Copeia, 2000(2), pp. 501–509
New Cnemidophorus (Squamata; Teiidae) from Coastal Rio de Janeiro State, Southeastern Brazil CARLOS FREDERICO D. ROCHA, ALEXANDRE F. BAMBERG ARAU´JO, DAVOR VRCIBRADIC, ELIZABETH MARIA MAMEDE DA COSTA
AND
A new lizard species of the genus Cnemidophorus (Teiidae) is described from Barra de Marica´, Rio de Janeiro State, southeastern Brazil. The species is characterized by having a narrow pale vertebral stripe, with two irregular narrow stripes [one dorsolateral (sometimes broken) and one lateral] on each side of it, and a sequence of small pale dashes on each side of the dorsal field. Its tail is bright blue-green, in contrast to the brownish tail of C. ocellifer. The new species has higher counts of femoral pores on each side (12–20) than the closely related species C. ocellifer (7– 12) and C. nativo (11–13) and differs further from those species in other morphometric and meristic characters, such as the number of subdigital lamellae on fingers and toes and transversal and longitudinal counts of ventral scale rows. Uma nova espe´cie de lagarto do geˆnero Cnemidophorus (Teiidae) e´ descrita de Barra de Marica´, no Estado do Rio de Janeiro, sudeste do Brasil. A espe´cie e´ caracterizada por uma listra vertebral clara e estreita e duas listras irregulares (uma dorsolateral [que pode ser interrompida] e uma lateral) de cada lado da mesma, com uma sequeˆncia de pequenos pontos claros de cada lado do campo dorsal. Sua cauda e´ verde-azulada, contrastando com a cauda marrom de C. ocellifer. A nova espe´cie possui mais poros femorais de cada lado (12–20) do que as espe´cies pro´ximas C. ocellifer (7–12) e C. nativo (11–13), e difere destas tambe´m em outros caracteres merı´sticos, como o nu´mero de lamelas subdigitais dos dedos e artelhos e as contagens transversais e longitudinais de escamas ventrais.
HE teiid genus Cnemidophorus is widely distributed in North, Central, and South America and the Caribbean and is the most speciose genus in the family. Maslin and Secoy (1986) listed 46 species for the genus, and Wright (1993) listed 45. Since then, five new species (four of them unisexual) have been described (Cole and Dessauer, 1993; Markezich et al., 1997; Rocha et al., 1997; Walker et al., 1997). Despite the high number of species in this genus occuring in North America (Wright, 1993), there are comparatively few species presently recognized in South America, all belonging to the lemniscatus group (sensu Wright, 1993). In Brazil, six species are currently recognized: the bisexual C. lemniscatus and C. gramivagus (but ´ vila-Pires, 1995) and the unisexual C. crypsee A tus, of the C. lemniscatus complex of the Amazon basin and Guianan region (Cole and Dessauer, ´ vila-Pires, 1995); the bisexual C. lacerto1993; A ides, found in southern Brazil and adjacent countries (Peters and Donoso-Barros, 1986; Lema, 1994; Feltrim, 1999); the unisexual C. nativo, a species of the C. ocellifer complex recently described from the northern coastal region of Espı´rito Santo State in southeastern Brazil (Rocha et al., 1997); and the bisexual C. ocellifer, which is widely distributed in South America,
T
south of Amazonia (Burt, 1931; Vanzolini et al., 1980; Peters and Donoso-Barros, 1986). Throughout its range, C. ocellifer is reported to occur over a wide range of habitats including the Cerrado (savannah-like vegetation of Central Brazil; Vitt, 1991; Arau´jo, 1994), the Caatinga (semiarid environments of northeastern Brazil; Vanzolini et al., 1980; Vitt, 1983, 1995), and the ‘‘restingas’’ (coastal dry sand dune habitats; Arau´jo, 1991; Rocha and Bergallo, 1997). However, the forms currently under the name C. ocellifer vary considerably in size, coloration, and morphological and meristic characters, which suggests that what is presently referred to as C. ocellifer may represent a complex of species (Rodrigues, 1987; Rocha et al., 1997). Recognition of nearly all Cnemidophorus populations south of the Amazon forest as a single taxon, C. ocellifer, probably reflects more the lack of a detailed revision of the forms of this genus in South America than the occurence of a single widely distributed species. At the restinga habitat of Barra de Marica´, in Rio de Janeiro State, southeastern Brazil, occurs a bisexual species of Cnemidophorus (Fig. 1) belonging to the ocellifer complex. The ocellifer complex is distinguished from the lemniscatus complex by the absence of preanal spurs in
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Fig. 1. Aspect of adult Cnemidophorus littoralis n. sp. in life.
males and by lower counts of femoral pores (see ´ vila-Pires, 1995), and from the lacertoides comA plex by the presence of supraorbital granular semicircles and by tongue shape (bilobed in the base) and scalation (lingual scales subimbricate; see Cei, 1993); the above species is referred to the ocellifer complex based on the presence of those characters. This species nevertheless differs markedly from the two previously recognized species of the ocellifer complex (C. ocellifer
Fig. 2. Comparison of color patterns among Cnemidophorus littoralis n. sp. (center), MNRJ 6620 (SVL 61 mm, adult male), Cnemidophorus ocellifer (left), MZUSP 26827 (SVL 58 mm, adult female), and the unisexual Cnemidophorus nativo (right), MNRJ 4736 (SVL 57 mm, adult female).
and C. nativo) in color pattern (Fig. 2), body proportions, scalation, number of femoral pores and other morphological features (Tables 1–2). Herein, we describe this population as a new species. MATERIAL
AND
METHODS
The new species was found in the restinga of Barra de Marica´, located approximately 38 km east of Rio de Janeiro City, Rio de Janeiro State, southeastern Brazil. Detailed descriptions of the area are given elsewhere (e.g., Vallejo and Vallejo, 1981; Arau´jo, 1991). Restingas are Quaternary coastal sand-dune habitats covered with herbaceous and shrubby xerophilous vegetation, belonging to the Atlantic Rainforest biome (Suguio and Tessler, 1984; Eiten, 1992). We performed meristic and morphometric analyses on 120 specimens from the type locality, after fixation in 10% formalin and preservation in 70% ethanol. Coloration in life was recorded for all specimens, and other characters were investigated after specimens were preserved. Meristic characters were always counted on the left side of the body (except for femoral pores, counted on both sides), under a stereomicroscope, and included the following: femoral pores (FP), ventral scales (V), subdigital lamellae under fourth finger (SLF), subdigital la-
HW/HH (%)
HL/HW (%)
HL/SVL(%)
F⫹TL (mm)
H⫹FiL (mm)
HiL (mm)
FL (mm)
HH (mm)
S-ATB (mm)
HW (mm)
HL (mm)
SVL (mm)
Characters
IN
65.4 62.6 18.6 16.9 10.4 9.1 19.5 17.7 8.4 7.5 13.6 12.5 22.4 20.9 11.4 10.7 26.1 24.7 0.283 0.273 1.802 1.880 1.231 1.206
⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ 9.5 7.8 2.6 1.7 2.1 1.1 2.6 1.7 1.5 0.8 1.9 1.7 3.4 2.4 1.4 1.1 2.9 2.1 0.013 0.016 0.141 0.118 0.078 0.074
C. littoralis
AS
77 43 74 42 76 43 76 43 77 43 77 43 77 43 76 43 77 43 74 42 74 42 76 43
n
55.1 52.2 15.5 13.7 8.8 7.4 16.5 14.3 7.7 6.6 10.8 9.7 17.6 15.8 9.5 8.6 20.3 18.4 0.276 0.264 1.722 1.844 1.152 1.136
⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ ⫾ 7.2 7.4 2.7 1.3 1.3 0.8 2.4 1.4 1.2 0.8 1.1 1.0 1.8 2.0 1.0 1.1 1.9 1.7 0.008 0.016 0.069 0.065 0.046 0.049
C. ocellifer
(45.6–66.0) (39.8–61.8) (13.0–18.7) (11.5–15.4) (7.3–11.0) (6.1–8.4) (13.6–20.7) (11.9–16.2) (6.3–9.5) (6.1–8.4) (9.4–12.4) (7.7–11.0) (15.1–20.0) (12.3–18.7) (8.2–11.1) (7.0–10.5) (18.0–23.4) (15.7–20.7) (0.26–0.29) (0.24–0.30) (1.64–1.84) (1.70–1.93) (1.08–1.21) (1.04–1.23)
MEAN ⫾ SD (FOLLOWED BY RANGE IN PARENTHESES) COMPARED TO C. ocellifer AND C. nativo.
(39.8–81.8) (43.9–74.8) (13.0–26.0) (13.6–20.6) (6.4–15.3) (6.9–11.7) (13.8–24.4) (13.7–20.5) (4.4–11.7) (5.8–9.7) (8.5–17.6) (8.9–15.8) (13.5–27.6) (14.8–24.4) (7.5–14.0) (7.9–12.3) (16.7–30.0) (19.2–27.7) (0.26–0.32) (0.25–0.31) (1.45–2.13) (1.63–2.13) (1.02–1.52) (1.07–1.44)
MORPHOMETRIC CHARACTERS GIVEN
Males Females Males Females Males Females Males Females Males Females Males Females Males Females Males Females Males Females Males Females Males Females Males Females
TABLE 1. VARIATION
9 18 8 18 9 18 8 18 9 18 8 14 9 18 8 17 9 18 8 18 8 18 9 18
n
OF
AND
C. nativo
— (47.4–69.5) — (12.4–17.6) — (6.6–9.8) — (13.6–18.2) — (5.2–7.6) — (9.4–12.5) — (15.1–21.6) — (8.0–10.2) — (17.3–23.5) — (0.24–0.29) — (1.63–1.95) — (1.13–1.55)
— 36 — 35 — 35 — 26 — 33 — 26 — 25 — 26 — 26 — 35 — 35 — 33
n
FEMALE Cnemidophorus littoralis N.
— 54.8 ⫾ 5.0 — 14.4 ⫾ 1.1 — 8.0 ⫾ 0.7 — 15.3 ⫾ 1.2 — 6.3 ⫾ 0.6 — 10.8 ⫾ 0.8 — 18.4 ⫾ 1.5 — 9.3 ⫾ 0.7 — 21.2 ⫾ 1.5 — 0.264 ⫾ 0.011 — 1.814 ⫾ 0.074 — 1.258 ⫾ 0.090
MALE
SP.,
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TABLE 2. VARIATION IN MERISTIC CHARACTERS GIVEN AS MODE VALUE (FOLLOWED BY RANGES IN PARENTHESIS) OF Cnemidophorus littoralis N. SP., COMPARED TO C. ocellifer AND C. nativo. % FD ⫽ percentage of individuals with a divided frontonasal. Characters
C. littoralis
n
FP (on left) FP (on right) V (transverse rows) V (longitudinal rows) SLF SLT SO % FD
16 (11–19) 17 (14–20) 34 (30–39) 10 (9–11) 19 (16–21) 33 (28–39) 4 (4) 35.6
119 120 118 118 119 120 120 118
mellae under fourth toe (SLT), supraocular scales (SO). The following measurements were taken to the nearest 0.1 mm with digital calipers: snout–vent length (SVL), head length (HL), head width (HW), distance from snout to anterior tympanum border (S-ATB), head height (HH), front leg length (FL), hind leg length (HiL), hand plus finger length (fourth finger; H ⫹ FiL) and foot plus toe length (fourth toe; F ⫹ TL). Based on all the above characters, we compared our specimens with the type series of C. nativo (at the Museu Nacional do Rio de Janeiro; MNRJ) and with 28
C. ocellifer
8 8 29 8 16 28 4 0
(7–9) (7–10) (27–30) (8) (15–18) (26–31) (4)
n
28 27 28 28 28 28 27 28
C. nativo
12 12 31 8 16 31 4 0
(11–13) (11–13) (29–32) (8) (16–17) and 32 (29–33) (4)
n
34 34 34 34 35 35 35 35
specimens of C. ocellifer (all from the collection of the University of Sa˜o Paulo Museum of Zoology; MZUSP) from the region of Salvador, in Bahia State, the type locality of that species [we decided to use only topotypes because of the problematic current situation of the status of C. ocellifer (sensu lato)]. We also used data from the original description of C. nativo by Rocha et al. (1997) and from descriptions of C. ocellifer given in Vanzolini et al. (1980) and Cei (1993) in our comparisons with the new species. Terminology of Cnemidophorus color patterns throughout the work follows Walker et al. (1997). The type series of our new species was deposited at the Museu Nacional do Rio de Janeiro (MNRJ) and on the Colec¸a˜o Herpetolo´gica of the Universidade de Brası´lia (CHUNB). Cnemidophorus littoralis n. sp. Figures 1–3 Cnemidophorus ocellifer: Arau´jo, 1991; Ribas et al., 1995; Teixeira-Filho et al., 1995; Rocha and Bergallo, 1997 Holotype.—MNRJ 6536, an adult male collected on 5 February 1995 at the restinga of Barra de Marica´ (22⬚57⬘S, 43⬚50⬘W), municipality of Marica´, Rio de Janeiro State, by Carlos Frederico Duarte Rocha. Paratopotypes.—All from the same locality, collected between January 1989 and April 1992 by C. F. D. Rocha and various coworkers: MNRJ 6537-6641; CHUNB 03230; 03232; 03235-6; 03238; 03241; 03245; 03249; 03251; 03254; 03257; 03259.
Fig. 3. Holotype of Cnemidophorus littoralis (MNRJ 6536): lateral, dorsal and ventral views of head.
Diagnosis.—A moderate-sized Cnemidophorus species (up to 82 mm SVL in adult males and 75 mm SVL in adult females), bisexual (only fe-
ROCHA ET AL.—NEW CNEMIDOPHORUS FROM BRAZIL males in C. nativo), characterized by a narrow, straight, white middorsal (vertebral) line (a pair of diffuse, wavy, thin white paravertebral lines in C. ocellifer; a broad light salmon stripe, straight from nape to midbody and changing to wavy from midbody to tail base in C. nativo; Fig. 2); an irregular (sometimes broken) dorsolateral white stripe on either side of (and as narrow as) the vertebral line (two diffuse white lines in C. ocellifer; stripes broader, vivid white and well defined in C. nativo; Fig. 2); between the vertebral and the dorsolateral lines, the dorsal field is black (in juveniles and young adults) or gray (in large adults), with a longitudinal sequence of white dashes or spots which can sometimes give the appearance of a dashed line [no dorsal dashes or spots in the dorsal fields of either C. ocellifer or C. nativo (but C. ocellifer sometimes with indefinite, subtle spots); Fig. 2]; one irregular white lateral stripe on each side of body, and of the same width as the dorsolateral ones (similar to C. ocellifer; lateral stripes broad and straight in C. nativo); between the dorsolateral and lateral lines, the field is solid black, sometimes with one to three tiny white dots on the posterior third of the body (a row of rounded white or pale blue spots in C. ocellifer; dark field solid black in C. nativo; Fig. 2); tail predominantly bright blue-green (brown to olivaceous in C. ocellifer; also blue-green in C. nativo); dorsal surface of hind limbs black mottled with blue-green (brown with black mottling in C. ocellifer; gray to olive-brown with black freckling in C. nativo); supraocular granules not extending beyond the posterior half of the second supraocular, with that scale always in contact with frontal (second supraocular totally separated from frontal by supraocular granules in C. ocellifer; granules also not extending beyond the posterior half of the second supraocular in C. nativo); nostril usually in suture between anterior and posterior nasal plates (same as in C. ocellifer; nostril completely inside anterior nasal in C. nativo); frontonasal frequently divided [frontonasal sometimes divided in C. ocellifer (undivided in all specimens examined from Salvador) and consistently not divided in C. nativo]; 12–20 (usually 16 or 17) femoral pores along each thigh (7–12 in C. ocellifer and 11–13 in C. nativo); usually 10 longitudinal rows of ventral scales (always eight in C. ocellifer and C. nativo); 30–39 transverse rows of ventral scales (29–32 in C. nativo; 27–30 among examined specimens of C. ocellifer). Description of holotype.—Adult male. Measurements (in mm): SVL: 74.5; HL: 20.7; HH: 10.2; TL: 143.8 (presenting evidence of regeneration
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at the point 95.5 mm from the tail base). Snout moderately pointed; rostral large, clearly visible from above, about as high as wide, separated from frontonasal by anterior nasals; nostril within anterior nasal plate, bordering posteriorly suture between anterior and posterior nasal plates, lateroposteriorly disposed, almost contacting suture with first supralabial inferiorly; frontonasal divided by oblique suture; prefrontals roughly trapezoidal, in broad contact; frontal approximately pentagonal, longer than wide and widest anteriorly, contacting first and second supraoculars; frontoparietals 2; parietals 4 (one on left side of interparietal and two on right side, the distal one considerably smaller); a concentration of irregular scales, subequal in size, behind parietals; supraoculars 4-4, posteriormost much smaller than others; supraciliaries 7-8, the anteriormost two approximately twice as long as wide, the third two-and-a-half times longer than wide, the fourth approximately oneand-a-half times longer than wide, and the fifth approximately as long as wide; only the anteriormost supraciliary in contact with supraocular, all others separated by a row of granular scales; first supraocular and anterior half of second in contact with frontal plate; supraocular granules extending to middle of second supraocular, isolating the posterior half of that scale from frontal plate and totally separating the third and fourth supraoculars from the frontoparietals; granules around supraocular plates 48 on left side, 47 on right; loreal single, large; preocular single, keeled, not in contact with labials; suboculars 3–4, anteriormost keeled, approximately pentagonal, in contact with fifth supralabial; second subocular keeled, longer than others, approximately rectangular, four times as long as high, in contact with fifth, sixth and seventh supralabials; third subocular smooth, approximately rectangular, three times longer than high (divided on left side of head); keels of preocular and first and second suboculars forming a continuous keel; supralabials 7-7; ear opening large, semicircular, higher than wide; mental anteriorly ellipsoid, posteriorly contacting first infralabial and postmental; postmental single, pentagonal, contacting first and second infralabials; infralabials 4-4; 5 pairs of chinshields in divergent rows, anteriormost pair in contact medially; only anteriormost pair of chinshields contacting infralabials, the others separated by a row of small scales which increase in size progressively from second to fifth chinshield; a field of small, circular to ovoid, smooth scales between chinshields, increasing in size progressively until middle of gular region; anterior and posterior gulars differentiated in size,
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with posterior gulars considerably smaller; a field of enlarged scales occurs in the posterior gular region. Scales on nape and sides of neck similar to dorsals; 188 dorsal scales along a middosal line from nape to base of tail; dorsal and flank scales similar, rounded, convex, smooth; flank scales conspicuously differentiated from ventrals resulting in a clear differentiation zone between the two regions; ventrals large, smooth, rectangular, wider than long, in 35 transverse rows and 10 longitudinal rows; preanal plate with large, octogonal median scale, in contact with two moderately large scales anteriorly and posteriorly, surrounded laterally by small scales. Antebrachial scales in 1–3 longitudinal rows of eight scales, scales of lateral row larger and approximately hexagonal to rectangular, scales of medial row of comparatively smaller size, mostly trapezoidal; scales on anteroventral surface of hind limb medium to large, smooth, hexagonal to trapezoidal; on anteroventral surface of thigh, seven transverse scale rows, each one with 3–4 scales, increasing in size distally; on anteroventral surface of tibia, two scale rows of six large trapezoidal tibials; femoral pores 16 on right side, 17 on left; five fingers on each hand; five toes on each foot; scales on dorsum of phalanges rectangular, single, wider than long; subdigital lamellae under fourth finger 18, subdigital lamellae under fourth toe 31; palmar scales round to oval, some few with central pits; plantar scales round to oval, granular. Scales on base of tail trapezoidal to rectangular, longer than wide, in transverse and longitudinal rows, scales smooth until approximately 20th transversal row from border of cloaca; remaining scales on dorsal and lateral surfaces of tail rectangular, longer than wide and keeled; scales in 110 transverse rows along the tail (57 along original portion and 53 along regenerated portion). Color in life.—Dorsal surface of head olive-gray. Dorsal field of body predominantly gray with a narrow white vertebral stripe (extending from nape to tail base, where it broadens and continues along the entire dorsal surface of tail) and one equally narrow white dorsolateral stripe on either side of it, each extending from upper edge of tympanum to tail base. Between the vertebral and each of the dorsolateral stripes, the dorsal field is maculated by a sequence of short, white dashes. One lateral stripe on each side of body, originating from suborbital region and extending to groin, and continuing behind hindlimb into first third of tail. Ground color between dorsolateral and lateral stripes (i.e., lateral field) solid black. Lower flanks (i.e., body field below lateral stripe) light gray, with faint,
diffuse, dashed pale stripe. Ventral surface of head, body and tail predominantly light blue or white, but distal ventral scales, anteroventral surface of thigh, tibia and dorsum of foot deep blue. Dorsal surface of front limbs olive-gray. Dorsal surface of hind limbs black mottled with blue-green. Dorsal surface of tail with bright blue-green middorsal line (actually, the extension of the vertebral white line) between two irregularly bordered black stripes; beyond these, on each side, are two diffuse lateral dark stripes; between the dark stripes the field is bright blue-green. Color in preservative (70% ETOH).—Blue-green portions faded to pale gray or pale blue. Some portions of deep blue coloration on distal ventral scales, surface of thigh and tibia may remain unchanged for some years. Coloration on other portions similar to that in life. Variation.—Morphometric and meristic data for the type series of Cnemidophorus littoralis (and for examined specimens of C. ocellifer and C. nativo for comparison) are presented in Tables 1 and 2, respectively. There are no apparent sexual differences in coloration, though in reproductively active adult males the ventral surfaces of the head and body may be predominantly orange-red or dark red (most of which fades in preservative). The dorsolateral stripes are usually continuous, but may be partially dashed in some individuals (see Fig. 2). In some individuals the supraocular granules do not reach the second supraocular, leaving that scale wholly in contact with the frontal. Males appear to have a divided frontonasal more frequently than females: of 76 males examined, 36 (47.4%) had the frontonasal divided, but such condition was present in only five of 41 females (12.2%). Number of parietal scales vary from three to five, with five being the usual. Etymology.—The name littoralis (inhabitant of coastal area, in Latin), a noun in apposition, refers to the presence of the species only in coastal sand dune habitats. Distribution.—Known only from the type locality and from the region of Carapebus Beach (22⬚17⬘S, 41⬚41⬘W; see Discussion), located some 10 km from Macae´ city, in the state of Rio de Janeiro, southeastern Brazil (Fig. 4). Ecological remarks.—Cnemidophorus littoralis is considerably abundant at the restinga habitat of Barra de Marica´, which is characterized by a xeromorphic vegetation, present mainly in dense
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locality include three nematodes (the commonest being the Physalopterid Physaloptera retusa; Ribas et al., 1995) and one cestode (Oochoristica ameivae; C. R. Bursey, unpubl. data). DISCUSSION
Fig. 4. Map of Rio de Janeiro State, Brazil, indicating the localities (stars) at which Cnemidophorus littoralis is known to occur.
or sparse shrubs separated from each other by wide stretches of open sand. Plants of the families Erythroxylaceae, Gramineae, Leguminoseae, Bromeliaceae, and Clusiaceae predominate. At this restinga, the preferred microhabitat of active C. littoralis is the edge of shrubs, where it forages and basks on the leaf litter or on adjacent open sand (Teixeira-Filho et al., 1995). In a series of 120 individuals from Barra de Marica´, we found a sex ratio biased to male (1.79:1). It is a diurnal species, being typically active between 0800 h and 1400 h, and mostly from 1000 h to 1200 h (Teixeira-Filho et al., 1995). Like other congeners, C. littoralis is a heliothermic lizard species that basks in the sun and forages actively. Mean (⫾ 1 SD) body temperature taken from 80 individuals by TeixeiraFilho et al. (1995) at Marica´ was 38.7 ⫾ 2.0 C. Stomach content analysis of 50 individuals from Marica´ indicated that the diet is composed of arthropods and that termites, present in 94% of the stomachs, are the predominant food items (96.2% by number; 67.7% by volume), followed by insect larvae (mainly Coleopteran and Lepidopteran; 2.0% by number; 16.7% by volume) and hemipterans (0.7% by number; 2.6% by volume), although blattarians, spiders, ants, adult coleopterans, chilopods, homopterans, thysanurans, pseudoscorpionids, and one fruit were also found in the diet. Arau´jo (1991) also found termites and larvae to be the most important items in the diet of this species, based on a sample of 30 individuals from the same area. Arau´jo (1985) observed predation on this species by the colubrid snake Liophis miliaris at Barra de Marica´. Endoparasites of this species at the type
The recognition of Cnemidophorus littoralis as a new taxon, together with the recent description of C. nativo (Rocha et al., 1997), illustrates the need for a revision of C. ocellifer (sensu lato) throughout its extensive distribution. Rodrigues (1987:224) suspected that what was called C. ocellifer actually comprised several distinct species and cited specimens he considered markedly different from C. ocellifer (sensu Spix, 1825) in coloration, scalation and number of femoral pores. One of those specimens, at the collection of the University of Sa˜o Paulo Museum of Zoology (MZUSP), had been collected at an unspecified coastal sand-dune area of Rio de Janeiro State. This specimen is almost certainly MZUSP 54904 (from Carapebus Beach, Rio de Janeiro State, collected by P. E. Vanzolini on 14 Dec. 1979; labeled as C. ocellifer), since it is the only specimen of Cnemidophorus in that collection from Rio de Janeiro State, as verified by one of us (DV) during a recent visit to that museum. The specimen, an adult male, agrees with our description of C. littoralis in both color pattern and meristic characters and is therefore assigned to this species. Rodrigues (1987) further speculated that the speciation of the mainland South American Cnemidophorus may have been concomitant (and underlain by similar phenomena of microgeographic isolation) with that of the Tropidurus torquatus group (which was itself partitioned into 13 different species after his revision; see also Rodrigues, 1988). Cnemidophorus littoralis is the second species of the genus known from coastal southeastern Brazil (the other is C. nativo, which also has a bluegreen tail) and, apparently, the only one found in the state of Rio de Janeiro. We base this assumption on the results of Rocha and Bergallo’s (1997) extensive field survey of lizard fauna from southeast Brazilian restingas (which, incidentally, did not include the Macae´ area): those authors failed to find any Cnemidophorus in Espirito Santo state south of the Doce River (which lies more than 200 km north of the border with Rio de Janeiro state), whereas C. nativo (CFDR, pers. obs.) was present north of that river and south of Salvador. Other species of reptiles and amphibians endemic to the restingas of Rio de Janeiro include the lizard Liolaemus lutzae (Tropiduridae) and the frogs Leptodactylus
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marambaiae (Leptodactylidae) and Scinax littoreus (Hylidae). Females of C. littoralis grow larger than individuals of the all-female species C. nativo (see Table 1). Both male and female C. littoralis were also larger than topotypic C. ocellifer examined by us, but the large proportion of young specimens in that sample (13 of 28 individuals were smaller than 55 mm SVL) weakens the reliability of the comparison. Indeed, specimens of C. ocellifer from other localities in northeastern Brazil grow even larger than C. littoralis (Vanzolini et al., 1980; Vitt, 1983, 1995), but it must be cautioned that, as already pointed out, a careful revision of this species complex is needed to assess whether such populations are conspecific with those from Salvador. Several other Brazilian species of Cnemidophorus south of the Amazon region are apparently awaiting formal description. Among them are two populations from the northeast (one montane and one coastal) mentioned by Rodrigues (1987:208, 224). The author also mentions a second coastal population (p. 224), but that one most probably represents C. nativo, judging by the area of occurrence (see Rocha et al., 1999). Other populations representing still unnamed species include that of a cerrado area of southern Rondo ˆ nia state (Vitt and Caldwell, 1993), a stripeless form from a dry caatinga locality in Piauı´ state (P. R. Manzani, pers. comm.), and a highland form from the southern state of Rio Grande do Sul (Lema, 1994; Feltrim, 1999). There are also other mainland South American species (most of which had been recently described or revalidated) that are not known from Brazil (i.e., C. longicaudus, C. serranus, and C. leachi from Argentina, C. arenivagus from northernmost Venezuela and Colombia, and the unisexual C. pseudolemniscatus from Suriname and French Guiana; Cei, 1993; Cole and Dessauer, 1993; Markezich et al., 1997). We find it likely that additional research will eventually prove that the genus Cnemidophorus is considerably more speciose in South America than was previously thought. SPECIMENS EXAMINED Cnemidophorus littoralis.—MZUSP 54904 (Carapebus, Rio de Janeiro). Cnemidophorus nativo.—MNRJ 4698 (holotype); MNRJ 4699–4737 (paratopotypes, Linhares, Espı´rito Santo). Cnemidophorus ocellifer.—MZUSP 191; 195–6; 201; 203; 205–206; 6150; 6155; 8468; 26827–43; 65762 (topotypes, Salvador, Bahia).
ACKNOWLEDGMENTS This study represents part of the results of the ‘‘Programa de Ecologia, Conservac¸a˜o e Manejo de Ecossistemas do Sudeste Brasileiro’’ and the Southeastern Brazilian Vertebrate Ecology Project (Vertebrate Ecology Laboratory), both of the Setor de Ecologia, Instituto de Biologia, Universidade do Estado do Rio de Janeiro. We thank P. F. Teixeira-Filho, L. F. L. Fonseca, L. N. B. Martins, M. Cunha-Barros, H. L. T. Zaluar, S. R. Silveira, and S. C. Ribas for their help in collecting the lizards. We also thank M. G. N. Oliveira for laboratorial assistantship. M. Van Sluys, H. G. Bergallo, J. P. Pombal Jr., U. Caramaschi, L. L. Grismer, J. J. Wiens, C. J. Cole, and T. C. Avila-Pires reviewed the manuscript and gave valuable suggestions. We also thank M. T. Rodrigues and P. E. Vanzolini for allowing examination of material housed at the University of Sa˜o Paulo Museum of Zoology (MZUSP). This study was partially supported by the Conselho Nacional de Desenvolvimento Cientı´fico e Tecnolo´gico (CNPq) through grants to CFDR (Process 300819/94-3 and 400115/97-2) and to DV (Process 143607/98-7). We thank the Universidade do Estado do Rio de Janeiro for travel support to Barra de Marica´. LITERATURE CITED ARAU´JO, A. F. B. 1985. Partilha de recursos em uma guilda de lagartos de restinga (Sauria). Unpubl. master’s thesis. Dpto. de Zoologia, Universidade Estadual de Campinas. . 1991. Structure of a white sand-dune lizard community of coastal Brazil. Rev. Brasil. Biol. 51: 857–865. . 1994. Comunidades de lagartos brasileiros, p. 39–57. In: Herpetologia do Brasil. I. A. T. Bernardes, L. B. Nascimento, and G. A. Cotta (eds.). Pontifı´cia Universidade Cato´lica de Minas Gerais, Belo Horizonte, Brasil. A´VILA-PIRES, T. C. S. 1995. Lizards of Brazilian Amazonia (Reptilia: Squamata). Zool. Verh. Leiden 299: 1–706. BURT, C. E. 1931. A study of the teiid lizards of the genus Cnemidophorus with special reference to their phylogenetic relationships. Bull. U.S. Natl. Mus. 154:viii ⫹ 1–286. CEI, J. M. 1993. Reptiles del Noroeste, Nordeste y Este de la Argentina. Herpetofauna de las Selvas Subtropicales, Puna y Pampas. Monogr. XIV, Mus. Reg. Sci. Nat., Torino. COLE, C. J., AND H. C. DESSAUER. 1993. Unisexual and bisexual whiptail lizards of the Cnemidophorus lemniscatus complex (Squamata: Teiidae) of the Guiana region, South America, with descriptions of new species. Am. Mus. Novit. 3081:1–30. EITEN, G. 1992. Natural Brazilian vegetation types
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(CFDR,DV) SETOR DE ECOLOGIA, DEPARTAMENTO DE BIOLOGIA ANIMAL E VEGETAL, INSTITUTO DE BIOLOGIA, UNIVERSIDADE DO ESTADO DO RIO DE JANEIRO, 20550-011, RUA SA˜O FRANCIS˜ , RIO DE JANEIRO, CO XAVIER 524, MARACANA RIO DE JANEIRO, BRAZIL; AND (AFBA,EMMC) DEPARTAMENTO DE ECOLOGIA, INSTITUTO DE BIOLOGIA, UNIVERSIDADE DE BRAS´ıLIA, CAMPUS ASA NORTE, BRAS´ıLIA, D.F., BRAZIL. E-mail: (CFDR)
[email protected]. Submitted: 23 April 1999. Accepted: 12 Oct. 1999. Section editor: A. H. Price.
