Chaparana (Chaparana) aenea ( Smith, 1922 )

Dubois, Alain & Ohler, Annemarie, 2005, Taxonomic notes on the Asian frogs of the tribe Paini (Ranidae, Dicroglossinae): 1. Morphology and synonymy of Chaparana aenea (Smith, 1922), with proposal of a new statistical method for testing homogeneity of small samples, Journal of Natural History 39 (20), pp. 1759-1778 : 1761-1774

Descriptions of specimens referred to the species Chaparana (Chaparana) aenea ( Smith, 1922)

Holotype of Rana aenea Smith, 1922 ( Figure 1a–c )

Smith (1922) described Rana aenea on the basis of two juvenile female specimens: (1) MAS 5821 (SVL 35 mm), which was clearly designated as ‘‘type’’ (we would now write ‘‘holotype’’) and shown in his Figure 1 (reproduced in Bourret 1942, p 277), and (2) MAS 5822 (SVL 38 mm), designated as paratype. Both specimens were later deposited in the collection of the British Museum (Natural History), now the Natural History Museum, London. Following is a detailed redescription of the holotype.

Holotype: BMNH 1947.2.2.31 (ex BMNH 1922.7.4.4, ex MAS 5821), juvenile female, collected in May 1920 by a native collector at Doi Chang (19 ° 23'N, 98 ° 52'E), on the border of Chiang Mai and Mae Hong Son Provinces , Thailand (then ‘‘northern Siam’’), about 1500 m altitude. This specimen has been dried up and is in rather bad condition.

Size and general aspect. (1) Specimen of rather small size (SVL 35.2 mm), body rather slender.

Head. (2) Head of medium size, slightly longer (HL 14.4 mm; MN 11.5 mm; MFE 8.9 mm; MBE 5.3 mm) than wide (HW 14.2 mm), convex. (3) Snout rounded, very slightly protruding, its length (SL 5.53 mm) longer than horizontal diameter of eye (EL 4.51 mm). (4) Canthus rostralis rounded, loreal region slightly concave, obtuse in cross-section. (5) Interorbital space flat, smaller (IUE 2.63 mm) than upper eyelid (UEW 3.19 mm) and internarial distance (IN 4.50 mm); distance between front of eyes (IFE 6.3 mm) more than one-half of distance between back of eyes (IBE 10.5 mm). (6) Nostrils closer to eye (EN 2.38 mm) than to tip of snout (NS 3.25 mm). (7) Pupil horizontal oval, very dilated. (8) Tympanum (TYD 2.63 mm) distinct, rounded, much wider than half of eye; tympanum–eye distance (TYE 1.25 mm) less than half its diameter. (9) Pineal ocellus absent. (10) Vomerine ridges present, bearing teeth, between posterior parts of choanae, with an angle of 45 ° to body axis, slightly closer to each other than to choanae, slightly longer than distance between them. (11) Tongue not examined. (12) Supratympanic fold distinct, from eye to shoulder. (13) Parotoid glands absent. (14) Cephalic ridges absent. (15) Co-ossified skin absent.

Forelimbs. (16) Arm short (FLL 7.5 mm), shorter than hand (HAL 8.2 mm), not enlarged. (17) Fingers long, rather thin (TFL 3.78 mm). (18) Relative length of fingers, shortest to longest: I5II,IV,III. (19) Tips of fingers slightly swollen, without discs. (20) Fingers without dermal fringes; webbing absent. (21) Subarticular tubercles prominent, rounded, single, all present. (22) Prepollex oval, very prominent, exceptionally developed laterally (with a digit aspect); palmar tubercles indistinct; supernumerary tubercles absent.

Hindlimbs. (23) Shank more than four times longer (TL 21.1 mm) than wide (TW 4.8 mm), longer than thigh (FL 18.9 mm) and than distance from base of internal metatarsal tubercle to tip of toe IV (FOL 18.9 mm). (24) Toes long, thin; toe IV rather long (FTL 14.4 mm), more than one-third of distance from base of tarsus to tip of toe IV (TFOL 29.4 mm). (25) Relative length of toes, shortest to longest: I,II,V,III,IV. (26) Tips of toes swollen, without discs. (27) Webbing moderate: I 0—2 II 0—2 M III 0— 2 K IV 2 K —1 V (WTF 3.38 mm; WFF 3.81 mm; WI 3.00 mm; WII 2.88 mm; MTTF 9.16 mm; MTFF 9.89 mm; TFTF 6.98 mm; FFTF 7.56 mm). (28) Dermal fringe along toe V well developed, from tip of toe to almost level of first subarticular tubercle. (29) Subarticular tubercles prominent, oval, simple. (30) Inner metatarsal tubercle long, prominent; its length (IMT 2.19 mm) about twice length of toe I (ITL 4.25 mm). (31) No tarsal ridge. (32) No outer metatarsal tubercle; no supernumerary tubercles; no tarsal tubercle.

Skin. (33) Dorsal and lateral parts of head and body smooth, with very small ridges (blood vessels) on flanks. (34) Dorsolateral folds present, not prominent, narrow, continuous until the rear of back. (35) Dorsal parts of limbs smooth, with very small ridges (blood vessels). (36) Ventral parts of head, body and limbs smooth, with some transverse folds on venter. (37) No macroglands.

Coloration in alcohol. (38) Dorsal and lateral parts of head and body: uniform brown with several darker bands and spots (a band between eyes, spots on anterior part of eyelids, a band from eye to snout, a band enclosing tympanum and around it, and a band along anterior third of latero-dorsal fold); clearer spots on upper lip, and clear triangle on tip of snout; flanks darker in their upper than in their lower part, with distinct small darker spots everywhere. (39) Dorsal parts of limbs: brown, with two darker bands on forelimb and four narrow bands on each hindlimb. (40) Ventral parts: throat orange with brown marblings and numerous darker spots on margins; chest and belly yellow orange; thighs orange; back of thighs brown with clearer spots and with darker spots similar to those of flanks; webbing marbled brown and cream.

Female sex characters. (41) Oviduct not seen. (42) Ovary juvenile, with white ovocytes (diameter roughly 0.5 mm).

Topotype of Rana (Chaparana) fansipani Bourret, 1939 ( Figure 1d–f )

MNHN 1999.5821, adult male, collected by the field staff of Frontier Vietnam on 23 November 1997, in Yi Li Ho (1460 m; 22 ° 18'N, 103 ° 50'E) on Fan Si Pan , Lao Cai Province, Vietnam.

Size and general aspect. (1) Specimen of rather large size (SVL 74.6 mm), body robust.

Head. (2) Head of large size, wider (HW 30.1 mm) than long (HL 27.3 mm; MN 23.1 mm; MFE 18.0 mm; MBE 11.4 mm), flat. (3) Snout rounded, very slightly protruding, its length (SL 11.3 mm) longer than horizontal diameter of eye (EL 7.7 mm). (4) Canthus rostralis rounded, loreal region concave, obtuse in cross-section. (5) Interorbital space flat, smaller (IUE 4.7 mm) than upper eyelid (UEW 5.4 mm) and internarial distance (IN 8.4 mm); distance between front of eyes (IFE 11.2 mm) more than one-half of distance between back of eyes (IBE 17.9 mm). (6) Nostrils closer to eye (EN 4.5 mm) than to tip of snout (NS 7.1 mm). (7) Pupil not observed. (8) Tympanum indistinct. (9) Pineal ocellus absent. (10) Vomerine ridges present, bearing few teeth, between posterior parts of choanae, with an angle of 40 ° to body axis, slightly closer to each other than to choanae, slightly longer than distance between them. (11) Tongue large, rounded, emarginate. (12) Supratympanic fold prominent, from eye to above shoulder. (13) Parotoid glands absent. (14) Cephalic ridges absent. (15) Co-ossified skin absent.

Forelimbs. (16) Arm short (FLL 20.2 mm), longer than hand (HAL 17.0 mm), enlarged. (17) Fingers short, rather thin (TFL 7.6 mm). (18) Relative length of fingers, shortest to longest: II,I,IV,III. (19) Tips of fingers slightly swollen, without discs. (20) Fingers II and III with dermal fringes; webbing absent. (21) Subarticular tubercles prominent, rounded, single, all present. (22) Prepollex oval, very prominent, exceptionally developed laterally (with a shovel aspect); one flat, oval palmar tubercle; supernumerary tubercles absent.

Hindlimbs. (23) Shank four times longer (TL 48.4 mm) than wide (TW 12.8 mm), longer than thigh (FL 42.0 mm) and than distance from base of internal metatarsal tubercle to tip of toe IV (FOL 42.8 mm). (24) Toes long, rather thin; toe IV rather long (FTL 23.7 mm), more than one-third of distance from base of tarsus to tip of toe IV (TFOL 62.2 mm). (25) Relative length of toes, shortest to longest: I,II,V,III,IV. (26) Tips of toes swollen, without discs. (27) Webbing moderate: I 0—2 II 0—2 M III 0—2 IV 2—0 V (WTF 9.74 mm; WFF 9.21 mm; WI 8.68 mm; WII 8.42 mm; MTTF 20.9 mm; MTFF 21.4 mm; TFTF 15.4 mm; FFTF 17.7 mm). (28) Dermal fringe along toe V well developed, from tip of toe to beyond level of first subarticular tubercle. (29) Subarticular tubercles prominent, oval, simple. (30) Inner metatarsal tubercle long, prominent; its length (IMT 5.40 mm) half of length of toe I (ITL 10.40 mm). (31) No tarsal ridge. (32) No outer metatarsal tubercle; no supernumerary tubercles; no tarsal tubercle.

Skin. (33) Dorsal and lateral parts of head smooth, back smooth with indistinct glandular warts, flanks granular. (34) Dorsolateral folds present, not prominent, narrow, continuous until the rear of back. (35) Dorsal parts of limbs smooth. (36) Ventral parts of head, body and limbs smooth bearing horny spinules and with some transverse folds on belly. (37) No macroglands.

Coloration in alcohol. (38) Dorsal and lateral parts of head and body: greyish brown with several darker bands and spots (a mid-dorsal chevron, a band from eye to snout, a band enclosing tympanum and around it, and a band along anterior third of latero-dorsal fold); clearer spots on upper lip, and clear triangle on tip of snout; flanks darker in their upper than in their lower part, with some darker spots. (39) Dorsal parts of limbs: brown, with indistinct darker bands; posterior part of thigh dark brown with white marbling. (40) Ventral parts: throat whitish with brown spots on margins; chest, belly and thighs whitish; webbing greyish brown.

Male secondary sex characters. (41) Large-sized, blackish nuptial spines present on prepollex, fingers I–III, chest and throat, forming two separate pads on finger I and prepollex and two widely separated plates of tightly connected spines on chest. Table I gives the numbers of spines in these different places. (42) Forearms distinctly enlarged. (43) Vocal sacs and slits absent.

Other specimens referred to the species Chaparana aenea ( Smith, 1922)

To our knowledge, beside the two specimens described in detail above, only 11 other specimens referable to this species are known, which are briefly discussed below. Table II gives the measurements of the 13 specimens here discussed, and Figure 2 shows the known localities of this species in China, Thailand and Vietnam, as well as areas of known occurrence of the closely related species Chaparana unculuanus (see below).

Specimens from Thailand

BMNH 1974.1113 (ex MAS 5822). Juvenile (SVL 38.6 mm), paratopotype of Rana aenea from Doi Chang. This specimen, slightly larger than the holotype, has similar characters.

MNHN 1989.0712 ( Figure 3a–c ). Non-breeding adult male (SVL 77.4 mm) from Doi Inthanon (18 ° 35'N, 98 ° 28'E), Chiangmai Province, Thailand. Except for its absence of secondary sex characters, this specimen is very similar to the three adult males from Fan Si Pan mentioned above and below. The major differences between them are: the complete absence of nuptial spines on fingers, breast and throat and the absence of enlargement of the forearms, as in the Vietnamese specimen MNHN 1999.5818. The ventral colour pattern is similar in all specimens, but the specimen from Doi Inthanon has a rather clear fawn dorsal colour, whereas the males from Fan Si Pan are dark brown on the back: this difference may be related to nycthemeral colour variation, as specimens collected at night are darker than specimens collected during the day. This specimen is clearly adult, and its absence of spines on the fingers and chest indicates that it was not collected during the breeding season. However, its first finger is notably enlarged, and this finger and the prepollex show structures which correspond to basements of spines.

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MNHN 1989.0713. Juvenile female (SVL 23.7 mm) from same locality as previous frog. This specimen is very similar in general aspect to both holotypes of Rana aenea and Rana fansipani .

MNHN 1989.0714. Juvenile male (SVL 40.5 mm) from same locality as both previous specimens, and very similar to them.

Specimens from Vietnam

MNHN 1948.0139 ( Figure 3d–f ). Juvenile male (SVL 54.5 mm), holotype of Rana fansipani , collected in August 1938 by René Bourret on Fan Si Pan mountain (22 ° 19'N, 103 ° 47'E), Sa Pa District , Lao Cai Province, Vietnam. This specimen was described in detail by Dubois (1977), who wondered whether this frog, clearly a male, was juvenile, subadult or adult. It is very similar in all its characters to the three adult males collected in 1997–1998 on the same mountain, except for its absence of male secondary sex characters: given its much smaller size (SVL 54.5 mm versus 64.7–74.6 mm in the three adult males), it is now clear that it is not adult, nor even subadult, but juvenile, hence its absence of any secondary sex character. This specimen is also very similar in all its characters to the lectotype and paralectotype of Rana aenea mentioned above.

MNHN 1999.5820. Adult male (SVL 64.7 mm), topotype of Rana fansipani Bourret, 1939 , collected on 17 November 1997 at Cat Cat (1280 m; 22 ° 19'N, 103 ° 49'E), on Fan Si Pan mountain, Vietnam. This second specimen is very similar to the specimen MNHN 1999.5821 described in detail above. The major differences between them are the secondary sex characters that are less developed in MNHN 1999.5820: the spines on the breast are large and blackish, but isolated from each other and less numerous ( Table I ); on the throat, blackish spines are distinct, but small; the forearms are distinctly enlarged.

MNHN 1999.5818 ( Figure 4a–c ). Adult male (SVL 69.1 mm), topotype of Rana fansipani Bourret, 1939 , collected on 1 August 1998 at Ban Khoang (1500 m; 22 ° 23'N, 103 ° 48'E), on Fan Si Pan mountain , Vietnam. This specimen is similar to the other two adult males mentioned above, but in a better condition of preservation. The secondary sex characters are similar to those of MNHN 1999.5820, but the blackish horny covering of the nuptial spines is lost on the breast and on the fingers, and the spines are less numerous ( Table I ). No spines are present on the throat, and the forearms are not enlarged.

MNHN 1999.5819. Juvenile male (SVL 28.4 mm) from O Qui Ho (1625 m; 22 ° 23'N, 103 ° 47'E), on Fan Si Pan mountain, Vietnam. Its size is smaller but its morphology and colour pattern are similar to those of the adults from the same mountain.

MNHN 1999.5998. Juvenile male (SVL 49.8 mm) from Fan Si Pan mountain (1700 m; 22 ° 23'N, 103 ° 47'E), Vietnam. This specimen is similar in colour pattern to the other specimens.

MNHN 2000.9018. Juvenile female (SVL 38.8 mm) from Pu Hoat (1300 m; 19 ° 25'N, 104 ° 37'E), Nghe An Province, Vietnam. This specimen was collected much further south, in the Annamite mountain range . In general morphology and coloration it resembles the topotypes from Fan Si Pan.

Specimen from China

MNHN 2001.0277. Juvenile male (SVL 39.2 mm) from Luchun Xian (altitude about 2050 m; 23 ° 01'N, 104 ° 21'E), Huang Lian Shan Nature Reserve , Yunnan, China. This specimen, the only one of this species known from China, is very similar in aspect and coloration to the Vietnamese specimens. It has a very conspicuous chevron on the anterior part of the back.

Morphometric comparisons

Intraspecific comparisons within Chaparana aenea ( Smith, 1922)

Despite the low number of specimens referable to the species Chaparana aenea available, we tried to carry out comparisons between them. As mentioned above, in their overall aspects and colours these 13 specimens are very similar. Part of the variation observed is related to age (size) and sex (especially in breeding adult males). However, the shapes and proportions of these specimens, as studied through 32 measurements (Table II), do not show any clear geographical trend, as shown in Figure 5 which presents the distribution of the 13 specimens according to the first two regression factor scores provided by the PCA. Before and after application of Bonferroni correction, Mann– Whitney U tests based on the first three regression factor scores of PCA (Table III) do not show significant differences between subgroups of these 13 specimens, based either on sex (males versus females) or on main geographic origin (mountains of northern Vietnam and southern Yunnan versus mountains of northern Thailand). However , prior to Bonferroni correction, when ontogenetic stages (juveniles versus adults) are considered, the first regression factor scores of juveniles are statistically different from those of adults. After Bonferroni correction the value is slightly too high for significance. The existence of significant differences between different ontogenetic stages is meaningful as it points to allometric growth in this species, as is usual in frogs, and especially in this group of torrent frogs ( Dubois 1976). From these limited available data, we conclude that, for the time being, no significant difference is documented between populations from Thailand, Vietnam and China referred to this species, and we consider the name Rana fansipani Bourret, 1939 as a junior subjective synonym of Rana aenea Smith, 1922 . Further data, based on more specimens and using additional information (molecular data, bioacoustics, tadpole morphology) will be necessary to confirm or refute this conclusion.

Comparisons between Chaparana aenea ( Smith, 1922) and Chaparana unculuanus (Liu, Hu and Yang, 1960)

Besides Chaparana aenea ( Smith, 1922) and its synonym Chaparana fansipani ( Bourret, 1939) , Dubois (1992) included a third nominotypical species in his subgenus Chaparana (Chaparana) , namely Chaparana unculuanus (Liu, Hu and Yang, 1960) . The latter ( Figure 4d–f ) is indeed very similar to Chaparana aenea in general morphology, and both species occur in southern Yunnan, although in different areas ( Figure 2 ). However , these two species differ markedly in their male secondary sex characters: as discussed above, breeding males of Chaparana aenea have enlarged forelimbs and black cornified spines on the fingers and chest, but no spines around the vent. In contrast, breeding males of Chaparana unculuanus are devoid of spines on fingers and chest and their forelimbs are not enlarged, but the skin around their vent is distended and bears spines. We examined and measured two adult males and four adult females of Chaparana unculuanus from Jingdong Xian (Yunnan, China) (Table IV). Univariate comparisons by Mann– Whitney U tests of measurements (expressed as ratios of SVL, except for SVL itself) between the four adult specimens of Chaparana aenea listed in Table II and the six adult specimens of Chaparana unculuanus listed in Table IV show that the two samples differ statistically in seven measurements out of 27 (HL, FLL, TFL, TL, IN, TYE, IMT). If these 10 specimens are included in a PCA, the two samples appear as two well-separated groups ( Figure 6 ) and the second regression factor scores for the two samples are significantly different ( Table V ). These data confirm the morphological distinctness of the two samples, independently from their male secondary sex characters, and they support the interpretation that the two samples represent different biological species. This is consistent with the result of the analysis of partial sequences of mitochondrial 12S and 16S rRNA genes provided elsewhere (Jiang et al. forthcoming), which confirms that the two species have distinct, although closely similar, sequences.

Taxonomic consequences of these observations at the generic level

Beside the synonymization of Rana aenea and Rana fansipani , the observations presented above have taxonomic consequences at the generic level. The nominotypical species Rana fansipani being the type-species of Chaparana Bourret, 1939 , the status of the latter name needs to be re-evaluated. Dubois (1992) recognized two genera in the tribe Paini Dubois, 1992 of the subfamily Raninae Rafinesque-Schmaltz, 1814 : Chaparana Bourret, 1939 (with four subgenera) and Paa Dubois, 1975 (with four subgenera). Based on unpublished molecular data, this tribe was transferred to the subfamily Dicroglossinae of the Ranidae Anderson, 1871 by Dubois et al. (2001), a taxonomy followed by Dubois (2003) and Roelants et al. (2004). Furthermore, following the results of the work of Jiang and Zhou (2003), the genus Nanorana Günther, 1896 , placed by Dubois (1992) in the Ranini, must be included in the tribe Paini .

In Dubois’s (1992) taxonomic arrangement, breeding males of all species of the genus Chaparana were stated to be devoid of breeding spines both on the forelimbs and on the breast, whereas most (but not all) species of Paa had such spines, at least on the first finger. Besides, breeding males of several species referred to Chaparana were known to have a specialized zone of skin, either swollen or bearing spines, around the vent. It is now known that the type-species of Chaparana is devoid of this latter character but has spines on the fingers and chest, thus fitting with the genus Paa as understood by Dubois (1992). This species, but this species alone, must be removed from the group called Chaparana by Dubois (1992), and placed in the group called Paa by this author. The name Chaparana has priority to designate this genus.

However, two significant differences remain between Chaparana aenea and all species referred by Dubois (1992) to the subgenus Paa (Paa) : (1) whereas, in breeding males of the former, both sides of the chest are covered by a well-delimited patch of densely packed spines, whose general shape recalls that of a bean or kidney ( Figure 7 ), in breeding males of those species of Paa that have breast spines (i.e. only a part of the species of this group), the two patches are composed of unequally spaced spines and have variable limits, so that these patches do not have a common general shape (see e.g. Dubois 1976, p 52, 89, 105, 121); (2) in Chaparana , the first finger is usually longer than the second (both fingers may be subequal in some juveniles), whereas in Paa the first finger is usually shorter than the second (in some large adults, both fingers may be subequal, or the first one may be exceptionally longer in the species Paa liebigii (Günther, 1860) : see Dubois 1976, p 65): in this character, Chaparana aenea is similar to the frogs placed by Dubois (1992) in the subgenus Paa (Quasipaa) . This suggests that the relationships between Chaparana and Paa should be explored further. As for the other groups referred by Dubois (1992) to several other subgenera of Chaparana and Paa , their taxonomy and nomenclature will also have to be modified as a result of a re-evaluation of the significance of some characters following the results presented above. To start clarifying this matter, we recently proposed a preliminary cladistic analysis of the Paini based on partial sequences of mitochondrial 12S and 16S rRNA genes of 17 species of the clade (Jiang et al. forthcoming), but this analysis only provides partial information. Therefore we explored the phylogenetic relationships in this clade on the basis of external morphological characters of adults and of a few other characters as available. These results and their implications will be presented elsewhere.

Figures

Figure 1. Chaparana aenea (Smith, 1922). (a–c) BMNH 1947.2.2.31 (ex BMNH 1922.7.4.4, ex MAS 5821), holotype of Rana aenea Smith, 1922, juvenile female from Doi Chang, Thailand: (a) whole body in dorsal view; (b) whole body in ventral view; (c) head in right lateral view. (d–f) MNHN 1999.5821, topotype of Rana (Chaparana) fansipani Bourret, 1939, adult male from Fan Si Pan, Vietnam: (d) whole body in dorsal view; (e) whole body in ventral view; (f) head in right lateral view.

Figure 2. Map showing the localities of specimens studied of the species Chaparana aenea (Smith, 1922) (circles) and Chaparana unculuanus (Liu, Hu and Yang, 1960) (square). 1, Doi Chang, Thailand; 2, Doi Inthanon, Thailand; 3, Fan Si Pan, Vietnam; 4, Pu Hoat, Vietnam; 5, Luchun, China; 6, Jingdong, China.

Figure 3. Chaparana aenea (Smith, 1922). (a–c) MNHN 1989.0712, non-breeding adult male from Doi Inthanon, Chiangmai Province, Thailand: (a) whole body in dorsal view; (b) whole body in ventral view; (c) head in right lateral view. (d–f) MNHN 1948.0139, holotype of Rana (Chaparana) fansipani Bourret, 1939, juvenile male from Fan Si Pan, Vietnam: (d) whole body in dorsal view; (e) whole body in ventral view; (f) head in right lateral view.

Figure 4. (a–c) Chaparana aenea (Smith, 1922), MNHN 1999.5818, topotype of Rana (Chaparana) fansipani Bourret, 1939, adult male from Fan Si Pan, Vietnam: (a) whole body in dorsal view; (b) whole body in ventral view; (c) head in right lateral view. (d–f) Chaparana unculuanus (Liu, Hu and Yang, 1960), MNHN 2001.0280, adult female from Jingdong Xian, Yunnan, China: (d) whole body in dorsal view; (e) whole body in ventral view; (f) head in right lateral view.

Figure 5. Plots of factors 1 and 2 of principal component analysis based on varimax rotated coefficients for logtransposed characters (32 measurements) for 13 specimens of Chaparana aenea (Smith, 1922) from China, Thailand and Vietnam (see Table II).

Figure 6. Plots of factors 1 and 2 of principal component analysis based on varimax rotated coefficients for logtransposed characters (25 measurements) for four adult specimens of Chaparana aenea (Smith, 1922) (see Table II) and six adult specimens of Chaparana unculuanus (Liu, Hu and Yang, 1960) (see Table IV).

Figure 7. Chaparana aenea (Smith, 1922), MNHN 1999.5821, topotype of Rana (Chaparana) fansipani Bourret, 1939, adult male from Fan Si Pan, Vietnam: ventral view of chest and throat showing shape of patch of nuptial spines.