Three New Species of theTropidurus spinulosusGroup (Squamata: Tropiduridae) from Eastern Paraguay

2016·0.79 MB·English

by  André Luiz G. Carvalho

A M ERIC AN MUSEUM NOVITATES
Number 3853, 44 pp. February 20, 2016
Three new species of the Tropidurus spinulosus group
(Squamata: Tropiduridae) from Eastern Paraguay
ANDRÉ LUIZ G. CARVALHO1
ABSTRACT
Tropidurus Wied, 1825, is one of the most ubiquitous lizard genera endemic to South
America. Herpetologists from different regions of the continent have progressively mapped new
populations, including undescribed species hidden under widely distributed nominal taxa. Cur-
rently, four monophyletic species groups are recognized in Tropidurus (T. bogerti group [mono-
typic], T. semitaeniatus group [four species], T. spinulosus group [five species], and T. torquatus
group [16 species]), but none have been comprehensively revised taxonomically. During a col-
lection expedition carried out in Paraguay in 2013, I recognized three new, distinct morphot-
ypes among populations of the Tropidurus spinulosus group formerly assigned to T. guarani
Alvarez et al., 1994. To delimit these new taxa, I analyzed coloration patterns, and quantified
meristic and morphometric variables, comparing freshly collected samples with specimens
housed in five museum collections. In this paper, I describe and illustrate the allopatric T.
lagunablanca, n. sp., T. tarara, n. sp., and T. teyumirim, n. sp., and provide notes on their dis-
tribution limits, natural history, and conservation status.
INTRODUCTION
The Tropidurus spinulosus species group was formally delimited by Frost et al. (2001) as
one of the four main clades constituting the South American lizard genus Tropidurus Wied,
1825. Currently, it includes five nominal species—T. callathelys Harvey and Gutberlet, 1998, T.
guarani Alvarez et al., 1994, T. melanopleurus Boulenger, 1902, T. spinulosus (Cope, 1862), and
1 Richard Gilder Graduate School and Division of Vertebrate Zoology (Herpetology), American Museum of
Natural History.
Copyright © American Museum of Natural History 2016 ISSN 0003-0082
2 AMERICAN MUSEUM NOVITATES NO. 3853
T. xanthochilus Harvey and Gutberlet, 1998—inhabiting open environments of the cis-Andean
tropical and subtropical South America, between 12° and 32° S. All together, their ranges
encompass territories within five countries, including areas in northern Argentina (T. spinulo-
sus and T. melanopleurus), eastern Bolivia (T. callathelys, T. melanopleurus, T. spinulosus, and
T. xanthochilus), central Brazil (T. aff. guarani and T. callathelys), eastern and western Paraguay
(T. guarani and T. spinulosus), and the southernmost part of Peru (T. melanopleurus); see Car-
valho (2013) for an updated review of the distribution of Tropidurus species.
In contrast with the Tropidurus torquatus and T. semitaeniatus species groups,2 whose taxo-
nomic diversity has been progressively expanded by recent morphological and molecular stud-
ies (Passos et al., 2011; Kunz and Borges-Martins, 2013; Werneck et al., 2015; Carvalho et al.,
2016), species diversity in the T. spinulosus group remain unrevised for almost two decades
(Alvarez et al., 1994; Frost et al., 1998; Harvey and Gutberlet, 1998). In Paraguay, Frost et al.
(1998) conducted the most comprehensive taxonomic investigation focused on species of the
T. spinulosus group to date,3 and detected high levels of morphological and molecular diver-
gence among populations of the then subspecies T. spinulosus guarani Alvarez et al., 1994, and
T. spinulosus spinulosus (Cope, 1862). That study also gathered compelling phylogenetic evi-
dence supporting the existence of at least two diagnosable evolutionary lineages east and west
of the Paraguay River. Consequently, T. s. guarani, distributed in the mesic eastern Paraguayan
region, and T. s. spinulosus, endemic to the xerophilous Chacoan districts west of the Paraguay
River, were elevated to the species rank as T. guarani and T. spinulosus, corroborating prelimi-
nary findings of Alvarez et al. (1994).
Although Frost et al.’s (1998) results did not reject the possibility of independent lineages
coexisting on the eastern side of the Paraguay River, the taxonomic diversity comprised by popu-
lations of the Tropidurus spinulosus group distributed in eastern Paraguay has not been reassessed
since. After analyzing numerous specimens collected during a three-month expedition carried
out in Paraguay between August and October 2013, I was able to distinguish four allopatric
morphotypes among populations previously assigned to T. guarani (Alvarez et al., 1994; Frost et
al., 1998). Of those, three were found to represent new species and the fourth morphotype was
associated to nominal T. guarani (fig. 1). To delimit these new taxa, I examined coloration pat-
terns, and quantified meristic and morphometric variation in a geographic scale, comparing
freshly collected material with specimens housed in five museum collections. In this paper, I
describe and illustrate T. lagunablanca, n. sp., T. tarara, n. sp., and T. teyumirim, n. sp., and pro-
vide notes on their distribution limits, natural history, and conservation status.
2 The Tropidurus bogerti group is monotypic and endemic to the Auyantepui, Venezeuela (Roze, 1958; Frost
et al., 2001; Myers and Donnelly, 2008; Carvalho, 2013).
3 Until Frost et al. (1998), only Tropidurus spinulosus was known from Paraguay, with two subspecies, T. s.
guarani Alvarez et al., 1994, and T. s. spinulosus (Cope, 1862), distributed, respectively, on the eastern and
western sides of the country, separated by the Paraguay River (Alvarez et al., 1994).
2016 CARVALHO: THREE NEW SPECIES OF THE TROPIDURUS SPINULOSUS GROUP 3
FIGURE 1. Distribution map of Tropidurus guarani, T. lagunablanca, n. sp., T. tarara, n. sp., and T. teyumirim,
n. sp. For information on the distribution of additional species of the T. spinulosus species group, refer to
Carvalho (2013).
4 AMERICAN MUSEUM NOVITATES NO. 3853
MATERIAL AND METHODS
Samples: I analyzed 245 specimens representing four (of five) nominal species assigned
to the Tropidurus spinulosus species group by Frost et al. (2001)—namely T. guarani, T. mela-
nopleurus, T. spinulosus, and T. xanthochilus—and extracted data on T. callathelys from Har-
vey and Gutberlet (1998) and Morais et al. (2014). In addition to individuals freshly collected
in numerous localities from Paraguay, Bolivia, and Argentina, my samples included speci-
mens housed in five museum collections. In the field, specimens were collected with the aid
of rubber bands or nooses between 05 August and 13 December 2013, euthanized with an
overdose of 2% lidocaine, preserved with 10% unbuffered formalin, and then transferred to
70% ethyl alcohol solution. All specimens collected and analyzed for this study are housed
in the following museum collections: American Museum of Natural History, New York
(AMNH); Colección Zoológica Para La Tierra, Santa Rosa del Aguaray, Paraguay (CZPLT);
Museo de Historia Natural Alcide d’Orbigny, Cochabamba, Bolivia (MNHNC); Museo
Nacional de Historia Natural del Paraguay, Asunción, Paraguay (MNHNP); and U.S. National
Museum–Smithsonian Institution, Washington D.C. (USNM). A complete list of examined
material is provided in appendix.
External morphology: I adopted the general terminology revised by Carvalho et al.
(2016) for description of external morphological structures. However, because that study dealt
exclusively with species of the Tropidurus torquatus group, I extended their protocol to account
for additional morphological features and scale counts found in species of the T. spinulosus
group. All species in the T. spinulosus group have a midvertebral row of protruding or spinelike
scales forming a dorsal crest that extends from the postoccipital region to the proximal, medial,
or distal portion of the tail. Scales forming the dorsal crest, hereafter referred to as vertebrals,
were counted from the first protruding, or spinelike scale on the postoccipital area, extending
to the level of the anterior margin of hind limb insertion. The dorsal crest is significantly more
conspicuous in males than females (Frost, 1992; Harvey and Gutberlet, 1998). Paravertebrals
correspond to the series of dorsal scales immediately adjacent to vertebrals and were counted
following a line drawn just to the right of the vertebral crest.
Tropidurine lizards have marked skin folds and granular mite pockets distributed in dif-
ferent areas of the body. Frost (1992) identified 10 types of skin folds. Of those, antegular,
antehumeral, dorsolateral, gular, longitudinal neck, oblique neck, postauricular, and rictal folds
were observed during my analysis of specimens of the T. spinulosus group. Frost (1992)
described four types of mite pockets with a number of morphological states, namely antegular-
oblique neck, axillary, antehumeral, and inguinal mite pockets. The antegular-oblique neck
mite pocket is the only one found among species of the T. spinulosus group; refer to morpho-
logical descriptions and figures 22–24 in Frost’s (1992) study for details and schematic repre-
sentations of the location of skin folds and mite pockets. Tufts of spines on the side of the neck
are additional structures shared by species of the T. spinulosus group. These tufts of spines are
usually more pronounced in males and vary ontogenetically, with adults showing bigger tufts
than juveniles and subadults. Their number, size, and morphology also show strong geographic
variation (Frost et al., 1998).
2016 CARVALHO: THREE NEW SPECIES OF THE TROPIDURUS SPINULOSUS GROUP 5
TABLE 1. Mean ± standard deviation, minimum and maximum values of morphometric measurements (in
mm) of adult males.
Abbreviations: SVL, snout–vent length; BH, body height; HH, head height; HW, head width; EOS, ear
opening–snout distance; AL, arm length; FAL, forearm length; HDL, manus length; THL, thigh length; SL,
shank length; FOL, pes length; TL, tail length. The number of measured individuals is followed (between
parentheses) by the number of individuals with fully grown tails. See Material and Methods for details on
treatment of individuals with broken, regrown, or missing tails.
T. guarani N = 5 (3) T. lagunablanca, n. sp. T. tarara, n. sp. T. teyumirim, n. sp.
N = 6 (4) N = 11 (5) N = 37 (18)
SVL 97.94 ± 9.46 104.02 ± 9.37 110.28 ± 8.65 84.34 ± 6.08
(87.73–109.44) (89.35–113.09) (98.04–122.82) (67.82–94.41)
TL 118.35 ± 22.01 146.88 ± 16.44 142.00 ± 13.13 127.23 ± 12.60
(97.05–141.00) (136–171) (124–155) (93.96–144.99)
HH 13.76 ± 1.69 13.91 ± 1.14 14.58 ± 1.68 10.97 ± 0.85
(12.05–16.32) (12.03–15.55) (11.77–17.52) (9.42–12.48)
EOS 22.00 ± 1.27 23.73 ± 2.25 24.70 ± 1.60 19.82 ± 1.19
(20.25–23.40) (20.25–25.95) (22.38–27.18) (16.58–21.68)
HL 27.22 ± 2.67 28.16 ± 2.82 29.65 ± 2.88 23.47 ± 1.42
(24.44–30.55) (24.22–31.11) (25.90–34.42) (19.57–26.56)
HW 20.42 ± 2.00 20.88 ± 2.08 22.43 ± 2.13 17.17 ± 1.16
(18.24–22.85) (17.76–23.86) (18.39–25.40) (14.03–19.14)
AL 16.76 ± 2.38 16.59 ± 2.23 17.06 ± 1.08 13.31 ± 1.21
(14.57–19.84) (13.03–18.94) (15.82–18.87) (10.82–15.34)
FAL 13.29 ± 1.27 14.29 ± 1.59 15.00 ± 1.10 11.43 ± 1.03
(12.13–15.38) (11.96–16.05) (13.39–17.20) (9.08–13.24)
HDL 18.10 ± 1.89 19.42 ± 1.20 20.26 ± 1.32 16.16 ± 0.98
(15.43–19.90) (17.80–20.97) (17.85–22.44) (13.84–17.57)
THL 20.99 ± 2.88 20.95 ± 2.37 22.03 ± 1.94 17.49 ± 1.53
(17.54–24.44) (17.29–23.51) (17.60–24.38) (14.37–20.28)
SL 18.04 ± 1.88 18.60 ± 1.66 18.51 ± 1.68 15.43 ± 1.20
(15.62–20.06) (16.25–20.16) (16.27–20.96) (12.43–17.17)
FOL 27.61 ± 2.66 29.82 ± 2.21 30.19 ± 1.58 24.69 ± 1.29
(25.02–30.66) (26.13–32.47) (26.55–32.05) (20.75–26.40)
Sex determination: Species of the Tropidurus spinulosus group are highly sexually dimor-
phic (Laurent, 1980; Alvarez et al., 1994; Harvey and Gutberlet, 1998). I determined the sex of
the specimens based on the observation of colored patches of glandular scales located on ventral
surface of the thighs and precloacal flap of adult males; adult females lack ventral colored patches
(Frost, 1992; Alvarez et al., 1994). Adult males have wider heads and thinner bodies than females
of the same body size and also have a marked middorsal crest; the dorsal crest is weakly marked
or nearly absent in females (Frost, 1992; Alvarez et al., 1994; Harvey and Gutberlet, 1998). In
several cases, I was unable to determine the sex of juveniles or subadults based on their external
morphology alone. Therefore, in some cases, I inspected gonads to determine sex.
Morphometric and meristic data: Morphometric measurements from the right side of
101 adult males and 111 adult females were taken with aid of a digital caliper (to the nearest
0.1 mm; tables 1–2). I followed the morphometric protocol described in Carvalho et al. (2016),
6 AMERICAN MUSEUM NOVITATES NO. 3853
TABLE 2. Mean ± standard deviation, minimum and maximum values of morphometric measurements (in
mm) of adult females.
Abbreviations: SVL, snout–vent length; BH, body height; HH, head height; HW, head width; EOS, ear
opening–snout distance; AL, arm length; FAL, forearm length; HDL, manus length; THL, thigh length; SL,
shank length; FOL, foot length; TL, tail length. The number of measured individuals is followed (between
parentheses) by the number of individuals with fully grown tails. See Material and Methods for details on
treatment of individuals with broken, regrown, or missing tails.
T. guarani N = 7 (6) T. lagunablanca, n. sp. T. tarara, n. sp. T. teyumirim, n. sp.
N = 2 (1) N = 4 (1) N = 49 (25)
SVL 70.59 ± 7.54 85.03 ± 13.51 88.96 ± 3.41 70.17 ± 4.24
(62.53–86.19) (75.48–94.59) (86.35–93.97) (61.41–80.79)
TL 102.09 ± 11.87 115.74 127.76 106.83 ± 6.73
(89.12–117.56) (92.08–121.95)
HH 9.28 ± 1.60 10.89 ± 2.33 11.77 ± 0.97 8.96 ± 0.53
(7.77–12.70) (9.25–12.54) (10.61–12.89) (7.89–10.10)
EOS 16.12 ± 1.78 18.92 ± 2.29 19.78 ± 0.69 16.54 ± 0.86
(14.69–19.84) (17.30–20.54) (18.82–20.36) (14.82–18.80)
HL 19.40 ± 2.18 22.98 ± 2.70 23.56 ± 1.18 19.47 ± 0.89
(17.45–24.07) (21.07–24.89) (22.72–25.23) (17.20–21.60)
HW 14.07 ± 1.49 16.82 ± 2.38 16.99 ± 1.43 14.01 ± 0.83
(12.59–16.93) (15.14–18.50) (15.4–18.64) (12.19–15.85)
AL 11.18 ± 1.31 12.39 ± 1.18 13.84 ± 0.71 10.87 ± 0.87
(9.96–13.74) (11.56–13.23) (13.24–14.69) (9.01–13.08)
FAL 9.54 ± 1.02 11.42 ± 1.53 12.12 ± 0.75 9.38 ± 0.67
(8.29–11.53) (10.34–12.5) (11.23–12.81) (7.88–10.95)
HDL 13.94 ± 1.56 16.21 ± 0.13 16.46 ± 0.87 13.71 ± 0.88
(12.55–17.22) (16.12–16.30) (15.73–17.48) (11.87–15.68)
THL 14.64 ± 2.01 17.41 ± 1.89 18.19 ± 0.75 14.51 ± 0.83
(13.31–19.06) (16.07–18.74) (17.14–18.85) (12.16–16.24)
SL 12.68 ± 1.11 15.06 ± 1.29 15.19 ± 0.78 12.40 ± 0.71
(11.53–15.03) (14.15–15.97) (14.50–16.00) (10.85–14.16)
FOL 20.71 ± 1.29 23.95 ± 2.01 25.24 ± 0.84 20.34 ± 1.03
(18.44–22.49) (22.53–25.37) (24.35–26.37) (17.92–23.14)
adding the following variables: AGD (armpit to groin distance), from the base of the armpit to
groin; BH (body height), distance between belly and dorsum measured at a point between the
insertion of forelegs and hind legs; BW (body width), distance between flanks measured at a
point between the insertion of forelegs and hind legs; HL (head length), from tip of snout to
posterior limit of the occipital region.
Variations in scale counts among species were investigated to detect taxonomically infor-
mative characters (tables 3–4). See External Morphology and Carvalho et al. (2016) for details
on scale counts (i.e., topography, scale nomenclature). In total, I analyzed 87 male and 61
female specimens for meristic data. I failed to identify consistent correlations between SVL (as
a proxy for age) and scales counts (results not shown), and for that reason I pooled juveniles,
subadults, and adults of each species for meristic comparisons. All statistical procedures were
performed for males and females separately, using the statistical platform R version 3.0.2 (R
Core Team, 2013).
2016 CARVALHO: THREE NEW SPECIES OF THE TROPIDURUS SPINULOSUS GROUP 7
TABLE 3. Mean ± standard deviation, minimum and maximum values of scale counts of male lizards.
Abbreviations: VB, vertebrals; PB, paravertebrals; VT, ventrals; GU, gulars; MD, scales around midbody;
TI, tibials; LF, subdigital lamellae under fourth finger; LT, subdigital lamellae under fourth toe; SL, supral-
abials; IL, infralabials.
T. guarani (N = 7) T. lagunablanca, n. sp. T. tarara, n. sp. T. teyumirim, n. sp.
(N = 10) (N = 18) (N = 18)
VB 60.43 ± 5.53 (51–68) 60.10 ± 2.96 (56–64) 56.61 ± 4.50 (48–65) 70.06 ± 7.33 (60–83)
PB 111.71 ± 6.24 (100–117) 121.60 ± 3.10 (116–125) 111.83 ± 6.27 (97–124) 117.17 ± 5.44 (107–128)
VT 85.71 ± 2.43 (82–89) 84.20 ± 2.97 (78–88) 82.22 ± 4.87 (71–91) 21.50 ± 0.71 (21–22)
GU 73.14 ± 4.78 (65–80) 76.00 ± 2.87 (71–80) 69.94 ± 7.60 (61–92) 75.50 ± 5.27 (67–86)
MB 99.57 ± 4.35 (92–103) 106.10 ± 4.18 (100–113) 99.89 ± 5.85 (92–112) 101.44 ± 3.70 (94–107)
TI 22.29 ± 1.38 (21–24) 25.60 ± 0.97 (24–27) 23.28 ± 2.16 (20–28) 21.22 ± 1.06 (19–23)
LF 20.43 ± 1.27 (19–22) 20.50 ± 1.08 (19–22) 19.94 ± 1.06 (17–22) 20.11 ± 1.23 (18–22)
LT 25.86 ± 0.69 (25–27) 26.30 ± 1.25 (24–28) 24.94 ± 1.43 (23–29) 24.94 ± 1.55 (22–28)
SL 4.57 ± 0.53 (4–5) 5.50 ± 0.53 (5–6) 5.50 ± 0.62 (5–7) 4.72 ± 0.57 (4–6)
IL 5.00 ± 0.00 (5–5) 5.20 ± 0.42 (5–6) 5.72 ± 0.46 (5–6) 4.89 ± 0.58 (4–6)
SPECIES ACCOUNTS
TROPIDURIDAE BELL, 1843
TROPIDURUS WIED, 1825
My analyses revealed three new species of the Tropidurus spinulosus group distributed in
eastern Paraguay. The species treated below may be diagnosed as members of the T. spinulosus
group by having: (1) an enlarged middorsal scale row forming a dorsal crest (absent in other
species groups); (2) yellow, orange, or cream thigh flash marks in the precloacal region and on
ventral surface of the thighs of males (black in males of other species groups); (3) a strong
antegular fold (absent in other species groups); (4) poorly defined postmental scale series (well
defined in other species groups); (5) usually two rows of circumorbital scales (a single row in
other species groups); and (6) body not extremely depressed (primarily found in the T. semi-
taeniatus species group and, more moderately, in T. bogerti).
Tropidurus lagunablanca, n. sp.
Figures 2A–F, 3A–F
Holotype: CZPLT-H 402 (adult male; 23° 48′ 43.20″ S 56° 17′ 40.92″ W, WGS84 system;
~207 m above sea level; geographic coordinates not specified in the CZPLT catalog, estimated
based on field observations of the restricted local distribution of the species in the type locality
in September 2013), collected at Reserva Natural Laguna Blanca, Santa Rosa del Aguaray, San
Pedro, Paraguay.
Paratypes: AMNH-R 176291 (juvenile male), AMNH-R 176292 (adult female), collected at
Reserva Natural Laguna Blanca, Santa Rosa del Aguaray, San Pedro, Paraguay (23° 48′ 43.20″ S
56° 17′ 40.92″ W, WGS84 system; ~207 m above sea level) by André L. Gomes de Carvalho,
8 AMERICAN MUSEUM NOVITATES NO. 3853
TABLE 4. Mean ± standard deviation, minimum and maximum values of scale counts of female lizards.
Abbreviations: VB, vertebrals; PB, paravertebrals; VT, ventrals; GU, gulars; MD, scales around midbody;
TI, tibials; LF, subdigital lamellae under fourth finger; LT, subdigital lamellae under fourth toe; SL, supral-
abials; IL, infralabials.
T. guarani (N = 7) T. lagunablanca, n. sp. T. tarara, n. sp. T. teyumirim, n. sp.
(N = 2) (N = 4) (N = 20)
VB 94.43 ± 3.41 (90–98) 77.00 ± 14.14 (67–87) 79.00 ± 7.12 (69–85) 96.90 ± 8.97 ± (79–112)
PB 113.86 ± 9.60 (94–121) 120.50 ± 3.54 (118–123) 119.25 ± 3.95 (116–124) 118.35 ± 5.51 (109–129)
VT 89.57 ± 2.82 (84–92) 88.50 ± 0.71 (88–89) 80.00 ± 2.45 (77–82) 85.30 ± 4.16 (77–93)
GU 76.43 ± 2.64 (74–81) 75.00 ± 2.83 (73–77) 64.25 ± 5.19 (58–69) 74.10 ± 5.30 (67–88)
MB 103.14 ± 5.52 (98–111) 110.50 ± 4.95 (107–114) 99.00 ± 4.24 (95–104) 103.10 ± 5.25 (94–112)
TI 22.71 ± 1.50 (21–25) 23.50 ± 0.71 (23–24) 21.50 ± 2.65 (19–25) 21.35 ± 1.14 (20–24)
LF 20.00 ± 1.73 (18–23) 20.50 ± 0.71 (20–21) 18.00 ± 1.41 (16–19) 19.55 ± 1.57 (17–22)
LT 25.29 ± 1.50 (23–27) 26.50 ± 0.71 (26–27) 23.00 ± 1.83 (21–25) 24.25 ± 1.68 (22–27)
SL 4.43 ± 0.53 (4–5) 5.00 ± 0.00 (5–5) 5.25 ± 0.50 (5–6) 4.65 ± 0.59 (4–6)
IL 5.00 ± 0.58 (4–6) 5.50 ± 0.71 (5–6) 5.25 ± 0.50 (5–6) 5.00 ± 0.46 (4–6)
Frederick Bauer, Nicolás Martínez, Viviana Espínola, and Marcelo Dujak in 7 September 2013.
AMNH-R 176293 (adult female), collected at Reserva Natural Laguna Blanca, Santa Rosa del
Aguaray, San Pedro, Paraguay (23° 48′ 43.20″ S 56° 17′ 40.92″ W, WGS84 system; ~207 m above
sea level) by André L. Gomes de Carvalho, Rodrigo Ayala, and Johanna López in 29 September
2013. CZPLT-H 157, CZPLT-H 406, CZPLT-H 409, CZPLT-H 461 (adult males), CZPLT-H 405
(juvenile male), collected at Reserva Natural Laguna Blanca, Santa Rosa del Aguaray, San Pedro,
Paraguay. MNHNP 11463 (adult male), collected at Laguna Blanca, San Pedro, Paraguay, by Nor-
man J. Scott in 18 February 2010. MNHNP 11755 (juvenile male), MNHNP 11756 (subadult
male), collected at Reserva Natural Laguna Blanca, Santa Rosa del Aguaray, San Pedro, Paraguay
(23° 48′ 43.20″ S 56° 17′ 40.92″ W, WGS84 system; ~207 m above sea level), by André L. Gomes
de Carvalho, Rodrigo Ayala, and Johanna López in 29 September 2013.
Morphological diagnosis: Tropidurus lagunablanca, n. sp., may be distinguished from all
congeners by the following combination of characteristics: (1) adult large and robust, reaching
113.09 and 94.59 mm SVL in males and females, respectively; (2) iris golden brown; (3) strongly
projecting, serrate vertebral crest in males, reduced in females; (4) 1–2 canthals; (5) preocular not
contacting canthal; (6) two rows of circumorbitals; (7) moderate tufts of neck spines; (8) high num-
ber of scales around midbody (100–113 in males and 107–114 in females); (9) sexes not extraordi-
narily dimorphic in body proportions, scutellation, and color pattern; (10) bright yellow lips in
males; (11) orange coloration on the neck, posterior and lateral head (small greenish-blue spots,
usually 2–3 scales in size, distributed frontally), extending posteriorly to form a vertebral stripe in
males; (12) dorsum and flanks with gray background decorated with numerous greenish-blue light
specks one scale in size or smaller, and irregular dark spots; (13) venter of head with dark and/or
burnt orange spots, never exhibiting dark bars on the chin connected with a dark medial patch; (14)
creamy yellow flash marks on the underside of the thighs and precloacal flap; (15) arboreal habits.
Comparison with Other Species: Tropidurus lagunablanca, n. sp., is likely to be con-
founded with T. tarara, n. sp., but differs from that species by having males with orange color-
2016 CARVALHO: THREE NEW SPECIES OF THE TROPIDURUS SPINULOSUS GROUP 9
FIGURE 2. Holotype of Tropidurus lagunablanca, n. sp. (CZPLT-H 402). A–C. Dorsal, lateral, and ventral
views of the head. D–F. Dorsal, lateral, and ventral views of the body.
10 AMERICAN MUSEUM NOVITATES NO. 3853