habitats over the 95% threshold (failing to meet the B1b(iii) subcriteria), were assigned to the Near Threatened (NT) since a reduction in forest cover could pose a threat in the near future. This was the case of E. brachycephala , E. fischerorum , E. keyi , and E. orcesi . Finally, the only species satisfying the B1ab(iii) criteria was E. panamensis , the reason why we assessed it as Vulnerable (VU).

Figures

FIGURE 2. A: Updated geographic distribution for the Echinosaura species. The stars indicate type localities, while white dots depict locations with genetic sampling. Black arrow in the main map points the border between the Atrato (A) and San Juan (SJ) rivers basin. To reduce point clusters and enhance the map displays, we thinned the records using an 18.5 km spacing buffer (a full list of records, their localities, georeferences, and vouchers can be found in Supplementary Files I). B: Violin plots showing the altitudinal distribution for the Echinosaura species.

FIGURE 3. Holotype of Echinosaura embera sp. nov. (MHUA-R 12591) in preservative.

FIGURE 4. Echinosaura embera in life, A: MHUA-R 12591 (holotype) from Chigorodó, Antioquia, Colombia, B: CBUCES- D 1867 (paratype) from Carmen de Atrato, Chocó, Colombia. Photos: Juan M. Daza and Juan D. Vásquez-Restrepo, respectively.

FIGURE 5. Dorsal and lateral view of scutellation tail patterns in E. palmeri (MHUA-R 12293) and E. embera (MHUA-R 12591). Scale bars = 2 mm. The yellow ovals and arrows show the major differences in scutellation patterns between E. palmeri and E. embera. Circles show the enlarged scales only in the posterior part of each autotomic segment, while arrows, the enlarged rows of scales forming a V-shaped pattern towards the anterior portion of each segment. Modified from Vásquez-Restrepo et al. (2020) with the permission of Oxford University Press.

FIGURE 6. Echinosaura hemipenial morphological patterns for the northern clade of species sensu Vásquez-Restrepo et al. (2020), as described in Sales-Nunes (2011). E. panamensis (KU 80584) from Panama province, Panama. E. embera sp. nov. (KU 113622) [paratype] from Cerro Pirre, Darien province, Panama. White arrows highlight the main differences between the two patterns (see text). Note that some spines in Pattern 2 may appear slightly smaller due to a thin layer of tissue covering them.