Cnemidophorus duellmani, Mccranie, James R. & Hedges, S. Blair, 2013

Mccranie, James R. & Hedges, S. Blair, 2013, A review of the Cnemidophorus lemniscatus group in Central America (Squamata: Teiidae), with comments on other species in the group, Zootaxa 3722 (3), pp. 301-316 : 308-312

Cnemidophorus duellmani sp. nov.

( Fig. 6 )

Cnemidophorus lemniscatus lemniscatus: Burt 1931: 30 (part). Cnemidophorus lemniscatus lemniscatus: Breder 1946: 428 . Cnemidophorus lemniscatus: Echternacht 1968: 153 (part). Cnemidophorus lemniscatus: Jaramillo et al. 2010: 621 .

Holotype. KU 80542, an adult male from El Real de Santa María, Darién Province, Panama, collected 9 January 1964 by William E. Duellman and Charles J. Cole. Anonymous (1969) gave the coordinates 8°08’N, 77°43’W for El Real de Santa María.

Paratypes (14). KU 80552, 80562, 80564, 80566, 80568 (all adult males), KU 80558, 80572, 80574, 80576– 77, 80580 (all adult females), KU 80581 (subadult male), KU 80582–83 (both juveniles), all with the same locality data as the holotype, except some collected 10 January 1964.

Geographic distribution. Cnemidophorus duellmani is known from the environs of the Río Tuira and its tributaries in Darién Province, on the Pacific versant of eastern Panama ( Fig. 7 ; also see Remarks).

Diagnosis. Cnemidophorus duellmani can be distinguished from all other species in the C. lemniscatus species group (see Introduction, Harvey et al. 2012) by being the only species to have ten transverse ventral plates ( Fig. 8 ) at the level of the fifteenth, sixteenth, and/or seventeenth ventral row (see Harvey et al. 2012: 105 who gave eight ventrals for species in the C. lemniscatus group). In addition, it differs further from C. ruatanus , the only other Central American species of the group, in having broad contact between the precloacal pair of enlarged plates, without a small scale extending anteriorly between those two plates ( Fig. 9 ), in having 8–15 (both sides combined) small scales bordering the posterior edge of supraocular four, in having a mean of 6.2 scales (both sides combined) between supraocular four and the outer parietal, and 8–15 (11.8 ± 2.0) circumorbitals; (versus paired precloacal plates frequently entirely separated by a small scale [ Fig. 5 ], or separated at least one-third of its distance by a small scale, 4–9 small scales bordering supraocular four, a mean of 4.0 small scales between supraocular four and outer parietal, and 2–9, x = 4.2 ± 1.4 circumorbitals in C. ruatanus ).

Description of holotype. An adult male with a snout-vent-length (SVL) of 92.2 mm; snout-ear length (head length) 24.2 mm; snout length (snout to anterior border of eye) 11.1 mm; head width 15.0 mm; head depth 15.7 mm; shank length 19.3 mm; foot length 32.7 mm; hand length 13.7 mm; tail length 206.0 mm, tail length/SVL 2.23.

Dorsal head scales enlarged, smooth, platelike, with paired prenasal scales with short median sutures, a single frontonasal, paired prefrontals, a single frontal, paired frontoparietals, two pairs of parietal scales, and a single interparietal scale; rostral not contacting frontonasal; postnasal not contacting prefrontal; prefrontal and first superciliary in narrow contact on one side, separated by posterior extension of loreal on other side; frontal ridge absent; posterior frontal suture aligned near midlength of supraocular 3; scales in frontoparietal region smooth, flat; interparietal narrower than flanking parietals; 2–2 scales between fourth supraocular and inner-most parietal; 3–3 scales between fourth supraocular and outer-most parietal; 4–4 supraoculars plus one smaller supraocular lateral to region of seam between supraoculars 2–3 on each side; median pair of occipital scales distinctly enlarged, much larger than first dorsal scale row; 26 occipitals (occipitals of Harvey et al. 2012 plus scales bordering frontoparietals); supratemporals moderately enlarged, separated from parietals; short rostral groove present; nostril centered in nasal suture; nostril opening subtriangular; first supraocular contacting second supraocular; 1.0 to 2.0 rows of lateral supraocular granules, 21–23 granules on each side; anterior extent of circumorbital semicircles in single row reaches posterior third of supraocular 3, that of double row reaching anterior to seam between supraoculars 3–4; 7–8 circumorbitals; 5–5 superciliaries, second elongated; 4–4 suboculars, first on each side entire, upper edge contacting first superciliary, lower edge contacting supralabial 3; subocular keel present; patch of enlarged scales located in front of auditory meatus; no auricular or preauricular flaps or folds; 1–1 loreals; 6–6 supralabials; 6–5 infralabials; first supralabial straight ventrally, longer than second supralabial on one side, both supralabials subequal on other side; lingual sheath absent; moveable eyelids present; pupil rounded; first pair of chinshields contacting infralabials, those chinshields separated only at posterior edge; intergular sulcus absent; 19 anterior gulars; 11 posterior gulars; gular patch of distinctly enlarged scales absent; intertympanic sulcus absent; sharp transition from anterior gulars to smaller posterior ones; mesoptychial scales moderately enlarged, bordered anteriorly by sharp transition to small scales; edge of gular fold not serrated; dorsal scales conical, 232 middorsals between occipitals and first enlarged, keeled caudal scale; 122 granules around midbody; middorsal scales subequal to lateral scales; chest scales large, flat; pectoral sulcus absent; ventral body scales large, platelike, squarish, juxtaposed, smooth, in 32 longitudinal rows, in 10 transverse rows at midbody; scales immediately lateral to outside ventral plate on each side small and granular; paired enlarged terminal scales forming precloacal plate; paired scales forming precloacal plate in broad contact medially, without small scale making indentation between those two scales; one larger, sub-diamond-shaped, scale anteriorly bordering precloacal terminal pair; 11 scales bordering all three enlarged plates; 3–3 smaller rounded scales between precloacal plate and precloacal spur; 6 precloacal scales; postcloacal buttons absent; pair of slightly enlarged postcloacal scales present; caudal annuli complete, tail lacking crests or dorsolateral row of serrated scales; third row of proximal subcaudal scales keeled; preaxial and postaxial brachial scales separated by band of continuous moderately enlarged scales; largest scales on preaxial and postaxial brachial surfaces 1.5 to over 2 times as wide as long, both sets of enlarged scales smooth, extending well beyond centers of arm; postaxial brachial scales in continuous enlarged row with preaxial brachial scales; postaxial antebrachial scales slightly enlarged; 29 combined subdigital lamellae on fourth finger; subarticular lamellae of Fingers III–IV homogeneous in size, entire; 61 combined subdigital lamellae on fourth toe, distal ones smooth; subarticular lamellae of Toes III–IV divided, each scale smaller than other lamellae; no row of distinctly enlarged scales between Toes IV–V; small scales separating supradigital scales from subdigital lamellae continuous, or nearly continuous; denticulate fringe absent along postaxial edge of outer toes; fifth toe not reduced, claw extending beyond level of articulation of Toes III–IV; 6 prefemoral scales on left side; heels without expanded scales; tibiotarsal shields present; tibiotarsal spurs absent; 41 total femoral-abdominal pores; no gap between femoral and abdominal pores; 4 scales separating each femoral-abdominal pore series; HL/SVL 0.26; HW/SVL 0.16; HD/SVL 0.17; HW/HD 0.96; SL/SVL 0.12; SHL/SVL 0.21; foot length/SVL 0.36; hand length/SVL 0.15.

Color in alcohol: middorsal longitudinal band, single, complete, pale bluish brown, extending from posterior edge of head onto base of tail; middorsal band on each side with a narrow dark brown band located inside middorsal band; dorsolateral ground color pale brown, becoming pale blue on flanks; three rows of about 8–10 white spots per row between axilla and groin; dorsal surface of head brown without distinct markings; anterior supralabials pale blue, those below eye pale blue with brown tinge; infralabials pale blue; dorsal surfaces of forelimbs brown, those of hind limbs bluish black with pale brown spots; middorsal pale band and dark brown band extending onto tail for about one-third of its length, lateral surface of tail in that area blue with scattered white spots, tail becoming brown at about midlength and then pale brown for distal third; all ventral surfaces pale blue, except palms and soles pale brown and subcaudal surface changing to bluish brown at about midlength and then brown on distal third; one row of widely scattered blue to white spots present on some outer ventral plates.

Variation. The 12 adult specimens of the type series have the following measurements and proportions and scale counts (bilateral counts combined, unless otherwise stated): SVL 61.3–92.2 (81.4 ± 11.4) mm in males, 56.2– 74.9 (70.2 ± 7.3) in females; TAL/SVL 2.23–2.61 in five males, 2.33–2.53 in three females; HL/SVL 0.24–0.27 in males, 0.22–0.24 in females; SHL/SVL 0.21–0.23 in males, 0.18–0.23 in females; foot length/SVL 0.34–0.38 in males, 0.32–0.37 in females; hand length/SVL 0.13–0.15 in males, 0.12–0.14 in females; SL/SVL 0.10–0.12 in males, 0.10–0.11 in females; HW/SVL 0.13–0.16 in males, 0.12–0.15 in females; HD/SVL 0.13–0.17 in males, 0.11–0.13 in females; HW/HD 1.10–1.13 in males, 1.03–1.12 in females; femoral pores 41–44 (43.0 ± 1.3) in males, 36–44 (41.0 ± 3.0) in females; 104–132 (116.3 ± 10.4) granules around midbody; 27–32 (30.5 ± 1.4) longitudinal ventrals; 210–236 (222.2 ± 10.1) middorsal scales between interparietal and first enlarged and keeled scale at base of tail; 25–44 (31.5 ± 5.7) lateral supraoculars; 55–63 (59.7 ± 2.9) combined subdigital lamellae on fourth toe in 11; 27–37 (32.2 ± 2.7) combined subdigital lamellae on fourth finger; 25–29 (26.6 ± 1.4) occipitals (including scales bordering frontoparietals); 8–15 (11.5 ± 2.0) circumorbitals; 5–7 (6.2 ± 0.6) scales between supraocular 4 and outer parietal; 4–5 (4.1 ± 0.3) scales between supraocular 4 and inner parietal; 5–7 (5.8 ± 0.6) prefemoral scales on right side only; 4–5 (4.1 ± 0.3) precloacals; 17–22 (19.1 ± 1.8) anterior gulars; 8–14 (11.8 ± 1.7) posterior gulars; 9–11 (9.7 ± 0.7) scales around three enlarged precloacal plates.

Color in life: Duellman recorded in his field notes the following about a series of Cnemidophorus duellmani : “Adult male: throat, chin, and anterior surfaces of forearms blue; yellowish green dorsolateral stripe bordered above by dark brown stripe; middorsum buffy tan; flanks orange-tan; tail green laterally, blue below, brown above.”

Color in alcohol of the male paratypes is similar to that recorded for the holotype. However, Cnemidophorus duellmani is sexually dichromatic, thus female color in alcohol is described as follows: middorsal band pale brown with two slightly darker brown stripes placed laterally inside large band, middorsal band bordered laterally by black, broad stripe, middorsal band and bordering stripes extending from posterior end of head onto tail, both continuous; two white dorsolateral stripes on each side, those pale stripes separated by broader dark brown to black stripe; lateral field with dark brown to black stripe bordering lowest of paired dorsolateral white stripes, lateral field below that dark stripe pale brown, with or without 1–3 rows of pale spots; two most lateral ventral plates dark blue; dorsal surfaces of forelimbs bluish brown without distinct markings; dorsal surfaces of hind limbs bluish brown, usually with indistinct pale spots; tail bluish gray anteriorly, becoming some shade of brown on distal third; belly paler blue than outer ventral plates; remainder of color in alcohol similar to that of adult males. KU 80583 (SVL 33.3 mm), a juvenile, has eight longitudinal white stripes present; middorsal longitudinal band pale brown, complete, extending from posterior edge of head onto tail; throat and chin white; belly very pale blue; under surfaces of forelimbs pale brown, those of hind limbs dirty white; subcaudal surface pale brown on anterior third, becoming dark brown for posterior half. Another juvenile (KU 80582; SVL 33.1 mm) was similar to that described for KU 80583, whereas that of a subadult male (KU 80581; SVL 48.2 mm) is also similar to KU 80583, except that the ventral surfaces are pale brown.

Habitat. Duellman recorded in his field notes that the type series of Cnemidophorus duellmani was “found in open areas.” Breder (1946: 428) recorded the species was “Common along the lower reaches [of the Río Chucunaque and Río Chico valleys] in open fields.” Sexton et al. (1964: 293) studied this species at a site along the Río Chucunaque and stated that the “ Cnemidophorus is commonly associated with areas having a low percentage of vegetation coverage as well as plants low in height.”

Breder (1946: 382) recorded the region encompassing the Panamanian Río Tuira and Río Chucunaque region and tributaries where Cnemidophorus duellmani occurs as in the “Lower Arid Zone,” whereas Jaramillo et al. (2010: 621) considered C. duellmani as occurring in the “Lowland Wet/Moist Forest.” Curiously, none of the authors who presented ecological data on this species mentioned the presence or absence of “foot waving,” which is usually present in members of the C. lemniscatus group.

Conservation notes. Jaramillo et al. (2010: 621) characterized Cnemidophorus duellmani (as C. lemniscatus ) as a species of medium vulnerability because of the former perceived wide geographic distribution of C. “ lemniscatus .” With our study, C. duellmani is now a Panamanian endemic species with vulnerability score of 13, thus placing the species in the high vulnerability category. However, in this case, that score is misleading since C. duellmani is a species that benefits from human disturbance of habitat (see Sexton et al. 1964, Heatwole 1966). Thus, C. duellmani appears to not be threatened in the foreseeable future, and in fact will likely benefit from continued clearing of lands along the river valleys in the region where it occurs.

Remarks. Burt (1931) recorded Cnemidophorus l. lemniscatus from two localities in the Panamanian Canal Zone (MCZ specimens from Ancón and Obispo Station), otherwise all of Burt’s localities for this species are from the vicinities of the Río Tuira and tributaries in Darién Province. The occurrence of Cnemidophorus in the Canal Zone needs to be investigated, but Ibáñez et al. (1995) did not list it as occurring in the Canal Zone. It is also possible that Burt (1931) misidentified specimens of Holcosus undulatus (Wiegmann) in the MCZ from Ancón, Panama as Cnemidophorus since there is a series of H. undulatus in the MCZ from Ancón (Echternacht 1971).

Etymology. We take pleasure in naming this new species after William E. Duellman of the University of Kansas. Bill was not only one of the collectors of the type series, but has also produced two significant studies of Mexican and Guatemalan species groups of lizards, which at those times were considered to be members of the genus Cnemidophorus (now Aspidoscelis ).

Figures

FIGURE 6. Dorsal surfaces of the holotype of Cnemidophorus duellmani sp. nov. (KU 80542).

FIGURE 7. Map of eastern Panama and northern Colombia showing the localities for Cnemidophorus duellmani sp. nov. (circle), Cnemidophorus gaigei comb. nov. (square = specimens examined, open square = type locality), and the Islas de Providencia and San Andrés (upper left hand corner) where Cnemidophorus espeuti occurs.

FIGURE 9. Cloacal plate region of the holotype of Cnemidophorus duellmani sp. nov. (KU 80542) showing the broad medial contact between the enlarged terminal precloacal plates.

FIGURE 8 A. Lateral surface of the holotype of Cnemidophorus duellmani sp. nov. (KU 80542) showing the extra ventral plate. 8 B. Lateral surface of Cnemidophorus ruatanus comb. nov. (KU 101329) showing the absence of the extra ventral plate.

FIGURE 5 A. Cloacal plate region of Cnemidophorus ruatanus comb. nov. (KU 101329) showing the small scale separating the enlarged terminal precloacal plates; 5 B Cloacal plate region of Cnemidophorus gaigei comb. nov. (USNM 117471) showing the enlarged terminal precloacal plates in broad contact medially.