Anolis binotatus Peters, 1863

Anolis bitectus — Cope, 1864; Boulenger, 1885; Peters, 1967; Peters & Donoso-Barros, 1970; Williams et al., 1995; Torres-Carvajal, 2001; Poe, 2004; Miyata, 2013. Type specimens: BMNH 1946.8.5.83 and 1946.8.20.11.

Anolis binotatus —Bocourt, 1873; Peters, 1967; Peters & Donoso-Barros, 1970; Williams et al., 1995; Torres-Carvajal, 2001; Poe et al., 2017.

Anolis lemniscatus — Boulenger, 1898; Peters & Donoso-Barros, 1970; Williams et al., 1995; Gemel et al., 2019. Type specimens: BMNH 1946.8.8.35–39, MCZ 16783, NMW 12812:1–3.

Anolis tropidogaster —Peters, 1967.

Norops binotatus — Nicholson, 2002; Nicholson et al., 2012.

Norops bitectus — Nicholson, 2002.

Norops lemniscatus — Nicholson, 2002.

Holotype.— ZMB 4685, Guayaquil, Guayas province, Ecuador.

English common names: Two-marked anoles, roof anoles, west ecuadorian anoles, white-ribbon anoles. Spanish common names: Anolis de dos marcas, anolis de dorso cubierto, anolis de cinta blanca.

Diagnosis

Anolis binotatus belongs to the Draconura, Norops, and Digilimbus clades of Poe et al., 2017 based on work presented here and the phylogenetic tree of Poe et al. (2017). Within Draconura, A. binotatus forms a clade with A. datzorum Köhler et al., 2007, A. gracilipes Boulenger, 1898, A. notopholis Boulenger, 1896 and A. salvini Boulenger, 1885. Anolis binotatus is readily distinguished from these forms by its enlarged middorsal scale rows (middorsals not distinctly enlarged in A. datzorum and A. salvini); its whitish-brown dorsum, with a straight brown stripe behind eye extending above tympanum, and by having 7−14 enlarged middorsal scale rows (greenish brown dorsum with an irregular brown stripe behind eye extending above tympanum, and 9−17 enlarged middorsal scale rows in A. gracilipes); and its lack of a black line from eye to nostril below canthus rostralis (present in A. notopholis).

Anolis binotatus is a brown, small anole (maximum recorded SVL 54.1 mm in males, 55.2 mm in females). Compared to other anoles belonging to the Draconura clade from western Ecuador, it can be distinguished from A. granuliceps Boulenger, 1898 (character states in parentheses) by having a larger size, keeled ventral scales, and a large dewlap past arms well on to chest in males (maximum SVL 49 mm, smooth or very faintly keeled ventral scales, and a small dewlap to arms in males of A. granuliceps). Anolis binotatus differs from A. lynchi Miyata, 1985 in having larger head scales, 6−15 scales across the snout between the second canthals (smaller head scales, 16−29 between second canthals in A. lynchi). Anolis binotatus differs from A. lyra Poe et al., 2009 in having an orange dewlap in males and no dewlap in females (male dewlap red with dark central blotch; female dewlap present, usually blue or white with dark central blotch in A. lyra), and in possessing fewer toe lamellae, 10−15 (16−21 in A. lyra). Anolis binotatus differs from A. maculiventris Boulenger, 1898 in having keeled ventral scales (smooth in A. maculiventris), and from A. parvauritus Williams, 1966 in its brown dorsum (green in A. parvauritus). Anolis binotatus is most similar morphologically to A. gracilipes but differs from it in having 7−14 enlarged middorsal scale rows (9−17 in A. gracilipes); larger ventral scales, 3−7 in 5% SVL (5−11 in 5% SVL in A. gracilipes); a straight brown stripe behind eye extending above tympanum (irregular brown stripe behind eye extending above tympanum in A. gracilipes), a “V” shaped band on the snout, pointing backward (band absent in A. gracilipes), and dorsum of body whitish brown (greenish brown in A. gracilipes; see also genetic distances in the Phylogenetic Relationships section above). Finally, A. sagrei Duméril & Bibron, 1837 (Trachypilus clade), a species recently introduced in Ecuador (Amador et al. 2017) is similar to A. binotatus in having a brown body dorsum, and ventral scales keeled and larger than dorsals. However, A. sagrei differs from A. binotatus in lacking enlarged middorsal scale rows, having an orange, yellow, or red dewlap with a yellow border, and possessing a laterally more strongly compressed tail.

Description and variation

SVL in males 45.0– 54.1 mm, and 45.3–55.2 mm in females. Variation in ratios: head length/SVL 0.22–0.30; head width/SVL 0.12–0.16; HH/SVL 0.10–0.14; interorbital length/SVL 0.02–0.05; interparietal length/SVL 0.02– 0.05; ear height/SVL 0.01–0.03; humerus length/SVL 0.13–0.22; ulna length/SVL 0.11–0.26; hand length/SVL 0.13–0.19; femur length/SVL 0.27–0.36; tibia length/SVL 0.23–0.31; foot length/SVL 0.29–0.41; 4 th toe length/ SVL 0.17–0.33; 4 th toe width/SVL 0.01–0.02. Head scales in supraocular disc and frontal region keeled (single or multiple), rugose, or smooth; 6−15 scales between second canthals; 4−8 scales bordering rostral posteriorly; divided anterior nasal in contact or separated by 1–2 scales from rostral, or anterior nasal in contact or separated by one scale from rostral, or circumnasal separated by one or two scales from rostral; supraorbital semicircles separated by 1–3 scales; supraocular disc with 1–8 gradually enlarged scales; supraocular edge continuous or broken; 1–4 elongate superciliaries followed by gradually smaller scales; 5−8 loreal rows;>15 loreal scales; mid-snout without or with parallel scale rows; rostral with smooth or cleft dorsal edge; frontal region of head with a depression; rostral overlapping the mental or even; interparietal larger than ear opening, separated by 1−4 scales from semicircles; ear round with normal edge or nub; transparent scales in lower eyelids absent; preoccipital scale usually absent; suboculars in contact with supralabials or separated by one scale; 7−10 supralabials counted up to a point below center of eye; 4−7 postmentals; no enlarged sublabials or 1−2 in contact with infralabials; mental divided partially or completely; mental extending further back posteriorly than rostral along edge of mouth, or rostral extending further than mental or even; posterior edge of mental dented, straight or concave; no nuchal crest; dorsal scales keeled; 7−14 middorsal scale rows larger than flank scales; 4−8 middorsal scales in a longitudinal segment representing 5% of SVL; flank scales smooth, homogeneous in size, barely separated by skin; ventral scales larger than dorsals, keeled or slightly keeled, imbricate, and arranged in diagonal or transversal rows; 3−7 midventral scales in a longitudinal segment representing 5% of SVL; inconspicuous axillary pocket present or absent; toepads overlapping distal phalanx in all toes; 10−16 lamellae under phalanges II and III of fourth toe; supradigitals with multiple keels; tail crest absent or serrate; tail laterally compressed or round, with a single or double row of middorsal scales; insolitus tail (Poe 2004) absent; enlarged postcloacal scales absent (in males and females); when adpressed against body the hindlimb reaching posterior to ear, or to ear, or to eye, or between ear-eye, or between eye-snout, or to snout, or past snout.

Nuchal and dorsal folds absent in preserved specimens but often present in males in life; dewlap large, extending posteriorly behind forelimbs (in males), with 6−7 longitudinal rows of one elongate scale each, same size as ventrals, and separated by naked skin. Intraspecific variation in morphological characters in Anolis binotatus is presented in Tables 4 and 5.