Taxonomic revision of the geckos of the Gonatodes concinnatus complex (Squamata: Sphaerodactylidae), with description of two new species Author Sturaro, Marcelo José Author Avila-Pires, Teresa C. S. text Zootaxa 2011 2869 1 36 journal article 10.5281/zenodo.277513 c499f937-b2bb-4333-8504-958183e30f73 1175-5326 277513 Gonatodes concinnatus ( O’Shaughnessy, 1881 ) ( Figs.1 A, 4, 5, 6) Goniodactylus concinnatus O’Shaughnessy, 1881 : 237 ( syntypes BMNH 1046.9.7.10–12, formerly BMNH 80.12.8.29–31, type-locality: Canelos, Ecuador , collected by Buckley). Goniodactylus buckleyi O’Shaughnessy, 1881 : 238 ( syntypes BMNH 1046.9.7.13–14, formerly BMNH 80.12.8.32–33, typelocality: Canelos; BMNH 1946.9.7.15, formerly BMNH 80.12.8.34, type-locality: Pallatanga [probably in error , see Rivero-Blanco, 1968 : 105 , 1979: 92]; all in Ecuador , collected by Buckley). Gonatodes concinnatus ; Boulenger, 1885 : 61 –62; Burt & Burt, 1933 : 2 ; Vanzolini, 1955: 123, 1968: 25 (part); Wermuth, 1965 : 44 ; Mechler, 1968 : 331 (part); Rivero-Blanco, 1968 : 104 (part); Peters & Donoso-Barros, 1970 : 132 (part); Dixon & Soini, 1975 : 19 , 1986: 23; Duellman, 1978 : 195 ; Rodrigues, 1980 : 313 ; Duellman & Mendelson III, 1995 : 358 ; Moravec, Tuanama & Burgos, 2001 : 51 ; Bartlett & Bartlett, 2003 : 158 (part); Powell & Henderson, 2005 : 714 ; Cole & Kok, 2006 : 4 ; Gamble, Simon, Colli & Vitt, 2008 : 271 . Gonatodes concinnatus concinnatus ; Rivero-Blanco, 1979 : 92 (part). FIGURE 3. (A) Geographic distribution of Gonatodes concinnatus complex. (B) Detailed geographic distribution of Gonatodes concinnatus . Red = Gonatodes concinnatus (Pattern A). Cyan-green = Gonatodes ligiae (Pattern B). Lime-green = Gonatodes tapajonicus (Pattern C). Yellow = Gonatodes nascimentoi sp. nov. (Pattern D). Blue = Gonatodes riveroi sp. nov. (Pattern E). Open symbols = type-localities; half-black symbol = literature record; question mark = Pallatanga, Ecuador, probably in error ; triangle = locality referred to only as ‘Rio Napo’. Numbers refer to localities cited under each species. One number may represent more than one locality. Material examined (numbers in bold between parentheses refer to Fig. 3 ): ECUADOR . “Pallatanga” ( 1 ): Photos of BMNH 1946.9.7.15 (formerly BMNH 80.12.8.34, syntype of Goniodactylus buckleyi ), leg. Buckley. SUCUMBÍOS. Shushufindi, Rio Aguarico ( 2 ): USNM 234497, female. Santa Cecilia ( 3 ): KU 105290–91, KU 109572–73, KU 146643, KU 146645, KU 147588–89, KU 152496, 4 males and 5 females . Lago Agrio ( 4 ): KU 126683, 126685–86, 126688, 2 males and 2 females ; KU 297992, female, leg. J. Lee. Limoncocha ( 5 ): LACM 73293–300, a juvenile, 5 males and 2 females , leg. W.R. Heyer; MZUSP 54655, male, leg. K. Miyata; KU 144378, MCZ 86414, a juvenile and a male. Reserva de Producción Faunística de Cuyabeno (RPF Cuyabeno), Estación Biológica de la Pontificia Universidad Catolica del Ecuador ( 6 ): OMNH 36406–11, 36424, 3 males and 4 females , leg. L.J. Vitt. Reserva de Producción Faunística de Cuyabeno (RPF Cuyabeno), Saladero de Dantas ( 6 ): OMNH 36412–14, a juvenile, a male and a female, leg. L.J. Vitt. Corporación Estatal Petrolera Ecuatoriana ( CEPE ) oil field (near Tarapoa, near Lago Agrio), Cuyabeno ( 7 ): MCZ 163233, male. SUCUMBIOS–ORELLANA. Rio Napo (undefined locality - 8 ): UMMZ 90813, female, leg. W.M. Clark. NAPO . George Kiederle Hacienda, southern bank Rio Napo, 2 km W Puerto Napo ( 9 ): USNM 166135–37, 2 males and a female, leg. J.A. Peters. Rio Misahualli ( 10 ): USNM 234490, female, leg. M. Olalla. 1 km E Jatun Sacha ( 11 ): OMNH 36377–78, a male and a female. San José de Sumaco ( 12 ): AMNH 28879, female, leg. C. Ollalo. ORELLANA. Between Loreto and Concepcion ( 13 ): CAS-SU 16529–30, a male and a female, leg. J. Olalla and C.A. Olalla. Loreto ( 14 ). MZUSP 3382–83, 2 females , leg. J. Olalla; USNM 234494, male, leg. J. Ollala. Rio Punino, affluent of Rio Payamino above Coca ( 15 ): MCZ 131046 , juvenile. Rio Payamino ( 16 ): USNM 234491–93, a male and 2 females , leg. M. Olalla. Mouth of Rio Coca ( 17 ): USNM 234495, male, leg. J. Olalla. PASTAZA. Rio Pindo ( 18 ): USNM 234500, male, leg. R. Olalla. Canelos ( 19 ): Photos of BMNH 1046.9.7.10–12 ( syntypes of Goniodactylus concinnatus , formerly BMNH 80.12.8.29–31), 3 males ( Fig. 4 ), Photos of BMNH 1046.9.7.13–14 ( syntypes of Goniodactylus buckleyi , formerly BMNH 80.12.8.32–33), 2 females , leg. Buckley. CAS-SU 15814–15, CAS-SU 16531, a male and 2 females , leg. A. Proana. Sarayacu, ( 20 ): MCZ 37703–04, KU 121088 –89, a juvenile and 3 females . Rio Villano ( 21 ), USNM 234499, female, leg. C. Estrella. Nuevo Golandrina, on trail W toward Rio Curaray, 130 km S of Coca ( 22 ): USNM 321064, female, leg. R.P. Reynolds and W. Lamar. Tiguino (UNOCAL Base Camp), 130 km S of Coca ( 22 ): USNM 321057, female, leg. R. P. Reynolds and W. Lamar. Rio Conambo ( 23 ): USNM 234502, female, leg. R. Olalla. MORONA-SANTIAGO. Cusuime, Rio Cusuime, 60 km airline SE Macas ( 24 ): AMNH 113666–67, 2 males , leg. B. Malkin. Taisha, Macas ( 25 ). USNM 234498, male, leg. M. Ollala; USNM 283922, USNM 283924, a juvenile and a female, leg. C. M. Fugler. PERU : LORETO. San Jacinto ( 26 ): KU 222138, female, leg. R. A. Leschen. Moropón ( 27 ): TCWC 36665–71, TCWC 36743–46, TCWC 38994, TCWC 41213–20, TCWC 41902–04, TCWC 41906–08, TCWC 42706–13, TCWC 42828, TCWC 43332, TCWC 44230, TCWC 44283, a juvenile, 25 males and 12 females ; leg. P. Soini; TCWC 41763, juvenile, leg. J.R. Dixon; MZUSP 28248–49, MZUSP 28260–63, MZUSP 28273–79, MZUSP 28311–13, MZUSP 28319, MZUSP 28375, MZUSP 39218, 3 juveniles , 12 males and 4 females , leg. P. Soini. Explorama Lodge, junction Rio Yanamono and Rio Amazonas ( 28 ): KU 220370 , juvenile, leg. W. E. Duellman. Iquitos ( 29 ): TCWC 41137–38, a juvenile and a female, leg. P. Soini. ACEER on Quebrada Grande, close to junction Rio Sucusari and Rio Napo ( 30 ): KU 220371 , male, leg. W. E. Duellman. Yanamono ( 31 ): MZUSP 28354–55, a juvenile and a female, leg. P. Soini. Rio Orosa ( 32 ). MZUSP 56657, male, leg. P. Soini. Estirón, Rio Ampiyacu ( 33 ): MZUSP 13458, male, leg. B. Malkin. Diagnosis. A relatively large Gonatodes , with maximum SLV of 52.6 mm . Scales around midbody 109–131. Ventral scales in a longitudinal row 49–61. Proximal subdigital lamellae as wide as digits, in total 17–22 under fourth finger, 21–27 under fourth toe. Three or four lateral rows of scales on distal portion of fingers and toes. Tail with midventral scales distinctly wider than long, forming a repetitive sequence of two single midventrals (one after the other), each in contact with one laterodistal scale per side, followed by a divided (only on proximal portion of tail, when present) or single midventral in contact with two laterodistal scales per side (respectively 1’1’2” and 1’1’1”). A white suprahumeral bar, bordered by black, present both in males and females, although more conspicuous in males; it reaches dorsally between the dorsolateral and middorsal regions. Males with head dorsally without vermiculations; back and limbs with a vermiculated pattern of dark and light spots; no dark streaks on gular region (at least in preservative). Description. Maximum SVL in males of 52.6 mm (TCWC 36669), in females of 49.2 mm (KU 146645). Tail round in cross section, tapering toward tip, 1.0–1.4 (1.15 ± 0.09, N =37) times SLV. Head length 0.21–0.27 (0.24 ± 0.01, N =137) times SLV, 1.3–1.6 (1.45 ± 0.06, N =136) times as long as wide, 1.2–1.7 (1.42 ± 0.08, N =135) times as wide as high. Snout round, moderately elongate ( Fig. 5 A), gently sloping toward top of head. Neck slightly narrower than head and anterior portion of body. Body cylindrical. Limbs well developed, lower arm 0.11–0.15 (0.14 ± 0.01, N =139) times SVL, lower leg 0.13–0.17 (0.15 ± 0.01, N =139) times SVL. Rostral convex, 1.6–2.9 (2.07 ± 0.16, N =139) times as wide as high; posterior margin with a shallow depression medially and slightly indented by 1–3 medial postrostrals, with or without a median cleft extending anteriorly. Postrostrals 3 (72.0%), 4 (19.4%), or 5 (8.6%), laterals ones (supranasals) distinctly larger than medial one(s) ( N =139) ( Fig 5 A). Nasal bordered by rostral, first supralabial (only a narrow contact in some specimens), 3 (92.1%) or 4 (7.9%) postnasals ( N =139), and supranasal; internostril distance 0.20–0.28 (0.24 ± 0.02, N =136) times head width. Supranasal roughly oval or semicircular, 0.8–1.5 (1.16 ± 0.13; N =139) times as wide as long. Postnasals slightly larger than, or similar in size to, adjacent loreals. Scales on snout convex, hexagonal to round, juxtaposed, relatively uniform in size. Canthus rostralis rounded. Loreal region with scales slightly more elongate than those on snout, largest on row adjacent to supralabials, 9–13 (10.4 ± 0.8, N =138) loreals on a line between postnasals and anterior corner of eye. Top and posterior portion of head, as well as supraorbital region, with granular scales. A short supraciliary flap present, anteriorly with 6–12 (8.4 ± 1.1, N =133) enlarged and flattened scales, among which 0–7 (2.0 ± 1.3; N =133) small, conical spines. Pupil round, eye diameter 0.19–0.26 (0.21 ± 0.01, N =129) times head length. Scales on temporal region similar to those on top of head. Ear-opening much smaller than eye, oval, posterior to, and at same level of, commissure of mouth. Supralabials 5–7 (6.2 ± 0.5; N =139), distinctly enlarged anteriorly and decreasing in size posteriorly, one or two of them posterior to centre of eye, followed to corner of mouth by small scales. Mental large, distinctly wider anteriorly than posteriorly, with posterior margin forming a wide angle, 0.9–1.4 (1.1 ± 0.1; N =137) times as wide as long; 1 (1.4%), 2 (82.7%), 3 (12.9%) or 4 (2.9%) postmentals ( Fig. 5 B). Scales on chin flat, smooth, polygonal, juxtaposed, larger anteriorly, decreasing in size posteriorly. Infralabials 4–7 (5.5 ± 0.7; N =138), distinctly enlarged anteriorly and decreasing in size posteriorly; 0–3 of them posterior to centre of eye, followed to corner of mouth by small scales. Scales on nape small and granular, becoming slightly larger on sides of neck. Scales on throat anteriorly granular, posteriorly flat, smooth, hexagonal or round, increasing in size toward the posterior region, with a short transitional zone between the anterior and posterior parts. Dorsals granular, increasing in size toward the flanks. Ventrals larger than dorsals, roughly hexagonal, flat, smooth, imbricate, in oblique rows; 49–61 (56.1 ± 2.1; N =119) scales along the midventral line between anterior margin of forelimbs and vent; 17–20 (18.5 ± 0.9; N =126) scales in a transverse line at midbody, with a short transitional zone between ventrals and scales on flanks. Scales around midbody 109–131 (120.7 ± 4.6; N =110). Scales on preanal plate similar to ventrals, except for those bordering vent, which are very small. Escutcheon present in males on posterior portion of belly and on 3–5 (mostly four) rows (body-knee direction) on ventral surface of thighs. Scales on anterodorsal surface of forelimbs flat, smooth, roundish or rhomboidal, imbricate, largest close to the wrist; on posterodorsal and ventral surface of forelimbs convex, smooth, rhomboidal, juxtaposed, relatively small. Scales on anterodorsal surface of thighs and ventral surface of hind limbs flat, smooth, roundish or rhomboidal, imbricate and relatively larger; on posterodorsal surface of thighs and dorsal surface of lower legs smaller, convex, smooth, rhomboidal, juxtaposed. Lamellae under second ( II ) through fourth ( IV ) fingers (proximal lamellae in parentheses): II : 14–18 (5–7), III : 16–21 (5–7), and IV : 17–22 (5–9) ( Fig. 5 C). Lamellae under second through fourth toes (proximal lamellae in parentheses): II : 14–18 (5–7), III : 16–22 (5–8), and IV : 21–27 (9–13) ( Fig. 5 D). Claws exposed, non-retractile, between two basal scales. Fingers and toes with three, occasionally four or two (USNM 234497; USNM 234494 and TCWC 42706), lateral scales distally, between the fourth (counted from the claw towards the hand) subdigital lamella and the dorsal scale. Scales on tail dorsally and laterally relatively small, rhomboidal, flat, smooth, imbricate. On ventral surface of tail scales smooth, flat, imbricate, increasing in size toward midventral line. Midventral scales, except close to the base of the tail, distinctly wider than long, forming a repetitive sequence of two single midventrals (one after the other), each in contact laterodistally with one scale per side, followed by a divided (only on proximal portion of tail, when present) or single midventral scale in contact laterodistally with two scales per side – respectively 1’1’2” and 1’1’1” in the codification of Avila-Pires (1995: Figure 2 ) ( Fig. 5 E). Color in preservative. In males ( Figs. 1 A, 4), dorsal surface of head beige, without markings. Back and flanks, base of tail and hind limbs with relatively large, beige and brown vermiculations. A large, conspicuous, white suprahumeral bar, bordered by black, extending dorsally at least up to the dorsolateral region, in some cases almost reaching the middorsal region; never in the form of an ocellus, but Vitt & Torre (1996) present the photo of a specimen with the white bar divided into two spots by a transversal black band. Ventral surface of head, gular region and chest beige or reddish-brown, without oblique streaks; belly gray; underside of limbs beige. Tail brown and/or black dorsally, white and/or brown ventrally. Escutcheon area (belly and thighs) light gray. In females, dorsal surface of head and limbs with brown and black irregular spots; back gray with dorsolateral pairs of black spots and, in some specimens, pairs of beige spots; flanks gray with black and brown spots. Suprahumeral bar conspicuous, white with black margins, similar in extension but thinner than that of males. Ventral surface of head and gular region white with dark oblique streaks, in contact or not at midventral line; belly and underside of limbs light gray. Tail brown and/or black dorsally, white and/or brown ventrally; original tail distally with white bands that form complete rings around the tail. FIGURE 4. Gonatodes concinnatus (syntypes, BMNH 1946.9.7.10–12). (A) Dorsal, (B) ventral views. Scale bar = 50 mm. (Photo by Colin McCarthy). FIGURE 5. Gonatodes concinnatus (MZUSP 54655). (A) Dorsal and (B) ventral views of head; ventral views of (C) right FIGURE 6. Gonatodes concinnatus from Cuyabeno, Sucumbios, Ecuador. (A) Adult male (LJV 5014) and (B) adult female (OMNH 36409; LJV 5046) (Photos by Laurie Vitt). Color in life. Color in life has been described by Duellman (1978) and Vitt & de la Torre (1996) . In males ( Fig. 6 A) head, neck and forelegs orange or reddish brown with cream spots; suprahumeral bar white bordered by black; body olive-green or brown, with reddish brown or yellowish white vermiculations that may present a black margin. Throat orange with cream streaks, belly yellowish gray, and tail black with white rings, or the entire belly and tail black. In females ( Fig. 6 B), head, body and tail grayish tan or drab gray with irregular crossbands, brown or black anteriorly and white posteriorly; distally the tail becomes black and white banded. Throat cream with dark streaks; venter creamy tan or yellow. Iris reddish brown with a cream circle around the pupil. Distribution. Western Amazonia, in Ecuador and northern Peru ( Fig. 3 A–B). Remarks. Gonatodes ligiae Donoso-Barros, 1967 , from Venezuela , is considered in this paper as a valid species. Material from Colombia previously identified as G. concinnatus is here described as a new species, G. riveroi .