Morphometric Analysis of Three Draco Lizard Species (D. Blanfordii, D

Kasetsart J. (Nat. Sci.) 48 : 383 - 402 (2014)

Morphometric Analysis of Three Draco Lizard Species (D. blanfordii, D. maculatus, and D. taeniopterus) (Squamata: Agamidae) from Thailand

Nattawut Srichairat1, Pattanee Jantrarotai1,*, Prateep Duengkae2 and Yodchaiy Chuaynkern3

ABSTRACT

Two hundred sixty-seven preserved specimens of three Draco lizard species from Thailand consisting of D. blanfordii (17 females and 70 males), D. maculatus (58 females and 79 males) and D. taeniopterus (13 females and 30 males) were examined using 21 morphometric characters and 25 character ratios to assess morphometric differences. The results of univariate analysis showed that there were 12 morphometric characters and 2 character ratios that can be used for discrimination of the three Draco species regardless of sexual differences. Among these three species, it was found that 13 morphometric characters and 7 character ratios and 15 morphometric characters and 8 character ratios can be used to discriminate female and male lizards, respectively. The results of cluster analysis produced a dendrogram of the relationships among the three species and revealed that the three Draco species were divided into two morphometric clusters at a level of 0% of information remaining in which the D. blanfordii cluster was distinctly different from the D. maculatus and D. taeniopterus cluster. Keywords: morphometric analysis, Draco, D. blanfordii, D. maculatus, D. taeniopterus

INTRODUCTION Musters (1983) and Nabhitabhata et al. (2000) reported there are nine species of Draco lizards The agamid lizard genus Draco is consisting of: D. blanfordii, D. fimbriatus, D. the most remarkable and successful of gliding haematopogon, D. maculatus, D. melanopogon, vertebrates. It is well-known as a flying lizard D. obscurus, D. quinquefasciatus, D. taeniopterus or flying dragon due to its ability to glide long and D. volans. The three commonly found Draco distances between trees using its wing-like patagial lizard species—D. blanfordii, D. maculatus and membrane, supported by elongated ribs or rib-like D. taeniopterus—are distributed in many regions dermal structures (McGuire, 1998; McGuire and of Thailand, with Draco blanfordii distributed Dudley, 2011). The habitats of Draco lizards are from Northern to Southern Thailand, whereas, D. large, tall and thick forest with climbing sites, maculatus and D. taeniopterus are distributed in open areas for gliding and sheltered areas for all regions (Musters, 1983). hiding (Mori and Hikida, 1994). In Thailand,

1 Department of Zoology, Faculty of Science, Kasetsart University, Bangkok 10900, Thailand. 2 Department of Forest Biology, Faculty of Forestry, Kasetsart University, Bangkok 10900, Thailand. 3 Department of Biology, Faculty of Science, Khon Kaen University, Khon Kaen 40002, Thailand. * Corresponding author, e-mail: [email protected] Received date : 21/11/13 Accepted date : 25/04/14 384 Kasetsart J. (Nat. Sci.) 48(3)

Morphometric analysis is the quantitative may be used as a diagnostic tool to discriminate analysis of form and is commonly used in research among the three Draco species. to characterize the pattern of morphological variation of organisms (Rohlf, 1990; Ljubisavljevic MATERIALS AND METHODS et al., 2010). In Draco lizards, morphometric analysis has been used for the identification and In total, 267 specimens were inspected description of organisms as well as being used to that had been preserved in 70% ethanol and study morphological variation in the population were housed in the collections of the Thailand (Inger, 1983; Musters, 1983). However, there is Natural History Museum and the National Science no reported usage of morphometric analysis to Museum, Pathum Thani, Thailand. Specimens discriminate Draco lizard species in Thailand. were collected from 1967 to 2012. Eighty seven Therefore, the objective of this study was to individuals of D. blanfordii (DBl, 17 females and analyze the morphometric difference among the 70 males), 137 individuals of D. maculatus (DMa, three Draco lizard species—D. blanfordii, D. 58 females and 79 males) and 43 individuals of maculatus and D. taeniopterus. Furthermore, the D. taeniopterus (DTa, 13 females and 30 males) differences in the morphometric characters of were measured for 21 morphometric characters to males and females among the three Draco species the nearest 0.05 mm using a set of vernier calipers were also analyzed. The results from this study (Figure 1). The measurements were modified

Figure 1 Snout-vent length (SVL), snout-forelimb length (SFL), forelimb length (FL), axilla-groin length (AGL), tail length (TaiL), cloacal length (CL), head width (HW), internarial distance (ID), head length (HL), head depth (HD), mouth length (ML), dewlap length (DeL), snout to nostril (SN), distance between nostril to anterior edge of eye (DBNE), distance between posterior edge of eye to anterior edge of tympanum (DBET), eye length (EL), snout to anterior edge of eye (snout length 1(SL1)), snout to posterior edge of eye (snout length 2 (SL2)), snout to anterior edge of tympanum (ST), tympanum diameter (vertical) (TDV), and tympanum diameter (horizontal) (TDH). The figure is modified from Musters (1983) and Stebbins (2003). Kasetsart J. (Nat. Sci.) 48(3) 385 from Taylor (1963), Inger (1983), and Musters All morphometric characters and (1983). There were three sets of characters in this character ratios of the three Draco lizard species study: external morphology, head characters, and and the morphological difference of females and the ratio of characters (Table 1). Morphological males were analyzed using univariate analysis differences among females and among males of which consisted of the mean, the 95% confidence the three Draco species were measured according intervals for the mean and one-way analysis of to the characters described above. variance followed by Duncan’s post-hoc test

Table 1 External morphology, head characters and character ratios examined in 267 specimens of three Draco lizard species from Thailand. External morphology - Snout-vent length (SVL): the tip of the snout to the vent - Snout-forelimb length (SFL): the tip of the snout to the shoulder - Forelimb length (FL): the left side from the axilla to the tip of the fourth finger when extended at right angles - Axilla-groin length (AGL): the axilla to the anterior edge of hindlimb at its insertion into the body - Tail length (TaiL): the vent to the tip of the tail - Cloacal length (CL): the broadest part at the level of the cloacal opening Head characters - Head width (HW): the widest part of the head - Internarial distance (ID): across the nostrils at the snout - Head length (HL): the tip of the snout to the posterior margin of the jaw - Head depth (HD): the deepest part of the head - Mouth length (ML): the tip of the snout to the mouth corner - Dewlap length (DeL): maximally extended, from the ventral edge of the lower jaw to the tip of the dewlap - Snout to nostril (SN): the tip of the snout to the anterior edge of the nostril - Distance between nostril to anterior edge of eye (DBNE): the anterior edge of the nostril to the anterior edge of the eye - Distance between posterior edge of eye to anterior edge of tympanum (DBET): the posterior edge of the eye to the anterior edge of the tympanum - Eye length (EL): the anterior edge of the orbit to the posterior edge of the orbit - Snout to anterior edge of eye (snout length 1(SL1)): the tip of the snout to the anterior edge of the eye - Snout to posterior edge of eye (snout length 2 (SL2)): the tip of the snout to the posterior edge of the eye - Snout to anterior edge of tympanum (ST): the tip of the snout to the anterior edge of the tympanum - Tympanum diameter (vertical) (TDV): the dorsal edge of the tympanum to the ventral edge of the tympanum - Tympanum diameter (horizontal) (TDH): the anterior edge of the tympanum to the posterior edge of the tympanum Character ratios - SFL/SVL, FL/SVL, AGL/SVL, TaiL/SVL, CL/SVL, HW/SVL, ID/SVL, HL/SVL, HD/SVL, ML/ SVL, DeL/SVL, SN/SVL, DBNE/SVL, DBET/SVL, EL/SVL, SL1/SVL, SL2/SVL, ST/SVL, TDV/ SVL, TDH/SVL, FL/AGL, HW/HL, HD/HW, HD/HL and DeL/HL 386 Kasetsart J. (Nat. Sci.) 48(3) using the software SPSS (version 16; SPSS Inc.; DeL/HL. Chicago, IL, USA). The relationships among the Discrimination of males among the three Draco lizard species and for the females three Draco species is shown by the values of and males were determined on the basis of morphometric characters and character ratios in morphometric characters and character ratios using Tables 7 and 8. There was a significant difference cluster analysis according to Jaccard and Ward’s in 15 morphometric characters—SFL, FL, TaiL, method in the software program PC-ORD (version HW, ID, HL, HD, ML, SN, DBNE, DBET, SL1, 5.10; MJM Software Design; Gleneden Beach, SL2, ST and TDH—and 8 character ratios—FL/ OR, USA) in the form of a dendrogram. AGL, CL/SVL, HD/SVL, DeL/SVL, SL1/SVL, SL2/SVL, TDH/SVL and DeL/HL. RESULTS AND DISCUSSION Therefore, to discriminate the three Draco species regardless of sexual differences, Univariate analysis there are 12 suitable morphometric characters— The mean values, 95% confidence FL, TaiL, HW, HL, HD, ML, SN, DBET, SL1, intervals for the mean and significance levels SL2, ST and TDH—and 2 character ratios—HD/ (P < 0.05) of the morphometric characters and SVL and SL2/SVL (Table 4). Furthermore, 12 of character ratios are shown in Tables 2 and 3. the morphometic characters and 2 character ratios The results showed significant differences in 21 mentioned above can be used to discriminate morphometric characters as well as in the character females among the three Draco species in addition ratios which also showed significant differences to another 6 characters—DeL, HW/SVL, DeL/ in most characters except for the ratio of DeL/HL. SVL, ST/SVL, HD/HW and DeL/HL (Table 5 and Most of the morphometric characters showed DBl 6). Similarly, the males among three Draco species having the highest values, followed by the values can be discriminated using another 9 characters of DMa and DTa. —SFL, ID, DBNE, FL/AGL, CL/SVL, DeL/SVL, There is no previous report on the use SL1/SVL, TDH/SVL and DeL/HL (Tables 7 and of morphometric characters to separate the three 8). Draco species in Thailand. However, on the basis In practical use, the morphometric of this study, there are 12 morphometric characters characters may not be appropriate for the and 2 character ratios—FL, TaiL, HW, HL, HD, identification of the threeDraco species. However, ML, SN, DBET, SL1, SL2, ST, TDH, HD/SVL and they can be used as an additional or diagnostic SL2/SVL—that can be used to distinguish species tool to support the qualitative characters in the as shown in Table 4. three Draco lizard species. Furthermore, the The 95% confidence interval for the morphometric characters can be used to describe mean were narrow for most of the morphometric the size of the three Draco species. The present characters and character ratios except for the TaiL study showed that DBl is the largest species character (Table 4). and this is consistent with Taylor (1963), Inger Discrimination of females among the (1983), and Musters (1983) while DMa and DTa three Draco species is shown by the values of were classified as moderate and small species, morphometric characters and character ratios in respectively (Taylor, 1963). Tables 5 and 6. There was a significant difference The 95% confidence interval for the mean in 13 morphometric characters—FL, TaiL, HW, showed the TaiL character had a wide confidence HL, HD, ML, DeL, SN, DBET, SL1, SL2, ST and interval. Therefore, it would not be a good TDH—and 7 character ratios—HW/SVL, HD/ character to use for determining the differences SVL, DeL/SVL, SL2/SVL, ST/SVL, HD/HW and among the three Draco species. Kasetsart J. (Nat. Sci.) 48(3) 387 value − P D. blanfordii Upper bound mean Draco species: 4.230.320.470.54 4.640.86 0.331.85 0.49 0.000 0.06 0.56 0.000 0.89 0.000 1.91 0.004 0.07 0.000 0.000 0.000 70.1022.9533.56 72.5038.32 23.90 34.70 0.000 39.89 0.000 0.000 0.000 bound Lower 131.11 136.73 0.000 (n = 43) DTa 95% Confidence interval for b b b b b c a a b a a b Mean Upper bound mean 5.020.330.410.55 5.270.74 0.331.47 0.420.07 4.44 0.56 0.33 0.76 0.48 1.52 0.55 0.07 0.87 1.88 0.06 72.0523.8229.9039.84 74.32 24.56 30.90 71.30 41.21 23.42 34.13 39.10 bound Lower 106.90 111.60 133.92 DMa (n = 137) 95% Confidence interval for b b c b c b a b b b b a Mean Upper bound mean < 0.05). P 5.300.320.410.56 5.650.72 0.331.84 0.420.05 5.15 0.57 0.33 0.75 0.42 1.89 0.55 0.06 0.75 1.49 0.07 98.1731.5640.59 104.0555.01 33.42 73.19 42.72 58.84 24.19 30.40 40.52 bound Lower 182.99 192.82 109.25 DBl (n = 87) 95% Confidence interval for a a a a a b b a b a c a Mean Mean, 95% Confidence interval for means and one-way analysis of variance of external morphology in three Thailand. ) from ( DBl ), D. maculatus DMa ) and taeniopterus DTa FL 41.65 CL 5.48 SFL 32.49 TaiL 187.90 SVL 101.11 AGL 56.92 FL/SVL 0.41 CL/SVL 0.05 FL/AGL 0.74 SFL/SVL 0.32 Character TaiL/SVL 1.87 AGL/SVL 0.56 Morphometric Table 2 Table superscripts with in different the = Values Cloacal length. length; CL Tail = TaiL Axilla-groin length; = AGL = = = Forelimb Snout-forelimb Snout-vent length; length; length; FL SFL SVL ( same row indicate a significant difference 388 Kasetsart J. (Nat. Sci.) 48(3) DBl ), P - value ( D. blanfordii Upper bound mean 7.391.826.61 7.76 1.981.36 0.000 6.883.45 0.000 2.645.77 0.000 1.474.80 3.649.84 2.81 0.000 5.981.58 0.000 5.05 10.261.29 0.000 0.10 0.000 0.03 0.000 1.73 0.000 0.19 1.41 0.11 0.000 0.03 0.000 0.19 0.000 0.000 0.000 Draco species: 11.41 11.93 0.000 13.2410.17 13.7816.69 10.62 0.000 20.91 0.000 0.000 bound Lower (n = 43) DTa 95% Confidence interval for c c c c c c c c b c c c b c b b c b Mean Upper bound mean 8.662.727.32 8.92 2.82 7.58 1.74 7.50 1.90 3.653.24 6.74 5.82 1.835.59 3.78 3.35 1.41 5.94 3.54 1.87 5.77 2.73 1.54 5.87 0.12 4.92 0.04 1.970.21 1.61 0.12 1.66 0.04 1.35 0.21 0.11 0.03 0.19 11.46 11.76 10.40 14.9819.96 15.36 13.51 22.70 18.80 10.7012.89 10.97 13.2 10.05 11.67 bound Lower DMa (n = 137) 95% Confidence interval for b a b b b b b b b b b b b a b a a a Mean Upper bound mean 2.498.75 2.692.10 9.12 2.77 5.103.92 7.41 7.18 2.257.20 5.42 4.13 1.78 7.43 3.72 1.86 7.63 3.29 1.70 5.88 0.11 5.68 0.02 2.020.19 1.86 0.11 1.92 0.03 1.57 0.19 0.12 0.04 0.21 18.9014.00 19.8824.35 14.69 15.17 27.38 11.61 21.33 12.9716.36 13.55 17.17 10.83 13.04 10.47 11.05 8.79 bound Lower DBl (n = 87) 95% Confidence interval for a b a a a a a a a a a a a a a b b b Mean Means, 95% confidence intervals for means and one-way analysis of variance of head characters in three ) from Thailand. ) from D. maculatus ( DMa ) and taeniopterus DTa ID 2.59 ST 16.76 EL 7.31 SN 2.17 HL 19.39 HD 8.93 ML 14.35 HW 10.76 SL1SL2 7.42 13.26 DeL 25.86 TDV 1.94 TDH 1.78 DBET 4.03 DBNE 5.26 ID/SVL 0.03 HL/SVL 0.19 character HW/SVL 0.11 Morphometric Table 3 Table Kasetsart J. (Nat. Sci.) 48(3) 389 P - value Upper bound mean 0.090.140.23 0.100.02 0.150.05 0.290.04 0.000 0.020.08 0.000 0.050.07 0.019 0.040.14 0.000 0.080.16 0.009 0.070.02 0.000 0.140.02 0.000 0.170.55 0.000 0.020.88 0.000 0.020.49 0.000 0.57 0.000 0.91 0.000 0.51 0.000 0.000 0.000 bound Lower (n = 43) DTa 95% Confidence interval for b b ab b b b a c b b b b b a a Mean Upper bound mean 0.100.160.27 0.100.02 0.160.05 0.31 0.09 0.04 0.02 0.15 0.08 0.05 0.26 0.08 0.05 0.02 0.15 0.08 0.05 0.18 0.08 0.04 0.03 0.15 0.08 0.02 0.18 0.07 0.57 0.03 0.14 0.84 0.02 0.16 0.48 0.59 0.02 0.86 0.02 0.49 0.56 0.89 0.50 bound Lower < 0.05). DMa (n = 137) 95% Confidence interval for a a a a ab a a a a a a a a b b Mean Upper bound mean 0.090.140.24 0.090.02 0.140.05 0.27 0.10 0.04 0.02 0.16 0.07 0.05 0.29 0.07 0.04 0.02 0.13 0.07 0.05 0.16 0.07 0.05 0.02 0.13 0.08 0.02 0.17 0.08 0.55 0.02 0.15 0.82 0.02 0.18 0.46 0.56 0.03 0.85 0.02 0.47 0.58 0.85 0.49 bound Lower DBl (n = 87) 95% Confidence interval for 95% Confidence interval c b b b a b b b c b c c b b c Mean (Continued) HD/HL 0.46 HW/HL 0.56 ST/SVL 0.17 EL/SVL 0.07 DeL/HL 1.33 1.26 1.40 1.40 1.31 1.48 1.39 1.24 1.55 0.575 HD/HW 0.84 SN/SVL 0.02 HD/SVL 0.09 ML/SVL 0.14 character SL1/SVLSL2/SVL 0.07 0.13 DeL/SVL 0.26 TDV/SVLTDH/SVL 0.02 0.02 DBET/SVL 0.04 DBNE/SVL 0.05 Morphometric Table 3 Table = HW Head width; ID = Internarial = distance; Head HL length; HD = Head = depth; Mouth = ML length; Dewlap DeL length; SN = Snout to nostril; DBNE = Distance between nostril to anterior edge of eye; = DBET Distance between posterior edge of eye to anterior = edge Eye of length; tympanum; SL1 EL = Snout to anterior edge of eye (snout length 1); SL2 = Snout to length. Snout-vent = SVL (horizontal); diameter Tympanum = TDH (vertical); diameter Tympanum = TDV tympanum; of edge anterior to Snout = ST 2); length (snout eye of edge posterior ( P superscripts in the same row indicate a significant difference with different Values 390 Kasetsart J. (Nat. Sci.) 48(3) ) DTa ( P - value Upper bound D. taeniopterus mean < 0.05). 7.396.61 7.761.36 0.000 6.882.644.80 0.000 1.479.84 2.81 0.000 5.051.29 10.26 0.000 0.09 0.000 0.14 0.000 1.41 0.10 0.000 0.14 0.000 0.000 11.41 11.93 0.000 33.56 34.7013.24 0.000 10.17 13.78 0.000 10.62 0.000 DMa ) and bound Lower 131.11 136.73 0.000 (n = 43) DTa 95% Confidence interval for b b c c c c c c c c c c b b Mean D. maculatus ( DBl ), Upper bound mean D. ( blanfordii 8.667.32 8.921.74 7.503.24 7.58 5.59 1.83 6.74 3.35 5.771.54 1.41 0.10 2.73 0.15 4.92 1.61 0.10 0.15 1.35 0.09 0.14 11.46 11.76 10.40 29.90 30.90 34.13 14.98 15.36 13.51 10.7012.89 10.97 13.20 10.05 11.67 bound Lower 106.90 111.60 133.92 DMa (n = 137) 95% Confidence interval for interval Confidence 95% c c b b b b b b b b b b a a Draco species: Mean Upper bound mean 8.752.10 9.123.927.20 2.25 7.41 4.13 7.631.70 1.78 0.09 3.29 0.13 5.68 1.86 0.09 0.13 1.57 0.10 0.15 40.59 42.72 30.40 10.4718.90 11.0514.00 19.88 8.79 14.69 15.17 12.97 11.61 16.36 13.55 17.17 10.83 13.04 bound Lower 182.99 192.82 109.25 DBl (n = 87) 95% Confidence interval for a a a a a a a a a a a a c c Mean Thailand regardless of sexual difference. from Morphometric Morphometric characters used to discriminate three FL 41.65 ST 16.76 SN 2.17 HL 19.39 HD 8.93 ML 14.35 HW 10.76 SL1SL2 7.42 13.26 TaiL 187.90 TDH 1.78 DBET 4.03 HD/SVL 0.09 character SL2/SVL 0.13 Morphometric Table 4 Table FL = Forelimb length; TaiL = Tail length; HW = Head width; HL = Head length; HD = Head depth; ML = Mouth = length; Head SN length; = = HD Snout Distance = to between Head nostril; posterior = depth; DBET edge Head ML of width; length; HL HW Tail = TaiL = Forelimb length; FL TDH tympanum; of edge anterior to Snout = ST 2); length (snout eye of edge posterior to Snout = SL2 1); length (snout eye of edge anterior to Snout = SL1 tympanum; of edge anterior to eye ( P superscripts in the same row indicate a significant difference with different Values = Snout-vent length . diameter (horizontal); SVL Tympanum = Kasetsart J. (Nat. Sci.) 48(3) 391 P - value Draco lizard Upper bound mean 4.240.47 4.960.83 0.027 1.83 0.490.06 0.000 0.89 1.94 0.000 0.07 0.000 0.000 68.1022.3933.07 73.0038.05 24.41 0.000 35.04 0.000 41.49 0.000 0.000 bound Lower 127.67 137.99 0.000 (n = 13) DTa 95% Confidence interval for b b b b b b a a a a Mean Upper bound mean 4.840.41 5.290.73 0.42 4.60 1.440.07 0.75 0.48 1.50 0.07 0.86 1.89 0.07 69.6022.9528.92 72.6139.12 23.96 70.55 30.26 23.40 40.97 34.05 39.77 bound Lower 101.87 107.02 132.83 DMa (n = 58) 95% Confidence interval for b c b c ab b b b a b Mean Upper bound mean < 0.05). P 4.930.41 6.020.711.82 0.43 5.06 0.05 0.77 0.42 1.91 0.06 0.74 1.47 0.07 93.1830.6139.54 105.6852.26 34.96 71.11 43.82 23.45 60.80 29.59 40.05 bound Lower 173.80 196.18 104.44 DBl (n = 17) 95% Confidence interval for a a a a a a b b a b Mean ) from Thailand. ) from ( DBl ), D. maculatus DMa ) and taeniopterus DTa species: D. blanfordii Means, 95% confidence intervals for means and one-way analysis of variance of external morphology in females among three FL 41.68 CL 5.47 SFL 32.79 TaiL 184.99 SVL 99.43 AGL 56.53 FL/SVL 0.42 CL/SVL 0.05 FL/AGL 0.74 character SFL/SVL 0.33 0.32 0.34 0.33 0.33 0.33 0.33 0.32 0.34 0.677 TaiL/SVL 1.86 AGL/SVL 0.57 0.55 0.58 0.56 0.56 0.57 0.56 0.55 0.58 0.699 Morphometric Table 5 Table superscripts with in different the = Values Cloacal length. length; CL Tail = TaiL Axilla-groin length; = AGL = = = Forelimb Snout-forelimb Snout-vent length; length; length; FL SFL SVL ( same row indicate a significant difference 392 Kasetsart J. (Nat. Sci.) 48(3) lizard P - value Draco Upper bound mean 7.31 8.13 0.000 1.796.509.89 2.128.301.28 7.12 0.000 10.963.302.57 9.76 0.000 0.000 5.67 1.484.69 3.73 0.000 9.53 2.86 0.000 6.16 0.000 1.58 5.24 0.000 10.571.19 0.000 0.10 0.000 0.000 0.03 1.960.19 1.48 0.11 0.016 0.03 0.000 0.20 0.000 0.000 0.000 13.00 14.24 0.000 10.99 12.16 0.000 bound Lower (n = 13) DTa 95% Confidence interval for c b c c c c c b c b c c c b c b b c Mean Upper bound mean 8.54 8.97 7.72 2.697.11 2.84 7.42 1.95 1.673.51 6.81 3.14 1.795.77 3.745.42 3.28 1.38 5.96 3.52 5.70 2.72 1.84 5.91 1.52 4.97 0.12 1.990.04 1.630.21 0.13 1.77 0.04 1.34 0.21 0.11 0.03 0.19 11.21 11.69 10.43 14.63 15.2412.18 13.62 13.20 9.03 10.4712.61 10.92 13.12 10.05 11.57 bound Lower DMa (n = 58) 95% Confidence interval for b a b b b b b b b b b b b ab b a a a Mean Upper bound mean 2.548.40 3.00 2.77 9.161.934.82 7.27 3.71 2.347.05 5.626.84 1.73 4.27 3.63 7.63 3.21 7.881.91 5.86 1.76 5.56 0.11 2.280.03 2.200.19 1.91 0.12 1.57 0.03 0.12 0.20 0.04 0.21 10.59 11.89 8.75 18.3913.69 20.8313.58 14.94 15.40 15.78 11.45 12.69 12.6015.86 13.92 17.85 10.70 12.86 bound Lower DBl (n = 17) 95% Confidence interval for a a a a a a a a a a a a a a a b b b Mean ) from Thailand. ) from ( DBl ), D. maculatus DMa ) and taeniopterus DTa species: D. blanfordii Means, 95% confidence intervals for means and one-way analysis of variance of head characters in females among three ID 2.77 ST 16.85 EL 7.34 SN 2.14 HL 19.61 HD 8.78 ML 14.54 HW 11.24 SL1SL2 7.36 13.26 DeL 14.68 TDV 2.09 TDH 1.98 DBET 3.99 DBNE 5.22 ID/SVL 0.03 HL/SVL 0.20 HW/SVL 0.11 characters Morphometric Table 6 Table Kasetsart J. (Nat. Sci.) 48(3) 393 P - value Upper bound mean 0.090.140.12 0.100.02 0.15 0.14 0.000 0.04 0.02 0.000 0.08 0.000 0.07 0.04 0.005 0.14 0.090.16 0.07 0.000 0.02 0.15 0.000 0.02 0.17 0.000 0.03 0.000 0.86 0.02 0.000 0.49 0.000 0.62 0.91 0.040 0.51 0.71 0.000 0.000 0.000 bound Lower (n = 13) DTa 95% Confidence interval for b b c b b a b b c a b a a c Mean Upper bound mean 0.100.160.17 0.100.02 0.16 0.180.04 0.10 0.020.08 0.15 0.08 0.13 0.050.15 0.02 0.080.18 0.080.03 0.04 0.150.02 0.08 0.18 0.07 0.030.82 0.14 0.020.48 0.16 0.82 0.02 0.85 0.02 0.49 0.88 0.88 0.50 0.66 bound Lower DMa (n = 58) 95% Confidence interval for < 0.05). a a a a a a a a a a a b a a Mean Upper bound mean 0.090.140.14 0.100.02 0.15 0.160.04 0.10 0.020.07 0.16 0.07 0.18 0.040.13 0.02 0.080.17 0.080.02 0.04 0.140.02 0.08 0.17 0.08 0.020.75 0.15 0.020.43 0.18 0.70 0.03 0.82 0.02 0.47 0.80 0.83 0.49 0.85 bound Lower DBl (n = 17) 95% Confidence interval for c b b b b b b c b b ab c b b Mean (Continued) HD/HL 0.45 HW/HL 0.57 0.56 0.59 0.59 0.58 0.59 0.57 0.55 0.58 0.101 ST/SVL 0.17 DeL/HL 0.75 EL/SVL 0.07 HD/HW 0.79 SN/SVL 0.02 HD/SVL 0.09 ML/SVL 0.15 character SL1/SVLSL2/SVL 0.07 0.13 DeL/SVL 0.15 TDV/SVLTDH/SVL 0.02 0.02 DBET/SVL 0.04 DBNE/SVL 0.05 0.05 0.05 0.05 0.05 0.05 0.05 0.05 0.05 0.447 Morphometric Table 6 Table = HW Head width; ID = Internarial = distance; Head HL length; HD = Head = depth; Mouth = ML length; Dewlap DeL length; SN = Snout to nostril; DBNE = Distance between nostril to anterior edge of eye; = DBET Distance between posterior edge of eye to anterior = edge Eye of length; tympanum; SL1 EL = Snout to anterior edge of eye (snout length 1); SL2 = Snout to length. Snout-vent = SVL (horizontal); diameter Tympanum = TDH (vertical); diameter Tympanum = TDV tympanum; of edge anterior to Snout = ST 2); length (snout eye of edge posterior ( P superscripts in the same row indicate a significant difference with different Values 394 Kasetsart J. (Nat. Sci.) 48(3) P - value Draco lizard Upper bound mean 4.100.320.47 4.630.54 0.330.86 0.49 0.000 1.84 0.55 0.000 0.06 0.90 0.000 1.91 0.000 0.07 0.000 0.000 0.000 70.1822.8833.42 73.0737.92 23.99 34.89 0.000 39.71 0.000 0.000 0.000 bound Lower 130.85 137.94 0.000 (n = 30) DTa 95% Confidence interval for b c b b b b a a b a a b Mean Upper bound mean 5.060.330.41 5.360.54 0.340.75 0.42 4.37 1.47 0.55 0.33 0.07 0.77 0.48 1.55 0.54 0.07 0.88 1.88 0.06 73.1424.2230.30 76.2939.89 25.23 71.62 31.69 23.44 41.86 34.16 38.81 bound Lower 109.33 116.22 134.39 DMa (n = 79) 95% Confidence interval for b b c b c a a b b b b a Mean Upper bound mean < 0.05). P 5.300.320.41 5.660.55 0.320.72 0.421.83 5.21 0.570.05 0.33 0.75 0.42 1.90 0.55 0.06 0.76 1.51 0.07 98.1331.3640.41 104.9154.83 33.47 74.71 42.89 24.72 59.21 30.99 40.88 bound Lower 183.01 194.21 112.78 DBl (n = 70) 95% Confidence interval for a a a a a a b b a c a c Mean Means, 95% confidence intervals for means and one-way analysis of variance of external morphology in males among three ) from Thailand. ) from ( DBl ), D. maculatus DMa ) and taeniopterus DTa species: D. blanfordii FL 41.65 CL 5.48 SFL 32.41 TaiL 188.61 SVL 101.52 AGL 57.02 FL/SVL 0.41 CL/SVL 0.05 FL/AGL 0.74 character SFL/SVL 0.32 TaiL/SVL 1.87 AGL/SVL 0.56 Morphometric SVL = Snout-vent length; SFL = Snout-forelimb length; FL = Forelimb length; AGL = Axilla-groin length; TaiL = Tail length; CL = Cloacal length. Values with different superscripts with in different the = Values Cloacal length. length; CL Tail = TaiL Axilla-groin length; = AGL = = = Forelimb Snout-forelimb Snout-vent length; length; length; FL SFL SVL ( same row indicate a significant difference Table 7 Table Kasetsart J. (Nat. Sci.) 48(3) 395 P - value Upper bound Draco lizard species: mean 7.301.78 7.726.56 1.97 0.000 0.000 6.871.363.45 0.000 2.62 1.505.75 3.664.76 0.000 2.849.82 0.000 5.97 0.000 5.051.54 10.28 0.000 1.29 0.000 0.10 0.000 1.680.02 1.420.18 0.000 0.11 0.000 0.03 0.000 0.19 0.000 0.000 11.41 12.02 0.000 13.1510.14 13.7722.10 0.000 10.64 23.96 0.000 0.000 bound Lower (n = 30) DTa 95% Confidence interval for c c c c c b c c c b c c c b c b b b Mean Upper bound mean 8.662.72 8.977.40 2.84 7.51 1.87 7.631.763.70 6.72 3.29 1.895.81 3.875.65 1.43 3.42 3.56 5.97 2.73 5.881.86 5.86 1.52 4.91 0.12 2.000.04 1.620.20 1.61 0.12 1.36 0.04 0.11 0.21 0.03 0.19 11.55 11.91 10.39 15.11 15.5726.80 13.46 28.55 23.03 10.7612.98 11.11 13.36 10.05 11.72 bound Lower DMa (n = 79) 95% Confidence interval for b a b b b a b b b b b b b a b a a a Mean Upper bound mean 2.438.75 2.66 9.192.10 2.78 5.093.92 7.51 2.277.15 5.457.19 4.15 1.82 7.44 3.79 7.681.81 3.36 1.64 5.89 0.10 5.77 1.990.02 1.820.19 0.11 1.93 0.03 1.57 0.19 0.12 0.04 0.21 10.32 10.97 8.82 18.7913.91 19.8827.42 15.34 14.69 29.74 11.73 27.68 12.9216.28 13.60 17.20 10.93 13.17 bound Lower DBl (n = 70) 95% Confidence interval for a b a a a a a a a a a a a a a b b b Mean ) from Thailand. ) from ( DBl ), D. maculatus DMa ) and taeniopterus DTa D. blanfordii Means 95% confidence intervals for means and one-way analysis of variance of head characters in males among three ID 2.55 ST 16.74 EL 7.30 SN 2.18 HL 19.34 HD 8.97 ML 14.30 HW 10.65 SL1SL2 7.43 13.26 DeL 28.58 TDV 1.90 TDH 1.73 DBET 4.04 DBNE 5.27 ID/SVL 0.03 HL/SVL 0.19 character HW/SVL 0.11 Morphometric Table 8 Table 396 Kasetsart J. (Nat. Sci.) 48(3) P - value Upper bound mean 0.090.140.31 0.100.02 0.150.05 0.33 0.000 0.04 0.02 0.000 0.08 0.05 0.000 0.07 0.04 0.000 0.14 0.08 0.011 0.16 0.07 0.000 0.02 0.14 0.000 0.02 0.17 0.000 0.55 0.02 0.000 0.88 0.02 0.000 0.49 0.57 0.000 1.66 0.91 0.000 0.51 0.000 1.76 0.002 0.000 0.000 bound Lower (n = 30) DTa 95% Confidence interval for a c a a b b b b b b b b b b b b Mean Upper bound mean 0.100.150.36 0.100.02 0.160.05 0.09 0.380.04 0.14 0.030.08 0.32 0.050.08 0.02 0.050.14 0.05 0.080.17 0.04 0.080.02 0.08 0.150.02 0.07 0.180.57 0.14 0.030.84 0.16 0.020.48 0.02 0.591.76 0.02 0.87 0.56 0.50 0.90 1.84 0.50 1.71 bound Lower DMa (n = 79) 95% Confidence interval for < 0.05). a a a a ab a a a a a a a a b a a Mean Upper bound mean 0.090.140.27 0.090.02 0.140.05 0.29 0.10 0.04 0.02 0.16 0.07 0.05 0.37 0.07 0.04 0.02 0.13 0.07 0.05 0.16 0.07 0.05 0.02 0.13 0.08 0.02 0.17 0.08 0.54 0.02 0.15 0.83 0.02 0.18 0.46 0.56 0.03 1.44 0.87 0.02 0.47 0.58 1.51 0.86 0.49 1.80 bound Lower DBl (n = 70) 95% Confidence interval for c a c c c c b b b b b b b b b b Mean (Continued) HD/HL 0.47 HW/HL 0.55 ST/SVL 0.17 EL/SVL 0.07 DeL/HL 1.48 HD/HW 0.85 SN/SVL 0.02 HD/SVL 0.09 ML/SVL 0.14 character SL1/SVLSL2/SVL 0.07 0.13 DeL/SVL 0.28 TDV/SVLTDH/SVL 0.02 0.02 DBET/SVL 0.04 DBNE/SVL 0.05 Morphometric Table 8 Table = HW Head width; ID = Internarial = distance; Head HL length; HD = Head = depth; Mouth = ML length; Dewlap DeL length; SN = Snout to nostril; DBNE = Distance between nostril to anterior edge of eye; = DBET Distance between posterior edge of eye to anterior = edge Eye of length; tympanum; SL1 EL = Snout to anterior edge of eye (snout length 1); SL2 = Snout to length. Snout-vent = SVL (horizontal); diameter Tympanum = TDH (vertical); diameter Tympanum = TDV tympanum; of edge anterior to Snout = ST 2); length (snout eye of edge posterior ( P superscripts in the same row indicate a significant difference with different Values Kasetsart J. (Nat. Sci.) 48(3) 397

Multivariate analysis SFL/SVL, SN/SVL, DBNE/SVL, TDH/SVL, The mean values of 21 morphometric AGL/SVL, HW/HL, TaiL/SVL, FL/SVL, FL/ characters and 25 character ratios were subjected AGL, HD/SVL, HD/HL, HD/HW, CL/SVL, EL/ to multivariate analysis using cluster analysis. The SVL, TDV/SVL, ID/SVL, DBET/SVL, DeL/SVL results of clustering produced a dendrogram which and DeL/HL (Figure 3). depicted a level of dissimilarity (difference). This The dendrogram for males among the study produced three dendrograms of the three three Draco lizard species showed that the DBl Draco species: 1) regardless of sexual differences; cluster was distinctly different from the cluster 2) females; and 3) males shown in Figures 2, 3, of DMa and DTa and was characterized by 19 and 4, respectively. morphometric characters—SVL, SFL, HL, ST, All three dendrograms showed two main SL1, DBNE, AGL, TaiL, FL, EL, HW, ML, HD, distinguishable clusters. The DBl cluster was SL2, SN, DBET, DeL, CL and TDH. The cluster easily distinguishable from the cluster of DMa of DMa and DTa was different from the DBl and DTa on the basis of morphometric characters cluster based on 1 morphometric character and 26 and character ratios at a level of 0% of information character ratios—ID, TDV/SVL, ID/SVL, DBET/ remaining. SVL, DeL/SVL, CL/SVL, TDV, HW/SVL, HL/ In the dendrogram of the three Draco SVL, HD/SVL, ST/SVL, SL1/SVL, ML/SVL, species without regard to sexual differences, SL2/SVL, DeL/HL, EL/SVL, SFL/SVL, SN/SVL, the DBl cluster was distinctly different from the DBNE/SVL, TDH/SVL, AGL/SVL, HW/HL, HD/ cluster of DMa and DTa and this difference was HL, HD/HW, FL/SVL, FL/AGL and TaiL/SVL supported by 18 morphometric characters—SVL, (Figure 4). SFL, DBNE, AGL, HL, ST, SL1, HW, DBET, The dendrograms produced by the current SN, TaiL, FL, EL, HD, SL2, ML, DeL and TDH. study were not consistent with the dendrogram of On the other hand, the cluster of DMa and DTa Musters (1983) as the current study revealed the was different from the DBl cluster based on three cluster of DBl was distinctly different from the morphometric characters—ID, CL and TDV; and cluster of DMa and DTa as the current study used 25 character ratios—HL/SVL, HD/SVL, DeL/ morphometrics only for clustering while Musters SVL, HW/SVL, ST/SVL, ML/SVL, SL2/SVL, (1983) reported the cluster of DMa was distinctly SL1/SVL, SFL/SVL, DeL/HL, HW/HL, SN/SVL, different from the cluster of DBl and DTa based on DBNE/SVL, TDH/SVL, AGL/SVL, EL/SVL, HD/ morphological characters consisting of quantitative HL, HD/HW, FL/SVL, FL/AGL, TaiL/SVL, ID/ and qualitative characters and geographic ranges SVL, DBET/SVL, CL/SVL and TDV/SVL (Figure for clustering. However, the dendrogram in the 2). current study was also not consistent with the study The dendrogram for females among by Honda et al. (1999) due to their clustering being the three Draco lizard species showed a clearly based on the 12S and 16S rRNA mitochondrial separate DBl cluster from DMa and DTa based on sequence (779 base pairs) which revealed the 18 morphometric characters—SVL, SFL, AGL, cluster of DTa was distinctly different from the DBNE, FL, TaiL, HL, ST, HW, SL1, ML, SN, cluster of DBl and DMa, while another clustering DBET, TDH, DeL, HD, SL2 and EL. The cluster of Honda et al. (1999) based on allozymic data of DMa and DTa was different from the DBl cluster (20 presumptive loci) revealed the cluster of DMa based on three morphometric characters—ID, CL, was distinctly different from the cluster of DBl and and TDV and 25 character ratios—HL/SVL, ST/ DTa which was consistent with Musters (1983). SVL, HW/SVL, ML/SVL, SL1/SVL, SL2/SVL, Furthermore, the dendrogram of the current study 398 Kasetsart J. (Nat. Sci.) 48(3) ) DTa ( D. taeniopterus D. ) and DMa ) D. maculatus ( D. DBl ), ( blanfordii D. lizard species regardless of sexual difference: difference: sexual of regardless species lizard Draco Dendrogram of three of Dendrogram from Thailand. The vertical line indicates a level of 0% of information remaining. See Figure 1 for definitions of the morphometric terms used terms morphometric the of definitions for 1 Figure See remaining. information of 0% of level a indicates line vertical The Thailand. from in this figure. Figure 2 Figure Kasetsart J. (Nat. Sci.) 48(3) 399 ) from Thailand. The The Thailand. from ) DTa D. taeniopterus ( D. ) and DMa ) D. maculatus ( D. DBl ), ( blanfordii D. lizard species: lizard Draco vertical line indicates a level of 0% information remaining. See Figure 1 for definitions the morphometric terms used in this figure. Dendrogram of females among three among females of Dendrogram Figure 3 Figure 400 Kasetsart J. (Nat. Sci.) 48(3) ) from Thailand. The The Thailand. from ) DTa ( taeniopterus D. ) and DMa ) ( maculatus D. ), DBl ( D. blanfordii D. lizard species: species: lizard Draco vertical line indicates a level of 0% information remaining. See Figure 1 for definitions the morphometric terms used in this figure. Dendrogram of males among three three among males of Dendrogram Figure 4 Figure Kasetsart J. (Nat. Sci.) 48(3) 401 was not consistent with the study of McGuire and SVL, HD/SVL, DeL/SVL, SL2/SVL, ST/SVL, Heang (2001) as their clustering was based on HD/HW and DeL/HL); and 15 morphometric mitochondrial DNA sequence data (1,120 base characters and 8 character ratios (SFL, FL, TaiL, positions) which revealed the cluster of DMa was HW, ID, HL, HD, ML, SN, DBNE, DBET, SL1, distinctly different from the cluster of DBl and DTa SL2, ST, TDH, FL/AGL, CL/SVL, HD/SVL, which was consistent with the clustering based on DeL/SVL, SL1/SVL, SL2/SVL, TDH/SVL and morphological characters and geographic ranges DeL/HL) can be used to discriminate female and of Musters (1983) and the clustering based on male lizards, respectively. The results of cluster allozymic data of Honda et al. (1999). analysis in all three dendrograms showed two main From the dendrogram of the three Draco distinguishable clusters. The D. blanfordii cluster species regardless of sexual difference, the females was distinguished from the cluster of D. maculatus and males among the three Draco lizard species and D. taeniopterus on the basis of morphometric were distinguishable based on the size of each characters and character ratios at a level of 0% of species in clustering which in the current study information remaining (100% dissimilarity). indicated that most of the morphometric characters of DBl had higher values than those of DMa and ACKNOWLEDGEMENTS DTa. However, the CL character in males among the three Draco species was not related to size The authors gratefully acknowledge (Figure 4) because during mating, this character the Department of Zoology, Faculty of Science, of male lizards may not be a character for sexual Kasetsart University, for research funding and the selection during evolution (Andersson, 1994). Thailand Natural History Museum and the National Science Museum, Pathum Thani, Thailand for CONCLUSION proving access to preserved specimens.

A sample of 267 preserved specimens LITERATURE CITED of three Draco lizard species from Thailand consisting of 87 individuals of D. blanfordii Andersson, M. 1994. Sexual Selection. Princeton (17 females and 70 males), 137 individuals of University Press, Princeton, NJ, USA. 624 D. maculatus (58 females and 79 males), and pp. 43 individuals of D. taeniopterus (13 females Honda, M., H. Ota, M. Kobayashi, J. Nabhitabhata, and 30 males) was examined based on 21 H. Yong and T. Hikida. 1999. Phylogenetic morphometric characters and 25 character ratios relationships of the flying lizards, genus to assess morphometric differences. The results Draco (Reptilia, Agamidae). Zoological of univariate analysis showed that there were Science 16(3): 535–549. 12 morphometric characters and 2 character Inger, R.F. 1983. Morphological and ecological ratios that can be used for discrimination among variation in the flying lizards (genus Draco). the three Draco species regardless of sexual Fieldiana: Zoology 18: 1–35. difference, being: FL, TaiL, HW, HL, HD, ML, Ljubisavljevic, K., D. Jovic and G. Dzukic. 2010. SN, DBET, SL1, SL2, ST, TDH, HD/SVL and Morphological variation of the common SL2/SVL. Among these three species, it was lizard (Zootoca vivipara Jacquin, 1787) in the found that 13 morphometric characters and 7 Central Balkans. Arch. Biol. Sci., Belgrade character ratios (FL, TaiL, HW, HL, HD, ML, 62(3): 791–799. DeL, SN, DBET, SL1, SL2, ST, TDH, HW/ 402 Kasetsart J. (Nat. Sci.) 48(3)

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