Systematics of the Calotes irawadi complex (Squamata, Agamidae) with two newly described species from Thailand Author Prakobkarn, Arpapan https://orcid.org/0009-0000-9047-1102 Ph. D. Program in Zoology, Department of Biology, Faculty of Science, Chulalongkorn University, Bangkok 10330, Thailand Author Zug, George R. 0000-0003-4922-5721 Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560, USA Author Tandavanitj, Nontivich Department of Biology, Faculty of Science, Chulalongkorn University, Bangkok 10330, Thailand Author Ngamprasertwong, Thongchai 0000-0003-3146-1849 Department of Biology, Faculty of Science, Chulalongkorn University, Bangkok 10330, Thailand text ZooKeys 2026 2026-06-03 1281 69 104 journal article 315339 10.3897/zookeys.1281.175455 1321a16e-cff9-4247-8200-6557d35c47d3 E54EA084-1419-444B-B507-C4646540A63B

Table 4. Morphological (mensural) characters of holotype, paratypes and all specimens of Calotes thailandensis sp. nov., C. maehongsonensis sp. nov., C. irawadi and C. wangi. Measurements are represented in mean ± SD (range) in millimeters (mm). Abbreviations are showed in Materials and methods.

Characters	C. thailandensis sp. nov.					C. maehongsonensis sp. nov.					C. irawadi		C. wangi
	Holotype	Paratypes		All specimens		Holotype	Paratypes		All specimens		All specimens		Huang et al. (2023)
	Male	Male (n = 5)	Female (n = 5)	Male (n = 26)	Female (n = 27)	Male	Male (n = 5)	Female (n = 5)	Male (n = 10)	Female (n = 10)	Male (n = 28)	Female (n = 14)	Male (n = 2)	Female (n = 2)
* TailL of male paratypes from C. thailandensis sp. nov. (n = 4) and C. maehongsonensis sp. nov. (n = 2); TailL of female paratypes from C. thailandensis sp. nov. (n = 5) and C. maehongsonensis sp. nov. (n = 4).
SVL	87.24	85.37 ± 1.22	77.08 ± 3.57	87.11 ± 4.80	74.93 ± 4.41	76.16	78.47 ± 1.28	74.51 ± 4.14	78.81 ± 2.48	73.36 ± 4.68	82.59 ± 8.32	77.40 ± 7.91
		(83.87–86.30)	(70.87–79.53)	(80.90–99.74)	(65.30–82.91)		(76.86–79.6)	(70.71–80.80)	(76.16–83.38)	(67.36–80.80)	(66.40–106.80)	(64.30–90.30)	(88.34–96.4)	(82–96.5)
EyeEar	6.25	6.36 ± 0.61	4.46 ± 0.23	6.39 ± 0.52	4.54 ± 0.36	4.8	5.35 ± 0.42	4.42 ± 0.23	5.32 ± 0.41	4.35 ± 0.20	5.76 ± 0.89	4.30 ± 0.60
		(5.63–7.10)	(4.20–4.80)	(5.24–7.28)	(3.98–5.53)		(4.7–5.8)	(4.19–4.74)	(4.70–5.90)	(4.14–4.74)	(4.20–7.90)	(3.10–5.50)	(5.97–6.30)	(5.06–6.17)
InterOrb	8.6	8.93 ± 0.36	7.43 ± 0.24	8.80 ± 0.49	7.33 ± 0.59	7.42	7.88 ± 0.46	6.93 ± 0.72	7.80 ± 0.39	6.89 ± 0.64	9.40 ± 0.91	8.33 ± 1.05
		(8.48–9.35)	(7.14–7.68)	(7.81–9.70)	(6.30–8.73)		(7.14–8.4)	(6.56–8.22)	(7.14–8.40)	(6.38–8.22)	(7.90–11.30)	(6.90–10.70)	(8.51–9.48)	(10.24–12.43)
HeadH	15.25	14.99 ± 0.83	11.49 ± 0.86	14.18 ± 1.26	11.23 ± 0.80	11.66	11.78 ± 0.66	10.93 ± 0.43	11.70 ± 0.53	10.75 ± 0.41	14.82 ± 2.19	12.96 ± 1.55
		(14.28–16.40)	(10.85–12.85)	(11.70–16.40)	(10.00–12.85)		(11.12–12.9)	(10.38–11.30)	(11.10–12.90)	(10.10–11.30)	(11.20–21.50)	(8.90–15.00)	(13.18–14.42)	(12.42–14.42)
HeadW	20.18	19.55 ± 2.61	13.71 ± 0.16	19.69 ± 1.83	13.19 ± 0.81	16.1	15.82 ± 1.89	12.70 ± 0.76	15.68 ± 1.43	12.63 ± 0.83	19.09 ± 3.67	14.31 ± 1.92
		(16.00–21.95)	(13.50–13.90)	(16.00–23.22)	(11.70–14.70)		(13.00–18.30)	(11.80–13.90)	(13.00–18.30)	(11.55–13.90)	(13.30–28.30)	(11.20–17.80)	(15.42–16.09)	(14.21–14.79)
JawW	14.21	13.85 ± 1.05	11.91 ± 0.47	14.41 ± 1.08	11.60 ± 0.81	12.24	12.54 ± 0.26	11.43 ± 0.89	12.56 ± 0.56	11.26 ± 0.78	15.85 ± 2.76	12.58 ± 1.54
		(12.74–15.05)	(11.30–12.38)	(12.40–16.82)	(10.20–13.46)		(12.31–12.97)	(10.70–12.80)	(11.54–13.71)	(10.60–12.80)	(11.70–22.50)	(9.50–15.20)	(16.06–16.14)	(13.47–16.84)
SnW	5.27	5.24 ± 0.35	4.63 ± 0.06	5.36 ± 0.34	4.74 ± 0.44	4.12	4.58 ± 0.23	4.59 ± 0.25	4.56 ± 0.23	4.47 ± 0.32	5.32 ± 0.51	5.18 ± 0.45
		(4.74–5.60)	(4.54–4.68)	(4.74–6.15)	(4.04–5.70)		(4.36–4.92)	(4.36–4.95)	(4.12–4.92)	(3.91–4.95)	(4.30–6.50)	(4.40–5.90)	(5.29–5.87)	(5.33–5.76)
SnN	3.65	3.86 ± 0.20	3.47 ± 0.20	3.93 ± 0.26	3.41 ± 0.27	3	3.43 ± 0.16	3.44 ± 0.34	3.37 ± 0.19	3.36 ± 0.32	4.19 ± 0.63	3.84 ± 0.55
		(3.62–4.10)	(3.23–3.78)	(3.53–4.54)	(2.92–4.09)		(3.23–3.6)	(3.10–3.92)	(3.00–3.60)	(3.04–3.92)	(2.80–5.40)	(2.60–4.70)
SnEye	8.7	8.68 ± 0.42	7.57 ± 0.34	8.83 ± 0.35	7.49 ± 0.44	7.55	7.94 ± 0.46	7.45 ± 0.59	7.88 ± 0.42	7.33 ± 0.61	8.50 ± 0.87	7.81 ± 0.79
		(8.08–9.04)	(7.14–7.93)	(8.08–9.48)	(6.73–8.50)		(7.38–8.5)	(6.66–8.20)	(7.38–8.50)	(6.43–8.20)	(6.90–10.70)	(5.90–9.10)	(7.85–8.82)	(7.91–9.0)
NarEye	4.48	4.78 ± 0.36	3.91 ± 0.19	4.85 ± 0.30	3.96 ± 0.28	4.34	4.47 ± 0.32	4.13 ± 0.32	4.39 ± 0.25	3.97 ± 0.31	4.32 ± 0.41	3.96 ± 0.37
		(4.42–5.37)	(3.70–4.16)	(4.28–5.37)	(3.52–4.57)		(4.05–4.86)	(3.92–4.70)	(4.05–4.86)	(3.58–4.70)	(3.70–5.30)	(3.30–4.50)	(4.49–4.58)	(4.96–5.69)
HeadL	20.95	20.57 ± 1.22	17.25 ± 0.87	21.26 ± 1.18	17.16 ± 1.13	17.87	18.68 ± 0.71	16.68 ± 1.53	18.68 ± 0.75	16.15 ± 1.32	20.03 ± 1.83	17.64 ± 1.83
		(19.16–22.18)	(16.10–18.46)	(19.50–23.52)	(15.37–19.87)		(17.88–19.36)	(14.42–18.44)	(17.87–19.65)	(14.42–18.44)	(16.90–24.90)	(13.90–21.40)	(24.29–24.47)	(23.39–23.93)
SnForeL	30.38	29.14 ± 2.23	21.74 ± 1.20	29.15 ± 1.74	21.73 ± 1.40	23.72	24.58 ± 1.14	21.57 ± 1.70	24.86 ± 1.18	21.20 ± 1.51	29.19 ± 2.24	26.55 ± 3.32
		(25.74–31.20)	(20.44–23.60)	(25.74–32.40)	(19.46–25.08)		(23.1–25.87)	(20.03–24.06)	(23.10–26.70)	(19.40–24.06)	(25.40–33.70)	(20.80–32.10)	(30.74–32.65)	(25.99–30.4)
4 FingLng	10.16	10.27 ± 0.87	8.47 ± 0.34	10.42 ± 0.66	8.36 ± 0.35	8.8	9.73 ± 0.53	8.46 ± 0.63	9.48 ± 0.49	8.19 ± 0.64	10.64 ± 1.01	9.91 ± 0.91
		(9.00–11.17)	(7.97–8.87)	(9.00–11.36)	(7.50–8.92)		(9.15–10.46)	(7.72–9.42)	(8.80–10.46)	(7.32–9.42)	(8.70–12.80)	(8.70–11.50)	(13.41–14.10)	(9.82–13.68)
ForefL	14.92	14.41 ± 1.19	11.89 ± 0.39	14.75 ± 1.14	11.96 ± 0.76	13.15	13.19 ± 0.63	11.61 ± 0.90	13.02 ± 0.55	11.47 ± 0.64	14.95 ± 1.44	13.92 ± 1.14
		(12.93–16.16)	(11.37–12.22)	(12.24–16.75)	(10.04–13.50)		(12.25–13.80)	(10.24–12.74)	(12.24–13.80)	(10.24–12.74)	(12.10–18.70)	(11.60–16.00)
LoArmL	15.66	15.01 ± 0.50	13.17 ± 1.33	14.99 ± 0.74	12.32 ± 0.92	12.03	13.95 ± 0.49	12.50 ± 0.66	13.85 ± 0.75	12.36 ± 0.62	15.83 ± 1.69	13.86 ± 1.50
		(14.54–15.80)	(11.36–15.00)	(13.62–16.80)	(10.30–15.00)		(13.21–14.36)	(11.50–13.16)	(12.03–14.38)	(11.41–13.16)	(13.10–21.10)	(10.10–16.00)	(13.59–14.34)	(13.6–15.11)
UpArmL	17.26	15.68 ± 0.86	12.82 ± 0.66	15.15 ± 1.05	12.16 ± 0.62	13.47	13.90 ± 0.58	12.29 ± 0.45	13.98 ± 0.51	12.03 ± 0.65	17.06 ± 2.03	15.19 ± 2.15
		(14.36–16.63)	(12.26–13.88)	(13.50–17.42)	(11.00–13.88)		(13.29–14.74)	(11.72–12.88)	(13.29–14.76)	(10.90–12.88)	(13.40–21.70)	(11.80–19.90)	(15.8–16.64)	(15.35–16.36)
4 ToeLng	16.4	16.27 ± 0.54	13.23 ± 0.54	16.17 ± 0.81	13.38 ± 0.54	13.8	14.29 ± 1.05	12.96 ± 0.90	14.34 ± 0.83	12.54 ± 0.82	16.27 ± 1.72	14.44 ± 0.86
		(15.68–17.13)	(12.72–14.00)	(14.24–18.00)	(12.54–14.20)		(13.33–15.89)	(11.96–14.11)	(13.33–15.89)	(11.56–14.11)	(13.80–20.70)	(13.30–16.00)	(19.04–21.57)	(15.36–17.97)
HindfL	25.92	26.04 ± 0.65	21.63 ± 0.50	25.87 ± 1.46	21.61 ± 0.98	21.02	23.11 ± 1.47	20.98 ± 1.64	23.26 ± 1.43	20.44 ± 1.33	26.83 ± 2.29	24.52 ± 1.59
		(25.19–26.65)	(21.09–22.36)	(22.80–28.80)	(19.76–23.64)		(21.48–25.12)	(18.80–23.18)	(21.02–25.20)	(18.80–23.18)	(23.00–34.10)	(21.00–26.50)
CrusL	19.18	19.20 ± 0.72	16.13 ± 0.68	18.94 ± 0.88	15.75 ± 0.73	16.72	17.51 ± 0.53	15.80 ± 0.88	17.71 ± 0.86	15.50 ± 0.85	19.45 ± 1.67	17.70 ± 1.38
		(18.56–20.30)	(15.10–16.90)	(17.33–20.85)	(14.16–17.35)		(16.74–18.21)	(14.86–17.07)	(16.72–19.53)	(14.33–17.07)	(16.50–24.80)	(14.00–19.40)	(16.87–20.26)	(17.38–18.08)
UpLegL	21.86	21.61 ± 0.81	18.39 ± 0.64	21.05 ± 1.10	17.77 ± 0.94	19.2	19.37 ± 0.38	17.44 ± 1.12	19.40 ± 0.70	17.25 ± 0.82	20.16 ± 1.95	18.12 ± 1.52
		(20.78–22.78)	(17.36–19.00)	(18.54–22.78)	(15.98–20.00)		(19.04–19.84)	(16.07–18.73)	(18.23–20.94)	(16.07–18.73)	(16.80–25.50)	(14.50–20.90)	(21.94–23.01)	(17.27–20.76)
TrunkL	42.93	39.87 ± 0.88	37.80 ± 2.66	40.97 ± 2.97	36.08 ± 2.74	34.54	36.80 ± 1.40	35.93 ± 2.41	36.76 ± 2.14	35.59 ± 3.27	41.67 ± 5.41	40.61 ± 4.34
		(38.75–40.79)	(33.26–40.20)	(37.27–49.10)	(30.16–40.51)		(35.48–38.78)	(33.26–38.53)	(34.24–41.42)	(31.14–39.73)	(31.70–56.90)	(32.10–44.90)	(43.82–49.82)	(40.81–47.02)
PectW	14.74	15.35 ± 1.48	12.18 ± 0.35	15.28 ± 1.48	11.81 ± 0.95	13.38	13.37 ± 0.91	11.90 ± 0.64	13.52 ± 0.94	12.05 ± 1.02	14.61 ± 2.45	12.28 ± 1.68
		(12.90–16.72)	(11.75–12.56)	(12.90–19.76)	(10.12–13.93)		(12.45–14.38)	(11.10–12.86)	(12.44–15.33)	(11.07–14.40)	(9.80–21.40)	(9.50–14.70)
PelvW	14.26	13.28 ± 0.67	11.31 ± 0.99	13.16 ± 0.93	11.27 ± 0.99	11.33	11.46 ± 0.91	11.61 ± 1.06	11.38 ± 0.67	11.52 ± 0.87	8.59 ± 0.98	8.15 ± 0.87
		(12.58–14.09)	(10.15–12.54)	(11.62–15.07)	(9.51–13.44)		(10.46–12.95)	(10.20–12.88)	(10.46–12.95)	(10.20–12.88)	(6.90–10.90)	(6.80–9.30)
TailH	13.35	12.56 ± 0.52	6.19 ± 0.33	12.41 ± 1.13	6.18 ± 0.59	10.72	11.12 ± 0.73	6.19 ± 0.43	11.48 ± 0.83	6.23 ± 0.48	10.11 ± 1.43	7.37 ± 1.48
		(12.12–13.38)	(5.82–6.54)	(9.55–14.83)	(5.34–7.30)		(10.05–11.79)	(5.70–6.62)	(10.05–12.74)	(5.37–6.84)	(7.80–14.20)	(4.70–10.20)
TailW	11.61	11.27 ± 0.72	9.34 ± 0.30	10.78 ± 0.85	8.73 ± 0.94	9.37	10.36 ± 0.65	8.51 ± 0.59	10.32 ± 0.68	8.72 ± 0.58	9.69 ± 1.28	6.94 ± 1.34
		(10.62–12.32)	(9.00–9.81)	(9.26–12.92)	(7.02–10.87)		(9.30–11.02)	(7.78–9.40)	(9.30–11.40)	(7.78–9.63)	(7.30–13.40)	(5.30–10.50)
TailL *	253	250.8 ± 4.7	224 ± 14.6	254.9 ± 11.2	212.30 ± 17.47	210	222.0 ± 15.6	188.5 ± 16.2	221.9 ± 10.5	186.8 ± 15.4	231.52 ± 20.73	211.4 ± 21.5
		(24 5–254)	(215–250)	(236–278)	(174–250)		(211–233)	(175–212)	(210–233)	(170–212)	(191–288)	(182–250)	(256.15–283.90)	(243.5–330)