Cristidorsa, Wang & Che & Lin & Deepak & Aniruddha & Jiang & Jin & Chen & Siler, 2019

Finally, it is important to note that our new classification aims to provide a suite of testable hypotheses for future phylogenetic work, and the new classifications for non-sampled species represent our best attempt at a more stable taxonomic structure. Many species, such as Japalura sagittifera View in CoL and Diploderma hamptoni , are known from poorly surveyed regions, and lack vouchered specimens and genetic samples. Therefore, our new classification of these understudied taxa is based on the best available morphological data alone, which may be subject to convergence and warrants additional investigations. Additionally, future phylogenomic studies are needed to better resolve the higher level relationships of Japalura s.l. To facilitate future taxonomic work, we provide updated morphological diagnoses to all genera of the subfamily Draconinae below [modified from Smith (1935) and Zhao et al. (1999)].

DIAGNOSTIC KEY TO ALL GENERA OF SUBFAMILY DRACONINAE 1a. Femoral pores present .................................................................................................................. Mantheyus

1b. Femoral pores absent ................................................................................................................................... 2

2a. Ribs much prolonged, supporting wing-like expansion ...................................................................... Draco

2b. Ribs not elongated into wing-like structure................................................................................................ 3

3a. Four toes on hind limbs; males with prominent, retractable dewlaps....................................................... 4

3b. Five toes on hind limbs................................................................................................................................. 5

4a. Enlarged scales present on posterior thigh, flank scales heterogeneous ......................................... Sitana

4b. Enlarged scales absent on posterior thigh, flank scales homogeneous ........................................... Sarada

5a. Distinct bony supra-orbital arch present; globular protuberance present on snout............ Lyriocephalus

5b. Above structures all absent.......................................................................................................................... 6

6a. Series of long, lanceolate spines present along medial line on gular; gular appendage projecting forward beyond snout .............................................................................................................................. Hypsicalotes

6b. No large spine series along medial line on gular; gular appendage absent or cannot extend forward beyond snout ................................................................................................................................................. 7

7a. Distinct, elongated rostral appendage present (always in males, sometimes a mere tubercle in females) ......................................................................................................................................................... 8

7b. No rostral appendage, or appendage short, not elongated ....................................................................... 10

8a. Dorsal body scales heterogeneous.............................................................................................. Ceratophora

8b. Dorsal body scales mostly homogenous....................................................................................................... 9

9a. Dorsal and nuchal crests tall; dorsal crest scales enlarged in distinct, large triangles......... Harpesaurus

9b. Nuchal crest low; dorsal crest indistinct ....................................................................... Thaumatorhynchus

10a. Three parallel longitudinal gular folds on each side of medial throat, curving and converging posteriorly into U-shape pattern ................................................................................................................ Ptyctolaemus

10b. Gular folds not in above shape................................................................................................................... 11

11a. Dorsal body scales homogeneous ............................................................................................................... 12

11b. Dorsal body scales heterogeneous.............................................................................................................. 15

12a. Lateral skin fold present from posterior jaw to shoulder on each side; tail long TAL>300% SVL..................................................................................................................................................... Bronchocela

12b. Lateral skin folds absent from posterior jaw to shoulder; tail relatively short, TAL <300% SVL mostly .......................................................................................................................................................... 13

13a. Body compressed dorsally; dorsal crest feeble ..................................................................... Psammophilus

13b. Body not compressed dorsally; dorsal crests relatively well developed................................................... 14

14a. Lateral gular sac pockets present; granular scutellation of gular sac..................................... Complicitus

14b. Lateral gular sac pockets absent; scales of gular not in granular shape......................................... Calotes

15a. Tail markedly swollen at base, flattened above, with dorsolateral keels formed by enlarged, angular scales ............................................................................................................................................ Phoxophrys

15b. Tail not markedly swollen at basee, or swollen but rounded above......................................................... 16

16a. Post-occipital and post-orbital spines present; tympanum exposed ........................................................ 17

16b. Spines on dorsal head absent; tympanum exposed or concealed ............................................................. 18

17a. Nuchal and dorsal crest short; tail long, TAL>185% SVL; lateral gular fold weakly developed ....................................................................................................................................................... Oriocalotes

17b. Nuchal and dorsal crest much longer; tail short, TAL<180% SVL mostly; lateral gular fold strongly developed.................................................................................................................................. Acanthosaura

18a. Enlarged dorsal body scales arranged in regular rows or ridges............................................................. 19

18b. Enlarged dorsal body scales arranged randomly...................................................................................... 21

19a. Enlarged scales in dorsolateral ridges or rows only (except in D. swinhonis , which also in V-shape rows) ..................................................................................................................................................... Diploderma

19b. Enlarged scales in both lateral and V-shaped ridges or rows .................................................................. 20

20a. Pair of enlarged, conical scales present on nape; head relatively wide and robust, HW>70% HL mostly ..................................................................................................................................... Cristidorsa gen. nov.

20b. Enlarged conical scale pair on nape absent; head relatively narrow and slender, HW <70% HL mostly .......................................................................................................................................................... Japalura

21a. Ventral body scales heterogeneous in size and shape ............................................................... Mictopholis

21b. Ventral body scales homogeneous in size and shape mostly .................................................................... 22

22a. Nuchal crest scales low, only slightly differentiated; body and limbs slender; neck distinct and long .. 23

22b. Nuchal crest scales tall, elongated in lanceolate spines; body and limbs relatively robust; neck robust and relatively short .................................................................................................................................... 24

23a. Toe V not longer than Toe I; males usually with distinctively developed dewlaps extending to anterior venter ............................................................................................................................................. Otocryptis

23b. Toe V longer than Toe I; males with much smaller lateral gular folds...................................... Aphaniotis

24a. Suborbital scales in multiple rows, subequal in size; dorsal scales fine, smaller than ventrals; distinct transverse gular fold present; nuchal crest separated from nuchal scales by one or more rows of enlarged, supporting scales ..................................................................................................................... Gonocephalus

24b. Suborbital scales in singular row, or multiple rows but one row distinctively enlarged; dorsal scales larger or smaller than ventrals; transverse gular fold absent or indistinct; nuchal crest not separated from nuchals by enlarged, supporting scales ............................................................................................ 25

25a. Majority of dorsal body scales smaller than ventrals ............................................................................... 26

25b. Majority of dorsal body scales larger than ventrals ................................................................................. 29

26a. Lateral gular fold strongly developed with rounded tip, extending to anterior region of venter when folded; dorsal body scales muricate ....................................................................................... Malayodracon

26b. Lateral gular fold less developed without an obvious tip, not extending beyond throat when folded; dorsal body scales more heterogeneous with rhombic scales ........................................................................ 27

27a. Head tall, robust, distinctively raised on occipital region with distinct cranial ridges ........ Dendragama

27b. Head more compressed dorsally, much more slender, not distinctively raised on occipital region ........ 28

28a. Tympana exposed .................................................................................................................... Pseudocalotes

28b. Tympana concealed ............................................................................................................... Pseudocophotis

29a. Tail prehensile, short, TAL <135% SVL; tympanum concealed ..................................................... Cophotis

29b. Tail not prehensile, long, TAL>160% SVL; tympanum exposed.............................................................. 30

30a. Dorsal body scales less heterogeneous; dorsal body scales more rounded; tail short, TAL <175% SVL.................................................................................................................................................... Lophocalotes

30b. Dorsal body scales more heterogeneous; dorsal body scales more elongated; tail long, TAL>200% SVL................................................................................................................................................................ Salea

SPECIES-LEVEL RELATIONSHIPS AND TAXONOMY