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Acanthosaura titiwangsaensis WOOD, GRISMER, GRISMER, AHMAD, ONN & BAUER, 2009

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Higher TaxaAgamidae (Draconinae), Sauria, Iguania, Squamata (lizards)
Subspecies 
Common NamesE: Malayan Mountain Horned Agamid, Titiwangsa Horned Tree Lizard 
SynonymAcanthosaura titiwangsaensis WOOD, GRISMER, GRISMER, AHMAD, ONN & BAUER 2009
Acanthosaura titiwangsaensis — GRISMER & QUAH 2019 
DistributionW Malaysia (Pahang)

Type locality: Bishop’s Trail, Fraser’s Hill, Pahang, Peninsular Malaysia (3º43.230'N; 101º44.887'E), 1,112 m elevation.  
Reproduction 
TypesHolotype: ZRC 2.6805, adult male, collected on 28 June 2008 by Jesse L. Grismer, L. Lee Grismer, Perry L. Wood Jr., at 2330 h.; Paratypes: ZFMK and ZRC 
DiagnosisDiagnosis. Maximum SVL of adult males 118.4 mm, maximum SVL of females 112.7 mm; head triangular, snout short (SL/HL 46–52%); eye moderate (EYE/HL 28–30%); no enlarged elliptical scales below orbit; scales on the canthus rostralis-supraorbital ridge 14 or 15; large rectangular rostral scale; a short cylindrical spine above posterior margin of eye surrounded by a rosette of four smaller keeled scales at base; spine on occiput between tympanum and nuchal crest present; tympanum naked, oblong, taller than wide; dewlap present in males and females; body triangular in cross section; scales of flanks intermixed with randomly arranged clusters of larger keeled scales; nuchal crest present; 11–13 scales in nape diastema; vertebral crest composed of enlarged, pointed scales beginning at shoulder region and tapering to a series of enlarged keeled scales on anterior portion of tail; crest scales largest at base of nape; tail 140–150% of SVL; black collar present; posterior 2/3 of tail black to dark brown; greenish-yellow to light orange spots encircled by gray on body, gular region orangish yellow; no yellow medial stripe on dewlap.

Comparisons. Table 4 shows a comparison of morphometric and meristic data for all currently recognized species of Acanthosaura and Acanthosaura tongbiguanensis sp. nov. It is based mostly on the interspecific comparison tables provided by Pauwels et al. (2015: table 2), Nguyen et al. (2018: table 3) and Nguyen et al. (2019: table 3).
Acanthosaura tongbiguanensis sp. nov. can be distinguished from A. armata by hav- ing 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 la- mellae 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 (Wood et al. 2009).
From Acanthosaura brachypoda, Acanthosaura tongbiguanensis sp. nov. can be dif- ferentiated 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 tong- biguanensis 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 subdigi- tal 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), pres- ence 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 back- ward 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’ descrip- tion and photographs in Wood et al. 2010), besides, Acanthosaura tongbiguanensis sp. nov. has fewer or no spots on the dorsal surface of the body, whereas A. cardamomensis has more spots on the dorsal surface of the body.
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 sub- digital 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 postor- bital 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 Günther 1861 and expanded descriptions by Günther 1864; Boulenger 1885).
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 tongbi- guanensis 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 Acantho- saura 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 pos- torbital 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 Acantho- saura 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 Nguyen et al. 2018).
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 men- tal (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. ex- tends 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 inser- tions while it usually does so in A. nataliae (see species’ description and photographs in Orlov et al. 2006).
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 Nguyen et al. 2019), 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. phongdienensis.
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 Pauwels et al. 2015) and Acanthosaura tongbiguanensis sp. nov. has fewer or no spots on the dorsal surface of the body, whereas A. phuketensis has more spots on the dorsal surface of the body.
From Acanthosaura titiwangsaensis, the new species can be distinguished by its rela- tively 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 gu- lar 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 (Wood et al. 2009). 
Comment 
EtymologyThis specific epithet titwangsaensis is in reference to the mountain range, Banjaran Titiwangsa, to which we believe this species is likely to be restricted. 
References
  • GRISMER, L. LEE; & EVAN S. H. QUAH 2019. An updated and annotated checklist of the lizards of Peninsular Malaysia, Singapore, and their adjacent archipelagos. Zootaxa 4545 (2): 230–248 - get paper here
  • Grismer, L.L. 2011. Lizards of Peninsular Malaysia, Singapore and their adjacent archipelagos. Edition Chimaira, Frankfurt, 728 pp. [review in Herp. Rev. 43: 155] - get paper here
  • Liu S, Rao D 2019. A new species of the genus Acanthosaura from Yunnan, China (Squamata, Agamidae). ZooKeys 888: 105-132 - get paper here
  • PAUWELS, OLIVIER S. G.; MONTRI SUMONTHA, KIRATI KUNYA, AWAT NITIKUL, PHAMON SAMPHANTHAMIT, PERRY L. WOOD, JR. & LEE L. GRISMER 2015. Acanthosaura phuketensis (Squamata: Agamidae), a new long-horned tree agamid from southwestern Thailand. Zootaxa 4020 (3): 473–494 - get paper here
  • WOOD JR., PERRY L.;JESSE L. GRISMER, L. LEE GRISMER, NORHAYATI AHMAD, & CHAN KIN ONN & AARON M. BAUER 2009. Two new montane species of Acanthosaura Gray, 1831 (Squamata: Agamidae) from Peninsular Malaysia. Zootaxa 2012: 28-46 - get paper here
  • Wood, P.L.; Grismer, L.L.; Grismer, J.L.; Neang, T.; Chav, T. & holden, J. 2010. A new cryptic species of Acanthosaura Gray, 1831 (Squamata: Agamidae) from Thailand and Cambodia. Zootaxa 2488: 22–38 - get paper here
 
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