Leiolepis ngovantrii, Grismer, Jesse L. & Grismer, Lee, 2010

Grismer, Jesse L. & Grismer, Lee, 2010, Who’s your mommy? Identifying maternal ancestors of asexual species of Leiolepis Cuvier, 1829 and the description of a new endemic species of asexual Leiolepis Cuvier, 1829 from Southern Vietnam, Zootaxa 2433, pp. 47-61 : 52-58

Leiolepis ngovantrii sp. nov.

Ngo Van Tri’s Lady Butterfly Lizard Figure 3 a.

Holotype. Adult female ( LSUHC 9234) collected on 21 June 2009 from Binh Chau – Phuoc Buu Nature Reserve, Xuyen Moc district, Ba Ria Vung Tau Province, Vietnam (10° 32.418N 107° 32.177E; 30 m a.s.l.).

Paratypes. The collection data for the paratypes (all females) LSUHC 9230–33 and LSUHC 9235 are the same as the holotype.

Diagnosis. Leiolepis ngovantrii differs from all sexual species of Leiolepis in lacking males; it differs from all asexual species in having a combination of nine rows of enlarged keeled scales across the forearm; 37–40 subdigital lamellae beneath the fourth toe; a relatively narrow head (0.71–0.83); 50–53 dorsal scales between the dorsolateral stripes; absence of a lateral, caudal stripe; having 24 or 25 gular scales between the enlarged, mandibular scales; the ventrolateral stripe not reaching the axillary region; absence of a black, Yshaped marking in the pectoral region, a vertebral stripe, and eyespots bearing white centers; presence of transverse bars on the flanks; presence of thin, transverse, dorsal, caudal bars, and the absence of dorsal, caudal blotches.

Description of the holotype. Head moderate in size, (HL 21mm; HL/SVL 0.17) obtusely rounded in lateral profile, triangular in dorsal profile (HW 17.5mm; HW/HL 0.83); interorbital, frontal region, and rostrum high, convex (HH 13.5 mm; RH 8.5mm; RH/HH 0.63), sloped anteriorly, covered with small, undifferentiated keeled scales; occipital and supraorbital regions covered with, 22 keeled, granular scales half the size as head scales; canthus rostralis short, rounded, composed of one or two enlarged scales; dorsal head scales strongly keeled, seven scales across the frontal bone between supraorbital regions; rostral large, triangular, bordered laterally by first supralabials, posteriorly by seven smaller scales; external nares large, set wide apart (DN 4.5 mm; DN/HW 0.26) rounded, directed laterally, set in single, oval, nasal scale surrounded by several small scales; an elongate, keeled, suborbital scale extends from beyond anterior margin of eye to just beyond midorbital region, seven enlarged, keeled suborbitals extend from anterior margin of eye to posterior margin of eye; superciliary scales elongate, keeled, imbricate, continuous with canthal scales; eyelid scales granular; tympanum naked, deeply set, surrounded by granular scales; temporal scales small, slightly raised; nine rectangular supralabials whose contact with one another produces an undulating labial margin; mental longer than wide, pointed posteriorly, larger than adjacent infralabials; two large postmentals in contact medially, being first of a series of eight enlarged scales along the angle of jaw, separated from all but first infralabials by smaller scales; 11L,10R rectangular infralabials, first infralabial largest; gular scales small, rounded, granular; 24 gular scales between enlarged, mandibular scales; anterior and posterior gular folds present; dewlap absent; antehumeral fold continuous with posterior gular fold.

Body elongate (AG 60.5 mm; AG/SVL0.50) somewhat dorsoventrally compressed; body scales small, granular, slightly keeled; 52 scales between dorsolateral stripes; 16 scale rows between ventrolatral stripe and dorsolateral stripes at widest point of trunk; scales of flanks abruptly transition into much larger, flat scales of belly and pectoral region; 43 scales across middle of belly contacting apex of the umbilical scar; precloacals much smaller, juxtaposed; limbs relatively short, robust (FA 15 mm; FA/FL 0.52; FL/SVL 0.23); dorsal surface of forelimbs and posterior surface of brachia covered with large, keeled, imbricate scales; nine rows of enlarged, keeled scales across forearm; scales of forearms less keeled than those of brachia; ventral surface of forelimbs and posterior surface of forearms covered with granular scales, those of forearms raised; plantar scales small, raised; subdigital lamellae of fingers tricarinate, transversely elongate; claws long; hind limbs relatively long (TIB 23 mm; HB 43 mm; TIB/HB 0.53; HB/SVL 0.35); dorsal scales on hind limbs small, weakly keeled; scales on anterior surface of thighs large, flat, weakly keeled, imbricate; those on forelegs slightly enlarged, keeled; postfemoral scales small, granular; 11L,10R longitudinal rows of large, flat, imbricate subtibial scales; 42 femoral pores in total; each pore set in larger scale; 13 non-pore-bearing scales between pore-bearing-femoral scales; plantar scales small, raised; subdigital lamellae of toes bicarinate, 39 beneath fourth toe; 32 enlarged, plate-like scales along dorsal surface of the fourth toe; four enlarged, triangular scales on posterior surface at base of third toe, (ESL 1.2mm; ESL/TE 0.09); 10 enlarged, plate-like scales along length of first toe; tail dorsoventrally compressed, noticeably swollen at base, constricted at its contact point with body, covered dorsally with small, keeled scales grading ventrally into larger, flat, weakly keeled, subcaudals; caudal scales in transverse rows.

LSUHC 9234 LSUHC 9233 LSUHC 9235 LSUHC 9232 LSUHC 9230 LSUHC 9231 continued next page Coloration in life ( Fig. 3 a). Dorsal ground color of head, body, limbs, and tail light brown to dull yellow; no light spotting on top of head; light marking on anterior corner of eye and below eye; posterior portion of lower eyelid white; series of small, closely spaced, yellowish spots forming broken, postorbital stripes that extend across temporal regions to above shoulders where they become continuous, turn downward, and extend onto forelimb insertions; postorbital stripe counter shaded ventrally by black on neck and shoulders; lower temporal region beige to cream colored; white spots on side of neck; the stripe itself being bright yellow; large, prominent, olive to dull yellow eye spots on back; dorsal eye spots enclosed within a reticulum of thin, black lines, 4–5 eyespots between dorsolateral stripes at widest point on trunk; wide, prominent, yellow, dorsolateral stripe extends from base of tail onto shoulder region; a yellow ventrolateral stripe fades anteriorly and contacts axillary region; faint, white spots on forelimbs; dorsal pattern of body extends to dorsal surface of hind limbs; smaller, olive to dull yellow, diffuse spots on middorsal region of tail enclosed in black reticulum; lateral caudal stripe present, subcaudal region immaculate; posterior region of tail bearing thin, transverse, black, dorsal bars; gular region beige bearing faint to bold, irregularly shaped, oblique, dark lines; throat gray; pectoral and abdominal regions and underside of limbs beige, generally immaculate; faint, dark reticulum occasionally present in pectoral region.

Variation. Differences in scale counts and measurements are presented in TABLE 2 . The paratypes closely approach the holotype in aspects of coloration and pattern. The tail tip of LSUHC 9232 is regenerated and lacks a pattern. The remaining paratypes are juveniles and differ slightly in having a lateral, caudal stripe; a prominent, vertebral stripe, and a more contrasting color pattern that is better defined.

Distribution. Leiolepis ngovantrii is known only from the type locality at Binh Chau-Phuoc Buu Nature Reserve, Xuyen Moc District, Ba Ria Vung Tau Province, Vietnam ( Fig. 1 ).

Natural history. Leiolepis ngovantri occurs in coastal areas and areas dominated by Melaleuca forests ( Fig. 4 a) where loose, sandy soils provide ample opportunity for burrow construction and sparse, scattered vegetation offer areas for basking. Lizards were most common in ecotonal areas between the coastal dune areas ( Fig. 4 b) dominated by L. guttata and the mangrove regions. Here, L. ngovantrii was most active during sunny morning hours and would intermittently retreat into burrows during the hotter periods of the afternoon. Leiolepis ngovantrii builds complex, interconnected tunnels that are significantly deeper than those of its sexual congeners (pers. observ.). In fact, we were told stories about people being killed by the dune faces collapsing on them because they had to dig so deep to catch the lizards. We hypothesize the depth of these burrows may be due to the fact that this is an all female species and female Leiolepis of all species usually dig deeper tunnels in which to lay eggs and house hatchlings ( Cox et al., 1998).

Etymology. This species is named in honor of Mr. Ngo Van Tri whose extensive field work in southern Vietnam resulted in the discovery of this new, asexual population as well as many other new species of lizards.

Comparisons. Leiolepis ngovantrii differs from all asexual species except L. triploida in having a narrower head (HW/HL 0.71–0.83 vs. 0.66–0.69 and 0.65–0.68 in L. boehmei and L. guntherpetersi ), 50–53 dorsal scales between the dorsolateral stripes vs. 42–48 and 38–42 in L. boehmei and L. guentherpetersi , respectively; and lacking a light lateral, caudal stripe. Leiolepis ngovantrii differs further from all asexuals except L. boehmei in lackinga black, Y-shaped, pectoral marking, a vertebral stripe, and eyespots bearing white centers. Leiolepis ngovantrii differs further from all asexuals except L. guentherpetersi in having more gular scales (24–25 vs. 18–19 in L. boehmei and 17–19 in L. triploida ) between the enlarged, mandibular scales; the ventrolateral stripe not reaching the axillary region; the presence of transverse bars on the flanks and thin, transverse, dorsal caudal bars; and lacking, dorsal caudal blotches. Leiolepis ngovantrii differs further from L. boehmei in having fewer supralabials (9–10 vs. 11–12) and from L. guentherpetersi in having more supraorbital scales across dorsal surface of supraorbital region (6–7 vs. 4) and fewer scale across the ventral side of the tibia (10–11 vs. 14).

Phylogenetic Analysis. The maximum parsimony (MP) and maximum liklihood (ML) analyses support the monophyly of Leiolepis , however, they returned differing topologies ( Fig 5 a&b). The MP analysis ( Fig. 5 a) recovered a clade of L guentherpetersi that formed a sister group relationship to a clade of northern L. guttata , which in turn, is the sister group to a clade containing L. triploida , L. boehmei , L. ngovantrii and southern L. guttata . Those two clades are sister to the remaining sexual species, with L. belliana being sister to L. peguensis plus L. reevesii . The ML analysis returns a different topology. Leiolepis reevesii and L. peguensis are sister species and form a polytomy with L. belliana and L. guttata plus all the asexual species. Although the ML and MP topologies ( Fig. 4 b) conflict for three of the sexual species, the relationships between the asexuals and populations of L. guttata are the same.

Figures

FIGURE 1. Distribution of all species of Leiolepis, with colored circles representing the samples used in the phylogenetic analysis. * indicate asexual species.

FIGURE 3. (a) Leiolepis ngovantrii (b) female L. guttata, and (c) male L. guttata from Binh Chau – Phuoc Buu Nature Reserve, Ba Ria Vung Tau Province, Vietnam.

FIGURE 4. Top is ecotonal habitat between mangrove forest, where Leiolepis ngovantrii is most common and costal dune habitat (bottom).

FIGURE 5. (a) The results from the MP analysis. (b) The results from the ML analysis. The red square indicates the topological difference between the MP and ML trees. MY = Malaysia, TH = Thailand, MR = Myanmar, CH = China, NV = Northern Vietnam, and SV = Southern Vietnam.

Tables

TABLE 2. Descriptive measurements (mm) and characters of the type series. ^ indicates juvenile, and / indicates character does not apply.

Holotype	Paratype	Paratype	Paratype	Paratype^	Paratype^
Sex F	F	F	F	F	F
SVL 122	126.5	117	95.2	52.2	65.8
HL 21	24.5	24	21	10.2	15.5
HL/SVL 0.17	0.2	0.2	0.22	0.2	0.23
HW 17.5	17.5	17	14.8	6	11
HW/HL 0.83	0.71	0.71	0.71	0.59	0.71
DN 4.5	4.5	4.5	4	2.1	2.6
DN/HW 0.26	0.26	0.26	0.27	0.35	0.24
HH 13.5	13.5	13.5	11	7.2	8
RH 8.5	8.5	8	6.7	3.8	4.7
RH/HH 0.63	0.63	0.6	0.61	0.53	0.59
HE 5	5.5	5	4.3	3	3.4
HE/HH 0.37	0.4	0.37	0.39	0.41	0.42
SL 9	11L–10R	10L–9R	9L–10R	9	10L–9R
IL 11L–10R	10L–9R	10	10	10	10
FB 6	6	7	7	6	7
PR 7	6	6	7	6	6
SO 22	20	21	23	22	21
SBO?L–7R	7L–8R	7	7	8L–7R	7
ESM 8	10	9	10	9L–10R	10
GBM 24	24	24	25	23	23
VMR +	+	+	+	+	+
FA 15	15	13.9	12	7.6	8.3
FA/FL 0.55	0.57	0.56	0.57	0.59	0.56
FL/SVL 0.23	0.2	0.21	0.22	0.24	0.22
KSFA 9	9	9	9	9	9
AG 60.5	62	56	46.3	25	32
AG/SVL 0.5	0.49	0.48	0.48	0.48	0.49
BY –	–	–	–	–	–
VS –	–	–	–	–	–
DSS 52	50	50	53	49	52
ESS 4	5	5	4	4	5
VNS +	+	+	+	+	+
VNSA +	+	+	+	+	+
SVNVL 16	18	18	18	18	19
TB –	–	–	–	–	–
HB 43	44	40	34.5	23	25.1
HB/SVL 0.35	0.34	0.35	0.36	0.44	0.38
TIB 23	24	21.6	19.3	12.7	13.3