Research Article |
Corresponding author: Shuo Liu ( liushuo@mail.kiz.ac.cn ) Corresponding author: Dingqi Rao ( raodq@mail.kiz.ac.cn ) Academic editor: Thomas Ziegler
© 2019 Shuo Liu, Dingqi Rao.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Liu S, Rao D (2019) A new species of the genus Acanthosaura from Yunnan, China (Squamata, Agamidae). ZooKeys 888: 105-132. https://doi.org/10.3897/zookeys.888.38491
|
A new species of Acanthosaura from Yunnan, China is described based on unique morphometric and meristic external characters and a very distinctive color pattern. The fourteenth species recorded of this genus, Acanthosaura tongbiguanensis sp. nov., was previously considered A. lepidogaster although it more closely resembles A. crucigera. It can be separated from all other species of the genus by having different numbers of subdigital lamellae on the fourth finger and toe, and a different shape of the black eye patch. The new species differs genetically from investigated congeners by percentage distance of 14.46% to 23.27% (cytochrome b gene).
crucigera, Dehong, lepidogaster, Tongbiguan
The genus Acanthosaura (Gray, 1831) includes thirteen currently recognized species: A. armata (Hardwicke & Gray, 1827); A. lepidogaster (Cuvier, 1829); A. capra (Günther, 1861); A. coronata (Günther, 1861); A. crucigera Boulenger, 1885; A. nataliae Orlov et al., 2006; A. bintangensis Wood et al., 2009; A. titiwangsaensis Wood et al., 2009; A. cardamomensis Wood et al., 2010; A. brachypoda Ananjeva et al., 2011; A. phuketensis Pauwels et al., 2015; A. murphyi Nguyen et al., 2018; and A. phongdienensis Nguyen et al., 2019. It has a very wide distribution, and phylogenetic studies have shown that the genus was in need of revision as it included several undescribed and cryptic species as revealed by molecular data (
During our field research in Dehong Autonomous Prefecture, Yunnan Province, China, we discovered some lizards that looked superficially like Acanthosaura lepidogaster. According to
Specimens were collected by hand. Photographs were taken to document color pattern in life prior to euthanasia. Liver tissues were stored in 99% ethanol and lizards were preserved in 75% ethanol. Specimens were deposited at Kunming Natural History Museum of Zoology, Kunming Institute of Zoology, Chinese Academy of Sciences.
Forty-nine meristic and mensural characters were noted for each adult specimen of the type series, but only meristic characters were noted on juvenile specimens (see Table
BEP presence (1) or absence (0) of a black eye patch;
CS number of canthus rostralis-supraciliary scales, counted from the nasal scale to the posterior end of the ridge at the posterior margin of the orbit;
DIAS length of the diastema, measured from the posterior end of the nuchal crest to the anterior end of the dorsal crest;
DIASN number of scales in the vertebral crest scale diastema, counted from the posterior end of the nuchal crest to the anterior end of the dorsal crest;
DS maximum length of the largest spine in the dorsal crest, measured from the base to the tip;
DSL longest dorsal scale, measured at the base below the dorsal crest;
ESBO presence (1) or absence (0) of elliptical scales below the orbit;
EYE eye diameter, measured from the posterior to the anterior edge of the eye;
FI number of subdigital lamellae on the fourth finger;
FOREL forelimb length, measured from axilla to the proximal edge of the palmar region;
GP size of gular pouch, scored as absent (0), small (1), medium (2), large (3) or very large (4);
HD maximum head height, measured across the parietal region;
HINDL hindlimb length, measured from groin to the proximal edge of the plantar region;
HL head length, measured from posterior edge of the lower jaw to the tip of the snout;
HW head width, maximum head width, the width at the level of the tympanum;
INFRAL number of infralabials;
LKP presence (1) or absence (0) of light knee patch;
MH mental height;
MW mental width;
NCS number of scales between the fifth canthals;
ND presence (1) or absence (0) of a black, diamond shaped, nuchal collar;
NR number of scales between the nasal and the rostral;
NS number of scales between the nasals;
NSCSL number of scales from the fifth canthal to the fifth supralabial;
NSL maximum length of the largest spine in the nuchal crest measured from the base to the tip;
NN
number of nuchal crest spines (in addition to characters abbreviations listed in
NSSLC number of scales between the seventh supralabial and the sixth canthal;
NSSOS number of scales surrounding the occipital spine;
NSSPS
number of scales surrounding the postorbital spine (in addition to characters abbreviations listed in
OF presence (1) or absence (0) of oblique humeral fold;
ORBIT orbit diameter, measured from the posterior to the anterior edge of the orbit;
OS length of the occipital spine, measured from the base to the tip;
PM number of scales bordering the mental;
PS postorbital spine length, measured from the base to the tip of the spine;
RH rostral height;
RS number of scales bordering the rostral scale;
RW rostral width;
SL snout length, measured from the anterior edge of the orbit to the tip of the snout;
SUPRAL number of supralabials;
SVL snout-vent length, measured from the tip of the snout to the tip of the vent;
TL tail length, measured from the posterior margin of the vent to the tip of the tail;
TBW tail base width, maximum width at tail base;
TD tympanum diameter, measured horizontally from the anterior to the posterior border of the tympanum;
TN scales absent on tympanum (0) or present (1);
TO subdigital lamellae on the fourth toe;
VENT number of ventral scales, counted at the midline from the anterior edge of the shoulders to the edge of the vent;
WNC maximum width of the spines in the nuchal crest, measured at the base;
WDS maximum width of the largest dorsal scale below the dorsal crest, measured at the base;
YAS presence (1) or absence (0) of a Y-shaped arrangement of enlarged scales on the snout.
We compared the characters of the new collection with the characters of all currently recognized species of Acanthosaura (
The character DIAS of Acanthosaura brachypoda is given both as 4.5 and 1.9 mm in the original description, which is based on a single specimen, so this character is not used here for comparisons. The methodology for taking FOREL and HINDL was insufficiently described in the original description of A. brachypoda and thus could not be compared here; CS, NCS, NR, NSCSL, NSSLC and NSSOS were not provided in the original description of A. brachypoda; NSSLC, PM, ND, LKP, ESBO and OF were not provided in the original description of A. murphyi; SL, ORBIT, WNC, FOREL, HINDL, VENT, OS, NSSOS, CS, RS, NS, NSC, NSCSL,NR, NSSLC, PM, YAS, BEP, ESBO and GP were not provided in the original description of A. phongdienensis.
Molecular data were generated for three specimens and all available homologous sequences obtained from GenBank, all new sequences have been deposited in GenBank. According to
Species | Locality | Voucher no. | GenBank no. |
---|---|---|---|
Acanthosaura armata | Pulau Pinang, Pinang, Malaysia | PCUM | AY572871 |
Pulau Pinang, Pinang, Malaysia | PCUM | AY572872 | |
No data | NSMT-H4595 | AB266452 | |
No data | No data | NC_014175 | |
Acanthosaura coronata | Krong Pa, Gia Lai, Vietnam | ROM31985 | AY572896 |
Dong Nai, Cat Tien, Dong Nai, Vietnam | ROM42240 | AY572897 | |
Dong Nai, Cat Tien, Dong Nai, Vietnam | ROM37083 | AY572898 | |
Dong Nai, Cat Tien, Dong Nai,Vietnam | ROM42241 | AY572899 | |
Acanthosaura crucigera | Bago Division, Bago Yoma, Myanmar | CAS206626 | AY572888 |
Bago Division, Bago Yoma, Myanmar | CAS208426 | AY572895 | |
Acanthosaura lepidogaster | Chi Linh, Hia Duong, Vietnam | ROM31954 | AY572901 |
Chi Linh, Hia Duong, Vietnam | ROM31957 | AY572902 | |
Chi Linh, Hia Duong, Vietnam | ROM31960 | AY572911 | |
Chi Linh, Hia Duong, Vietnam | ROM35038 | AY572903 | |
Tam Dao, Vinh Phu, Vietnam | ROM30503 | AY572906 | |
Tam Dao, Vinh Phu, Vietnam | ROM30720 | AY572907 | |
Tam Dao, Vinh Phu, Vietnam | ROM30694 | AY572908 | |
Tam Dao, Vinh Phu, Vietnam | ROM30693 | AY572927 | |
Quang Thanh, Cao Bang, Vietnam | ROM36073 | AY572909 | |
Quang Thanh, Cao Bang, Vietnam | ROM36075 | AY572910 | |
Nakai, Khammouane, Laos | FMNH255488 | AY572920 | |
Nakai, Khammouane, Laos | FMNH255487 | AY572921 | |
Thaphabat, Bolikhamxay, Laos | FMNH255491 | AY572919 | |
Hainan, China | MD001 | KR092427 | |
Sa Pa, Lao Cai, Vietnam | ROM38117 | AY572924 | |
Sa Pa, Lao Cai, Vietnam | ROM38115 | AY572925 | |
Sa Pa, Lao Cai, Vietnam | ROM38116 | AY572926 | |
Acanthosaura nataliae | Krong Pa, Gia Lai, Vietnam | ROM31983 | AY572873 |
Krong Pa, Gia Lai, Vietnam | ROM32167 | AY572874 | |
Krong Pa, Gia Lai, Vietnam | ROM32160 | AY572875 | |
Krong Pa, Gia Lai, Vietnam | ROM31984 | AY572876 | |
Krong Pa, Gia Lai, Vietnam | ROM32154 | AY572877 | |
Krong Pa, Gia Lai, Vietnam | ROM32155 | AY572878 | |
Krong Pa, Gia Lai, Vietnam | ROM32160 | AY572879 | |
Tram Lap, Gia Lai, Vietnam | ROM30627 | AY572880 | |
Tram Lap, Gia Lai, Vietnam | ROM30628 | AY572881 | |
Krong Pa, Gia Lai, Vietnam | ROM32161 | AY572882 | |
Krong Pa, Gia Lai, Vietnam | ROM32152 | AY572883 | |
Krong Pa, Gia Lai, Vietnam | ROM32162 | AY572884 | |
Krong Pa, Gia Lai, Vietnam | ROM32143 | AY572885 | |
Krong Pa, Gia Lai, Vietnam | ROM32166 | AY572886 | |
Acanthosaura cf. phongdienensis | Khe Moi River, Nghe An, Vietnam | ROM26328 | AY572900 |
Annam, Vu Quang, Ha Tinh, Vietnam | ZISP20753-1 | AY572904 | |
Annam, Vu Quang, Ha Tinh, Vietnam | ZISP20753-2 | AY572905 | |
Boualapha, Khammouane, Laos | FMNH255481 | AY572912 | |
Con Cuong, Nghe An, Vietnam | FMNH255582 | AY572913 | |
Con Cuong, Nghe An, Vietnam | FMNH255583 | AY572914 | |
Tuong Duong, Nghe An, Vietnam | FMNH255585 | AY572915 | |
Tuong Duong, Nghe An, Vietnam | FMNH255587 | AY572916 | |
Con Cuong, Nghe An, Vietnam | FMNH255581 | AY572917 | |
Tuong Duong, Nghe An, Vietnam | FMNH255584 | AY572918 | |
Vieng Tong, Huaphan, Laos | FMNH255489 | AY572922 | |
Khao Yoi, Thailand | PCUM | AY572923 | |
Acanthosaura cf. phuketensis | Kao Yoi, Phetchaburi, Thailand | No data | AY572887 |
Khao Lak, TakuaPa, Phang Nga, Thailand | PCUM | AY572889 | |
Khao Lak, TakuaPa, Phang Nga, Thailand | PCUM | AY572890 | |
Khao Lak, TakuaPa, Phang Nga, Thailand | PCUM | AY572891 | |
ThaiMuang, Phang Nga, Thailand | IRSNB15141 | AY572892 | |
Malaysia | No data | AY572893 | |
Malaysia | No data | AY572894 | |
Acanthosaura sp. 1 | Myanmar | HLMD-RA2969 | AY572929 |
Acanthosaura sp. 1 | Myanmar | HLMD-RA2970 | AY572930 |
Acanthosaura sp. 2 | Ngoc Linh, Kon Tum, Vietnam | ROM37082 | AY572928 |
Calotes versicolor | Vietnam | HLDM57 | AY572870 |
Pseudocalotes brevipes | Pac Ban, Tuyen Quang, Vietnam | ROM30515 | AY572869 |
Acanthosaura tongbiguanensis sp. nov. | Tongbiguan, Dehong, Yunnan, China | KIZL201801 | MN604012 |
Tongbiguan, Dehong, Yunnan, China | KIZL201802 | MN604013 | |
Tongbiguan, Dehong, Yunnan, China | KIZL201803 | MN604014 |
Sequences were aligned using CLUSTAL X v1.83 (
The obtained sequence alignment is 795 bp in length. Both Bayesian inference and Maximum likelihood analyses recovered this lineage of the new samples as the sister to the clade consisting of Acanthosaura crucigera and A. cf. phuketensis with weak support (Figures
Average uncorrected p-distances (%) between investigated members of Acanthosaura and outgroups calculated from cytb gene sequences.
Species | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 |
1 Acanthosaura tongbiguanensis sp. nov. | |||||||||||
2 A. armata | 14.80 | ||||||||||
3 A. coronata | 23.27 | 22.20 | |||||||||
4 A. crucigera | 15.52 | 15.11 | 22.71 | ||||||||
5 A. lepidogaster | 15.18 | 16.11 | 22.48 | 16.52 | |||||||
6 A. nataliae | 14.94 | 14.90 | 22.15 | 15.45 | 13.72 | ||||||
7 A. cf. phongdienensis | 15.88 | 14.43 | 22.63 | 15.23 | 12.32 | 14.77 | |||||
8 A. cf. phuketensis | 14.46 | 14.83 | 22.65 | 11.17 | 15.35 | 15.02 | 14.04 | ||||
9 Acanthosaura sp. 1 | 22.94 | 21.81 | 16.18 | 22.51 | 23.04 | 23.77 | 23.62 | 23.86 | |||
10 Acanthosaura sp. 2 | 15.35 | 14.08 | 23.90 | 16.47 | 14.82 | 15.82 | 14.93 | 15.79 | 23.78 | ||
11 Pseudocalotes brevipes | 22.85 | 24.43 | 26.92 | 24.32 | 23.66 | 24.00 | 24.79 | 24.44 | 26.39 | 24.67 | |
12 Calotes versicolor | 27.25 | 25.49 | 28.40 | 27.60 | 27.75 | 27.11 | 26.39 | 26.38 | 28.56 | 29.33 | 28.00 |
Acanthosaura lepidogaster:
Acanthosaura lepidogaster:
Holotype. KIZL201804, an adult male, 22:18 02 Sept 2018, leg. Shuo Liu, Tongbiguan Township (24°36’51.24”N, 97°35’1.88”E, 1170.24 m elevation), Yingjiang County, Dehong Autonomous Prefecture, Yunnan, China.
Paratypes. KIZL201801, an adult male, 22:53 01 Sept 2018, leg. Shuo Liu, same locality as holotype; KIZL201802 and KIZL201803, two juveniles, 21:00–22:00 02 Sept 2018, leg. Shuo Liu, same locality as holotype; KIZL201805, adult female, 22:40 02 Sept 2018, leg. Shuo Liu, same locality as holotype; 74I0039 and 74I0040, two gravid females, old specimens in the specimen collection room of Kunming Institute of Zoology, Chinese Academy of Sciences, Aug 1974, leg. Longchuan County, Dehong Autonomous Prefecture, Yunnan, China.
The name refers to Tongbiguan Nature Reserve, the locality where the new species was found.
A medium-sized (maximum SVL 115.6 mm) agamid lizard with two pairs of spines: postorbital (supraciliary) spines and spines on occiput between tympanum and nuchal crest; tympanum naked; moderately developed gular pouch; scales on flanks randomly intermixed with medium and large scales; nuchal crest present and strongly developed; diastema between the nuchal and dorsal crests present; dorsal crest slightly developed, composed of enlarged, pointed scales beginning at shoulder region and decreasing regularly in size; tail 1.56–1.85 times SVL; black nuchal collar present; black eye patch present; black oblique folds anterior to the fore limb insertions present.
The new species can be separated from all congeners by having different numbers of subdigital lamellae on the fourth finger (19–21) and toe (25–28), and a different shape of the black eye patch, that extends from posterior margin of nostrils through orbit posteriorly and downwards beyond the posterior end of the tympanum but neither meeting the diamond shaped black nuchal collar on nape nor black oblique humeral fold.
Adult male. SVL 110.8 mm. TL 205.0 mm, tail complete. Head length 31.1 mm; head moderately long (HL/SVL 28%), somewhat narrow (HW/SVL 18%), not tall (HD/HL 52%), triangular in dorsal and lateral profile. Snout short (SL/HL 31%); interorbital and frontal regions and rostrum wide, steeply sloping anteriorly. Canthus rostralis prominent, forming a large projecting ridge extending above eye, composed of 11/13 enlarged scales; the ridge terminates with a notch anterior to the postorbital spine. Rostral moderate in size, rectangular; nasal concave, nostrils surrounded by a circular scale. Eye relatively large (EYE/HL 22%), orbit very large (ORBIT/HL 35%). Prefrontal and frontal scales slightly keeled and larger than scales between supralabials; scales on occiput weakly keeled. Moderately elongate epidermal spine above posterior margin of eye, straight, surrounded by 5/4 enlarged scales. A notch present between the supraciliary edge and postorbital spine. Moderately elongate epidermal spine on occipital region, straight, surrounded by a rosette of 5/4 short spiny scales. Tympanum exposed, oblong, surrounded by small scales. Supralabials 13/13, rectangular, scales in center of series largest; mental squarish above, becoming triangular below, larger than first pair of INFRAL; five scales contacting the mental; infralabials 13/12, rectangular, scales in center of series largest; gulars sharply keeled and spinose. Dewlap extensible, gular pouch moderate. Nuchal crest composed of four very elongate, lanceolate, laterally compressed scales and one moderately elongate, lanceolate, laterally compressed scale bordered on each side by one row of enlarged, spinose scales; nuchal crest followed by a diastema at base of nape. Dorsal body crest slightly developed, extending from posterior margin of diastema onto base of tail; vertebral crest composed of enlarged, epidermal, laterally compressed, spinose scales, bordered by a single row of smaller paravertebral spinose scales; vertebral crest tapers slightly to base of tail, then fades progressively. Body slightly short, triangular in cross-section. Dorsal scales small, mixed with large scales indistinctly arranged in slanted forward and downward rows from the midline of the back, keels projecting posterior wards; scales of pectoral region and abdomen larger than dorsal scales, keeled, more or less arranged in transverse rows; keeled scales anterior to vent not enlarged. Limbs relatively long (FOREL/SVL 39%, HINDL/SVL 56%); dorsal and ventral scales of forelimbs keeled, spinose, about the same size. Five digits on manus; subdigital scales keeled, subdigital lamellae under fourth finger 20/21. Scales of hind limbs keeled and spinose; postfemoral scales small, interspersed with larger spinose scales. Five digits on pes; subdigital scales keeled, subdigital lamellae under fourth toe 26/27. Tail length 1.85 times SVL, tail covered with keeled spinose scales, keels on subcaudals directed posteriorly; subcaudals much longer than supracaudals; base of tail 13.1 mm wide.
Dorsal view (top) and ventral view (bottom) of type series of Acanthosaura tongbiguanensis sp. nov. in preservative. From left to right: male holotype (KIZL201804), female paratype (KIZL201805), male paratype (KIZL201801), female paratype (74I0039), female paratype (74I0040), juvenile paratype (KIZL201802), juvenile paratype (KIZL201803).
Dorsal surface of head black, dorsal surface of body and limbs orangish brown; black eye patch extending from posterior margin of nostrils through orbit posteriorly and downwards beyond the posterior end of the tympanum but neither meeting the diamond shaped black nuchal collar on nape nor black oblique folds anterior to the fore limb insertions; upper lip white, same as color of lateral and ventral sides of neck, lower lip white with small black speckle at posterior region; iris orangish brown; black nuchal collar extending downward to reach black oblique folds anterior to fore limb insertions, two white patches at lower back of black nuchal collar; gular region white; postorbital spines, occipital spines, nuchal crest spines and ridge of the rostral and orbit cream-colored; tongue and inside of mouth pink; few small black speckles and yellow diagonal stripes from midline of the back, irregular light colored spots on sides of body not obvious; stripes checkered with black and white on dorsal ground of limbs and tail; ventral sides of limbs and body white, front part white and back part dark on ventral side of tail. However, it should be noted that this species can change the color of its body within a certain range like most other members of the genus.
Morphometric and meristic data for the type series are provided in Table
Morphometric (in mm) and meristic data for the type series of Acanthosaura tongbiguanensis sp. nov. For character abbreviations see material and methods. Paired meristic characters are given left/right. NA = not applicable.
Adult males | Adult females | Juveniles | Range | Mean | |||||
---|---|---|---|---|---|---|---|---|---|
Holotype | Paratype | Paratypes | Paratypes | ||||||
KIZL201804 | KIZL201801 | KIZL201805 | 74II0039 | 74I0040 | KIZL201802 | KIZL201803 | |||
SVL | 110.8 | 108.1 | 107.2 | 93.0 | 115.6 | NA | NA | 93.0–115.6 | 106.9 |
TL | 205.0 | NA | 180.0 | 144.9 | 183.5 | NA | NA | 144.9–205.0 | 178.4 |
TL/SVL | 1.85 | NA | 1.68 | 1.56 | 1.59 | NA | NA | 1.56–1.85 | 1.7 |
TBW | 13.1 | 13.0 | 9.2 | 10.5 | 12.1 | NA | NA | 9.2–13.1 | 11.6 |
HL | 31.1 | 33.0 | 30.4 | 27.5 | 33.2 | NA | NA | 27.5–33.2 | 31.0 |
HW | 20.0 | 22.0 | 18.8 | 18.6 | 23.3 | NA | NA | 18.6–23.3 | 20.5 |
HD | 16.1 | 17.4 | 15.7 | 13.9 | 17.3 | NA | NA | 13.9–17.4 | 16.1 |
SL | 9.8 | 10.3 | 10.1 | 9.2 | 11.0 | NA | NA | 9.2–11.0 | 10.1 |
ORBIT | 11.0 | 10.6 | 10.2 | 7.7 | 10.6 | NA | NA | 7.7–11.0 | 10.0 |
EYE | 6.9 | 5.8 | 6.9 | 6.0 | 7.3 | NA | NA | 5.8–7.3 | 6.6 |
TD | 3.6 | 4.2 | 3.3 | 3.2 | 3.7 | NA | NA | 3.2–4.2 | 3.6 |
TD/HD | 0.22 | 0.24 | 0.21 | 0.23 | 0.21 | NA | NA | 0.21–0.24 | 0.2 |
TN | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
PS | 5.3 | 6.3 | 4.6 | 3.6 | 5.4 | NA | NA | 3.6–6.3 | 5.0 |
PS/HL | 0.17 | 0.19 | 0.15 | 0.13 | 0.16 | NA | NA | 0.13–0.19 | 0.2 |
NSSPS | 5/4 | 5/5 | 5/5 | 5/5 | 5/NA | 4/4 | 4/4 | 4/4–5/5 | 4.7/4.5 |
NSL | 6.5 | 6.7 | 5.4 | 4.0 | 6.2 | NA | NA | 4.0–6.7 | 5.8 |
NSL/HL | 0.21 | 0.20 | 0.18 | 0.15 | 0.19 | NA | NA | 0.15–0.21 | 0.2 |
DS | 3.7 | 4.2 | 3.7 | 2.4 | 3.8 | NA | NA | 2.4–4.2 | 3.6 |
DS/HL | 0.12 | 0.13 | 0.12 | 0.09 | 0.11 | NA | NA | 0.09–0.13 | 0.1 |
NN | 5 | 6 | 5 | 6 | 6 | 5 | 4 | 4–6 | 5.3 |
DSL | 1.9 | 1.9 | 1.8 | 1.7 | 2.6 | NA | NA | 1.7–2.6 | 2.0 |
WNC | 1.2 | 1.5 | 1.0 | 1.1 | 1.5 | NA | NA | 1.0–1.5 | 1.3 |
WDS | 1.4 | 1.5 | 1.5 | 1.4 | 2.1 | NA | NA | 1.4–2.1 | 1.6 |
DIAS | 6.0 | 5.6 | 5.5 | 6.1 | 3.9 | NA | NA | 3.9–6.1 | 5.4 |
DIAS/SVL | 0.05 | 0.05 | 0.05 | 0.07 | 0.03 | NA | NA | 0.03–0.07 | 0.1 |
DIASN | 10 | 8 | 9 | 8 | 6 | 7 | 9 | 6–10 | 8.1 |
FOREL | 43.2 | 42.5 | 42.4 | 34.7 | 41.3 | NA | NA | 34.7–43.2 | 40.8 |
HINDL | 62.1 | 63.9 | 62.5 | 54.1 | 62.9 | NA | NA | 54.1–63.9 | 61.1 |
SUPRAL | 13/13 | 14/14 | 13/13 | 11/11 | 13/13 | 11/11 | 12/12 | 11–14 | 12.4/12.4 |
INFRAL | 13/12 | 13/14 | 14/14 | 12/11 | 12/12 | 10/11 | 12/13 | 10–14 | 12.3/12.4 |
VENT | 66 | 59 | 62 | 52 | 53 | 53 | 60 | 52–66 | 57.9 |
FI | 20/21 | 20/19 | 21/20 | 19/19 | 21/20 | 20/20 | 20/21 | 19–21 | 20.1/20.0 |
TO | 26/27 | 25/26 | 28/27 | 27/26 | 26/25 | 26/27 | 25/26 | 25–28 | 26.1/26.3 |
OS | 6.1 | 7.0 | 6.9 | 4.5 | 6.3 | NA | NA | 4.5–7.0 | 6.2 |
OS/HL | 0.20 | 0.21 | 0.23 | 0.16 | 0.19 | NA | NA | 0.16–0.23 | 0.2 |
NSSOS | 5/4 | 5/5 | 4/4 | 5/5 | 5/5 | 5/5 | 4/4 | 4–5 | 4.7/4.6 |
CS | 11/13 | 14/15 | 13/13 | 10/10 | 12/11 | 11/10 | 13/11 | 10–14 | 12.0/11.9 |
RW | 3.7 | 4.5 | 3.4 | 3.3 | 3.5 | NA | NA | 3.3–4.5 | 3.7 |
RH | 1.6 | 1.7 | 2.0 | 1.0 | 1.9 | NA | NA | 1.0–2.0 | 1.6 |
RS | 9 | 9 | 7 | 7 | 7 | 6 | 6 | 6–9 | 7.3 |
NS | 9 | 8 | 8 | 8 | 8 | 8 | 8 | 8–9 | 8.1 |
NCS | 11 | 11 | 10 | 12 | 13 | 11 | 10 | 10–13 | 11.1 |
NSCSL | 9 | 8 | 9 | 7 | 8 | 9 | 8 | 7–9 | 8.3 |
NR | 2 | 2 | 2 | 2 | 2 | 2 | 2 | 2 | 2 |
NSSLC | 13 | 10 | 13 | 10 | 9 | 11 | 12 | 9–13 | 11.1 |
MW | 1.6 | 1.4 | 1.4 | 1.9 | 1.5 | NA | NA | 1.4–1.9 | 1.6 |
MH | 1.2 | 2.0 | 1.3 | 1.4 | 1.7 | NA | NA | 1.2–2.0 | 1.5 |
PM | 5 | 4 | 5 | 5 | 4 | 4 | 5 | 4–5 | 4.6 |
YAS | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
ND | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
LKP | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
BEP | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
ESBO | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
GP | 2 | 2 | 2 | 2 | 2 | 1 | 1 | 1–2 | 1.7 |
OF | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Acanthosaura tongbiguanensis sp. nov. is only recorded in Tongbiguan Nature Reserve including Yingjiang County, Longchuan County and Ruili City, the border region with northern Myanmar in western Yunnan, China, so it probably occurs in northern Myanmar.
The type series of Acanthosaura tongbiguanensis sp. nov. was collected at night while they were asleep on small trees in a primordial forest. However, we suppose that they forage for food on the ground during the day. At the type locality we found four other species of reptiles, namely Cyrtodactylus khasiensis (Jerdon, 1870), Pseudocalotes kakhienensis (Anderson, 1879); P. microlepis (Boulenger, 1887); Trimeresurus yingjiangensis Chen et al., 2019; and seven species of amphibians, Leptobrachella yingjiangensis (Yang et al., 2018); Limnonectes longchuanensis Suwannapoom et al., 2016; Megophrys feii Yang et al., 2018; M. glandulosa Fei et al., 1990; Raorchestes longchuanensis (Yang & Li, 1978); Theloderma moloch (Annandale, 1912); Zhangixalus smaragdinus (Blyth, 1852).
Table
Comparisons of morphometric (in mm) and meristic data for all currently recognized species of Acanthosaura and Acanthosaura tongbiguanensis sp. nov., “?” = data not available. (1) Acanthosaura tongbiguanensis sp. nov.; (2) A. armata; (3) A. bintangensis; (4) A. brachypoda; (5) A. capra; (6) A. cardamomensis; (7) A. coronata; (8) A. crucigera; (9) A. lepidogaster; (10) A. murphyi; (11) A. nataliae; (12) A. phongdienensis; (13) A. phuketensis; (14) A. titiwangsaensis.
(1) | (2) | (3) | (4) | (5) | (6) | (7) | (8) | (9) | (10) | (11) | (12) | (13) | (14) | |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
SVL | 93.0–115.6 | 72.4–138.0 | 83.9–142.0 | 117 | 94.0–137.9 | 82–149 | 66.0–86.1 | 92.2–127.0 | 76.5–101.1 | 103.7–127.3 | 106.7–158.0 | 58.5–77.4 | 69.2–123.5 | 91.8–118.4 |
TL | 144.9–205.0 | 96.6–190 | 112.8–206.0 | 185.4 | 133.6–182.1 | 103–188 | 86.3–105.0 | 130.0–174.0 | 130.6–144.1 | 159.3–195.8 | 171.0–287.0 | 94.6–137.2 | 107.0–205.6 | 136.0–174.0 |
HL | 27.5–33.2 | 6.6–33.7 | 16.9–25.4 | 30.3 | 16.3–38.9 | 16.3–42.4 | 14.4–16.3 | 18.7–23.6 | 18.9–29.7 | 29.1–36.8 | 25.2–43.6 | 18.6–23.8 | 19.7–31.4 | 20.0–24.3 |
HW | 18.6–23.3 | 15.3–23.0 | 17.5–23.4 | 20.6 | 16.8–27.0 | 16.4–27.7 | 13.6–17.5 | 16.0–22.3 | 13.4–19.1 | 20.3–24.6 | 20.2–27.8 | 13.1–15.9 | 14.4–22.8 | 17.5–23.4 |
HD | 13.9–17.4 | 12.2–18.9 | 15.0–19.2 | 17.2 | 14.8–24.3 | 12.6–21.7 | 11.9–16.8 | 15.7–22.5 | 12.0–12.5 | 18.5–20.6 | 16.9–24.9 | 10.4–13.6 | 10.9–18.6 | 15.7–20.2 |
SL | 9.2–11.0 | 6.3–16.6 | 7.9–11.3 | 12.2 | 7.6–16.6 | 8.6–18.7 | 6.9–8.4 | 8.7–12.1 | 9.3–10.2 | 10.3–15.3 | 12.0–19.9 | ? | 6.8–11.0 | 9.7–12.5 |
ORBIT | 7.7–11.0 | 5.4–12.2 | 8.4–12.6 | 8.3 | 7.6–11.6 | 6.0–12.7 | 6.9–7.5 | 8.9–10.8 | 4.7–9.1 | 9.9–12.3 | 7.2–10.9 | ? | 6.6–11.2 | 9.8–13.2 |
TD | 3.2–4.2 | 2.4–5.2 | 2.5–3.0 | 3.6 | 3.4–5.2 | 2.5–5.8 | 1.7–2.8 | 2.5–3.9 | 2.2–3.0 | 3.2–5.2 | 3.9–7.0 | 1.78–2.81 | 3.5–4.7 | 2.7–4.0 |
TD/HD | 0.21–0.24 | 0.19–0.28 | 0.16 | 0.21 | 0.21–0.23 | 0.20–0.27 | 0.14–0.17 | 0.14–0.21 | 0.18–0.24 | 0.17–0.28 | 0.23–0.28 | 0.17–0.22 | 0.22–0.33 | 0.17–0.20 |
TN | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0–1 | 1 | 0 | 0 | 0 | 0 |
PS | 3.6–6.3 | 4.9–9.9 | 1.9–4.2 | 3.2 | 5.2–10.2 | 3.2–12.7 | Absent | 1.9–7.8 | 1.5–2.5 | 5.6–11.8 | 7.7–17.8 | 1.18–2.07 | 4.6–11.8 | 3.3–4.4 |
PS/HL | 0.13–0.19 | 0.22–0.56 | 0.07–0.19 | 0.11 | 0.36 | 0.14–0.45 | 0 | 0.09–0.33 | 0.06–0.11 | 0.16–0.34 | 0.36 | 0.06–0.09 | 0.23–0.38 | 0.14–0.18 |
NSL | 4.0–6.7 | 5.5–11.2 | 1.3–4.7 | 4.7 | 4.2–14.7 | 3.8–17.4 | 0 | 3.1–8.9 | 2.9–3.4 | 7.0–14.9 | 8.5–23.8 | 1.24–4.18 | 4.1–12.2 | 2.7–4.4 |
NSL/HL | 0.15–0.21 | 0.22–0.51 | 0.17–0.21 | 0.16 | 0.42–0.43 | 0.17–0.66 | 0 | 0.14–0.38 | 0.12–0.15 | 0.24–0.43 | 0.58 | 0.07–0.18 | 0.21–0.39 | 0.11–0.18 |
DS | 2.4–4.2 | 4.9–11.3 | 1.8–2.2 | 1.9 | 3.5–6.8 | 2.0–14.2 | Absent | 2.0–5.5 | 1.5–2.7 | 2.6–10.5 | 6.0–17.7 | 0.58–1.65 | 2.3–8.3 | 1.7–2.1 |
DS/HL | 0.09–0.13 | 0.20–0.52 | 0.08–0.09 | 0.06 | 0.16–0.17 | 0.14–0.45 | 0 | 0.09–0.24 | 0.07–0.12 | 0.14–0.51 | 0.44 | 0.03–0.07 | 0.11–0.26 | 0.07–0.09 |
WNC | 1.0–1.5 | 1.0–2.2 | 1.6–2.1 | 1.6 | 2.3–4.1 | 1.8–4.2 | 0 | 1.3–3.4 | 1.5 | 2.9–4.8 | 3.1–4.8 | ? | 1.4–2.9 | 1.4–1.6 |
DIAS | 3.9–6.1 | 1.2–6.8 | 5.0–7.9 | ? | 2.0–6.7 | 2.7–8.3 | Absent | 4.9–8.4 | 6.3 | 2.6–4.8 | 2.5 | Absent | 3.6–7.6 | 5.1–7.6 |
DIASN | 6–10 | 1–8 | 11–15 | 7 | 4–7 | 6–15 | Absent | 9–25 | 10–12 | 4–8 | 10 | Absent | 12–17 | 10–13 |
DIAS/SVL | 0.03–0.07 | 0.01–0.06 | 0.04–0.07 | ? | 0.05 | 0.03–0.07 | Absent | 0.04–0.08 | 0.08 | 0.02–0.04 | 0.04 | Absent | 0.05–0.08 | 0.05–0.07 |
FOREL | 34.7–43.2 | 33.7–48.9 | 33.9–61.5 | ? | 54.2–83.8 | 31.7–56.8 | 30.2–35.3 | 35.6–49.8 | 33.0–37.1 | 49.8–56.6 | 60.0–85.0 | ? | 22.3–42.9 | 38.0–51.7 |
HINDL | 54.1–63.9 | 39.0–69.6 | 43.3–68.6 | ? | 78.5–107.2 | 42.0–77.1 | 38.4–47.8 | 48.8–65.0 | 49.4–50.4 | 60.4–68.4 | 85.0–129.7 | ? | 38.2–60.6 | 48.5–65.6 |
SUPRAL | 11–14 | 10–14 | 12 | 12–13 | 10 | 11–15 | 12–13 | 10–13 | 10–13 | 12–14 | 11 | 9–12 | 10–12 | 12–13 |
INFRAL | 10–14 | 12–15 | 11–12 | 11 | 12–13 | 10–14 | 11–13 | 10–12 | 9–13 | 12–14 | 11–12 | 10–11 | 10–12 | 11–12 |
VENT | 52–66 | 51–68 | 51–55 | 63 | 55–66 | 50–65 | 53–58 | 55–63 | 55–61 | 55–65 | 64–71 | ? | 57–67 | 47–57 |
FI | 19–21 | 13–17 | 23 | 18 | 16–17 | 15–20 | 13–14 | 16–18 | 17–19 | 15–18 | 16–21 | 14–17 | 15–17 | 20–21 |
TO | 25–28 | 19–26 | 26–28 | 24 | 22–24 | 20–25 | 17–19 | 21–26 | 22–23 | 21–23 | 20–27 | 19–23 | 21–24 | 23–27 |
TL/SVL | 1.56–1.85 | 1.2–1.6 | 1.3–1.4 | 1.58 | 1.2–1.5 | 1.2–1.6 | 0.6–1.0 | 1.1–1.8 | 1.6–1.9 | 1.48–1.54 | 1.2–1.5 | 1.5–1.9 | 1.4–1.7 | 1.1–1.5 |
OS | 4.5–7.0 | 4.0–9.4 | 1.2–2.6 | 1.0 | Absent | 4.1–13.3 | 0 | 2.5–4.9 | 3.2–3.4 | Absent | Absent | ? | 2.6–9.5 | 1.8–2.3 |
OS/HL | 0.16–0.23 | 0.16–0.38 | 0.10–0.11 | 0.03 | 0 | 0.24–0.56 | 0 | 0.11–0.50 | 0.14–0.15 | 0 | 0 | ? | 0.13–0.30 | 0.09–0.10 |
NSSOS | 4–5 | 4–6 | 6–7 | ? | Absent | 4–6 | 4–5 | 4–6 | 5 | Absent | Absent | ? | 4–5 | 4–5 |
CS | 10–14 | 11–15 | 14–15 | ? | 12–14 | 11–16 | 12–15 | 12–15 | 10–12 | 12–14 | 13 | 9–13 | 10–14 | 14–15 |
RW | 3.3–4.5 | 1.8–4.5 | 3.6–5.3 | 3.5 | 4.2–4.6 | 1.7–4.7 | 0.8–0.9 | 2.7–4.0 | 2.8–3.0 | 3.3–5.1 | 6.1 | 2.07–2.65 | 2.3–3.8 | 3.6–5.2 |
RH | 1.0–2.0 | 0.9–1.8 | 1.7–2.0 | 2.3 | 1.8–2.3 | 1.1–2.2 | 0.5–0.8 | 1.3–2.0 | 1.4–1.5 | 1.2–2.0 | 2.6 | 1.00–1.32 | 1.1–1.7 | 1.4–1.8 |
RS | 6–9 | 5–9 | 7–9 | 7 | 7–8 | 5–9 | 5 | 7–9 | 5–9 | 8–9 | 6 | ? | 5–8 | 9 |
NS | 8–9 | 7–10 | 8 | 9 | 9 | 7–10 | 7–9 | 7–9 | 7–8 | 7–8 | 8 | ? | 7–8 | 8 |
NCS | 10–13 | 10–17 | 10–11 | ? | 9 | 9–17 | 8–11 | 9–12 | 7–10 | 13–16 | 14 | ? | 12–13 | 11–12 |
NSCSL | 7–9 | 6–14 | 7–8 | ? | 7–8 | 7–12 | 5–6 | 7–11 | 8 | 7–10 | 8 | ? | 8–10 | 9–11 |
NR | 2 | 1–2 | 1 | ? | 1–2 | 1–2 | 3–4 | 1–2 | 1–2 | 3–4 | 1 | ? | 1–2 | 1–2 |
NSSLC | 9–13 | 10–22 | 9–12 | ? | 9–11 | 10–19 | 6–11 | 10–14 | 10 | ? | 16 | ? | 11–14 | 11–14 |
MW | 1.4–1.9 | 0.9–2.0 | 1.3–1.8 | 2.9 | 1.9–2.2 | 0.2–2.1 | 0.6–1.5 | 1.0–1.5 | 1.2–1.3 | 1.7–2.2 | 2.9 | 0.87–1.52 | 0.5–1.4 | 1.4–2.0 |
MH | 1.2–2.0 | 0.8–2.2 | 1.4–2.1 | 2.1 | 1.7–2.2 | 0.9–2.0 | 1.3–1.6 | 1.1–1.7 | 1.2–1.3 | 1.4–2.0 | 2.0 | 1.04–1.60 | 0.6–1.6 | 1.4–2.4 |
PM | 4–5 | 3–6 | 4–5 | 4 | 4 | 4–5 | 4–5 | 4 | 5 | ? | 4 | ? | 4 | 5 |
YAS | 1 | 0–1 | 1 | 1 | 1 | 0–1 | 0–1 | 1 | 1 | 0–1 | 1 | ? | 0–1 | 1 |
ND | 1 | 0–1 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | ? | 0 | 1 | 1 | 1 |
LKP | 1 | 1 | 0 | 1 | 1 | 1 | 1 | 1 | 1 | ? | 0 | 1 | 1 | 0 |
BEP | 1 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | 0–1 | 0–1 | 1 | ? | 1 | 1 |
ESBO | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | ? | 0 | ? | 0 | 0 |
GP | 1–2 | 1 | 3–4 | 0 | 3–4 | 1–4 | 0 | 1–2 | 0–1 | 4 | 4 | ? | 0–2 | 2–4 |
OF | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | ? | 1 | 1 | 1 | 1 |
Acanthosaura tongbiguanensis sp. nov. can be distinguished from A. armata by having more subdigital lamellae on the fourth finger (19–21 vs. 13–17) and the fourth toe (25–28 vs. 19–26), shorter postorbital spines (3.6–6.3 vs. 4.9–9.9 mm, PS/HL 0.13–0.19 vs. 0.22–0.56) and shorter occipital spines (4.5–7.0 vs. 4.0–9.4 mm, OS/HL 0.16–0.23 vs. 0.16–0.38), much shorter nuchal crest spines (4.0–6.7 vs. 5.5–11.2 mm, NSL/HL 0.15–0.21 vs. 0.22–0.51) and much shorter dorsal crest spines (2.4–4.2 vs. 4.9–11.3 mm, DS/HL 0.09–0.13 vs. 0.20–0.52), a higher number of scales in the diastema between the nuchal and the dorsal crests (6–10 vs. 1–8), a relatively longer tail (TL/SVL 1.56–1.85 vs. 1.2–1.6). Acanthosaura tongbiguanensis sp. nov. has a black eye patch (vs. absent) and an obvious black nuchal collar (vs. not obvious or absent); Acanthosaura tongbiguanensis sp. nov. has fewer or no spots on the dorsal surface of the body, whereas A. armata has more spots on the dorsal surface of the body.
The new species can be distinguished from Acanthosaura bintangensis by having a larger tympanum (3.2–4.2 vs. 2.5–3.0 mm, TD/HD 0.21–0.24 vs. 0.16), longer head (27.5–33.2 vs. 16.9–25.4 mm), longer postorbital spines (6.3 vs. 4.2 mm), higher maximal length of spines in the nuchal crest (6.7 vs. 4.7), longer spines in the dorsal crest (2.4–4.2 vs. 1.8–2.2 mm, DS/HL 0.09–0.13 vs. 0.08–0.09), less subdigital lamellae on the fourth finger (19–21 vs. 23), much longer occipital spines (4.5–7.0 vs. 1.2–2.6 mm, OS/HL 0.16–0.23 vs. 0.10–0.11), less scales surrounding the occipital spine (4–5 vs. 6–7), lower number of scales in the diastema between the nuchal and the dorsal crests (6–10 vs. 11–15), presence of a light knee patch (vs. absence), less developed gular pouch (1–2 vs. 3–4), absence of an enlarged row of keeled scales below orbit (vs. presence), absence of large yellow spots edged in blackish-brown arranged on body and base of tail (vs. presence); the black eye patch in Acanthosaura tongbiguanensis sp. nov. extends backward and downward beyond the posterior end of the tympanum while it never extends onto the head side in A. bintangensis (
From Acanthosaura brachypoda, Acanthosaura tongbiguanensis sp. nov. can be differentiated by having more subdigital lamellae on the fourth finger (19–21 vs. 18) and the fourth toe (25–28 vs. 24), longer postorbital spines (3.6–6.3 vs. 3.2 mm, PS/HL 0.13–0.19 vs. 0.11) and longer occipital spines (4.5–7.0 vs. 1.0 mm, OS/HL 0.16–0.23 vs. 0.03), much longer spines in the dorsal crest (2.4–4.2 vs. 1.9 mm, DS/HL 0.09–0.13 vs. 0.06), the presence of gular pouch (vs. absence). Acanthosaura tongbiguanensis sp. nov. does not have pairs of transverse creamy spots along both sides of spine forming a symmetrical pattern present as in A. brachypoda.
Acanthosaura tongbiguanensis sp. nov. can be distinguished from A. capra based on its smaller body size (93.0–115.6 vs. 94.0–137.9 mm) but longer tail (144.9–205.0 vs. 133.6–182.1 mm, TL/SVL 1.56–1.85 vs. 1.2–1.5), a higher number of subdigital lamellae on the fourth finger (19–21 vs. 16–17) and the fourth toe (25–28 vs. 22–24), lower maximal length of forelimb (43.2 vs. 83.8 mm) and hindlimb (63.9 vs. 107.2 mm), shorter postorbital spines (3.6–6.3 vs. 5.2–10.2 mm, PS/HL 0.13–0.19 vs. 0.36), nuchal crest spines (4.0–6.7 vs. 4.2–14.7 mm, NSL/HL 0.15–0.21 vs. 0.42–0.43) and dorsal crest spines (2.4–4.2 vs. 3.5–6.8 mm, DS/HL 0.09–0.13 vs. 0.16–0.17),lower width of the nuchal crest spines (1.0–1.5 vs. 2.3–4.1 mm), higher number of scales in the diastema between nuchal and dorsal crests (6–10 vs. 4–7), presence of occipital spines (vs. absence), a smaller gular pouch (1–2 vs. 3–4); the black eye patch in Acanthosaura tongbiguanensis sp. nov. extends backward and downward beyond the posterior end of the tympanum, while it usually extends backwards and upwards to reach the nuchal crests in A. capra.
From Acanthosaura cardamomensis, the new species can be separated based on a higher number of subdigital lamellae on the fourth finger (19–21 vs. 15–20) and the fourth toe (25–28 vs. 20–25), a longer tail (144.9–205.0 vs. 103–188 mm, TL/SVL 1.56–1.85 vs. 1.2–1.6), much shorter postorbital spines (3.6–6.3 vs. 3.2–12.7 mm, PS/HL 0.13–0.19 vs. 0.14–0.45), occipital spines (4.5–7.0 vs. 4.1–13.3 mm, OS/HL 0.16–0.23 vs. 0.24–0.56), nuchal crest spines (4.0–6.7 vs. 3.8–17.4 mm, NSL/HL 0.15–0.21 vs. 0.17–0.66) and dorsal crest spines (2.4–4.2 vs. 2.0–14.2 mm, DS/HL 0.09–0.13 vs. 0.14–0.45), a lower width of nuchal crest spines (1.0–1.5 vs. 1.8–4.2 mm); the black eye patch in Acanthosaura tongbiguanensis sp. nov. extends backward and downward beyond the posterior end of the tympanum but never reaches the dark nuchal marking on nape while it does so in A. cardamomensis (see species’ description and photographs in
Acanthosaura tongbiguanensis sp. nov. is distinguishable from A. coronata based on its much bigger body size (93.0–115.6 vs. 66.0–86.1 mm), much longer tail (144.9–205.0 vs. 86.3–105.0 mm, TL/SVL 1.56–1.85 vs. 0.6–1.0), higher number of subdigital lamellae on the fourth finger (19–21 vs. 17–19) and the fourth toe (25–28 vs. 22–23), relatively larger tympanum (TD/HD 0.21–0.24 vs. 0.14–0.17), bigger rostral (RW 3.4–4.5 vs. 0.8–0.9 mm, RH 1.0–2.0 vs. 0.5–0.8 mm), the presence of postorbital spines, occipital spines, nuchal and dorsal crests (vs. absence or not obvious), a diastema between nuchal crest and dorsal crest (vs. a continuous nuchal and dorsal crest), presence of a black nuchal collar (vs. absence), presence of a black eye patch (vs. absence), and the presence of a gular pouch (vs. absence) (see the original description by
Acanthosaura tongbiguanensis sp. nov. can be differentiated from A. crucigera by having more subdigital lamellae on the fourth finger (19–21 vs. 16–18) and the fourth toe (25–28 vs. 21–26), a relatively larger tympanum (TD/HD 0.21–0.24 vs. 0.14–0.21), a higher maximal length of tail (205.0 vs. 174.0 mm), a higher maximal length of occipital spines (7.0 vs. 4.9 mm), a lower number of scales in the diastema between the nuchal and the dorsal crests (6–10 vs. 9–25), a larger mental (MW 1.4–1.9 vs. 1.0–1.5 mm, MH 1.2–2.0 vs. 1.1–1.7 mm). Most obvious is the difference in the color pattern: the black eye patch in Acanthosaura tongbiguanensis sp. nov. extends back and downwards beyond the posterior end of the tympanum, while it only extends to the anterior edge of the tympanum in A. crucigera; additionally, Acanthosaura tongbiguanensis sp. nov. has fewer or no spots on the dorsal surface of the body, whereas A. crucigera has more spots on the dorsal surface of the body.
Acanthosaura tongbiguanensis sp. nov. can be separated from A. lepidogaster based on its higher number of subdigital lamellae on the fourth finger (19–21 vs. 17–19) and the fourth toe (25–28 vs. 22–23), its bigger body size (93.0–115.6 vs. 76.5–101.1 mm), longer postorbital spines (3.6–6.3 vs. 1.5–2.5 mm, PS/HL 0.13–0.19 vs. 0.06–0.11) and longer occipital spines (4.5–7.0 vs. 3.2–3.4 mm, OS/HL 0.16–0.23 vs. 0.14–0.15), longer nuchal crest spines (4.0–6.7 vs. 2.9–3.4 mm, NSL/HL 0.15–0.21 vs. 0.12–0.15) and longer dorsal crest spines (2.4–4.2 vs. 1.5–2.7 mm, DS/HL 0.09–0.13 vs. 0.07–0.12), much higher maximal length of tail (205.0 vs. 144.1 mm), lower number of scales in the diastema between the nuchal and the dorsal crests (6–10 vs. 10–12), much wider rostral (3.3–4.5 vs. 2.8–3.0 mm), and larger gular pouch (1–2 vs. 0–1). The black eye patch in Acanthosaura tongbiguanensis sp. nov. extends backwards and downwards beyond the posterior end of the tympanum but never backwards and upwards to reach the black nuchal collar, while it usually does so in A. lepidogaster; the black nuchal collar extends downwards to reach the black oblique humeral fold, while it rarely reaches the black oblique humeral fold in A. lepidogaster; besides, the tongue and the inside of the mouth are pink in Acanthosaura tongbiguanensis sp. nov., while they are bluish-grey or black in A. lepidogaster; the postorbital spines, occipital spines, nuchal crest spines, the ridge of the rostralis, and orbit are lighter in color in Acanthosaura tongbiguanensis sp. nov., whereas they are darker in color in A. lepidogaster.
Acanthosaura tongbiguanensis sp. nov. can be separated from A. murphyi based on its smaller body size (93.0–115.6 vs. 103.7–127.3 mm) but relatively longer tail (TL/SVL 1.56–1.85 vs. 1.48–1.54), a higher number of subdigital lamellae on the fourth finger (19–21 vs. 15–18) and the fourth toe (25–28 vs. 21–23), shorter forelimb (34.7–43.2 vs. 49.8–56.6 mm) and hindlimb (54.1–63.9 vs. 60.4–68.4 mm), much shorter postorbital spines (3.6–6.3 vs. 5.6–11.8 mm, PS/HL 0.13–0.19 vs. 0.16–0.34), nuchal crest spines (4.0–6.7 vs. 7.0–14.9 mm, NSL/HL 0.15–0.21 vs. 0.24–0.43) and dorsal crest spines (2.4–4.2 vs. 2.6–10.5 mm, DS/HL 0.09–0.13 vs. 0.14–0.51), much lower width of the nuchal crest spines (1.0–1.5 vs. 2.9–4.8 mm), higher number of scales in the diastema between nuchal and dorsal crests (6–10 vs. 4–8), presence of occipital spines (vs. absence), a smaller gular pouch (1–2 vs. 4); the black eye patch in Acanthosaura tongbiguanensis sp. nov. extends backward and downward beyond the posterior end of the tympanum, while it usually extends backwards and upwards to reach the nuchal crests in A. murphyi (see species’ photographs in
Acanthosaura tongbiguanensis sp. nov. can be separated from A. nataliae by its smaller body size (93.0–115.6 vs. 106.7–158.0 mm) and a lower maximal tail length (205.0 vs. 287.0 mm) but a relatively longer tail (TL/SVL 1.56–1.85 vs. 1.2–1.5), much shorter length of postorbital spines (3.6–6.3 vs. 7.7–17.8 mm, PS/HL 0.13–0.19 vs. 0.36), nuchal crest spines (4.0–6.7 vs. 8.5–23.8 mm, NSL/HL 0.15–0.21 vs. 0.58) and dorsal crest spines (2.4–4.2 vs. 6.0–17.7 mm, DS/HL 0.09–0.13 vs. 0.44), a lower width of the nuchal crest spines (1.0–1.5 vs. 3.1–4.8 mm), lower width of mental (1.4–1.9 vs. 2.9 mm), a lower number of ventral scales (52–66 vs. 64–71), lower maximal length of forelimb (43.2 vs. 85.0 mm) and hindlimb (63.9 vs. 129.7 mm), presence of occipital spines (vs. absence), much lesser development of gular pouch (1–2 vs. 4), presence of light knee patch (vs. absence) and presence of a black nuchal collar (vs. absence); the black eye patch in Acanthosaura tongbiguanensis sp. nov. extends backward and downward beyond the posterior end of the tympanum but never continues backward to reach the black oblique folds anterior to the fore limb insertions while it usually does so in A. nataliae (see species’ description and photographs in
Acanthosaura tongbiguanensis sp. nov. is distinguishable from A. phongdienensis based on its bigger body size (93.0–115.6 vs. 58.5–77.4 mm), longer tail (144.9–205.0 vs. 94.6–137.2 mm), higher number of subdigital lamellae on the fourth finger (19–21 vs. 14–17) and the fourth toe (25–28 vs. 19–23), longer postorbital spines (3.6–6.3 vs. 1.18–2.07 mm, PS/HL 0.13–0.19 vs. 0.06–0.09), longer nuchal crest spines (4.0–6.7 vs. 1.24–4.18 mm, NSL/HL 0.15–0.21 vs. 0.07–0.18) and longer dorsal crest spines (2.4–4.2 vs. 0.58–1.65 mm, DS/HL 0.09–0.13 vs. 0.03–0.07), a diastema between nuchal crests and dorsal crests (vs. a continuous nuchal and dorsal crest); the black eye patch in Acanthosaura tongbiguanensis sp. nov. extends backwards and downwards beyond the posterior end of the tympanum but never backwards and upwards to reach the black nuchal collar, while it does so in male A. phongdienensis (see species’ description and photographs in
Acanthosaura tongbiguanensis sp. nov. can be differentiated from A. phuketensis by having a higher number of subdigital lamellae on the fourth finger (19–21 vs. 15–17) and the fourth toe (25–28 vs. 21–24), a relatively longer tail (TL/SVL 1.56–1.85 vs. 1.4–1.7), much shorter postorbital spines (3.6–6.3 vs. 4.6–11.8 mm, PS/HL 0.13–0.19 vs. 0.23–0.38), nuchal crest spines (4.0–6.7 vs. 4.1–12.2 mm, NSL/HL 0.15–0.21 vs. 0.21–0.39) and dorsal crest spines (2.4–4.2 vs. 2.3–8.3 mm, DS/HL 0.09–0.13 vs. 0.11–0.26), a lower width of nuchal crest spines (1.0–1.5 vs. 1.4–2.9 mm), a lower maximal length of occipital spines (7.0 vs. 9.5 mm), a lower number of scales in the diastema between the nuchal and the dorsal crests (6–10 vs. 12–17), a bigger mental (MW 1.4–1.9 vs. 0.5–1.4 mm, MH 1.2–2.0 vs. 0.6–1.6 mm); the black eye patch in Acanthosaura tongbiguanensis sp. nov. never extends backward to reach the nuchal crest while it does so in male A. phuketensis (see species’ original description by
From Acanthosaura titiwangsaensis, the new species can be distinguished by its relatively larger tympanum (TD/HD 0.21–0.24 vs. 0.17–0.20), its longer tail (144.9–205.0 vs. 136.0–174.0mm, TL/SVL 1.56–1.85 vs. 1.1–1.5), higher maximal length of postorbital spines (6.3 vs. 4.4 mm) and nuchal crest spines (6.7 vs. 4.4 mm), higher length of dorsal crest spines (2.4–4.2 vs. 1.7–2.1 mm, DS/HL 0.09–0.13 vs. 0.07–0.09), much longer occipital spines (4.5–7.0 vs. 1.8–2.3 mm, OS/HL 0.16–0.23 vs. 0.09–0.10), lower number of scales in the diastema between the nuchal and the dorsal crests (6–10 vs. 10–13), presence of a light knee patch (vs. absence), less developed gular pouch (1–2 vs. 2–4), absence of medium-sized light orange spots edged in a faded black color on body and base of tail (vs. presence); the black eye patch in Acanthosaura tongbiguanensis sp. nov. extends backward and downward beyond the posterior end of the tympanum while it is restricted to the orbit and not extends into the postorbital region in A. titiwangsaensis (
Although Acanthosaura collections from Myanmar and other Southeast Asian countries were not available for comparative analyses, we could demonstrate that Acanthosaura tongbiguanensis sp. nov. is a distinct species using data available from literature (
Several morphometric characters of Acanthosaura tongbiguanensis sp. nov. overlap with some characters of other species in this genus, however, the new species can be differentiated from all other species of Acanthosaura by the black eye patch extending from the posterior margin of the nostrils through the orbit backwards and downwards to beyond the posterior end of the tympanum but neither meeting black nuchal collar nor the black oblique humeral fold (see Fig.
The Acanthosaura crucigera group is wide ranging and its morphological variation is conserved, it is not surprising to find cryptic diversity within the A. crucigera complex (
We would like to thank Decai Ouyang and Lei Ouyang for assistance in the field. Thanks also to our workmates for their help and advice. Thanks to the manager of the collection room of the Institute of Zoology, Chinese Academy of Sciences for helping to find specimens. We also thank the reviewers for their valuable comments on the manuscript. We owe thanks to the curator Weiwei Li for giving us the opportunity to conduct the field investigation; this work was supported by Kunming Natural History Museum of Zoology, Kunming Institute of Zoology, Chinese Academy of Sciences.