Dactyloa maia, Batista, Abel, Vesely, Milan, Mebert, Konrad, Lotzkat, Sebastian & Köhler, Gunther, 2015

Batista, Abel, Vesely, Milan, Mebert, Konrad, Lotzkat, Sebastian & Köhler, Gunther, 2015, A new species of Dactyloa from eastern Panama, with comments on other Dactyloa species present in the region, Zootaxa 4039 (1), pp. 57-84 : 65-70

Dactyloa maia sp. nov.

Figures 1 , 4 E–F, 5E–F, 6D, 8, 9, 10.

Holotype. SMF 97268, adult male ( Figs. 4 E, 5E, 6D, 9), from the ridge of the Serranía de Darién ( Fig. 1 ) along the trail that connects the Comarca Wargandí and the Comarca Guna Yala, about 10 km northeast of the village Nurra, 9.06142° N, 77.97961° W, 344 m asl., Corregimiento de Nurra, Comarca Wargandí, Panama; collected by Abel Batista and Milan Vesely on 0 3 October 2012; original field number AB 760.

Paratypes. All from Panama: SMF 97269, a male, from Cerro la Javillosa Ambroya, Torti, Chepo, Panama province, collected on 28 September 2012, 19:39 hrs, 8.92267° N, 78.62530° W, 851 m asl, collected by Abel Batista and Milan Vesely; MHCH 2782, a female, same collecting data as holotype; MHCH 2781 and MHCH 2783, females, respectively, from Cerro Pechito Parado, Bajo pequeño, Lajas blancas, Cémaco, Comarca Emberá- Wounáan, on 0 7 November 2012, 8.47911° N, 77.52799° W, 718 m asl, collected by Abel Batista; SMF 97270, a female from la Cascada trail, Burbayar private reservation, Cartí, Narganá, Comarca Guna Yala, on 26 November 2012, 9.31577° N, 79.00580° N, 322 m asl, collected by Abel Batista and Konrad Mebert.

Diagnosis. A moderate-sized species (SVL 62–78 mm) of the genus Dactyloa , D. latifrons species group (sensu Nicholson et al. 2012), that is most similar in external morphology to D. pupurescens , D. limon , and D. ibanezi , and according to molecular evidence is most closely related to D. pupurescens and D. ibanezi ( Fig. 7 ). These four species share a moderate adult size (SVL 62–88 mm); a large dewlap; a peculiar flank pattern in males, which is green with dark oblique bands, or blotches, or ocelli always arranged in oblique rows ( Fig. 5 ); enlarged postcloacal scales in males; and smooth ventrals. Dactyloa maia can readily be distinguished from these three species by its color pattern and morphology ( Figs. 3–6 ; Table 1–2), and from the remaining species of Dactyloa within the D. latifrons group by its moderate size (SVL <100 mm); and the orange male dewlap with an uninterrupted white margin. Dactyloa maia can be differentiated from D. limon by its male dewlap coloration which is orange with an uninterrupted white margin (vs. yellowish near the throat and tan on distal portion, or uniformly light tan in D. limon ; Fig. 4 ). Dactyloa maia further differs from D. purpurescens , D. limon and D. ibanezi in the color pattern of the flanks as follows: Dactyloa maia has oblique rows of turquoise ocelli or oblique dark green bands without sexual dimorphism ( Fig. 8 ); in D. limon , males have wide dark green bands on the flanks whereas females have diffuse dark green spots distributed evenly or randomly; males of D. purpurescens exhibit oblique rows of ocelli or blotches whereas females have dark green spots arranged in oblique rows; in D. ibanezi both sexes exhibit oblique thin black lines. The hemipenis of D. maia is a small, unilobate organ (slightly bilobate in D. ibanezi ; no information available for D. limon and D. purpurescens ). Also, Dactyloa maia differs from D. purpurescens , D. limon , and D. ibanezi in mean values of several morphological characters as follows (mean values for D. purpurescens , D. limon , and D. ibanezi presented in that order): HL/SVL 0.27 in Dactyloa maia vs. 0.25, 0.23 and 0.26,.LST 64.7 in Dactyloa maia vs. 62.8, 43.7, and 44.5; number of scales between first and second canthals 14.5/12.3 (first canthals/second canthals) in Dactyloa maia vs 12.9/10.2, 10/8.6, and 11.5/10.

Description of the holotype. Adult male as indicated by everted hemipenes, a pair of enlarged postcloacal scales, and presence of a large dewlap ( Figs. 3 E, 4E, 7D, 9); SVL 76 mm; tail length 178 mm (tail complete), tail length/SVL ratio 2.34; tail laterally compressed in cross section, tail height 2.2 mm, tail width 2.0 mm; axilla to groin distance 35.6 mm; head length 19.2 mm, HL/SVL ratio 0.25; snout length 9.3 mm; head width 9.7 mm; longest toe of adpressed hind limb reaching posterior margin of orbit; shank length 20.7 mm, shank length/SVL ratio 0.27, shank length/HL ratio 1.08; tip of longest finger of extended forelimb reaching tip of snout; longest finger of adpressed forelimb not reaching to anterior insertion of hind limbs; prefrontal ridges distinct, parietal ridges conspicuous; scales on snout mostly keeled; 6 postrostrals; 7 scales between nasals; scales in distinct prefrontal depression smooth; supraorbital semicircles differentiated, composed of smooth scales, separated by a minimum of 2 scales; supraorbital disc composed of 7 enlarged smooth scales; two elongated, smooth anterior superciliaries, followed posteriorly by a much smaller, elongate scale; about 5 rows of small keeled scales extending between enlarged supraorbitals and superciliaries; interparietal plate distinct, parietal eye visible; canthal ridge distinct, composed of 3 large (posterior) and 7 small (anterior) canthal scales; 11 scales present between second canthals; 13 scales present between posterior canthals; 78 loreal scales arranged in 7 horizontal rows; subocular scales flat, subocular row well-defined; 9 supralabials to level below center of eye; ear opening 0.73 x 1.6 mm (length x height); mental distinctly wider than long, almost completely divided medially, bordered posteriorly by 6 postmentals; 10 infralabials to level below center of eye; third and fourth sublabials posterior to mental slightly enlarged; keeled granular to elongate scales present on chin and throat; dewlap large, extending well onto body, anterior insertion is about halfway between nose and orbit, posterior insertion at a level between one-third and one half of the distance between axilla and groin, with about 4 gorgetal-sternal rows, each 2–3 scales wide, becoming less regular posteriorly; low nuchal crest present, dorsal crest barely visible; dorsum of body with keeled scales, 1–2 middorsal rows of prominently keeled, but not otherwise enlarged scales; about 50 medial dorsal scales in one HL; about 125 medial dorsal scales between levels of axilla and groin; lateral scales small, long and keeled; ventrals at midbody smooth, subimbricate; about 66 ventral scales in one HL; about 87 ventral scales between axilla and groin; about 164 scales around midbody; caudal scales strongly keeled, without whorls of enlarged scales, subcaudal scales with a single prominent keel; a pair of greatly enlarged postcloacal scales, larger one about 0.88 x 1.90 mm (length x width); tube-like axillary pocket not developed; scales on anterodorsal surface of thigh and on dorsal surface of forearm keeled; digital pads dilated, dilated pad about 3 times width of nondilated scales under distal phalanx; distal phalanx narrower than and raised from dilated pad; 31/31 (left/right) lamellae under phalanges ii to iv of 4th toe; 11/12 scales under distal phalanx of 4th toe; 21/19 lamellae under phalanges ii to iv of 4th finger; 10/9 scales under distal phalanx of 4th finger.

Hemipenis description: The completely everted hemipenis ( Fig. 10 ) of SMF 97269 is a small, unilobate organ; sulcus spermaticus bordered by well-developed sulcal lips, opening at base of apex into a small concave area; large asulcate processus and ridge present; a prominent fleshy fringe present on each lateral side of truncus; most of apex on asulcate side and distal portion finely calyculate, truncus with transverse folds..

Coloration in life of the holotype. ( Fig. 4 E) Dorsal and lateral ground color of body and limbs Light Grass Green (109); lateral surface of body with three Parrot Green (121) oblique bands directed backwards as well as downwards, each with six to seven Medium Blue (168) small oval spots; tail with Dark Green (136) transverse bands; eyelids Cream Yellow (82); iris Chrome Orange (74); ventral surfaces dirty white, suffused with Pale Cyan (157); dewlap Light Pratt’s Rufous (71) with three well defined longitudinal series of Emerald Green (143) scales, free margin of dewlap dirty white, anterior insertion of dewlap suffused with Beige (254).

Coloration of the holotype after approximately two years of preservation in 70% ethanol. ( Fig. 9 ) Dorsal and lateral ground color of body and limbs Lavender (195); lateral surfaces of body with three oblique rows of small oval Medium Water Blue (182) spots directed backwards as well as downwards; tail with Plumbeous (295) transverse bands; ventral surfaces dirty white, suffused with Medium Blue Gray (194); dewlap Pale Sulfur Yellow (92), its base suffused with Pale Neutral Gray (296).

Color variation. Another male (SMF 97269, 8E–F), at the moment of encounter, was uniform green with bluish spots on flanks arranged in oblique rows; after collection it exhibited metachrosis, and the coloration recorded was as follows: the dorsal ground color of body, head and limbs Salmon Color (58), grading into Yellow Green (103) toward the flanks; tail with Russet (44) transverse bands; four well-defined Olive Green (123) oblique bands between axilla and groin, each including four to seven irregular Cyan Black (153) blotches; ventral surfaces and dewlap as in holotype. A female (MHCH 2782, Figs. 4 F, 5F, 8A–B) lacked the small oval spots, and only the three Parrot Green (121) transverse bands were present with three small Sulphur Yellow (80) spots between each band; dewlap Medium Chrome Orange (75) with three well defined longitudinal series of Emerald Green (143) scales, free margin of dewlap dirty white, anterior insertion of dewlap suffused with Light Neutral Gray (297). Another female (SMF 97270; Fig. 8 G) agrees in general color pattern with female MHCH 2782. A third female (MHCH 2781; Fig. 8 C–D) has the same dorsal ground color as the holotype, but with six well defined Greenish Olive (125) oblique bands in the flanks (without spots); dewlap Olive (126), with three well defined longitudinal series of Lime Green (116) scales, anterior and external border of dewlap dirty white, anterior insertion of dewlap suffused with Light Neutral Gray (297).

Distribution and Natural history. As far as we know, Dactyloa maia is endemic to eastern Panama, inhabiting the foothills and ridges of the Majé, San Blas, Darién (as far south and east as to Río Tuquesa, thereafter apparently replaced by D. purpurescens ), and Piedras-Pacora (two specimens photographed by Angel Sosa- Bartuano, not plotted in Fig. 1 ) mountain ranges, where it occurs in the eastern Panamanian montane forest (Fund 2014) and the Isthmian-Atlantic moist forests (in the Piedras-Pacora and San Blas mountain range, Hogan & Fund 2014), at 322–852 m asl. All specimens of Dactyloa maia were found during night searches sleeping on branches or leaves 2 to 3 m above the ground.

Etymology. Abel Batista dedicates this beautiful new species to his recently born daughter, Maia . The name also comes from Greek mythology, where it is applied to the eldest of the Pleiades, sometimes called mountain nymphs, and are believed to live on the trees in mountains and groves as the guardians for that habitat, a role Dactyloa maia could also represent.

Figures

FIGURE 1. Distribution of Dactyloa species formerly referred to as D. chocorum in eastern Panama, based on specimens collected for this study, provided by A. Sosa, literature and online databases (http: // www. vertnet. org /).

FIGURE 3. Some species of Dactyloa found in Eastern Panama. A – B) D. purpurescens (female and male, MHCH 2635 – 36); C – D) D. kunayalae (male, SMF 97266); E) D. frenata (male, MHCH 2785); F) D. latifrons (young male, SMF 96575); G – H) D. chloris (female, SMF 97096).

FIGURE 4. Dewlaps of the Dactyloa spp. related to D. purpurescens (left column males, right column females). A – B) D. limon (MHUA 11760; holotype; photo by Juan Pablo Hurtado), female not collected (from; photo by Alejandro Montoya); C – D) D. purpurescens, (MHCH 2636, MHCH 2635); E – F) D. maia sp. nov. (SMF 97268, MHCH 2782); G – H) D. ibanezi (MHCH 2019, SMF 91475).

FIGURE 5. Flank pattern in species of Dactyloa related to D. purpurescens. Arrows indicate capitad direction for all males (left column) and females (right column). A) D. limon holotype MHUA 11760; B) D. limon MHUA 11248; C) D. purpurescens (MHCH 2636); D) D. purpurescens (SMF 91475); E) D. maia (SMF 97268); F) D. maia (MHCH 2782); G) D. ibanezi, MHCH 2184; H) D. ibanezi, SMF 91475. Scale bars equal 10 mm.

FIGURE 6. Lateral head views of the species of Dactyloa related to D. purpurescens. A) D. limon holotype MHUA 11760; B) D. ibanezi MHCH 2184; C) D. purpurescens SMF 97271; D) D. maia sp. nov. holotype SMF 97268. Scale bars equal 10 mm

FIGURE 7. Bayesian consensus tree of the genus Dactyloa based on 16 S and COI mtDNA. Outgroups: Norops capito, Norops limifrons, Norops tropidogaster, Norops poecilopus and Polychrus marmoratus. Asterisks on nodes indicate estimated posterior probabilities P ≥ 0.90. Scale bar refers to substitutions per site.

FIGURE 8. Color variation of Dactyloa maia sp. nov. in life. A – B) female MHCH 2782; C – D) female MHCH 2781; E – F) male SMF 97269; G) female SMF 97270.

FIGURE 9. Dactyloa maia sp. nov. holotype (SMF 97268), preserved specimen. A) head, laterally; B) head, dorsally; C) head, ventrally; D) venter; E) dorsum; F) flank; G) left hand; H) left foot.

FIGURE 10. Hemipenis of the Paratype (SMF 97269) of Dactyloa maia sp. nov. A) Sulcate view; B) asulcate view; C) lateral view.