Anolis benedikti, Lotzkat, Sebastian, Bienentreu, Joe-Felix, Hertz, Andreas & Köhler, Gunther, 2011

Lotzkat, Sebastian, Bienentreu, Joe-Felix, Hertz, Andreas & Köhler, Gunther, 2011, A new species of Anolis (Squamata: Iguania: Dactyloidae) formerly referred to as A. pachypus from the Cordillera de Talamanca of western Panama and adjacent Costa Rica, Zootaxa 3125, pp. 1-21 : 8-18

Anolis benedikti sp. nov.

Figures 1 F–H, 5–7

Anolis pachypus: Poe and Ibañez 2007 (in part); Norops pachypus: Savage 2002 (in part); Köhler 2008 (in part)

Holotype. SMF 90149 ( Figs. 1 H, 5A and F, 6), adult male, from the north slope of Cerro Pando, leaving the cattle trail to the right after following it to about 1000 m airline north of the large border monument ( Fig. 8 D), 8.9333°N, 82.7131°W, 2310 m asl, Parque Internacional La Amistad ( PILA), Bocas del Toro Province, Panama, close to the border with Costa Rica; collected by Andreas Hertz and Sebastian Lotzkat on 19 November 2009; original field number SL 529.

Paratypes. Collected by Sebastian Lotzkat and Andreas Hertz, if not indicated otherwise. All from the PILA in Panama. Bocas del Toro: north slope Cerro Pando: SMF 89744, male, near type locality, 8.9314°N, 82.7137°W, 2390 m asl, 19 April 2009; SMF 89745–6, male and female, same locality as holotype, 2330 and 2310 m asl, 20 April 2009; SMF 90148, juvenile male, near type locality, 8.9354°N, 82.7128°W, 2280 m asl, 19 November 2009; SMF 91508, male, SL 677–8, subadult male and female, along tributary to Río Changena, 8.9474°N, 82.7098°N, 1980–2000 m asl, 13 July 2010; SMF 91507, female, along tributary to Río Changena, 8.9524°N, 82.7093°N, 1960 m asl, 14 July 2010; SMF 91505–6, SL 686, 695, females, Río Changena, 8.9785°N, 82.6901°N, 1640–1660 m asl, 14–17 July 2010; SMF 91509, juvenile, between Río Clarito and Río Changena, 8.9887°N, 82.6749°N, 1820 m asl, 17 July 2010; Chiriquí: south slope Cerro Pando, Jurutungo: SMF 85276–7, males, 8.9083°N, 82.7168°W, 2010– 2100 m asl, collected by Gunther Köhler on 11 January 2006; SMF 85272–3, subadult males, 8.9109°N, 82.7144°W, 2060 m asl, collected by Gunther Köhler on 10 January 2006; SL 202, female, 8.9119°N, 82.7095°W, 2250 m asl, 6 July 2008; SMF 89505, female, 8.9111°N, 82.7159°W, 2020 m asl, 8 July 2008; SMF 89506, male, 8.9121°N, 82.7096°W, 2200 m asl, 8 July 2008; SMF 89507, male, 8.9114°N, 82.7129°W, 2070 m asl, 15 July 2008.

Referred specimens. See Appendix.

Diagnosis. A medium-sized species (maximum SVL 48.6 mm in males, 48.1 mm in females) of the genus Anolis sensu Poe (2004) , that is most similar in external morphology to A. magnaphallus , A. pachypus , A. pseudopachypus , and A. tropidolepis . These four species and A. benedikti share narrow toe pads, long legs (tip of fourth toe of adpressed hind limb reaches to a point anterior to eye), a dark interorbital bar, and usually at least the indication of a lyriform marking on the occipital region, keeled dorsal scales on head and body, weakly keeled ventral scales at midbody, and the lack of enlarged postcloacal scales in males. Anolis benedikti can be distinguished readily from all four aforementioned species by the coloration of the male dewlap which is red with a yellow anterior portion comprising just the anterior margin or, at most, the anterior third of the dewlap (versus solid purplish red in A. magnaphallus and A. tropidolepis , red with a central yellow blotch in A. pachypus , and solid yellow, sometimes grading into orange-yellow on the posterior portion in A. pseudopachypus ). In addition, A. benedikti differs from A. magnaphallus , A. pachypus , and A. tropidolepis in having more scale rows between the supraorbital semicircles (usually five or more in A. benedikti , versus four or fewer in A. magnaphallus , A. pachypus , and usually four or fewer, rarely five, in A. tropidolepis ), and more scales between supraorbital semicircles and interparietal plate (usually five or more, rarely four, in A. benedikti , versus four or fewer in A. magnaphallus , A. tropidolepis , and usually four or fewer, rarely five, in A. pachypus ). Furthermore, A. benedikti differs from A. magnaphallus , A. pachypus , and A. tropidolepis in having a very dark (almost blackish) gray tongue (versus a very light gray tongue in A. magnaphallus , A. pachypus , and A. tropidolepis ). At first sight, individuals of A. benedikti might be confused with similarly-colored A. humilis . Yet, regardless of sex or age, individuals of the latter species have at least eight longitudinal rows of greatly enlarged, visibly keeled median dorsal scales that are larger than the ventrals, whereas A. benedikti has a maximum of two slightly enlarged middorsal rows, the scales of which are smaller than the ventrals. Males of A. humilis have a red dewlap with a complete (i.e. bordering the whole dewlap) yellow margin, whereas males of A. benedikti exhibit yellow coloration only on the anterior portion of their otherwise solid red dewlap.

Description of the holotype. Adult male as indicated by everted hemipenes and presence of moderate-sized dewlap ( Fig. 1 H); snout-vent length 48.6 mm; tail length 81.7 mm, tail complete; tail slightly compressed in cross section, tail height 2.2 mm, tail width 1.8 mm; axilla to groin distance 18.7 mm; head length 11.9 mm, head length/ snout-vent length ratio 0.24; snout length 5.3 mm; head width 8.1 mm; longest toe of adpressed hind limb reaching to snout; shank length 12.6 mm, shank length/head length ratio 1.1; longest finger of extended forelimb reaching about 4.2 mm beyond tip of snout; longest finger of adpressed forelimb reaching to anterior insertion of hind limbs; scales on snout keeled; 5 postrostrals; 5 scales between nasals; scales in distinct prefrontal depression distinctly keeled; supraorbital semicircles differentiated, separated by a minimum of 7 scales; supraorbital disc composed of 14 slightly enlarged keeled scales; 3 elongated superciliaries; about 5 rows of small keeled scales extending between enlarged supraorbitals and superciliaries; interparietal plate slightly enlarged, 0.6 x 0.3 mm (length x width), surrounded by small scales; 10 scales present between interparietal plate and supraorbital semicircles; canthal ridge distinct, composed of 5 large (posterior) and 3 small (anterior) canthal scales; 15 scales present between second canthals; 17 scales present between posterior canthals; 93/95 keeled loreal scales in a maximum of 12 horizontal rows; subocular scales granular, subocular row ill-defined; 8 supralabials to level below center of eye; ear opening 0.8 x 1.5 mm (length x height); mental distinctly wider than long, almost completely divided medially, bordered posteriorly by 7 postmentals; 8 infralabials to level below center of eye; sublabials undifferentiated; slightly keeled granular scales present on chin and throat; dewlap moderate-sized, extending well onto chest, anterior insertion at level of anterior border of orbit, posterior insertion at level of axilla, with about 10 somewhat irregular, partially interrupted gorgetal-sternal rows of 20–26 scales per row; dorsum of body with keeled scales, 2 medial rows of slightly enlarged scales, largest dorsal scales subimbricate, about 0.40 x 0.35 mm (length x width); about 66 medial dorsal scales in one head length; about 112 medial dorsal scales between axilla and groin; lateral scales pointed, granular and homogeneous, average size 0.18 mm in diameter; ventrals at midbody weakly keeled, slightly mucronate, and subimbricate, about 0.30 x 0.40 mm (length x width); about 42 ventral scales in one head length; about 58 ventral scales between axilla and groin; about 162 scales around midbody; caudal scales strongly keeled, without whorls of enlarged scales; no enlarged postcloacal scales; tubelike axillary pocket not developed; scales on dorsal surface of forelimb strongly keeled, mucronate and imbricate, about 0.30 x 0.45 mm (length x width); digital pads dilated, dilated pad about 2 times width of non-dilated scales on distal phalanx; distal phalanx narrower than and raised from dilated pad; 23/22 lamellae under phalanges II–IV of fourth toe; 9/10 scales under distal phalanx of fourth toe; 15 lamellae under phalanges II–IV of fourth finger; 9 scales under distal phalanx of fourth toe.

The completely everted hemipenis is a stout bilobate organ; sulcus spermaticus bordered by well-developed sulcal lips and opening at base of apex into two broad concave areas, one on each lobe; a large asulcate ridge-like processus present; lobes strongly calyculate, truncus with transverse folds.

Coloration in life. The dewlap ( Fig. 1 H) was recorded as follows: Flame Scarlet (15) with a suggestion of Chrome Orange (16) on posterior three fourths, Orange Yellow (18) on anterior fourth and anterior margin; dewlap scales Sepia (119) and dirty white. Otherwise, no detailed notes of the holotype were taken; see Figures 5 A and F for photographs of the holotype in life.

Coloration after 18 months of preservation in 70% ethanol is similar to that in life, apart from that all reddish shades have faded, especially on the dewlap, which now appears dirty white.

Variation. The paratypes and referred specimens agree well with the holotype in terms of general morphology and pholidosis (see Table 1 ). Four of the 32 examined specimens (SMF 89507, 91505; SL 686, 695) have only four scales between the supraorbital semicircles. Two specimens (SMF 89507, 91505) have only four scales between the supraorbital semicircles and the interparietal plate. Among the 17 adult males included in statistical analyses, the variability in the nasal region ( Fig. 7 ) was recorded as follows: Type A one specimen (5.9 %); Type B two specimens (11.8 %); Type C four specimens (23.5 %); Type D five specimens (29.4 %); Type E three specimens (17.6 %); Type F two specimens (11.8 %).

Similar to what we observed in the other four members of the Anolis pachypus complex, the coloration of A. benedikti is rather variable among individuals, mostly concerning the extent and shape of color patterns, i.e., stripes, blotches, and bands on dorsal and lateral surfaces (compare Figs. 5 F– O). Common patterns include a broad middorsal band which is often interspersed with blotches or chevrons, flanked by lateral markings, or disintegrates into diamond-shaped markings posteriorly ( Figs. 5 F–J); a narrow light middorsal stripe, often bordered by narrow dark stripes (Figs. K–L); and series of triangular markings pointing laterally from middorsum (with or without narrow medial light stripe) which are either offset to form a zig-zag pattern or arranged symmetrically to form a series of diamonds ( Figs. 5 M– O). Most specimens exhibit contrasting markings on dorsal and lateral surfaces of the head, with postocular stripes and an interorbital bar being the most common.

Furthermore, these anoles are capable of metachrosis and, therefore, the coloration of a given individual is subject to physiological change according to time or situation. Most individuals encountered at night exhibited light tonalities, but could turn very dark during the day or when handled ( Figs. 5 K–L). Individuals spotted during the day all appeared dark. In the course of metachrosis, the conspicuousness of individual color patterns may vary considerably (as in the postocular stripes between the two females in Figs. 5 C–D).

The coloration of the male paratype SMF 89506 ( Figs. 1 G, 5B and G) was recorded as follows: Dorsal and lateral ground color Antique Brown (37) suffused with Light Drab (119C) and Tawny (38), laterally grading into Olive-Gray (42) mottled with Burnt Umber (22); a Mars Brown (223A) postorbital stripe mottled with Sepia (219) continues to form a dorsolateral band, which disintegrates into a series of blotches around midbody; a few Sepia (119) middorsal blotches present; dorsal and lateral surfaces of head Light Drab (119C); occipital region Mars Brown (223A) mottled with Sepia (219); a Mars Brown (223A) interorbital bar and a preorbital stripe bordered by Sepia (219); a dirty white with a suggestion of Yellow-Green (58) lateral postorbital stripe extending to shoulder; dorsal and lateral surfaces of limbs and tail Antique Brown (37), with irregular Sepia (219), Tawny (38), and Olive- Gray (42) transverse stripes; ventral surface of body and head Pearl Gray (81) suffused with Cinnamon-Rufous (40); ventral surface of tail with a Cinnamon-Rufous (40) midventral stripe; iris Brick Red (132A); tongue Plumbeous (78) with a suggestion of Indigo (73); gular region Flame Scarlet (15); dewlap Flame Scarlet (15) with a small Pearl Gray (81) region at posterior insertion and an Orange Yellow (18) anterior margin; dewlap scales dirty white bordered by Sepia (119) or Sepia (219).

continued next page The dewlap coloration of the juvenile male SMF 90148 from the vicinity of the type locality was recorded as follows: Flame Scarlet (15), with a suggestion of Chrome Orange (16), grading into Orange Yellow (18) on anterior margin; dewlap scales Sepia (119). As illustrated in Figs. 1 F–H, the extent of the yellow coloration on the anterior portion of the male dewlap of A. benedikti varies from covering just the basal part of the anterior margin ( Fig. 1 F) to covering the anterior third of the dewlap ( Fig. 1 H). The female dewlap ( Figs. 5 C–E) is much smaller than in males, and usually of a whitish color ( Fig. 5 C). Nevertheless, some female individuals bear reddish coloration around their gular area. These females tend to exhibit a reddish dewlap ( Figs. 5 D–E), sometimes even resembling that of males, like the female paratype SMF 89746 ( Fig. 5 E), that was recorded as follows: Flame Scarlet (15) posteriorly, grading into Orange Yellow (18) on anterior half and into dirty white with a suggestion of Chamois (123D) on posterior base; dewlap scales mostly Sepia (119), some dirty white.

Natural history notes. By far, most specimens were encountered at night sleeping on vegetation, i.e., on the upper surface of leaves or on twigs, usually 0.5–2 m above ground. During daytime, specimens were spotted when fleeing on the ground, mostly seeking shelter between tree buttresses or bushes. The vegetation at the type locality ( Figs. 8 E–F) is a lush ridgetop cloud forest rich in palms and with abundant growth of epiphytes that cover virtually every surface. Descending either slope of Cerro Pando, the trees become higher and the canopy cover denser. A detailed description of the forest environments around Cerro Pando was given by Myers (1969). Presently, especially below 2000 m elevation on either slope, major clearings for cattle farming disrupt the forest ( Figs. 8 A and G). In these pastures, Anolis benedikti can be found associated with bushes, single trees, or remnants of cut trees. Annual total precipitation at the type locality is approximately 2600 mm and mean annual temperature approximately 13.6 °C, indicating the presence of the Lower Montane Wet Forest life zone according to the Holdridge (1967) classification. Our datalogger recordings (N = 205) at the type locality yielded a temperature range of 11.1–16.7 °C, with a mean of 12.7 °C and standard deviation ± 1.1 °C. Without exception, our collecting localities of A. benedikti lie above 1600 m asl on the Caribbean slope and above 1900 m asl on the Pacific slope and receive more than 2000 mm of annual precipitation, i.e., are assignable to the Lower Montane Wet Forest life zone. In the immediate surroundings of the type locality, A. benedikti was the only reptile species we could find. Reptile species collected on the Caribbean versant along the trail descending the north slope of Cerro Pando ( Figs. 8 G–H) between 1640 and 2000 m asl were Anolis capito (Peters) , A. humilis (Peters) , A. kemptoni Dunn , A. microtus (Cope) , Ptychoglossus plicatus (Taylor) , and Sibon annulatus (Günther) . Reptile species collected on the Pacific versant between 2000 and 2400 m asl, in the high valley known as Jurutungo ( Figs 8 A–B) and along the trail ascending the south slope of Cerro Pando from Jurutungo, were A. kemptoni , A. pachypus, Sceloporus malachiticus Cope, Geophis godmani Boulenger , Rhadinaea calligaster (Cope) , and Bothriechis nigroviridis Peters.

Geographic distribution. As presently understood, Anolis benedikti inhabits montane elevations along the Talamancan highlands of eastern Costa Rica and extreme western Panama ( Fig. 9 ). All individuals we examined originated from between 82.728 °W and 82.425 °W. The species seems to dwell predominantly on the Caribbean versant of the Serranía de Talamanca, where we found it from the continental divide at up to 2390 m asl (Cerro Pando). Descending the Caribbean slope, we encountered it down to 1640 m asl at the Río Changena, 1740 m asl along the Culebra trail, and 1650 m asl along the Pianista trail. However, in at least three regions, this species is also found on the Pacific versant, where it occurs next to its close relatives: At Las Tablas, in the Costa Rican Province of Puntarenas next to the Panamanian border, SMF 92134 was found at 1960 m asl, less than a kilometer from the nearest collected A. pachypus at 1880 m asl. Just a few kilometers east, in Jurutungo (the headwaters of Río Candela in Chiriquí Province, Panama; Fig. 8 A), we found A. benedikti as low as 2020 m asl at certain places, less than 50 m away from the nearest collection site of A. pachypus . Further east, on the slopes of Volcán Barú north of Boquete, about one kilometer west of the ranger station of Alto Chiquero close to the upper Río Caldera, we collected four males of A. benedikti at 1920 m asl, as well as a single female farther south. These two sites are close to different collection sites (at both lower and higher elevations) as well as to the type locality of A. magnaphallus .

Etymology. The specific name is a patronym for Benedikt Stökl, Germany, in recognition of the financial support of taxonomic research provided by Karsten Lutz through the BIOPAT initiative.

Figures

FIGURE 1. Dewlaps of anoles related to Anolis pachypus from Costa Rica and western Panama, representative of the five OTUs (width of tweezers 3 mm): (A) OTU A (SMF 89729, Sendero La Cascada, Bajo Mono, Chiriquí, Panama); (B) OTU B (GK 2297, road from San José de la Montaña to Birrí, Heredia, Costa Rica); (C) OTU C (GK 2355, road from San Isidro de El General to Cerro de la Muerte, San José, Costa Rica); (D) OTU C (SL 439, Jurutungo, Chiriquí, Panama); (E) OTU C (SL 524, Jurutungo, Chiriquí, Panama); (F) OTU D (SMF 89507, Jurutungo, Chiriquí, Panama); (G) OTU D (SMF 89506, Jurutungo, Chiriquí, Panama); (H) OTU D (SMF 90149, N slope Cerro Pando, Bocas del Toro, Panama); (I) OTU E (SMF 89519, La Nevera, Comarca Ngöbe-Buglé, Panama); (J) OTU E (SL 510, La Nevera, Comarca Ngöbe-Buglé, Panama).

FIGURE 5. Type specimens of Anolis benedikti in life: (A) male holotype (SMF 90149); (B) male from Jurutungo, SMF 89506; (C – E) female dewlaps: (C) SMF 91506; (D) SL 695; (E) SMF 89746; (F – O) variation in dorsal patterns: (F) SMF 90149, male holotype; (G) SMF 89506, male; (H) SMF 89745, male; (I) SMF 91508, male; (J) SL 202, female; (K) SMF 89746, female, at night; (L) SMF 89746 the next day; (M) SMF 91507, female; (N) SL 686, female; (O) SMF 91505, female.

FIGURE 7. Designation of nasal scalation types. (A) Type A (SMF 90149); (B) Type B (SMF 92134); (C) Type C (SL 678); (D) Type D (SMF 85264); (E) Type E (SMF 89506); (F) Type F (85272). Scale bars equal 1 mm. See Text for details.

FIGURE 8. Habitats of Anolis benedikti: (A – C) Pacific slope in Chiriquí province: (A) view over Jurutungo north toward the continental divide (hidden in clouds), the ranges of A. benedikti and A. pachypus meet in the background; (B) forest trail at Jurutungo, one of the lowest collection sites of A. benedikti on the Pacific slope, 2070 m asl; (C) cloud forest near continental divide east of Cerro Pando, 2460 m asl; (D) international border monument on cattle trail along the continental divide, 2450 m asl; (E – H) Caribbean slope in Bocas del Toro province: (E) cloud forest at the type locality, 2310 m asl; (F) cloud forest near the type locality, 2390 m asl; (G) clearing for cattle along tributary to Río Changena, 1810 m asl; (H) Río Changena, the lowest collection site on the Caribbean slope, 1640 m asl.

FIGURE 9. Collection localities of Anolis benedikti (circles) and other members of the A. pachypus complex (A. magnaphallus: squares; A. pachypus: upright triangles; A. pseudopachypus: inverted triangles) in western Panama and adjacent Costa Rica. Hollow symbols represent type localities.

Tables

TABLE 1. Selected measurements, proportions and scale characters of Anolis benedikti, A. magnaphallus, A. pseudopachypus, A. pachypus, and A. tropidolepis. Range is followed by mean value and one standard deviation in parentheses. For abbreviations see text. Based on specimens with data on dewlap coloration plus additional specimens (males and females) for which the respective exact geographic origin and morphological key characters allowed for a precise species assignment.

	A. pachypus 3 11, Ƥ 5	A. pseudopachypus 3 16, Ƥ 5	A. magnaphallus 3 9, Ƥ 6	A. tropidolepis 3 10, Ƥ 0	A. benedikti 3 20, Ƥ 13
SVL 3	38.6–50.7 (45.5 ± 3.9)	30.0–45.5 (40.7 ± 4.6)	31.4–54.1 (47.5 ± 7.1)	41.4–52.3 (47.4 ± 3.5)	34.6–48.6 (41.6 ± 3.5)
Ƥ	47.6–51.1 (49.0 ± 1.3)	39.4–47.3 (45.0 ± 3.7)	46.5–58.9 (52.6 ± 5.4)		34.4–48.1 (42.5 ± 4.8)
Tail length / SVL 3	1.56–1.94 (1.81 ± 0.14)	1.61–2.06 (1.93 ± 0.12)	1.04–1.71 (1.58 ± 0.22)	1.72–2.18 (1.99 ± 0.18)	1.22–2.16 (1.76 ± 0.21)
Ƥ	1.64–1.72 (1.69 ± 0.04)	1.70–1.78 (1.74 ± 0.04)	1.38–1.57 (1.48 ± 0.08)		1.42–1.90 (1.67 ± 0.12)
Tail diameter vertical / 3 horizontal	1.11–1.31 (1.25 ± 0.07)	1.06–1.29 (1.17 ± 0.07)	1.08–1.45 (1.21 ± 0.10)	1.05–1.31 (1.19 ± 0.08)	1.00–1.43 (1.19± 0.10)
Ƥ	1.13–1.24 (1.18 ± 0.05)	1.13–1.36 (1.21 ± 0.12)	1.10–1.27 (1.18 ± 0.07)		1.12–1.27 (1.19 ± 0.06)
HL / SVL 3	0.24–0.28 (0.26 ± 0.01)	0.24–0.29 (0.26 ± 0.01)	0.24–0.28 (0.26 ± 0.01)	0.25–0.26 (0.26 ± 0.01)	0.24–0.31 (0.27 ± 0.02)
Ƥ	0.25–0.27 (0.26 ± 0.01)	0.24–0.27 (0.25 ± 0.01)	0.25– 0.26 (0.25 ± 0.01)		0.25– 0.29 (0.27 ± 0.02)
HL / HW 3	1.39–1.59 (1.50 ± 0.05)	1.43–1.65 (1.53 ± 0.06)	1.42–1.58 (1.52 ± 0.05)	1.47–1.61 (1.54 ± 0.04)	1.43–1.68 (1.52 ± 0.07)
Ƥ	1.45–1.56 (1.50 ± 0.05)	1.51–1.63 (1.56 ± 0.06)	1.47–1.58 (1.53 ± 0.04)		1.44–1.60 (1.52 ± 0.05)
Shank length / SVL 3	0.27–0.32 (0.28 ± 0.01)	0.29–0.31 (0.30 ± 0.01)	0.25–0.27 (0.26 ± 0.01)	0.24–0.29 (0.27 ± 0.01)	0.26–0.33 (0.29 ± 0.02)
Ƥ	0.26–0.28 (0.27 ± 0.01)	0.27–0.30 (0.28 ± 0.01)	0.22–0.27 (0.25 ± 0.02)		0.27–0.33 (0.29 ± 0.02)
Axilla-groin distance / 3 SVL	0.38–0.46 (0.41 ± 0.03)	0.34–0.43 (0.39 ± 0.02)	0.37–0.44 (0.40 ± 0.02)	0.37–0.40 (0.38 ± 0.01)	0.31–0.45 (0.39 ± 0.03)
Ƥ	0.37–0.45 (0.41 ± 0.03)	0.40–0.42 (0.41 ± 0.01)	0.37–0.45 (0.40 ± 0.03)		0.34–0.45 (0.41 ± 0.04)
Subdigital lamellae under phalanges II–IV of 4th toe	21–24 (22.6 ± 1.0)	21–24 (22.3 ± 1.0)	19–24 (22.3 ± 1.3)	20–22 (21.5 ± 0.7)	19–25 (22.2 ± 1.4)
Number of scales between SS	2–4 (3.2 ± 0.7)	4–8 (6.2 ± 1.2)	3–4 (3.4 ± 0.5)	3–5 (4.1 ± 0.6)	4–8 (5.8 ± 1.1)
Number of scales between IP and SS	3–5 (3.4 ± 0.6)	4–11 (7.6 ± 1.7)	2–4 (3.2 ± 0.5)	2–4 (3.5 ± 0.7)	4–10 (7.2 ± 1.5)
Number of SPL to level below center of eye	6–7 (6.1 ± 0.4)	6–8 (6.6 ± 0.7)	6–9 (7.1 ± 0.8)	5–8 (6.5 ± 0.9)	5–8 (6.3 ± 0.7)
Number of INL to level below center of eye	6–8 (6.9 ± 0.7)	6–8 (7.2 ± 0.8)	6–9 (7.8 ± 0.7)	5–7 (6.7 ± 0.7)	5–8 (6.7 ± 0.8)