Evidence for the recognition of two species of Anolis formerly referred to as A. tropidogaster (Squamata: Dactyloidae) Author Köhler, Gunther Author Batista, Abel Author Vesely, Milan Author Ponce, Marcos Author Carrizo, Arcadio Author Lotzkat, Sebastian text Zootaxa 2012 3348 1 23 journal article 10.5281/zenodo.281518 359289e5-7493-4abc-9117-b45c62a0354a 1175-5326 281518 Anolis tropidogaster Hallowell, 1856 Figures 1 a; 3a, b; 4b, d, f, h; 6a, b, c, g, h; 7a, b; 8 Anolis tropidogaster Hallowell 1856 :224 ; holotype (ANSP 7618) from “New Grenada ”. Dunn (1930) , Barbour (1934; in part.) , Barbour and Loveridge (1946) , Breder (1946) , Evans (1947) , Etheridge (1959) , Donoso-Barros (1968) , Peters and Donoso-Barros (1970; in part. ) , Williams (1976) , Ayala (1986) , Pefaur (1992) , Auth (1994; in part.) , Williams et al. (1995 ; in part.) , Young et al. (1999 ; in part.) , Ibáñez et al. (2001; in part.) , Moreno-Bejarano & Álvarez-León (2003) , Poe (2004; in part.) , Carvajal-Cogollo and Urbina-Cardona (2008) , Moreno-Arias et al. (2008) , Medina-Rangel (2011) . Anolis albi Barbour 1932 :101 ; holotype (MCZ 32301) from “Andagoya, Choco, western Colombia .” Anolis stigmosus Bocourt 1869 :43 ; syntypes (MNHN 2427 and 2427A) from “la Colombie et ont été recueillis près de la rivière de la Magdeleine.” Boulenger (1885) . Norops tropidogaster . Köhler (2003 , 2008 ; in part.) FIGURE 6. (a) Holotype of Anolis tropidogaster (ANSP 7618). (b) Anolis stigmosus syntype MNHN 2427. (c) Anolis stigmosus syntype MNHN 2427a. (d) Anolis gaigei paratype (UMMZ 48334) dorsal view. (e) Anolis gaigei (UMMZ 48322) hemipenis. (f) Anolis gaigei paratype (UMMZ 48334) cloacal region. (g) Anolis albi paratype (MCZ 32302) dorsal view. (h) MCZ 32302 cloacal region. Diagnosis. A medium-sized species (SVL in largest specimen examined 55 mm ) of the genus Anolis (sensu Poe 2004 ) that differs from all other Lower Central American beta anoles (sensu Etheridge 1967 ) in that it is longlegged (longest toe of adpressed hind leg reaches to at least center of eye, usually to a point between anterior border of eye and nostril); has strongly keeled ventral scales, a large almost uniformly purplish red (in life) colored dewlap in males; postcloacal scales not enlarged in the majority of males, some male with slightly enlarged postcloacal scales; a large bilobed hemipenis in males, and no tube-like axillary pocket. Anole species from Lower Central America that are somewhat similar in appearance to A. tropidogaster are A. cupreus , A. gaigei , A. osa , and A. polylepis . FIGURE 7. Adult individuals in life of (a) Anolis tropidogaster (SMF 93598), male from Laguna de Matusagaratí, Darién, Panama. (b) Anolis tropidogaster (SMF 93597), male from Cruce de Mono Station at Parque Nacional Darién, Darién, Panama. (c) Anolis gaigei (SMF 91918), male from Finca La Providencia, near Ponuga, Veraguas, Panama. (d) Anolis gaigei (SMF 91917), female from road from Interamericana to Horconcitos, Chiriquí, Panama. Anolis gaigei has a small, thin, unilobed hemipenis in males (large, bulbous and bilobed in A. tropidogaster ); a male dewlap that is orange yellow with a darker orange central area (uniform purplish red, sometimes with a paler orange central area in A. tropidogaster ); a pair of greatly enlarged postcloacal scales in males (these scales usually not differentiated in A. tropidogaster ); the dorsal head scales, especially in the parietal region large and flat (smaller and bearing minute tubercles in A. tropidogaster ); middorsal caudal scales distinctly enlarged and forming a regular series (only slightly enlarged and somewhat irregularly arranged in A. tropidogaster ); outer postmental scales greatly enlarged relative to medial ones (only slightly enlarged relative to medial ones in A. tropidogaster ). Anolis polylepis and A. osa have smooth ventral scales at midbody and a larger, mostly uniform orange male dewlap (in some parts of its range in Costa Rica , considerable variation was observed in male dewlap coloration of A. polylepis , see Köhler et al. 2010 ). The male dewlap of A. cupreus is brown to pink with an orange margin. For variation in selected morphometric and scalation characters of A. tropidogaster see Table 1 . Description. Anolis tropidogaster is a medium-sized anole (maximum recorded SVL 54.0 mm in males, 55.0 mm in females); dorsal head scales ( Fig. 8 ) in internasal region keeled, in prefrontal, parietal, and frontal areas rugose to tuberculate; deep frontal depression present, parietal depression absent; 5–8 (6.75 ± 0.69) postrostrals; anterior nasal usually single, occasionally divided, usually in contact with rostral and first supralabial ( Fig. 9 , Tab. 2 ); 6–11 (8.97 ± 0.94) internasals; canthal ridge sharply defined; scales comprising supraorbital semicircles weakly keeled, largest scale in semicircles about same size as largest supraocular scale; supraorbital semicircles well defined; 2–4 (3.06 ± 0.58) scales separating supraorbital semicircles at narrowest point; 2–5 (2.95 ± 0.73) scales separating supraorbital semicircles and interparietal at narrowest point; interparietal well defined, greatly enlarged relative to adjacent scales, surrounded by scales of moderate size, longer than wide, usually larger than ear opening; enlarged supraoculars not in contact with supraorbital semicircles; 2 elongate superciliaries, posterior one much shorter than anterior one; 2–3 enlarged canthals; 10–17 (13.44 ± 1.71) scales between second canthals; 12–20 (15.11 ± 1.86) scales present between posterior canthals; loreal region slightly concave, 33–64 (44.77 ± 8.80) mostly keeled (some smooth or rugose) loreal scales in a maximum of 5–8 (6.22 ± 0.83) horizontal rows; 6–9 (7.00 ± 0.93) supralabials to level below center of eye; suboculars keeled, separated from supralabials by one scale row; ear opening vertically oval; scales anterior to ear opening granular, similar in size to those posterior to ear opening; 6–10 (7.75 ± 0.97) postmentals, outer pair usually largest; keeled granular scales present on chin and throat; male dewlap large, extending onto chest; 6–8 horizontal gorgetal-sternal rows with 26–38 scales per row, rows somewhat irregular, some of them with pairs or triplets of scales, apical portion of dewlap between marginal pairs and last gorgetal-sternal row free of scales; modal number of marginal pairs 4–5; female dewlap small or absent; no nuchal crest or dorsal ridge; 2 middorsal scale rows slightly enlarged, weakly keeled, dorsal scales lateral to middorsal series gradually larger than granular lateral scales; no enlarged scales scattered among granular laterals; 42–73 (57.0 ± 7.53) dorsal scales along vertebral midline between levels of axilla and groin in males, 53–78 (61.8 ± 10.35) in females; 29–46 (37.6 ± 4.39) dorsal scales along vertebral midline contained in one head length in males, 30–42 (35.3 ± 4.12) in females; ventral scales on midsection about the same size as largest dorsal scales; ventral body scales moderately to strongly keeled, subimbricate to imbricate; 40–65 (51.9 ± 5.50) ventral scales along midventral line between levels of axilla and groin in males, 40–64 (49.0 ± 8.60) in females; 28–46 (36.0 ± 4.52) ventral scales contained in one head length in males, 27–36 (31.6 ± 2.92) in females; 112–144 (124.7 ± 7.90) scales around midbody in males, 122–162 (133.3 ± 12.6) in females; tubelike axillary pocket absent; preanal scales not keeled; postcloacal scales usually not enlarged, or, if differentiated, then only slightly enlarged; tail laterally compressed in cross section, tail height/tail width 1.07–1.53 (1.29 ± 0.11); basal subcaudal scales smooth; lateral caudal scales keeled, homogeneous; dorsal medial caudal scale row slightly enlarged, keeled, not forming a crest; most scales on lateral surface of antebrachium weakly keeled, unicarinate; 23–29 (25.78 ± 1.46) subdigital lamellae on Phalanges II–IV of Toe IV of hind limbs; SVL 42.0–54.0 (48.9 ± 2.96) mm in males, 40.0–55.0 (47.0 ± 5.20) mm in females; HL 11.3–14.6 (12.9 ± 0.72) mm in males, 11.1–14.1 (12.5 ± 1.00) mm in females; tail length 65.0–97.0 (83.9 ± 10.70) mm in males, 69.0–87.0 (80.0 ± 6.54) mm in females; shank length 12.2–15.9 (14.0 ± 1.06) mm in males, 11.2–16.0 (13.1 ± 1.65) mm in females; tail length/SVL 1.35–1.94 (1.75 ± 0.20) in males, 1.28–1.88 (1.68 ± 0.24) in females; HL/SVL 11.3–14.6 (12.9 ± 0.72) in males, 11.1–14.1 (12.5 ± 1.00) in females; shank length/SVL 0.25–0.33 (0.29 ± 0.02) in males, 0.25–0.30 (0.28 ± 0.01) in females; shank length/HL 0.94–1.22 (1.09 ± 0.07) in males, 0.90–1.16 (1.05 ± 0.08) in females. Of 45 specimens examined, the longest toe of the adpressed hind leg reaches to mid-eye in 3 individuals (6.7%), to anterior margin of eye in 29 individuals (64.4%), and to a point between eye and snout in 13 individuals (28.9%). FIGURE 8. Head scalation in Anolis tropidogaster (MHCH 1640). Scale bars = 1.0 mm. FIGURE 9. Designation of nasal scalation types. (a) Nasal type 1 (MHCH 1636); (b) Nasal type 2 (SMF 91901). Scale bars = 1.0 mm. See Text for details. FIGURE 10. Habitat of Anolis tropidogaster at Cruce de Mono Station, Darién, Panama. Coloration in life of an adult male (MHCH 2375) was recorded as follows: Dorsal ground color Drab (27) with Olive Brown (28) vertebral band, postorbital band Dark Drab (119B) extending to level of midbody; a longitudinal level Buff (124) stripe from tympanum to insertion of hind legs; dorsal surface of head Dark Drab (119B), with a medially interrupted Dark Brownish Olive (129) interorbital bar; forelegs and hind legs Drab (27) suffused with Olive Brown (28); dorsal surface of tail Drab (27) with indistinct Olive Brown (28) bands; ventral surfaces of head, body, and limbs Straw Yellow (57); dewlap Chrome Orange (16), grading into Spectrum Orange (17) at center; gorgetals dirty white; iris Sepia (119). Natural history notes. At the evergreen forest sites ( Fig. 10 ) in the Darién Province, Panama , visited by AB, Anolis tropidogaster was an uncommon species. AB and MP encountered it within the forest on low vegetation between 0.5 and 2.0 m above the ground. Occasionally, individuals were observed on the ground. At night, these animals sleep in the usual anole-like fashion on twigs or the upper surface of leaves. One adult male was collected at a forest edge at Matusagaratí Lake, a thin forest belt between the lake and pasture areas. Another individual was captured at the base of a mangrove tree (MHCH 1636) at a riverside at Caserete, Chepigana, Darién, Panama ; in the same region three individuals were seen in a cativo ( Prioria copaifera ) forest on low vegetation. Another individual (MHCH 1701) was captured during night sampling in a semideciduous forest on a bush at 1.5 m above ground at Sol Poniente, Chepigana, Darién, Panama . Some ecological observations on A. tropidogaster were published by Sexton et al. (1964) . Geographic Distribution. Anolis tropidogaster is distributed widely in Colombia and in eastern Panama ( Figs. 2 , 13 ). The claim that this species occurs in Ecuador seems to go back to Donoso-Barros (1968) , who based this view on the holotype of A. lemniscatus (from “Puente del Chimbo”, see Boulenger 1898 ), a taxon then placed in the synonymy of A . tropidogaster . We were unable to find additional evidence for the occurrence of A . tropidogaster in Ecuador and therefore remove this species from the list of known reptiles from this country. The documented vertical range of the species is from near sea level to about 1100 m . TABLE 1. Selected measurements, proportions and scale characters of Anolis gaigei and A. tropidogaster . Range is followed by mean value and standard deviation in parentheses. For abbreviations see text.

A. gaigei 3 49, Ƥ 30	A. tropidogaster 3 27, Ƥ 9
SVL 3	36.0–52.5 (45.6 ± 3.40)	42.0–54.0 (48.9 ± 2.96)
Ƥ	37.0–52.0 (44.7 ± 4.75)	40.0–55.0 (47.0 ± 5.20)
Tail length 3 Ƥ	59.0–115.0 (95.0 ± 11.63) 82.0–100.7 (92.3 ± 8.33)	65.0–97.0 (83.9 ± 10.70) 69.0–87.0 (80.0 ± 6.54)
HL 3	10.5–13.2 (12.1 ± 0.65)	11.3–14.6 (12.9 ± 0.72)
Ƥ	10.6–12.1 (11.5 ± 0.48)	11.1–14.1 (12.5 ± 1.00)
HW 3	6.5–7.9 (7.3 ± 0.37)	6.6–9.0 (7.8 ± 0.46)
Ƥ Shank length 3	6.5–7.7 (7.1 ± 0.37) 11.2–15.6 (13.6 ± 0.85)	6.4–8.7 (7.5 ± 0.66) 12.2–15.9 (14.0 ± 1.06)
Ƥ	10.5–14.0 (12.4 ± 0.82)	11.2–16.0 (13.1 ± 1.65)
Axilla–groin distance 3	11.2–19.5 (17.0 ± 1.90)	16.9–24.5 (19.9 ± 1.57)
Ƥ	13.6–24.0 (17.9 ± 2.64)	18.3–24.9 (20.1 ± 2.12)
Tail length / SVL 3	1.37–2.34 (2.09 ± 0.21)	1.35–1.94 (1.75 ± 0.20)
Ƥ Tail diameter vertical / horizontal 3 Ƥ	1.75–2.27 (2.10 ± 0.13) 1.05–1.47 (1.23 ± 0.10) 1.06–1.31 (1.17 ± 0.07)	1.28–1.88 (1.68 ± 0.24) 1.07–1.53 (1.29 ± 0.12) 1.13–1.36 (1.26 ± 0.08)
HL / SVL 3	0.26–0.30 (0.27 ± 0.01)	0.24–0.29 (0.26 ± 0.01)
Ƥ	0.24–0.29 (0.26 ± 0.02)	0.24– 0.32 (0.27 ± 0.02)
HL / HW 3	1.56–1.75 (1.66 ± 0.06)	1.49–1.89 (1.73 ± 0.09)
Ƥ	1.51–1.75 (1.63 ± 0.06)	1.60–1.75 (1.67 ± 0.05)
Shank length / SVL 3 Ƥ	0.28–0.32 (0.30 ± 0.01) 0.23–0.33 (0.29 ± 0.02)	0.25–0.33 (0.29 ± 0.02) 0.25–0.30 (0.28 ± 0.01)
Axilla–groin distance / SVL 3 Ƥ	0.24–0.43 (0.38 ± 0.04) 0.36–0.48 (0.40 ± 0.03)	0.34–0.46 (0.41 ± 0.03) 0.41–0.46 (0.44 ± 0.02)
Subdigital lamellae of 4th toe	20–28 (25.45 ± 1.65)	23–29 (25.78 ± 1.46)
Number of scales between SS	1–4 (2.10 ± 0.62)	2–4 (3.06 ± 0.58)
Number of scales between IP and SS Number of scales between SO and SPL	2–5 (3.03 ± 0.65) 0–1 (0.69 ± 0.47)	2–5 (2.95 ± 0.73) 1–1 (1.00 ± 0.00)
Number of SPL to level below center of eye Number of INL to level below center of eye	6–9 (7.31 ± 0.65) 6–9 (7.33 ± 0.61)	6–9 (7.00 ± 0.93) 5–9 (7.11 ± 0.85)
Total number of loreals	22–59 (40.25 ± 5.79)	33–64 (44.77 ± 8.80)
Number of horizontal loreal scale rows Number of postrostrals	5–8 (6.31 ± 0.60) 5–7 (6.06 ± 0.54)	5–8 (6.22 ± 0.83) 5–8 (6.75 ± 0.69)
Number of postmentals	5–9 (6.35 ± 0.80)	6–10 (7.75 ± 0.97)
Number of scales between nasals Number of scales between 2nd canthals	5–9 (7.03 ± 1.01) 8–14 (10.93 ± 1.14)	6–11 (8.97 ± 0.94) 10–17 (13.44 ± 1.71)
Number of scales between posterior canthals	10–16 (13.10 ± 1.32)	12–20 (15.11 ± 1.86)
Number of medial dorsal scales in one head length Number of medial ventral scales in one head length Number of scales around midbody	28–56 (37.95 ± 6.09) 25–48 (33.21 ± 4.87) 94 – 138 (117.60 ± 8.74)	29–46 (37.03 ± 4.38) 27–46 (34.89 ± 4.58) 112 – 162 (126.63 ± 9.67)
Numbers of medial dorsal scales between axilla and groin Numbers of medial ventral scales between axilla and groin	53 – 83 (62.72 ± 5.92) 40 – 58 (48.88 ± 4.01)	42 – 78 (58.06 ± 8.34) 40 – 65 (51.26 ± 6.32)

TABLE 2. Frequency distribution of nasal region and dorsal pattern types in Anolis gaigei and A. tropidogaster . Anolis gaigei A. tropidogaster 35 9 Nasal region Type A 3 (8.6%) 7 (20.6%) Nasal region Type B 32 (91.4%) 27 (79.4%)