Variation in iris and buccal mucosa pigmentation of U. fimbriatus and U. giganteus. Row (A) Iris colouration in U. giganteus (from left to right): Marojejy and Montagne d' Ambre (last). Row (B) Iris colouration in U. fimbriatus (from left to right): Masoala, Nosy Mangabe, Nosy Mangabe, Amboriana, Marolambo, and Ambodiriana. Row (C) Colouration of the buccal mucosa in U. giganteus (from left to right): Marojejy and Montagne d' Ambre (last). Row (D) Colouration of the buccal mucosa in U. fimbriatus (from left to right): Amboriana, Ambodiriana, Marolambo, Nosy Mangabe, Masoala. Row (E) Detailed photographs of the head colouration of U. giganteus from Marojejy. 

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... specimens from Montagne d' Ambre exhibited the following dorsal colouration (an overview is given in Fig. 7): The dorsal surface of unstressed specimens is mainly covered with brown, grey, and black, often remi- niscent of the bark of trees ( Fig. 7A and B). The coloura- tion of stressed individuals is much more contrasting: the brownish and greyish spots mostly have changed to yel- low and a black reticulation or marbling covers most dor- sal parts of back, tail, head and limbs (Fig. 7C). Most of the photographed individuals agree in having one large beige symmetrical patch in the neck, one at middorsum, and one in the sacral region. These patches can vary from poorly defined (e.g., Fig. 7D) to very distinct ( Glaw et al. 2006; Fig. 7B) and their borders can be well delimited by black or poorly recognizable. The light patches are often filled with brownish pattern and their shape is variable. The head col- ouration of U. giganteus from Montagne d' Ambre is rath- er characteristic: there are two distinct, chevron-shaped, blackish markings, both pointing posteriorly, one between the eyes and the other in front of the eyes. One, often poor- ly delimited, blackish spot is generally present between the nostrils and a further one posterior to the nostrils. The area between these spots and the two chevrons is usually beige, brown or yellow. Two black spots are present behind the posterior chevron, and the combined appearance often re- sembles a "sad smiley" symbol. The ventral surface is whit- ish. No dark colour was noted in the open mouth, and the anterior tip of the tongue is light red in the photographed individual (Fig. ...

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... most important and largely consistent difference between U. fimbriatus and U. giganteus is the colour of the Figure 5. Colour variation of U. fimbriatus form north-eastern populations. (A-B) adult specimen from Ambodivoangy, Makira; (C) adult female form Ambodiriana; (D) subadult from Ambodivoangy, Makira; (E) specimen from Mahasoa; (F) specimen from the island Nosy Boraha; (G) subadult from Ambodiriana; (H) defense behavior in an adult male from Ambodiriana. All photos by the authors. iris, as shown in Figure 9A. In U. giganteus it is whitish to beige, with brownish lines running vertically along the vertical pupil. These lines typically are chocolate brown without a distinct reddish-brown tone, thin and poorly defined, i.e., often interrupted and connected with each other in an irregular network pattern. In U. fimbriatus, the iris has a more beige-yellowish to olive greenish ground colour, and the vertical lines typically have a reddish- brown tone. Furthermore, these lines are more distinct, i.e., at least 2-3 of them are relatively wide and continu- ous; in some specimens, the pattern of wide and continu- ous vertical lines is so dense that it covers most of the iris ...

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... from within the range of the southern lineage, and thus assigned to U. fimbriatus, were available for 32 in- dividuals from 11 localities: Ambodiriana, Ambodivoangy (close to Makira), Kianjavato, Mahasoa, Masoala Peninsu- la (Anaravana, Cap Est, Hiaraka [photos examined but not included in the figures], Tampolo), Marolambo, Nosy Bo- raha, and Nosy Mangabe. Many of these corresponded to the specimens identified via molecular data (i.e., from all localities except Mahasoa, Marolambo, Nosy Boraha, and several sites in Masoala). The specimens exhibited the fol- lowing dorsal colouration (an exemplified overview is giv- en in Figs. 3-6). Dorsal ground colour varies between light beige to dark brown to black mainly covered with brown, grey and black patterns, often reminiscent of the bark of trees. Some specimens show a more distinct pattern with up to three well-delimited light patches on the neck, mid- dorsum and one in the sacral region (e.g., Figs 3B and 4D). Other individuals show only a poorly recognizable pattern of indistinct dark vertical, sometimes ruptured lines on greyish ground colouration (e.g., Figs 3G, 6C). Some in- dividuals show an irregular pattern of fine-grained beige- greenish to brown dots (e.g., Figs 3D, 5D, 5E). One individ- ual from the Masoala Peninsula showed three light white patches of variable shape well delimited by dark lines on the head and shoulders (Fig. 4C). The ventral surface is whitish. The colouration of the buccal mucosa in all U. fim- briatus populations is unpigmented. In some individuals the anterior tip of the tongue is light to dark red, but it is uncertain whether this is a stable individual character and if this colouration is caused by pigmentation or simply per- fusion of superficial blood vessels (Fig. ...

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... of U. giganteus from Marojejy differ clearly in terms of dorsal colouration from the Montagne d' Ambre individuals (an overview is given in Fig. 8): generally the Marojejy population seems to be much more variable in dorsal colouration and pattern. Some individuals resem- ble the general pattern as observed in Montagne d' Ambre (Fig. 8E) whereas others have an irregular pattern of in- distinct dark vertical, sometimes ruptured lines on grey- ish-brown ground colouration (e.g., Figs 8A, 8D, 8F). One individual showed several light white patches of variable shape well delimited by dark lines on the head, shoulders, dorsum, feet, and tail (Fig. 8D), very similar to the pattern observed in some U. fimbriatus (Fig. 4C). The head colour- ation of U. giganteus from Marojejy is rather heterogene- ous and chevron-shaped, blackish markings are absent or not as characteristic as in the Montage d' Ambre population (Fig. 9E). The ventral surface is whitish (Fig. 2). No dark colour of the buccal mucosa was noted; the anterior tip of the tongue is light red in one photographed individual and in another individual the middle part of tongue has a light yellowish colouration (Fig. ...

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... of U. giganteus from Marojejy differ clearly in terms of dorsal colouration from the Montagne d' Ambre individuals (an overview is given in Fig. 8): generally the Marojejy population seems to be much more variable in dorsal colouration and pattern. Some individuals resem- ble the general pattern as observed in Montagne d' Ambre (Fig. 8E) whereas others have an irregular pattern of in- distinct dark vertical, sometimes ruptured lines on grey- ish-brown ground colouration (e.g., Figs 8A, 8D, 8F). One individual showed several light white patches of variable shape well delimited by dark lines on the head, shoulders, dorsum, feet, and tail (Fig. 8D), very similar to the pattern observed in some U. fimbriatus (Fig. 4C). The head colour- ation of U. giganteus from Marojejy is rather heterogene- ous and chevron-shaped, blackish markings are absent or not as characteristic as in the Montage d' Ambre population (Fig. 9E). The ventral surface is whitish (Fig. 2). No dark colour of the buccal mucosa was noted; the anterior tip of the tongue is light red in one photographed individual and in another individual the middle part of tongue has a light yellowish colouration (Fig. ...

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... surface is whitish. The colouration of the buccal mucosa in all U. fim- briatus populations is unpigmented. In some individuals the anterior tip of the tongue is light to dark red, but it is uncertain whether this is a stable individual character and if this colouration is caused by pigmentation or simply per- fusion of superficial blood vessels (Fig. ...

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... spots and the two chevrons is usually beige, brown or yellow. Two black spots are present behind the posterior chevron, and the combined appearance often re- sembles a "sad smiley" symbol. The ventral surface is whit- ish. No dark colour was noted in the open mouth, and the anterior tip of the tongue is light red in the photographed individual (Fig. ...

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... well delimited by dark lines on the head, shoulders, dorsum, feet, and tail (Fig. 8D), very similar to the pattern observed in some U. fimbriatus (Fig. 4C). The head colour- ation of U. giganteus from Marojejy is rather heterogene- ous and chevron-shaped, blackish markings are absent or not as characteristic as in the Montage d' Ambre population (Fig. 9E). The ventral surface is whitish (Fig. 2). No dark colour of the buccal mucosa was noted; the anterior tip of the tongue is light red in one photographed individual and in another individual the middle part of tongue has a light yellowish colouration (Fig. ...

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... markings are absent or not as characteristic as in the Montage d' Ambre population (Fig. 9E). The ventral surface is whitish (Fig. 2). No dark colour of the buccal mucosa was noted; the anterior tip of the tongue is light red in one photographed individual and in another individual the middle part of tongue has a light yellowish colouration (Fig. ...

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... photos by the authors. iris, as shown in Figure 9A. In U. giganteus it is whitish to beige, with brownish lines running vertically along the vertical pupil. ...