Ameivula ocellifera ( Spix, 1825 )

Oliveira, Eliana F., Costa, Henrique C., São-Pedro, Vinícius A., Marques, Ricardo, Santana, Diego J., Colli, Guarino R., Mesquita, Daniel O. & Costa, Gabriel C., 2022, On the type locality of Ameivula ocellifera (Spix, 1825) (Squamata, Teiidae), with a neotype designation, Zoosystema 44 (19), pp. 475-491 : 482-487

Ameivula ocellifera ( Spix, 1825)

Tejus ocellifer Spix, 1825: 23 .

Cnemidophorus ocellifer – Peters 1877: 414.

Ameivula ocellifera – Harvey et al. 2012: 93 (type species of Ameivula ).

Cnemidophorus hygomi Reinhardt & Lütken, 1862: 231

(namebearing type: holotype, ZMB 4335 . Type locality: “ Maruim ”, state of Sergipe, Brazil).

Heterochresonymy

Tej[us] murinus (non Seps murinus Laurenti, 1768 ) – Boie 1826: 120 (partim). — Fitzinger 1827: 747 (partim).

Seps murinus (non Seps murinus Laurenti, 1768 ) – Wagler 1830: 154 (partim).

Teius (Ameiva) vulgaris (non Ameiva vulgaris Lichtenstein, 1823 ) – Gray 1831: 29 (partim).

Cnemidophorus murinus (non Seps murinus Laurenti, 1768 ) – Duméril & Bibron 1839: 107 (partim).

Cnemidophorus lacertoides (non Cnemidophorus lacertoides Duméril & Bibron, 1839 ) – Reinhardt & Lütken 1862: 146.

TYPE MATERIAL. — Original name-bearing types. Brazil • 2 syntypes; “ Bahia ”; ZSM 111/0 and a second unlabeled specimen, both lost. Neotype (here designated). Brazil • 1 ♂; Parque das Dunas ; 12°55’3”S, 38°19’12”W; 20 m a.s.l.; municipality of Salvador, state of Bahia; R. Marques, M. L. T. Oliveira, N. J. L. Santos, and M. S. Bonfim leg.; 15.VII.2018; ZUFMS-REP 04144 .

ORIGINAL TYPE LOCALITY. — “ Bahia ”.

TYPE LOCALITY (BY NEOTYPE DESIGNATION). — Parque das Dunas, 12°55’3”S, 38°19’12”W, 20 m a.s.l., municipality of Salvador, state of Bahia, Brazil ( Fig. 3 ).

ETYMOLOGY. — The specific name ocellifera (masculine ocellifer ) means “bearing little eyes”. It derives from the Latin ocellus (diminutive of oculus, i.e., “small eye”) and fero (“to carry”, “to bear”), in reference to the series of small bluish ocelli between the fore and hindlimbs.

DESCRIPTION OF THE NEOTYPE

Morphometry and lepidosis

Adult male in good condition; no evident scar or damage ( Figs 4 ; 5 ). Tail whole; small longitudinal incision on left thigh due to muscle sample collection. Snout-vent length, 61.08 mm; trunk length, 30.36 mm; tail length, 133.59 mm; head length, 18.53 mm; head width, 10.25 mm; head height, 8.41 mm; humeral length, 6.48 mm; forelimb length, 19.4 mm; femur length, 13.03 mm; tibia length, 12.68 mm; foot length, 22.10 mm; hindlimb length, 47.81 mm ( Fig. 4 ). Head distinct from neck ( Fig. 5 ), longer than wide (head length/head width, 1.80); head length/snout-vent length, 0.30. Trunk elongate (trunk length/ snout-vent length, 0.49); tail long, about 2.2 × snout-vent length. Snout moderately pointed. Rostral and frontonasal separated by pair of anterior nasals ( Figs 4 ; 5 , see details in Appendices 1; 2). Nostril anterior to and in contact with nasal suture. Posterior nasals contact pair of prefrontals. Prefrontal on each side separated from first supraciliary by loreal and first supraocular. Loreal, single, large, almost high as long, in contact with posterior nasal, prefrontal, first supraocular, first supraciliary, first and second suboculars, and third and fourth supralabials. Frontal entire, lacking ridges; posterior suture of frontal aligns with first half of third supraocular. Frontoparietals, two. Parietals, four (a pair on each side). Interparietal subequal in size to each flanking pair of parietals. Occipitals (c. 45) irregular, variable in size, larger than dorsals on first row, immediately posterior to them. Temporals irregular in size. Four enlarged supratemporals in longitudinal row behind supraciliaries. Supratemporals separated from parietals by occipitals. Circumorbitals reaching posterior border of first supraocular. Supraoculars, four on each side, 3 rd> 2 nd> 1 st> 4 th; first supraocular entire, contacting second supraocular, loreal, prefrontal, frontal, and first supraciliary; second to fourth supraoculars separated from supraciliaries by one row of lateral supraocular granules. Supraciliaries, five on each side; first supraciliary in contact with first subocular. Supralabials, six on each side, third largest. Infralabials, five on each side; third to sixth separated from chinshields by row of granules. Chinshields, four on each side; first pair with suture through almost their length. Gular region divided in two: anterior gular region with rounded and larger scales, in 20 irregular transverse rows; posterior gular region covered by smaller scales in 15 transverse rows. Interangular and intertympanic sulci absent; interauricular crease present. Dorsal and lateral head scales juxtaposed and smooth. Dorsals round, smooth, and sub-imbricate; 176 dorsals along mid-dorsum, from nape to base of tail; 88 around midbody. Ventrals smooth, wider than long, imbricated, in eight longitudinal rows and 26 transverse rows. Preanal plate with three enlarged scales (a central and two posterior scales), surrounded by seven smaller scales.Scales on base of tail rectangular, smaller than ventrals, in transverse rows; scales around tail, 28, longer and narrower posteriorly; tail scales keeled dorsally, smooth ventrally. Scales on flanks granular, round, smooth, sub-imbricate. Forelimbs with large, smooth, imbricate scales on dorsal surface of upper arms and anterodorsal forearms; elsewhere scales small and granular. Scales in the dorsal region of the forearm enlarged, in two longitudinal rows. Hindlimbs with large, smooth, imbricate scales on anteroventral surface of thighs and ventral surface of lower legs; elsewhere, scales small and granular; enlarged prefemoral scales, 10. Femoral pores in continuous row along each thigh; seven pores on right side and eight pores on left side. Abdominal pores, one on each side. Lamellae under fourth finger, 16 on each side; under fourth toe, 27 on each side. Fifth toe shortened. Tibiotarsal spurs absent.

Coloration in life

Dorsal parts of head, body, limbs, and tail medium brown ( Fig. 3A ). Lateral parts of the head light brown. Belly and ventral parts of the head, limbs, and tail, uniform creamy white. Vertebral stripe absent. Paravertebral field enlarged, light brown, bordered on both sides by lighter paravertebral stripes that run from nape to base of tail (see details in Appendix 1 ). Paravertebral stripes separated from dorsolateral stripe on each side by dark brown dorsolateral field, from nape to first third of tail. Dorsolateral stripe creamy white, from nape to first third of tail, separated from upper lateral stripe by upper lateral field. Upper lateral field with irregular black spots that tend to fuse anteriorly and series of small and irregularly spaced bluish ocelli (7-21 scales diameter) between fore and hindlimbs; ocelli close to hindlimbs in brown to greenish-yellow background; blue coloration along upper lateral stripe in some anterior caudal scales. Upper lateral stripe from subocular scales to first third of tail, creamy white anteriorly, light brown and greenish-yellow in middle of trunk, creamy white posterior to hindlimbs. Upper lateral stripe separated from creamy white lower lateral stripe by lower lateral field, enlarged anterior to forelimbs. Lower lateral stripe short, from below ear to axilla. Lower lateral field irregularly black-spotted, interspersed with whitish-blue ocelli (8-16 scales diameter) surrounded by greenish-yellow scales between fore and hindlimbs; some enlarged lateral scales and some outermost ventral scales blue.

Color in ethanol

Head brownish dorsally, brownish-white laterally, and bluishwhite ventrally. Dorsum brownish, stripes bluish-white, fields dark brown or black (except paravertebral field light gray), and ocelli faded bluish-white. Belly bluish-white. Ventral aspect of forelimbs and hindlimbs immaculate. Dorsal aspect of tail like in life, although faded, including few blue scales close to the base of tail. Ventral aspect of tail immaculate ( Fig. 4 ).

COMPARISON WITH OTHER AMEIVULA SPECIES

The following characters (see Table 2 and Table 3 ) distinguish A. ocellifera from its congeners (data in parentheses). Ameivula ocellifera differs from A. abalosi in having 82-141 scales around midbody (75-98), 26-38 subdigital lamellae under fourth toe (24-31), and whitish blue ocelli (light-blue or whitish green ocelli). Ameivula ocellifera differs from A. apipensis in having 14-28 femoral pores (12-17), 26-38 subdigital lamellae under fourth toe (24-27), vertebral stripes absent (present), whitish-blue ocelli (no ocelli), and immaculate throat (yellow). Although the description of A. apipensis indicates an intertympanic sulcus (absent in A. ocellifera ), the figures available indicate the contrary, i.e., the absence of such structure in A. apipensis . Ameivula ocellifera differs from A. cipoensis in having 13-20 subdigital lamellae under fourth finger (12-14), 26-38 subdigital lamellae under fourth toe (21-24), lower lateral stripes broken (absent), whitish-blue ocelli (yellow-greenish ocelli), and hindlimb spots present (absent). Ameivula ocellifera differs from A. confusioniba in having paravertebral and dorsolateral fields present (absent), paravertebral and lower lateral stripes present (absent), whitish-blue ocelli (bright yellow ocelli), and hindlimb spots present (absent). Ameivula ocellifera differs from A. jalapensis in having eight longitudinal rows of ventral scales (6-8), 14-28 femoral pores (10-16), 172- 248 dorsal scales (200-250), 35 gular scales (41-64), larger body size (maximum SVL, 56 mm), vertebral stripe absent (broken), upper and lower lateral fields spotted (uniform), whitish-blue ocelli (no ocelli), hindlimb spots present (absent), and immaculate throat (lime-green). Ameivula ocellifera differs from A. mumbuca in having eight longitudinal rows of ventral scales (6-8), 172-248 dorsal scales (194-271), 26-38 subdigital lamellae under fourth toe (24-32), 35 gular scales (38-49), larger body size (maximum SVL, 59 mm in males), lower lateral fields spotted (uniform), and whitish-blue ocelli (whitish-blue spots only in males). Ameivula ocellifera differs from A. nativo in having 25-31 transverse rows of ventral scales (29-32), vertebral stripe absent (present), paravertebral fields present (absent), paravertebral and lower lateral stripes present (absent), upper and lower lateral fields spotted (uniform), whitish-blue ocelli (no ocelli), hindlimb spots present (absent), immaculate throat (light blue), and bisexual (only females). Ameivula ocellifera differs from A. nigrigula in having 21-30 scales around tail (27-32), whitish-blue ocelli (bright turquoise ocelli), hindlimb spots present (absent), and throat region immaculate (black). Ameivula ocellifera differs from A. pyrrhogularis in having 172-248 dorsal scales (192- 255), 21-30 scales around tail (23-33), vertebral stripe absent (present in juveniles and females), whitish-blue ocelli (light blue or greenish-blue ocelli), and immaculate throat (orange in adult males). Ameivula ocellifera differs from A. xacriaba in having 26-38 subdigital lamellae under fourth toe (23-28), 35 gular scales (41-52), paravertebral stripes present (absent), whitish-blue ocelli (bright green ocelli), and hindlimb spots present (absent). Although the description of A. xacriaba indicates an intertympanic sulcus (absent in A. ocellifera ), the figures available reveal the contrary, i.e., the lack of such structure, but with an interauricular crease in A. xacriaba . Less marked differences between species can be found in Table 2 and Table 3.

A. ocellifera Character Neotype A. abalosi A. apipensis A. cipoensis A. confusioniba A. jalapensis A. mumbuca A. nativo A. nigrigula A. pyrrhogularis A. xacriaba VES absent absent present absent absent present, absent present absent present (juveniles absent broken and females) PVF present present present present absent present present absent present present present (juveniles) PVS present present (faint) present present absent present, present, broken absent present present absent broken (juveniles) DLF present present present present absent present present present present present present (juveniles) DLS ULF ULS LLF LLS LAS HLS THR SED present present present present present present present, broken present present present present

(occasionally) (juveniles)

spotted spotted spotted spotted spotted uniform uniform or spotted uniform spotted spotted spotted

present present present present present present present present present present present

spotted uniform or spotted spotted uniform or uniform uniform uniform spotted spotted spotted

spotted spotted

present, broken present present absent absent present, present, broken absent present present, broken present

broken

whitish- light blue or irregulars light yellow-greenish bright yellow absent bluish-white spots absent bright turquoiselight blue or bright green

blue ocelli whitish- green spots ocelli alternating ocelli (males) ocelli greenish-blue ocelli

alternating green ocelli (no ocellus) with irregular ocelli (adults),

with irregular alternating alternating black spots tan ocelli

black spots with irregular with black (juveniles)

black spots spots

present present present absent absent absent mostly present absent absent present absent

(frequently)

immaculate pearly white yellow white immaculate lime-green immaculate light blue black orange (adult white

males)

n/a absent present present (belly absent absent present partheno- present (stripes present (stripes present (flanks

(dorsolateral yellow in males, (anteriormost light genetic absent in barely visible bright green

fields dark bluish-white in areas bluish in (only males) and fields in adult

brown in females; ocelli males and whitish females) partially fade males,

adult males, from fore- to in females; lower away in adult bright yellow

bright green hindlimbs in lateral fields males) in adult

in females) males, 2-3 ocelli generally with females)

proximal to bluish-white spots the forelimb in in males, uniform females) in half of the females)

DISTRIBUTION

The current known geographic range of Ameivula ocellifera is in the southeastern portion of the Caatinga (states of Pernambuco, Alagoas, Sergipe, and Bahia) and part of the Atlantic Forest of eastern Bahia (including Salvador, its type locality), Sergipe, and Alagoas ( Arias et al. 2018). The identity of populations from central and north Caatinga (Paraíba, Rio Grande do Norte, Ceará, and Piauí) is under revaluation, and may reveal a distinct taxon in the near future. Records attributed to A. ocellifera in northern Brazil (Rondônia, Pará, and Tocantins), midwestern Brazil (Mato Grosso, Mato Grosso do Sul, Goiás, Distrito Federal, Minas Gerais, and São Paulo), and northeastern Brazil (Maranhão) ( Costa et al. 2022) represent candidate species close to A. ocellifera (e.g. “W-Ce clade” in Arias et al. 2018) or probably misidentifications.

Figures

FIG. 3. — A, Ameivula ocellifera (Spix, 1825) neotype male ZUFMS-REP 04144 in life; B, habitat of neotype at the Parque das Dunas, municipality of Salvador, state of Bahia, Brazil. Photos: Ricardo Marques.

FIG. 4. — Ameivula ocellifera (Spix, 1825) neotype male ZUFMS-REP 04144: A, dorsal view of the body; B, ventral view of the body; C, lateral view of the body. Scale bar: 20 mm.

FIG. 5. — Ameivula ocellifera (Spix, 1825) neotype male ZUFMS-REP 04144: A, dorsal, ventral, and lateral view of the head; B, ventral view of the right hand; C, ventral view of the right foot. Scale bar: 20 mm.

APPENDIX 1. — Neotype (ZUFMS-REP 04144, adult male) of Ameivula ocellifera (Spix, 1825): A, body in dorsal view, showing: two longitudinal rows of enlarged scales in the dorsal region of the forearm (EFA); dorsal scales (DS); paravertebral field (PVF); paravertebral stripe (PVS); dorsolateral field (DLF); dorsolateral stripe (DLS); upper lateral field (ULF); and upper lateral stripe (ULS); B, body in ventral view, showing: longitudinal rows of ventrals (VLR); transverse rows of ventrals (VTR); femoral pores (FP); abdominal pores (AP); enlarged prefemoral scales (EPF); and enlarged scales of preanal plate (EP); C, body in lateral view, showing: dorsolateral field (DLF); dorsolateral stripe (DLS); upper lateral field (ULF); upper lateral stripes (ULS); lower lateral field (LLF); and lower lateral stripe (LLS). Scale bar: 20 mm.

Tables

TABLE 2. — Comparison of meristic characters in the Ameivula ocellifera group: Ameivula ocellifera (Spix, 1825),Ameivula abalosi (Cabrera,2012), Ameivula apipensis Arias et al . , 2018 , Ameivula cipoensis Arias et al . , 2014 , Ameivula confusioniba (Arias et al., 2011), Ameivula jalapensis (Colli et al., 2009), Ameivula mumbuca (Colli et al., 2003), Ameivula nativo (Rocha et al., 1997), Ameivula nigrigula ( Arias et al . , 2011 ) , Ameivula pyrrhogularis (Silva & Ávila-Pires, 2013), Ameivula xacriaba Arias et al., 2014. Most values were taken from the original descriptions (values without symbols) and other studies ( values with symbols ) . Numbers indicate the range of each character, with mean in parentheses or mode value in brackets.Bold indicates data from the holotype. Italics indicates divergent data from the same source/ article ( text and tables ) . Abbreviations : VLR , longitudinal rows of ventrals; VTR, transverse rows of ventrals; FP, femoral pores on both sides; DS, dorsal scales; SAT, scales around tail; SAM, scales around midbody; L4 F , subdigital lamellae under fourth finger on both sides ; L4T, subdigital lamellae under fourth toe on both sides; GS, gular scales; see Appendices 1; 2. Symbols: †, raw data from Table 2 by Colli et al. (2003a) and Table 1 by Colli et al . ( 2009 ) , excluding specimens from Irecê and Ibipeba, state of Bahia, which may be Ameivula nigrigula; *, from Table 1 by Arias et al. (2011a) and Table 2 by Arias et al. (2011b); ¶, from Table 1 by Arias et al . ( 2014 a ) ; ‡ , from Table 2 by Arias et al . ( 2014 b ) ; § , from Table 2 by Silva & Ávila-Pires (2013);?, values not available.

A. ocellifera
Character This study Neotype	Colli et al. Arias et al. 2003a† 2011 a* n = 44 n = 52	A. abalosi n = 17 ♂; n = 6	A. apipensis n =?	A. cipoensis n = 40;	A. confusioniba A. jalapensis A. mumbuca A. nativo A. nigrigula n = 19; n = 51; n = 229; n = 34 or 35; n = 83;	A. pyrrhogularis n = 128	A. xacriaba n = 92
		♀; n = 22 or		n = 40‡	n = 34§; n = 38‡ n = 35‡ n = 19* n = 25‡
		28¶; n =?‡			n = 30‡
VLR 8	8 8	8; 8; 8¶	8	8	8; 8§ 6-8 (7.51); 6-8 6-8 (8); 8‡ 8 8	8	8
					(7.33)‡
VTR 26	25-31 (28) 26-28 (27.2) 27-31 (28.71);		28	28-30 (29.1)	27-29 (28.2); 24-29 (25.96); 24-29 (27.03);29-32 [31]; 26-29 (27.4)	26-31 (28.3)	27-29 (28)
		24-31 (28.33);			26-31 (28.8)§ 25-27 26-28 29-32
		27-30 (28.6)¶			(25.7)‡ (27.1)‡ (30.8)*
FP 15	16-28 (20.4) 14-17 (16)	16-19 (17.59);	12-17	17-20 (17.7)	16-21 (17.5); 11-16 (13.35); 14-20 (16.66);22-26 [24]; 15-20 (17)	16-22 (18.5)	14-17 (15)
		15-19 (17.17);			14-21 (17.6)§ 10-16 16-20 22-26 (24)*
		16-20 (17.5)¶			(13.1)‡ (18.3)‡
DS 176	203-248 172-188	184-212	184-230	202-225	188-211 (201.6); 200-250 194-271 192-212 190-220	192-255 (222.9)	191-216 (202);
	(222.5) (181)	(198.63); 190-		(213.6)	196-240 (215)§ (225.96); (229.57); (196)* (208.6)		184-216
		208 (201.1);			208-226 200-227
		185-208 (204)¶			(218)‡ (211.5)‡
SAT 28	21-30 (26.3) 25-30 (28)	23-26 (24.41);	31	25-29 (26.7); 22-28 (24.9); 19-26 (22.72); 19-27 (22.61);25-27 (26)* 27-32 (30)		23-33 (26.7)	23-29 (26.3);
		23-26 (24.67);		24-29	20-28 (23.8)§ 22-26 (24)‡ 22-28		24-29
		24-27 (25.3)¶		(27.2)	(25.1)‡
SAM 88	82-141 92-100	75-98 (84.6);	84-90	85-100	87-105 (95); 91-122 91-117 93-102 97-105	87-124 (104)	94-106 (99.3);
	(97.9) (94.6)	77-98 (85.83);		(92.6)	89-116 (104.04); (101.05); (95.5)* (99.8)		94-102
		84-95 (89)¶			(102.6)§ 89-100 95-102
					(94.3)‡ (98.8)‡
L4F 16	13-20 (16.7) 15-17 (16)	15-17 (16.06);	14-16	12-14 (12.5)	15-17 (15.6); 12-18 (15.49); 13-19 (16.26);16-17 [16]; 16-18 (17)	13-19 (16.1)	13-15 (14.1)
		14-16 (15.17);			14-18 (17)§ 14-16 15-17 (16)‡ 14-17 (16)*
		15-17 (15.9)¶			(14.7)‡
L4T 27	26-38 (31.6)28-31 (30)	26-31 (28.25);	24-27	21-24 (22.5)	29-35 (30.4); 22-37 (26.63); 24-32 (27.36);29-33 [31 30-32 (31.2)	24-34 (29.8)	23-28 (25.4)
		25-31 (27.67);			27-34 (31.1)§ 23-28 (26)‡ 25-29 and 32];
		24-30 (27.3)¶			(26.8)‡ 26-33 (31)*
GS 35	??	31-35‡	28-35	30-38‡;	29-39‡ 41-64 (51.3)‡ 38-49 (44.9)‡? 31-35 (33.3)‡?		41-52 (45.7);
				31-38‡			41-47