Gymnodactylus vanzolinii, Cassimiro, José & Rodrigues, Miguel T., 2009

Cassimiro, José & Rodrigues, Miguel T., 2009, A new species of lizard genus Gymnodactylus Spix, 1825 (Squamata: Gekkota: Phyllodactylidae) from Serra do Sincorá, northeastern Brazil, and the status of G. carvalhoi Vanzolini, 2005, Zootaxa 2008, pp. 38-52 : 40-49

Gymnodactylus vanzolinii sp. nov.

( Figs. 1–2 )

Holotype: MZUSP 68286 (adult female), field number 80.6069 (SVL 52.3 mm, TL 70 mm), from Serra do Sincorá, Chapada Diamantina, Mucugê municipality (13°09’S, 41°24’W), ca. 1,000 m elevation, State of Bahia, Brazil, collected 27–28 September 1987 by M. T. Rodrigues.

No. Character description

1 Sex: 3 = male, Ƥ = female, J = juvenile

2 Number of post-nasals

3 Internasals in contact (+) or no (–)

4 Number of granules between internasals

5 Number of supralabials

6 Number of infralabials

7 Number of granules in contact with supralabials, from first to fifth

8 Number of scales in contact with rostral

9 Number of interorbitals

10 Shape of granules in loreal region: flattened (F), conical (C), rounded (R)

11 Number of scales between postmentals and cloaca

12 Number of granules in contact with mental

13 Number of granules in contact of postmentals

14 Number of ventral granules between ears

15 Shape of gular scales: flattened (F), conical (C), rounded (R)

16 Number of color bands on head, nape, body, and tail

17 Number of scales between eye and ear (left side only)

18 Number of scales surrounding a dorsal tubercle (randomly counted for 5 tubercles)

19 Number of granules between the two middorsal paravertebrals tubercle rows counted at midbody

20 Number of granules between two transverse and adjacent rows of tubercles counted in paravertebral row at midbody

21 Number of granules between paravertebral and the immediately adjacent longitudinal row of tubercles, counted at midbody

22 Number of longitudinal rows of tubercles

23 Number of paramedian tubercles

24 Number of transverse rows of ventral scales at midbody

25 Number of subdigital lamellae on fourth finger

26 Number of subdigital lamellae on fourth toe

27 Number of subdigital lamellae on fifth toe

28 Number of subcaudals

29 Snout-vent length (SVL)

30 Distance between posterior level of arm and anterior margin of tight

31 Trunk length (TrunkL)

32 Tail length (TailL)

33 Head length (HeadL)

34 Head width (HeadW)

35 Head height (HeadH)

36 Orbital diameter (OrbL)

37 Ear length (EarL)

38 Internarial distance (InterNar)

39 Snout to eye distance (SnEye)

40 Nares to eye distance (EyeEar)

41 Eye to ear distance (EyeEar)

42 Interorbital distance (IntraOrb)

43 Forearm length (ArmL)

44 Tibia Length (CrusL)

continued.

Paratypes (N = 16): MZUSP 56290 (adult female) and MZUSP 56291 (adult female), 13°09’S, 41°24’W, 1000 m, collected on 6 December 1980 by M. T. Rodrigues, field numbers 80.1885 and 80.1886 respectively. MZUSP 68285 (adult female), collected 27–28 September 1987, by M.T. Rodrigues, field number 80.6020. MZUSP 73926, MZUSP 73927 (juveniles) and MZUSP 73928 (adult male) collected on 1 October 1990, by M.T. Rodrigues, field numbers 90.6026, 90.6027 and 90.6028 respectively. MZUSP 98245 (adult female), 13°01’06”S, 41°21’57”W, 1017 m, collected on 0 6 March 2005 by J. Cassimiro, F.S.F. Leite & L.E. Lopes, field number JC 1207. MZUSP 98246 (adult female), collected on 10 March 2005 by J. Cassimiro, field number JC 1209. MZUSP 98247 (adult female), 13°00’03”S, 41°21’57”W, 997 m, 12 March 2005 by J. Cassimiro, field number JC 1223. MZUSP 98248 (adult female), 13°00’02”S, 41°21’59W”, 1004 m, 13 March 2005 by J. Cassimiro, field number JC 1230. MZUSP 98249 (adult female), 13°00’03”S 41°21’58W”, 1010 m, by F.S.F. Leite & J. Cassimiro, field number JC 1235. MZUSP 98250 (adult female), 98251 (adult male), collected on 18 March 2005 by J. Cassimiro & F.S.F. Leite, field numbers JC 1238–9 respectively. MZUSP 98252 (adult female), 98253–4 (adult males), collected on 27 March 2005 by J. Cassimiro, field numbers JC 1280–2 respectively. All specimens collected in the same municipality as the holotype, in Mucugê, Chapada Diamantina , Bahia state, Brazil.

Etymology: The specific name is a homage to Dr. Paulo Emílio Vanzolini one of the most distinguished Brazilian zoologists. About Vanzolini see Heyer (2004).

Diagnosis: A small Gymnodactylus with a slender body, head distinct from neck, and tail longer than body. G. vanzolinii sp. nov. can be distinguished from all other species of the genus by the following combination of characters: 10–12 (n = 14) longitudinal rows of dorsal tubercles; 22–29 (n = 13) tubercles in a paramedian row; 18–22 (n = 11) transverse ventrals at midbody; 16–18 (n = 17) subdigital lamellae under fourth finger; 18–21 (n = 15) subdigital lamellae under fourth toe; color pattern with transverse dark bands along the dorsum.

Description of holotype: Adult female, SVL 52.3 mm, TL 70.6 mm, HL 12.9 mm, HW 9.3 mm, HH 6.8 mm. Head large, distinct from neck; covered with small granules and irregularly disposed, enlarged and scattered tubercles among them in parietal and occipital regions. Rostral scale of moderate size, 1.7 times wider than high, visible from above, indented and slightly concave dorso-medially. Supranasals wider than long, enlarged separated on midline by a granular scale and posteriorly bordered by three small granules, the exterior one larger and contacting the nostril. Snout sub-elliptical in dorsal view, rounded laterally. Top of snout, from rostral to level of anterior margin of orbit, covered with enlarged granules of different sizes, rounded, juxtaposed, slightly keeled or smooth, decreasing in size towards labial border and posterior part of head. Central and posterior part of head with slightly rounded, conic, juxtaposed granules much smaller than those from snout. A cluster of enlarged granules on top of head posterior to eyes. Superciliary region with a series of enlarged imbricate and smooth scales; anterior ones much larger, becoming progressively smaller and granular posteriorly. Loreal region with enlarged and juxtaposed granules similar to those on top of snout, those closer to labials longer than wide and flat. Temporal granules small, conical, juxtaposed, some keeled, similar to those on top of head. Enlarged, conical, and juxtaposed granules on the superior an anterior margin of ear opening, those posterior to ear opening smaller. External ear opening oblique, about half the size of the eye. Tympanum highly recessed. Nostril small and protruding, bordered by rostral, internasal, two enlarged postnasals, and two small granules separating nostril from first supralabial. Fourteen to 15 loreal granules in an imaginary row between postnasals and anterior margin of the orbit. Eye large, pupil vertical. About 22 interorbital granular scales. Seven to eight enlarged supralabials, fifth to seventh below center of eye, followed by small granules; two irregularly arranged rows of slightly enlarged flat scales between supralabials and eye. Mental sub-triangular, as long as broad, followed laterally by a pair of enlarged but much smaller sub-triangular postmentals. Postmentals in contact with first supralabials, broadly separated in the midline; in posterior contact with gular granules, the external ones larger. Six infralabials decreasing progressively in size posteriorly, first largest. A series of enlarged sublabials between gular scales and infralabials, longer than wider. Gular scales rounded, cycloid, increasing gradually in size posteriorly; those closer to sublabials slightly enlarged.

Neck granules conical, imbricate, juxtaposed, smaller dorsally, slightly enlarged laterally.

Dorsal and lateral parts of body covered with granules as small as or smaller than those on top of head. On each side five relatively regular longitudinal rows of enlarged and widely separated blunt conical tubercles. Twenty three paravertebral tubercles. Ventral scales large, much larger than the enlarged isolated dorsal tubercles, rounded, flat, subimbricate, cycloid, forming irregular longitudinal rows, 33 on the midline between the level of posterior edge of arm and anterior edge of thigh. Cloacal opening a straight transverse cleft, surrounded by granular scales.

Fore-limbs with enlarged, keeled, and imbricate scales dorsally; ventrally much smaller, almost granular and flat scales. Antero-dorsal and ventral parts of thigh with large, imbricate, cycloid scales those contacting scales of posterior part of thigh slightly keeled. Postero-dorsal part of thigh with much smaller, juxtaposed, conical granules, decreasing ventrally in size. Dorsal parts of tibia with enlarged juxtaposed slightly keeled granules. Ventral parts of tibia with cycloid, smooth, imbricate, flat scales, smaller than those on ventral part of thigh.

Palmar and plantar surfaces with juxtaposed rounded granules; an enlarged external and conspicuous palmar tubercle. Digits long, angulated distally, slightly compressed; claws relatively long, curve, protruding much beyond claw sheath; subdigital lamellae quadrangular, gradually decreasing in size distally, some, in the distal half, eventually substituted by a pair of granules. Fifteen infradigital lamellae under fourth finger; 19 under fourth toe.

Tail cylindrical, its dorsal scalation near the base identical to that present on dorsal parts of body. Distal part of the dorsal surface of the tail with enlarged, imbricate, slightly keeled or smooth scales that are as wide as long, gradually increasing in size and becoming elongated towards the tip of the tail. In the first third of dorsal part of tail, a pair of distinctive enlarged scales characterizes a series of regular whorls that become inconspicuous distally. Ventral part of base of tail with smooth, imbricate scales, similar to those of ventral part of body. Posterior part of ventral surface of tail with smooth, strongly imbricate scales; those from midventral area distinctly enlarged, at least twice as wide as long, laterally contacting much smaller scales.

Color in preservative (ca. 70% alcohol): Background color of dorsal and lateral parts of body light olivebrown. Head with a series of irregular dark brown spots, extending from the level of eyes to neck. A conspicuous dark brown stripe extending from nostril to eye is present both in loreal and canthal regions; these stripes extend behind the eyes on temporal region but become irregular on nuchal region. A series of nine irregular transverse dark brown bands, fading toward venter, extend from the level of arms to the level of legs. Ventral surface of body cream, immaculate. Three to four irregularly arranged melanophores on each gular scale. Dorsal parts of tail with dark brown irregular bands similar to those of dorsum, ventral parts immaculate anteriorly, becoming progressively dark-brown towards the extremity.

Color in life: Similar to colors observed in preserved condition, although more brightly colored.

Variation: Maximum snout-vent length in males 50.8 mm (N = 4); maximum snout-vent length in females 52.3 mm (N = 11); tail length 59.4–70.6 (N = 4); head length 8.2–12.9 (N = 17); head width 5.6–9.4 (N = 16); head height 3.6–6.8 (N = 16); number of longitudinal rows of tubercles 10–12 (N = 13); paravertebral tubercles 22–29 (N = 13); supralabial scales 5–8 (N = 16); internasals 0–1 (N = 16); ventrals 18–22 (N = 11); subdigital lamellae under finger IV 16–18 (N = 17); subdigital lamellae under toe IV 18–21 (N = 15).

The status of G. carvalhoi Vanzolini, 2005 and G. amarali Barbour, 1925 : Gymnodactylus amarali was described by Barbour in 1925 on the basis of a specimen from Engenheiro Dodt, state of Piauí ( Barbour 1925). In the course of a recent revisionary study Vanzolini (2005) described G. c a r v a l h o i, including under this name all specimens of Gymnodactylus from the Central Brazilian Cerrado currently attributed to Gymnodactylus geckoides amarali . Although Vanzolini did not examine the holotype of Barbour (1925), he redefined and redescribed G. amarali based on a single juvenile specimen he collected at Alto Parnaíba, state of Maranhão, in the surroundings of the type locality. Recognizing this specimen as the only other example of G. amarali he decided to allocate all other specimens from the Cerrado previously assigned to G. geckoides amarali , in his new species. We think that Vanzolini was in error and that G. carvalhoi is not a valid species.

Barbour was very clear when he said in the original description that the holotype of G. amarali , (MCZ 20682) had “about 15 longitudinal series” of “rows along the sides of the body” ( Barbour 1925). Later, the same specimen also was examined by Benjamin Shreve who found 14 rows of tubercles (In: Vanzolini 1953a). We do not understand why Vanzolini (2005) ignored this information, because he redefined G. amarali based on the presence of 10 longitudinal rows of dorsal scales in the juvenile specimen from Alto Parnaíba ( Vanzolini 2005). We re-examined this specimen (MZUSP 93075) and verified that its dorsal skin is partially damaged, probably leading Vanzolini to mistakenly count its scales. Actually the specimen has 14 longitudinal rows of dorsal scales instead of the 10 reported by Vanzolini (2005). The only species of Gymnodactylus with such a low number of longitudinal rows of dorsal tubercules are G. guttulatus and G. vanzolinii sp. nov., with values varying between 10 to 12 in both species (see Table 3 ). Vanzolini (2005) also admitted that the most important diagnostic characteristic of his new species was the color pattern, which was characterized by the presence of conspicuous and vivid ocelli with their centers situated on dorsal tubercles. He considered this ocellate pattern sufficient to separate G. amarali from all other samples he studied from the Cerrado. We disagree. Specimens from other areas in the Cerrado we examine exhibit a similar pattern of coloration. Live animals, particularly juveniles, show that “conspicuous and vivid” coloration described by him and Barbour. The “large tubercle that forms the center for each of the dorsal white spots” reported as diagnostic for G. amarali , is found in several specimens from the Cerrado examined and varies greatly in all samples. When Barbour described G. amarali he diagnosed his new species from G. geckoides found in the adjacent semi-arid Caatinga and emphasized color differences. In fact, color differences between G. geckoides and G. amarali are markedly contrasting: G. geckoides presenting commonly a less conspicuous color pattern, usually without ocelli, almost inconspicuous when compared with the vivid pattern of the Cerrado animals, especially in life or recently collected.

Vanzolini (2005) also estimated the number of tubercles in the paramedian row of his juvenile of “ amarali ” to be 66 tubercles. Using a similar method we estimate the number of tubercules in the damaged part of skin and counted those present in the remaining intact part, and the total number does not pass 40. This value is within the range for the samples of Gymnodactylu s from the Cerrado biome ( Table 3 ) examined by us.

Other characteristics pointed out by Vanzolini (2005) as diagnostic between the two species refers to the shape of the canthus rostralis and loreal, size and organization of the tubercles, shape of ear opening, and presence or absence of preanal patch. All these characteristics mentioned by him are highly variable and most of them are subjective, like “the dorsal tubercles of amarali are weaker and much less well organized”. Variation in the referred characters is so extensive in specimens from the Cerrado and in all other species of Gymnodactylus , that they are useless for diagnostical purposes.

Based on the considerations above, we consider that G. carvalhoi is identical to G. amarali and should be considered a synonym of that last name.

Comparison to other species: G. v a n z o l i n i i sp. nov. can be immediately distinguished from all other congeners by the presence of irregular transversal dark bands along the dorsum, absent or incomplete and interrupted in all other species. The most similar species seems to be G. guttulatus that also has irregular transverse band along the dorsum but they are incomplete and interrupted. G. guttulatus presents white dots scattered on the dorsum that are absent or faded in G. vanzolinii . In color pattern G. vanzolinii differs from G. darwinii in the absence of a nuchal dark collar (present in G. darwinii ), from G. amarali by the absence of light dorsal body ocelli, present in almost all individuals of G. amarali examined, and from G. geckoides by the absence of dark marblings or, less commonly, by the presence of diffuse, irregular and incomplete ocelli, forming incomplete transverse bands on the dorsum in G. geckoides (variation in G. geckoides is also extensive). G. vanzolinii can be distinguished from G. amarali , G. darwinii , and G. geckoides , by having frequently 10 (rarely 11 or 12) rows of longitudinal dorsal tubercles (mean = 10.43 ± 0.65; n = 14), instead of 12 or more in the other three species (see Table 3 ). The number of rows of longitudinal dorsal tubercles in G. guttulatus ranges from 11–14 (mean 12.4 ± 1.14, n = 5). Overlap in the number of rows of longitudinal dorsal tubercles with G. vanzolinii is slight but in these cases color pattern separates the two species. G. vanzolinii can be additionally distinguished from G. amarali and G. geckoides by having a 4th finger with 16–18 (n = 17) subdigital lamellae [11–14 (n = 14) in G. amarali , and 13–15 (n = 30) in G. geckoides ]. G. vanzolinii can be distinguished from G. darwinii by having less tubercles in a paramedian row [22–29 (n = 13) against 47–59 (n = 15) in G. darwinii ]. Finally, G. vanzolinii is distinctly larger than G. geckoides in body size ( Table 3 ).

Distribution and ecology: G. vanzolinii sp. nov. is presently known only from, and thought to be endemic to, the mountains of Serra do Sincorá, in the Chapada Diamantina , an area situated in the northern portion of Espinhaço mountain range ( Fig. 3 ). Situated in Bahia and Minas Gerais states between latitudes 10°– 20°35’S and longitudes 40°10’– 44°30’W, this high altitude phytogeographic province, reaches elevations of more than 2,000 m in some places. The area is characterized by a mosaic of vegetation types, of which “campos rupestres” [rocky fields in Magalhães’s (1966) connotation], a dominant open-rock pioneer vegetation with rock-dwelling plants, are most common. Nevertheless, various types of environments occur in the area, like marshes, gallery forests, “cerrado” (savanna-like), montane forests and semi-deciduous to deciduous forests ( Giulietti & Pirani 1988).

All specimens of G. vanzolinii were collected on rock outcrops in the proximity of the Mucugê municipality, Serra do Sincorá, Chapada Diamantina . Some were collected under solitary rocks or in rock crevices. Seven individuals were found inactive under rocks during the day. Active specimens were only observed at night, from early evening to about midnight. As most active specimens observed were foraging in deep fissures on large rocks we think that their apparently scarcity might be due to this microhabitat preference. Several active specimens observed under these circumstances were not collected because there were deeply inserted in the rock crevices.

Phyllopezus pollicaris and Hemidactylus brasilianus were collected syntopically with G. v a n z o l i n i i on the rocks, although they were also obtained in other microhabitats in the area. Other lizards recorded at Mucugê area [unpublished data, and Freitas & Silva (2007)] were: Hemidactylus mabouia (Gekkonidae) , Acratosaura mentalis , Acratosaura sp., Heterodactylus sp., Micrablepharus maximiliani , Psilophthalmus sp. ( Gymnophthalmidae ), Enyalius erythroceneus (Leiosauridae) , Polychrus acutirostris (Polychrotidae) , Mabuya heathi , Mabuya sp. ( Scincidae ), Ameiva ameiva , Cnemidophorus sp., Tupinambis merianae (Teiidae) , Eurolophosaurus sp., Tropidurus hispidus , Tropidurus mucujensis , and Tropidurus semitaeniatus (Tropiduridae) .

Figures

FIGURE 1. Gymnodactylus vanzolinii sp. nov., adult male paratype (MZUSP 98247); Serra do Sincorá, municipality of Mucugê, State of Bahia, Brazil. Photo by Felipe S. F. Leite.

FIGURE 2. Holotype of Gymnodactylus vanzolinii sp. nov. (MZUSP 68286). Dorsal (A), lateral (B), and ventral (C) views of the head (scale bar = 5 mm).

FIGURE 3. Type-locality of Gymnodactylus vanzolinii sp. nov., Mucugê municipality, State of Bahia (BA), Brazil, and distribution of G. guttulatus along the Espinhaço mountain range. Localities: 1. Mucugê; 2. Sopa, Diamantina, Minas Gerais (MG); 3. Extração, Diamantina (MG); 4. Guinda, Diamantina (MG); 5. Conselheiro Mata, Diamantina (MG).

Tables

TABLE 3. Comparative morphological data for Gymnodactylus vanzolinii sp. nov., and currently recognized species of the genus Gymnodactylus. Mean, standard deviation, range and sample size (n) are presented for each species. Data were taken from examined specimens (Appendix 1). * = estimated value.

Character	vanzolinii sp. nov.	guttulatus	Gymnodactylus amarali carvalhoi	geckoides	darwinii
Snout-vent Length	45.2 ± 6.18 (31.4–52.3; n = 17)	38.36 ± 8.73 (27.9–49.6; n = 7)	34.2 42.02 ± 7.27 (24.1–51.3; n = 13)	40.95 ± 2.18 (33.0–44.5; n = 30)	47.68 ± 4.45 (39.8–53.6); n = 16
Tail length	64.19 ± 5.03 (59.4–70.6; n = 4)	44,0 5 ± 5.16 (40.4–47.7; n = 2)	NA 42.5 ± 21.63 (27.2–57.8; n = 2)	45.35 ± 3.18 (43.1–47.6; n = 2)	55.3 ± 6.62 (46.7–66.8); n = 6
Head length	11.35 ± 1.36 (8.2–12.9; n = 17)	9.79 ± 1.65 (7.7–12.1; n = 7)	8.5 10.7 ± 1.63 (7.0–13.0; n = 13)	10.3 ± 0.57 (8.2–11.0; n = 30)	11.99 ± 0.88 (10.7–13.7); n = 16
Head width	8.225 ± 1.08 (5.6–9.4; n = 16)	7.19 ± 1.86 (4.0–9.1; n = 7)	6.7 8.32 ± 1.34 (5.5–10.7; n = 13)	7.62 ± 0.40 (6.3–8.2; n = 30)	8.79 ± 0.86 (7.4–10.4); n = 16
Head height	5.33 0.86 (3.6–6.8; n = 16)	4.85 ± 1.07 (3.4–6.1; n = 7)	3,95 5.24 ± 1.06 (3.7–7.3; n = 13	4.89 ± 0.36 (4.1–5.6; n = 30)	5.77 ± 0.68 (4.7–7.1)
Supralabial scales	7 ± 0.63 (5–8; n = 16)	6.14 ± 0.38 (6–7; n = 7)	5/6 5.92 ± 0.28 (5–6; n = 13)	6.13 ± 0.57 (5–7; n = 30)	6.44 ± 0.51 (6–7; n = 16)
Subdigital lamellae 4th finger	16.52 ± 0.87 (16–18; n = 17)	16.14 ± 0.90 (15–18; n =7)	13/12 12.61 ± 0.77 (11–14; n = 13)	13.70 ± 0.70 (13–15; n = 30)	18.44 ± 0.93 (17–20; n = 16
4th toe	19.52 ± 0.94 (18–21; n = 15)	19.86 ± 1.07 (18–21; n = 7)	15/14 16 ± 0.74 (15–17; n = 12)	17.5 ± 0.93 (15–19; n = 30)	21.94 ± 0.68 (21–23; n = 16)
Tubercles rows	10.43 ± 0.65 (10–12; n = 14)	12.4 ± 1.14 (11–14; n = 5)	14 14 ± 0 (14; n = 13)	12.06 ± 0.37 (12–14; n = 29)	14 ± 0 (14; n = 16)
Tubercles in a paramedian row	25.38 ± 2.29 (22–29; n = 13)	21.5 ± 1.64 (19–24; n = 6)	38* 31.75 ± 3.44 (27–38; n = 12)	34.48 ± 2.98 (27–40; n = 23)	51.53 ± 3.29 (47–59; n = 15)
Ventrals	20.27 ± 1.19 (18–22; n = 11)	23 ± 1.83 (20–26; n = 7)	19 21.08 ± 1.38 (19–24; n = 13)	21.07 ± 0.99 (19–24; n = 27)	18.75 ± 3.27 (15–24; n = 16)