Ameiva nodam Koch, Venegas, Rödder, Flecks & Böhme, 2013, sp. nov.
Creators: name: Koch, Claudia
name: Venegas, Pablo J.
name: Rödder, Dennis
name: Flecks, Morris
name: Böhme, Wolfgang
Publisher: name: Zenodo
Year: 2013
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Reptilia
Squamata
Teiidae
Ameiva
Ameiva nodam

description: Ameiva nodam sp. nov.
Figs. 1–2
Cnemidophorus divisus Fischer, Verhandlungen des Naturwissenschaftlichen Verein in Hamburg 3: 78–103. — 1879 (the original description refers to a Venezuelan taxon described by Fischer as C. divisus)
Ameiva bifrontata divisa— Ruthven, Occasional Papers of the Museum of Zoology, University of Michigan (155): 1–6. — 1924
Ameiva bifrontata divisa— Burt & Burt, Bulletin American Museum of Natural History, 61: 227–395. — 1931
Ameiva bifrontata divisa— Burt & Burt, Transactions of the Academy of Science of Saint Louis 28 (i): v–108. — 1933
Diagnosis and comparison. This comparatively small Ameiva is diagnosed by the following combination of characters: (1) maximum known SVL of 101mm; (2) lacking longitudinal ridge on frontal scale; (3) frontal plate divided in two subequal scales; (4) postnasals separated from prefrontals by frontonasals; (5) parietal scales usually five; (6) median gular scales not enlarged; (7) enlarged median mesoptychial scales slightly larger than largest gulars; (8) gulars posterior to the interauricular crease smaller than anterior gulars; (9) nasal suture passes centrally through nostril; (10) rostral contacting postnasals; (11) supranasals not contacting supralabials; (12) scales of circumorbital semicircle not extending to anterior margin of third supraocular; (13) 28–33 enlarged ventral scales between gular and vent; (14) 10 longitudinal rows of ventral plates, outermost often distinctly smaller; (15) 86–113 DOM; (16) 165–216 DL; (17) postbrachials dilated; (18) 25–35 LFT; (19) 27–32 SCF; (20) 10–19 FP; (21) five longitudinal yellows stripes on dorsum distinct in juveniles and females, less distinct in most males.
Ameiva nodam sp. nov. can be distinguished from all other described mainland congeners except for A. bifrontata and A. concolor by having a transversely divided frontal plate. From the latter two it differs in the color pattern and from A. concolor also by having the postbrachials dilated.
Especially due to the color pattern of female specimens with five longitudinal yellow dorsal stripes on a brown ground color the new species resembles Medopheos edracanthus, a teiid species that was just recently split of the genus Ameiva and is now considered as being monotypic. It differs from this species by having the frontal plate divided, by lacking preanal spurs and by having 10 instead of 8 longitudinal rows of enlarged ventral scales.
Holotype. An adult male (CORBIDI 1870, Figs. 1 A–F, 2A,B) from Bellavista, Province of Jaén, Region of Cajamarca, Peru (05°38’15.6’’S, 78°37’59.2’’W, 390–440 m above sea level), collected 0 9 May 2008 by P. Venegas and C. Koch.
Description of holotype: A large adult male with a SVL of 101 mm. Head 0.25 times SVL, 2.95 times longer than wide, 0.64 times as wide as high. Snout elongate, bluntly pointed; canthus rostralis distinct. Neck only slightly narrower than head, and body. Body cylindrical. Limbs well developed, forelimbs 0.3 times SVL, hindlimbs 0.58 times SVL, tibia 0.12 times SVL. Tail round in cross section, tapering toward the tip; 2.08 times SVL. Rostral in posterior part right-angled and bordered by supranasals, in anterior part laterally stretched and bordering supranasals; wider than high; smooth, without sutures; visible from above. Supranasals almost triangular, in short medial contact, not contacting supralabials, bordered posteriorly by hexagonal frontonasal. Postnasal almost triangular, in short contact with rostral and frontonasal and in broad contact with loreal and first and second supralabial. Oblique nasal suture passing centrally through oval nostril. Prefrontals paired and roughly pentagonal, with a short medial suture slightly longer than that between supranasals; laterally in contact with loreal, first supraocular and first supraciliary. Frontal plate divided in two subequal scales, the suture between both scales forming a straight line; anterior frontal pentagonal, laterally in contact with first and second supraoculars, distinctly larger than posterior frontal; posterior frontal pentagonal, laterally in contact with second and third supraoculars. Pair of pentagonal frontoparietals, longer than wide and with long medial suture; laterally in contact with third supraocular, and small circumorbital scales bordering posterior part of third supraocular. Interparietal pentagonal, higher than wide, slightly narrower than adjacent parietals, sutures with parietals straight; interparietal bordered at each side by two irregular parietals divided by oblique suture; outermost parietals slightly smaller than inner parietals; parietal series composed of five scales including interparietal. Supraoculars three at each side. Circumorbital semicircle formed by 12 scales at left side, 22 scales combining both sides, bordering posterior edge of third supraocular and separating it from frontoparietals by two thirds; third supraocular separated from parietals by two rows of circumorbital scales. Laterally, second and third supraocular separated from supraciliaries by a single row of small scales; 19 combining both sides. Supraciliaries seven (on one side), first highest, second longest, remaining ones shorter and subequal. Loreal very large, single, in contact with postnasal, frontonasal, prefrontal, first supraciliary, all three preoculars, first subocular, and second, third and fourth (narrowly) supralabials. Preoculars three, first and third very small and granular; second preocular larger, but distinctly smaller than suboculars. Suboculars three, all longer than wide and in contact with supralabials, first and third subocular almost equal in size, second subocular longest. A keel reaching from first through second subocular. Postoculars small, in two rows, first consisting of six and second of five scales. Enlarged supralabials seven, fifth below center of eye, second and third largest; followed to commissure of mouth by five small scales. No enlarged supratemporals distinguishable. Temporal region with polygonal or rounded scales, slightly smaller centrally. External auditory meatus large, vertically oval, bordered by granular scales, anterior margin semicircular, posterior one straight. Tympanum recessed. All dorsal and lateral head scales juxtaposed and smooth. Mental anteriorly ellipsoid, posteriorly straight, bordered by first infralabials and postmental. Postmental single and pentagonal, in contact with first and second infralabials, followed by six pairs of enlarged chinshields. First pair in broad medial contact and in contact with infralabials. Remaining chinshields separated from infralabials by one row of small, almost granular scales. Medial chin scales moderately small, convex, smooth, juxtaposed, oval or polygonal, in slightly oblique rows, all subequal in size. Enlarged infralabials six, fifth below center of eye; followed to commissure by six smaller scales. Gular region divided into two areas: anterior region with round or polygonal and flat scales in slightly oblique rows that usually remain subequal or rarely grade to slightly larger scales medially, delimited posteriorly by line uniting lower margin of ear openings. Posterior gular region covered by smaller polygonal or round scales in transverse rows. Mesoptychial scales moderately enlarged, slightly larger than anterior gular scales, in about three rows, hexagonal, flat, smooth, and juxtaposed. Scales on nape and sides of neck similar in size to dorsals. Dorsals and scales on flanks granular (slightly larger on dorsum than laterally), round, smooth, juxtaposed; 194 DL; 93 DOM. Ventrals large, smooth, rectangular, wider than long, in 10 longitudinal and 31 transverse rows; transition between ventrals and scales on flanks sharp. Preanal shield with three rows of enlarged scales. Preanal plate surrounded anteriorly and laterally by smaller scales; posteriorly by much smaller scales. 18 FP in continuous row along each thigh, with short gap medially. Each pore surrounded by four scales. Scales on tail dorsally rectangular, smaller than subcaudals, longer than wide, distinctly keeled, slightly imbricate; in transverse and approximately longitudinal rows, continuous with subcaudals around tail (except first few rows incomplete ventrally); 27 SCF. Distally caudals longer and narrower. Subcaudals rectangular, smaller than ventrals, wider than long, smooth, mostly juxtaposed. Forelimbs with row of very large, smooth, slightly imbricate, almost rectangular (distinctly wider than long) antebrachial scales on anterodorsal aspect of forearms and similar but smaller brachial scales on upper arms that extend almost to insertion of forelimbs. Antebrachials and brachials separated by smaller scales at elbow. Dorsoposterior, posterior, and ventral aspect of arms granular, slightly larger than dorsals, except for scales directly adjacent to brachials and antebrachials, which are slightly to moderately enlarged and irregular. Legs with large, smooth, imbricate scales on anterior and ventral aspects of thighs, and ventral aspect of shanks. Row of large, almost rectangular scales anteriorly on thigh, gradually becoming smaller and irregular toward pores. On ventral aspect of shanks, three rows of very large scales, anterior two more or less trapezoidal, posterior one rhomboidal, decreasing in size from anterior toward posterior row. Tibiotarsal spurs form a cluster of two rows, one of about four, and the other of about five sharply mucronate scales which are positioned along the postaxial edge of the distal end of the shank. Elsewhere on hindlimbs scales similar to dorsals. Subdigital lamellae transversely enlarged and single, moderately to distinctly tuberculate towards base. Lamellae of outer toe continuing to heel. On palms, lamellae of outer and inner fingers continuing to wrist only separated by few granules, tubercular and increasing in size towards it. 17 LFF; 32 LFT.
Measurements of holotype (in mm). SVL 101; HL 25.4; HH 13.4; HW 8.6; SL 10.5; ED 5.3; DSN 2.5; DNE 8.1; DEE 7.7; TL 210; AGL 45; BHM 13.6; BWM 17.3; FLL 30; HLL 59; TIL 12.2.
Coloration of holotype: In life (Fig. 2 A), the dorsal surface of the head of the male holotype is grayish-brown. The sides of the head are gray. The granules on the eyelids are white to grayish-white. Suboculars and adjacent supralabials are lighter gray than the scales in temporal region, anterior supralabials, postnasal and loreal. Supraciliaries grayish-green. Head ventrally white to grayish-white. Dorsal surface of body reddish-brown; five yellow stripes 2–3 scales in width in anterior part and 4–5 scales in width in posterior part, extending from the head to the base of the tail, one vertebral stripe beginning behind the interparietal, a dorsolateral stripe on each side beginning in the supraciliary region, and a lateral stripe beginning in the anterior upper edge of the ear opening (and being interrupted by the opening); in the posterior part the vertebral stripe is bordered on both sides by an irregular dark blackish-brown line, starting from behind the insertion of the forelimbs, very soft in the beginning and becoming up to four scales in width in direction of the base of the tail; ground color interspaces irregularly intermixed with some blackish-brown granules. Lateral body parts between armpit and groin gray with eleven small turquoise dots (4–6 granules big) arranged in a dotted line. Outermost ventrals gray, first three outer rows of ventrals with an irregular pattern of turquoise dots distinctly larger than the turquoise dots of the lateral row. Central rows of ventrals white. Forelimbs grayish-brown dorsoposterior region and brownish-gray in anteroventral region. Hindlimbs grayish-brown in dorsoposterior region, intermixed with some small dark and larger pale white spots; brownish-gray in anteroventral region. Tail brown with dark spots in anterior third, a lateral dark stripe extends from behind the hind limbs over the first sixth of the tail, adjacent and below to it is a white stripe of almost the same length. Subcaudals white.
In preservative (Fig. 2 B), the general dorsal color is mainly brown or grayish-brown; five dorsal stripes are gray to grayish-brown; interspaces between dorsolateral and lateral stripes are dark brown; dots between armpit and groin are pale blue. Remaining body parts are of the same pattern and coloration as in life.
Variation. Paratypes (59): An adult male (CORBIDI 1869) and an adult female (CORBIDI 1873) with the same data as the holotype; An adult female (CORBIDI 5762) from Bellavista, Province of Jaén, Region of Cajamarca (05°31’S, 078°31’W, 390–444 m above sea level), collected 30 March 2009 by E. Hoyos Granda and C. Koch; an adult male (MCZ 18134) and three adult females (MCZ 18135–7) from the same region, collected 1916 by G.K. Noble. A Male and a juvenile (ZFMK 88732–33) from Gotas de Agua, Province of Jaén, Region of Cajamarca (05°42’28.2’’S, 78°47’18.4’’W, 719 m above sea level), collected 0 5 June 2008 by A. Garcia Bravo and C. Koch; an adult male (CORBIDI 1910) from almost the same region (05°41’17.7’’S, 78°46’02.0’’W, 717 m above sea level) collected 0 5 July 2008 by W.A. Garcia Bravo, J. Novoa Cova and C. Koch. An adult male (ZFMK 88734) from Pucará Province of Jaén, Region of Cajamarca (06°01’51.2’’ S, 79°07’32.5’’ W, 1054 m above sea level), collected 10 July 2008 by W.A. Garcia Bravo, J. Novoa Cova and C. Koch; an adult female and a juvenile (ZFMK 90862, CORBIDI 5761) from almost the same region (06°03’ S, 79°05’ W, 970 m above sea level), collected 26 March 2009 by E. Hoyos Granda and C. Koch. An adult female (CORBIDI 5763) and a juvenile (ZFMK 90863) from Perico, Province of Jaén, Region of Cajamarca (05°20’30.7’’S, 78°47’52.1’’W, 490 m above sea level), collected 0 5 April 2009 by E. Hoyos Granda and C. Koch; an adult male (BM 1924.11.19.10), an adult female (BM 1924.11.19.11) and a juvenile (BM 1924.11.19.12) from the same region, collected 1924 collector not further determined; an adult male (MCZ 18132), two adult females (MCZ 18130–31), a juvenile (MCZ 18133) and an undefined adult specimen (ZMB 29752) from the same region, collected 1916 by G.K. Noble. A juvenile (KU 209520) from Fonda Atapaca (near Rio Chinchipe, East of San Ignacio, 450 m), Province of San Ignacio, Region of Cajamarca, collected on 4 October 1986 by CRS. Five adult males (KU 134835–8, 134841), six adult females (KU 134839–40, 134842–5) and a juvenile (KU 134834) from 7 km North of Jaén, Province of Jaén, Region of Cajamarca (730 m above sea level), collected 5 May 1970 by T.H. Fritts. An adult male (ROM 16273) from 52 km North of Jaén, Province of Jaén, Region of Cajamarca, collected 7 July 1986 by L.D. Wilson. An adult male (MHNG 2260.19), an adult female (MHNG 2260.18) and an subadult female (MHNG 2260.17) and two juveniles (MHNG 2260.16, 2260.20) from Jaén, Province of Jaén, Region of Cajamarca, collection date and collector not further determined. An adult female (ZFMK 90864) from Puerto Malleta, Province of Cutervo, Region of Cajamarca (06°03’57.1’’S, 78°36’21.1’’W, 509 m above sea level), collected 12 December 2009 by A. Garcia Bravo and C. Koch. Four males (CORBIDI 1874, 1900, 1902, ZFMK 88735), one subadult female (CORBIDI 1898) and six juveniles (CORBIDI 1786, 1833, 1853, 1872, 1905, ZFMK 88736) from Bagua Chica, Province of Bagua, Region of Amazonas (05°38’06.9’’S, 78°32’27.7’’W, 500 m above sea level), collected 20 July 2008 by J. Novoa Cova and C. Koch. Three adult males (CORBIDI 5769, ZFMK 90865–66) and two adult females (CORBIDI 5767–68) from Cumba, Province of Cumba, Region of Amazonas (05°56’14.6’’S, 78°39’50.4’’W, 465 m above sea level), collected 16–18 December 2009 by A. Garcia Bravo and C. Koch.
Description of paratypes: Maximum SVL in male paratypes 88.3 mm (KU 134837), maximum total length in male paratypes 280.7 mm (KU 134838); maximum SVL in female paratypes 78 mm, maximum total length in female paratypes 244 mm (both CORBIDI 05762). Shape of head, body, limbs and tail as in holotype. HL 0.25– 0.29 (0.27 ± 0.01, n = 15) times SVL in males; 0.24–0.28 (0.26 ± 0.01, n = 13) times SVL in females; HH 0.11– 0.14 (0.13 ± 0.01, n = 16) times SVL in both sexes; HW 0.09–0.13 (0.11 ± 0.01, n = 21) times SVL in males, 0.09– 0.13 (0.11 ± 0.01, n = 18) times SVL in females; SL 0.62–0.68 (0.66 ± 0.02, n = 10) times the HL; ED 0.20–0.28 (0.26 ± 0.02, n = 10) times the HL; DSN 0.08–0.13 (0.12 ± 0.02, n = 10) times the HL; DNE 0.26–0.31 (0.29 ± 0.02, n = 10) times the HL; DEE 0.22–0.26 (0.24 ± 0.02, n = 10) times the HL; TL 1.34–2.48 (1.87 ± 0.39, n = 13) times SVL in males; 1.42–2.9 (1.95 ± 0.39, n = 13) times SVL in females; AGL 0.38–0.52 (0.42 ± 0.04, n = 16) times SVL; FLL 0.33–0.38 (0.35 ± 0.01, n = 10) times SVL; HLL 0.66–0.72 (0.69 ± 0.02 n = 10) times SVL; TIL 0.17–0.2 (0.19 ± 0.01, n = 10) times SVL; foot 0.35–0.41 (0.38 ± 0.02, n = 10) times SVL.
Arrangement, shape and surface of scales as in holotype except for the following variations: Rostral in posterior part right-angled or nearly so. Frontonasal pentagonal or hexagonal. Prefrontals laterally contacting first supraciliary in most specimens, separated from it by the first supraocular and/or the loreal in some specimens. Median suture of prefrontals as short or slightly longer than that between supranalsals. Frontoparietals, posteriorly contacting interparietal and adjacent parietals in most specimens, some individuals with a small medial scale between the posterior suture of the frontoparietals and the interparietal. Interparietal subequal, with or without a short medial suture in anterior part; slightly narrower or slightly wider than adjacent parietals, sutures with parietals slightly oblique or straight; outermost parietals subequal or slightly smaller (rarely larger) to inner parietals. Parietal series normally composed of 5 (4 in one specimen: ZFMK 90862) scales including interparietal. Supraoculars three or four at each side, fourth much smaller than others, third supraocular largest. Circumorbital semicircle formed by 4–14 scales at left side, 8–27 scales combining both sides, bordering posterior edge of fourth (if present) and third supraocular and separating it from frontoparietals by more than half, not extending to anterior margin of third supraocular. Last supraocular separated from parietals by 2–4 rows of circumorbital scales. Laterally, all supraoculars except first separated from supraciliaries by a single or double row of small scales; 21– 34 combining both sides. Supraciliaries 5–7 (on one side). Loreal very large, single, in contact with postnasal, frontonasal, prefrontal, first supraciliary, prefrontals, first subocular, second and third and occasionally fourth supralabials and in few specimens with first supraocular. Preoculars normally one, two or three in some specimens. Postoculars small, in two rows, first consisting of 3–5 and second of 3–4 scales. Enlarged supralabials five or six (mostly six), second, third and fourth largest; followed to commissure of mouth by 4–7 small scales. Five to eight slightly enlarged supratemporals distinguishable or not from other scales. Postmental followed by three to 6–8 pairs of enlarged chinshields. Enlarged infralabials 5–7; followed to commissure by 5–8 smaller scales. 165–216 (195 ± 14.67, n = 15) DL; 86–113 (95 ± 7.64, n = 12) DOM. Ventrals in ten longitudinal and 28–33 (30 ± 1.41, n = 60) transverse rows; ventrals of outermost longitudinal row smaller than other ventrals, very small in some specimens. Preanal shield with 2–3 rows of enlarged scales. Postbrachials dilated. 10–19 (16 ± 1.88, n = 36) FP; hardly or even not visible in some female and juvenile specimens. 27–32 (29 ± 1.86, n = 12) SCF. In males, a cluster of 2–3 rows, each of about 3–6 sharply mucronate scales are positioned along the postaxial edge of the distal end of the shank; females and juveniles without such tibiotarsal spurs. 13–19 (16 ± 1.32, n = 59) LFF, 25–35 (31 ± 2.08, n = 60) LFT.
Color variation: This species exhibits conspicuous ontogenetic color changes. In life, juveniles (Fig. 2 D) show a dark brown dorsal ground color on head, body and limbs; five longitudinal yellow stripes on dorsum very distinct; in some juveniles a more or less distinct yellow dot is present in the temporal region adjacent to the preoculars and anterior to the lateral stripe, from which it is separated by several small scales; an additional longitudinal, yellowish-white, entire or dotted line is present between the insertion of the forelimbs and the groin, little above the ventrals; in some juveniles the dark brown ground color interspaces between the vertebral and dorsolateral stripes has yellow spots in the posterior part of the body that are arranged in a longitudinal line along the center of each interspace (e.g. ZFMK 88733); forelimbs and hindlimbs dorsally with striking yellow spots; vertebral and dorsolateral stripes continuing on former third or less of the tail, where the color of the stripes grade from yellow to white; a distinct white stripe extends from the posterior insertion of the hindlimbs on the sides of the tail to the former third or less of the tail; at least last two thirds of unregenerated tail dorsally and ventrally turquoise-blue, gaining in saturation towards the tip of the tail; pale turquoise dots laterally between axilla and groin. The smaller the specimen, the larger is the contrast between the dark ground color and the yellow dorsal stripes and dots, and the more intensive is the turquoise color of the tail. In some adult males the brown interspaces between the dorsolateral and lateral yellow stripes are darker brown than the ground color; in some males the brown ground color interspaces between the yellow vertebral and dorsolateral stripes possess yellow more or less distinct spots in the posterior part of the body that are arranged in a longitudinal line along the center of each interspace; dotted turquoise line on lateral body parts between armpit and groin varies between males in size and number of dots (8–16) and may be very distinct or hardly visible; five longitudinal yellow stripes on dorsum may be distinct or very pale and almost not visible; in some males the lateral stripe is only yellow and entire in the anterior part but suspended in the posterior part consisting of turquoise dots that are parallel to the lateral turquoise dotted line; 3–4 outer rows of ventrals of each side gray, in some individuals central row of ventrals also intermixed with gray, irregular pattern of turquoise dots on 1–3 outermost rows of ventrals. Females (Fig. 2 C) have a similar color pattern than males but mostly without turquoise dots on the outer ventrals and instead of lateral turquoise dotted lines they normally exhibit a yellow entire or dotted line between armpit and groin; 5 longitudinal yellow stripes on dorsum normally distinct.
In preservative, the general dorsal color is mainly brown or grayish-brown; five dorsal stripes are gray to grayish-brown and less contrasted to ground color interspaces than in living individuals; interspaces between dorsolateral and lateral stripes of most males and females dark brown or blackish; ground color of juveniles dark brown or blackish; tail of juveniles grayish-blue in posterior part; dots of males between armpit and groin pale blue; dilated scales on forelimbs brownish-gray on hindlimbs bluish-gray; outermost rows of ventrals grayish-blue to blackish-blue. Remaining body parts are of the same color as in life.
Etymology. The specific epithet is an agglutination of the exclamation “no dam”. We chose this peculiar name to protest against the possible dam construction activities for four hydroelectricity projects along the Marañón river, between the Regions of Cajamarca and Amazonas. With this we are trying to call attention to the fact that the unique dry forest habitat of this and other endemic species, several of them have only recently been discovered (Koch et al. 2006, 2011; Venegas et al. 2008), is gravely threatened by human interventions.
Distribution and natural history. This species was collected in the northern Peruvian Andes (Fig. 3), from the Canyons of the Chinchipe, Chamaya, Huancabamba, Utcubamba and Marañón rivers from Fonda Atapaca (near Chinchipe River, East of San Ignacio, 450 m), Province of San Ignacio, Region of Cajamarca to Puerto Malleta, Province of Cutervo, Region of Cajamarca (near Rio Marañón, 509 m above sea level). The lowest distributional level of the species was in Bellavista, Province of Jaén, Region of Cajamarca (near Rio Marañón, 390–440 m above sea level) and highest distributional level was in Pucará, Province of Jaén, Region of Cajamarca (near Rio Huancabamba, 1054 m above sea level, Fig. 4). The species was visually detected by the first author (could not be collected) in La Balza, Province of San Ignacio, Region of Cajamarca (04°59’ S, 79°07’ W, 930 m), which is a small village at the border to Ecuador and just separated from it by the small Canchis river. It is thus most likely that this species can also be found in the dryforest regions of the Southern Ecuadorian Andes.
This new species is endemic to the Equatorial Dry Forest ecoregion (Brack 1986), a habitat which receives less than 500 mm of annual rain fall (Duellman & Pramuk 1999).
Individuals of this species were found moving quickly on the ground in low vegetation during daytime or sleeping under stones during nighttime. Air temperature during the active hours of the species was between 31.3°C and 40.8°C, substrate temperature of the ground was between 32°C and 46.2°C. Ameiva nodam sp. nov. was found in the same habitat together with the tropidurid lizard Microlophus stolzmanni.
description: Published as part of Koch, Claudia, Venegas, Pablo J., Rödder, Dennis, Flecks, Morris & Böhme, Wolfgang, 2013, Two new endemic species of Ameiva (Squamata: Teiidae) from the dry forest of northwestern Peru and additional information on Ameiva concolor Ruthven, 1924, pp. 263-295 in Zootaxa 3745 (2) on pages 267-274, DOI: 10.11646/zootaxa.3745.2.6, http://zenodo.org/record/247483