Research Article
Research Article
The validity of Rana bannanica Rao & Yang, 1997 (Anura, Ranidae)
expand article infoShuo Liu, Tan Van Nguyen§, Nikolay A. Poyarkov|, Qiaoyan Wang#, Dingqi Rao, Song Li
‡ Chinese Academy of Sciences, Kunming, China
§ Duy Tan University, Da Nang, Vietnam
| Lomonosov Moscow State University, Moscow, Russia
¶ Joint Russian-Vietnamese Tropical Research and Technological Center, Hanoi, Vietnam
# Research Institute of Xishuangbanna National Nature Reserve, Jinghong, China
Open Access


One specimen of frog was collected from Mohan Town, Mengla County, Xishuangbanna Prefecture, Yunnan Province, China, which is the type locality of Rana bannanica Rao & Yang, 1997, currently considered as a junior synonym of Hylarana milleti (Smith, 1921). This specimen well agrees with the original description of R. bannanica. In regard to morphology, R. bannanica and H. milleti are, indeed, very similar, but can still be distinguished from each other. Phylogenetic analysis, based on mitochondrial gene sequences, showed that this specimen and H. milleti are different species. Hence, we remove R. bannanica from the synonymy of H. milleti.

Key Words

16S, COI, cytb, Hylarana milleti, resurrection, synonym, systematics, taxonomy


The systematics and taxonomy of Golden-backed frogs of the genus Hylarana Tschudi, 1838 have long been chaotic (Dubois 1992; Oliver et al. 2015; Chan et al. 2020; Dubois et al. 2021; Reilly et al. 2022). Previously, it had been divided into 10 genera, namely Hylarana, Hydrophylax Fitzinger, 1843, Amnirana Dubois, 1992, Chalcorana Dubois, 1992, Humerana Dubois,1992, Papurana Dubois, 1992, Pulchrana Dubois, 1992, Sylvirana Dubois, 1992, Abavorana Oliver, Prendini, Kraus & Raxworthy, 2015, and Indosylvirana Oliver, Prendini, Kraus & Raxworthy, 2015 (Oliver et al. 2015; Chan et al. 2020). Later, except for Abavorana, the other nine genera were placed back into Hylarana, relegating those ranks as subgenera (Dubois et al. 2021; Reilly et al. 2022). At present, the website Amphibian Species of the World ( also adopts this viewpoint (Frost 2023).

Rana bannanica Rao & Yang, 1997 is a poorly-known ranid species described in 1997 from Xishuangbanna Prefecture, Yunnan Province, China and its type locality is in Mohan Town, Mengla County, Xishuangbanna Prefecture, Yunnan Province, China, close to the border with Laos (Rao and Yang 1997). Since this species was described, there have been no formal collection reports of this species from its type locality.

Rana milleti Smith, 1921 was described from the Langbian Plateau, Lam Dong Province, central-southern Vietnam and, subsequently, this species was reported from Thailand and Cambodia (Chuaynkern et al. 2004; Stuart and Emmett 2006). Thereafter, Ohler (2007) proposed R. bannanica as a junior synonym of R. milleti by morphological data, based on several specimens collected from Phongsaly, Laos, as she considered that these specimens resemble both R. bannanica and R. milleti.

Currently, Rana milleti is regarded as Hylarana milleti and considered to be distributed in Vietnam, Thailand, Cambodia, Laos, southern China and peninsular Myanmar and R. bannanica is considered to be a synonym of H. milleti (Ohler 2007; Chan et al. 2020; Poyarkov et al. 2021; Zug 2022).

During our field surveys in Yunnan Province, China and in southern Vietnam from 2019 to 2023, we collected one frog specimen from the type locality of Rana bannanica, Xishuangbanna and six specimens of Hylarana milleti from its type locality, Langbian Plateau, Lam Dong Province (Fig. 1). We collected molecular sequence data of these specimens in order to reconstruct a molecular phylogeny and evaluate the phylogenetic position of them. Our phylogenetic analysis of molecular data demonstrated that the specimen from Xishuangbanna and the specimens from Langbian Plateau are not the same species. Thus, we resurrect R. bannanica and re-assess its taxonomic status.

Figure 1. 

Map showing the type locality (black star) of Hylarana bannanica in Xishaungbanna, Yunnan, China and the type locality (black square) of H. milleti in Langbian Plateau, Lam Dong, Vietnam and other confirmed distributions (black dots) of H. bannanica in Phongsaly, Laos, Chanthaburi, Thailand and Tanintharyi, Myanmar, respectively, based on molecular data.

Materials and methods

The specimen from China was collected from Manzhuang Village, Mohan Town, Mengla County, Xishuangbanna Prefecture, Yunnan Province and the specimens from Vietnam were collected from Da Lat and Di Linh, Lam Dong Province. Liver or muscular tissue samples were preserved in 99% ethanol for molecular analysis. The specimen from China was deposited in Kunming Institute of Zoology, Chinese Academy of Sciences (KIZ) and the specimens from Vietnam were deposited in Zoological Museum of Moscow University (ZMMU).

Measurements were taken with a digital calliper to the nearest 0.1 mm. We measured: snout-vent length (SVL), head length (HL), head width (HW), snout length (SL), eye diameter (ED), interorbital distance (IND), tympanum diameter (TD), forearm and hand length (FHL), total arm length (ARM), tibia length (TL), tarsus and foot length (TFL), foot length (FL) and total leg length (LEG).

Total genomic DNA was extracted from liver or muscular tissues. A fragment of the 16S rRNA (16S) gene was amplified and sequenced using the primers L2188: 5’–AAAGTGGGCCTAAAAGCAGCCA–3’ and 16H1: 5’–CTCCGGTCTGAACTCAGATCACGTAGG–3’ (Hedges 1994; Matsui et al. 2006), a fragment of the cytochrome c oxidase subunit I (COI) gene was amplified and sequenced using the primers Chmf4: 5’–TYTCWACWAAYCAYAAAGAYATCGG–3’ and Chmr4: 5’–ACYTCRGGRTGRCCRAARAATCA–3’ (Che et al. 2012) and a fragment of the cytochrome b (cytb) gene was amplified and sequenced using the primers cytb-c: 5’–CTACTGGTTGTCCTCCGATTCATGT–3’ and CB-J-10933: 5’–TATGTTCTACCATGAGGACAAATATC–3’ (Bossuyt and Milinkovitch 2000). All new sequences have been deposited in GenBank. Other sequences were obtained from GenBank (Table 1).

Table 1.

Sequences used in this study.

Species Voucher 16S COI cytb
Abavorana luctuosa FMNH 273219 KR264088 KR264180
Hylarana albolabris MVZ 234147 JX564871 JX564871 JX564871
Hylarana amnicola USNM 584214 KY080042 KY079510
Hylarana annamitica AMNH A-161290 MG606336
Hylarana arfaki BPBM19463 KR264048 KR264139
Hylarana asperrima AMNH 117606 MG552350
Hylarana attigua FMNH 258259 MG606313
Hylarana aurantiaca BNHS 5813 KM068913 KM069020 KM069128
Hylarana aurata RG6548 KR264101 KR264192
Hylarana bahuvistara BNHS 5921 KT281144
Hylarana bannanica KIZ2019BN PP034378 PP050559 PP035221
Hylarana bannanica 0281Y KR827825 KR087730
Hylarana bannanica 2004.0373 KR827824 KR087729
Hylarana bannanica 2004.0375 KR827823 KR087728
Hylarana bannanica USNM 594329 MT608886 MT608217
Hylarana bannanica USNM 594330 MT608885 MT608216
Hylarana baramica BNP024 DQ835348
Hylarana caesari BNHS 5842 KM068915 KM069022 KM069130
Hylarana centropeninsularis DWNP1189 EU604198
Hylarana chalconota MVZ 239431 KR264095 KR264187
Hylarana cubitalis 2005.0228 KR827781 KR087688
Hylarana daemeli BPBM36025 KR264057 KR264148
Hylarana darlingi PEMA6989 KR264121 KR264208
Hylarana doni BNHS 5819 KM068928 KM069035 KM069143
Hylarana elberti USNM 579404 OL960068
Hylarana erythraea USNM 587708 MT608883 MT608214
Hylarana eschatia USNM 587721 MT608717 MT608044
Hylarana faber 2003.0419 KR827801 KR087706
Hylarana flavescens BNHS 5845 KM068931 KM069038 KM069146
Hylarana florensis MVZ 292869 OL960058
Hylarana fonensis ZMBMH0145 MG552337
Hylarana galamensis MVZ 245225 KR264096 KR264188
Hylarana garritor BPBM15488 KR264042 KR264133
Hylarana glandulosa KUHE 53618 AB719223
Hylarana gracilis DZ 1173 KM068939 KM069046 KM069154
Hylarana grandocula PNM 7588 KP298046
Hylarana guentheri LC2018092202 MN248533 MN248533 MN248533
Hylarana humeralis USNM 583171 MG935823 MG935527
Hylarana indica BNHS 5856 KM068951 KM069058 KM069165
Hylarana intermedia BNHS 5831 KM068960 KM069067 KM069174
Hylarana jimiensis BPBM22832 KR264053 KR264144
Hylarana krefftii KM247362 KM247362 KM247362
Hylarana lacrima USNM 583124 MG935997 MG935703
Hylarana lateralis MBM-JBS19852 MG935831 MG935537
Hylarana laterimaculata KUHE 17594 AB719229
Hylarana latouchii LSU20200422001ZL MT702387 MT702387 MT702387
Hylarana leptoglossa ZHRAB OR058745 OR058745 OR058745
Hylarana lepus USNM 584215 KY080044 KY079512
Hylarana longipes CAS 254205 MG552472
Hylarana macrodactyla 0095Y KR827808 KR087713
Hylarana magna BNHS 5857 KM068964 KM069071 KM069178
Hylarana malabarica BNHS 5880 KM068968 KM069075 KM069182
Hylarana malayana USNM 587709 MT609321 MT608642
Hylarana maosonensis K742 KR827809 KR087714
Hylarana megalonesa FMNH 268981 KR264085 KR264176
Hylarana melanomenta ELR165 KF477693
Hylarana milleti ZMMU NAP-12372 PP034384
Hylarana milleti ZMMU NAP-12373 PP034383
Hylarana milleti ZMMU NAP-12374 PP034382
Hylarana milleti ZMMU NAP-12862 PP034381
Hylarana milleti ZMMU NAP-12863 PP034380
Hylarana milleti ZMMU NAP-12864 PP034379
Hylarana milleti DL35 OR095101
Hylarana milleti DL38 OR095092
Hylarana milleti DL40 OR095093
Hylarana milleti DL51 OR095094
Hylarana milleti ROM 34429 KR264108 KR264199
Hylarana cf. milleti ROM 7240 AF206490
Hylarana milneana BPBM15749 KR264044 KR264135
Hylarana miopus 0313Y KR827778 KR087686
Hylarana montana BNHS 5866 KM068978 KM069085 KM069192
Hylarana montosa NCSM 76398 MG606619
Hylarana mortenseni FMNH 263303 KR264076 KR264166
Hylarana nigrovittata USNM 583176 MG936001 MG935707
Hylarana papua LSUMZ97639 KR264091 KR264183
Hylarana parvaccola FMNH 268599 EF487450
Hylarana raniceps FMNH 267958 DQ650408
Hylarana roberti CAS 243913 MG606593
Hylarana rufipes FMNH268575 KR264081 KR264172
Hylarana serendipi DZ 1145 KM068980 KM069087 KM069194
Hylarana siberu BJE203 KF477742
Hylarana signata FMNH 273117 KR264086 KR264178
Hylarana similis KU 306511 KF477775
Hylarana spinulosa MVZ 236683 KR264093 KR264185
Hylarana sreeni BNHS 5872 KM068994 KM069102 KM069208
Hylarana supragrisea BPBM24218 KR264055 KR264146
Hylarana taipehensis USNM 595412 OM387135 OM420406
Hylarana temporalis DZ 1092 KM068995 KM069103 KM069209
Hylarana tytleri USNM 583137 MG935833 MG935539
Hylarana urbis BNHS 5841 KM069008 KM069116 KM069221
Hylarana volkerjane RG7636 KR264104 KR264195
Hylarana waliesa BPBM16384 KR264046 KR264137

Sequences were aligned using MAFFT 7.471 (Katoh and Standley 2013) with default parameters. Phylogenetic analyses were constructed, based on the concatenated sequences of 16S, COI and cytb genes. The best fit models were selected using the corrected Akaike Information Criterion Akaike Information Criterion (AICc) in ModelFinder which was implemented in IQ-TREE 1.6.12 (Kalyaanamoorthy et al. 2017). Bayesian Inference (BI) was performed in MrBayes 3.2.7 (Ronquist et al. 2012), based on the GTR+F+I+G4 substitution model for all partitions and the Markov chains were run for 5,000,000 generations and sampled every 100 generations. Maximum Likelihood (ML) analysis was performed in IQ-TREE 1.6.12 (Nguyen et al. 2015), based on the GTR+F+R4 substitution model for 16S, the TIM2+F+I+G4 substitution model for COI and the TVM+F+I+G4 substitution model for cytb and branch support was assessed using 1,000 ultrafast bootstrap replicates.


Morphologically, the specimen from the type locality of Rana bannanica well agrees with the original description of R. bannanica in that all measurements and the relative lengths of body parts of this specimen are within the ranges of those in the type series of R. bannanica (n = 8) (Table 2). Therefore, we consider this specimen to belong to the species described by Rao and Yang (1997). However, the ratio of head width to head length in males of R. bannanica (n = 9) is relatively smaller than that in males of H. milleti (n = 5) (HW/HL 0.31–0.35 vs. 0.35–0.36), the ratio of snout length to snout-vent length in males of R. bannanica is relatively larger than that in males of H. milleti (SL/SVL 0.17–0.18 vs. 0.15–0.17) and the ratio of total leg length to snout-vent length in males of R. bannanica is relatively smaller than that in males of H. milleti (LEG/SVL 1.53–1.69 vs. 1.70–1.78) (Table 2). In addition, R. bannanica differs from H. milleti by having relatively larger discs on tips of fingers and toes, having relatively larger outer metatarsal tubercle, having tarsal folds and having vocal sacs in males.

Table 2.

Morphological comparison between the type and topotypic specimens of Hylarana bannanica and the type specimens of H. milleti. Data for the type specimens of H. bannanica and H. milleti were obtained from the original descriptions by Rao and Yang (1997) and Smith (1921), respectively.

Hylarana bannanica Hylarana bannanica Hylarana milleti Hylarana milleti
n = 8 n = 1 (KIZ2019BN) n = 5 n = 5
Holotype and paratypes Topotype Syntypes Syntypes
SVL 38.0–43.0 40.4 36–39 45–48
HL 10.5–17.0 16.2 14–14.5 16–17
HW 12.0–14.0 13.5 12.5–14 15–16
HL/SVL 0.28–0.44 0.40 0.36–0.39 0.33–0.37
HW/SVL 0.31–0.35 0.33 0.35–0.36 0.31–0.35
HW/HL 0.76–1.14 0.83 0.89–1.00 0.94
SL 7.0–7.5 6.8 6 7–8
SL/HL 0.41–0.67 0.42 0.41–0.43 0.41–0.47
SL/SVL 0.17–0.18 0.17 0.15–0.17 0.15–0.17
ED / 5.0 4–4.5 5
ED/HL / 0.31 0.29–0.32 0.29–0.31
ED/SVL / 0.12 0.10–0.13 0.10–0.11
IND 3.8–4.2 3.9 3–4 4–5
TD 4.1–5.0 4.1 3.4–4 4
TD/HL 0.25–0.43 0.25 0.24–0.29 0.24–0.25
TD/SVL 0.10–0.12 0.10 0.09–0.11 0.08–0.09
FHL 18–20 19.2 / /
ARM / 25.2 21–24 27–29
TL 19.8–23.0 21.2 20–21 25–27
TL/SVL 0.52–0.56 0.52 0.54–0.56 0.53–0.60
TFL 29.0–33.0 32.7 / /
FL 20.0–23.0 22.9 20–21 25–27
FL/SVL 0.51–0.58 0.57 0.54–0.56 0.53–0.58
LEG 58–67 68.2 62–67 79–85
LEG/SVL 1.53–1.69 1.69 1.70–1.78 1.49–1.80

In phylogenetic analyses of mtDNA sequences, BI and ML methods yielded similar results. The newly-generated sequences of Hylarana milleti, as well as the sequences of the specimens of H. milleti from Lam Dong and Gia Lai provinces in Vietnam and the newly-generated sequence of the specimen from the type locality of Rana bannanica, as well as the sequences of the specimens previously identified under the name H. milleti from Phongsaly Province in Laos, Chanthaburi Province in Thailand and Tanintharyi Region in Myanmar, formed two distant, separate clades within the genus Hylarana (Fig. 2). Clearly, the specimens previously considered to be H. milleti from China, Laos, Thailand and Myanmar and the specimens of H. milleti from Vietnam belong to different species, although their phylogenetic positions within the genus have not been resolved. Therefore, we formally remove Rana bannanica Rao & Yang, 1997 from the synonym of Hylarana milleti Smith, 1921.

Figure 2. 

Bayesian phylogenetic tree of Hylarana inferred from the concatenated 16S, COI and cytb sequences. Numbers before slashes indicate Bayesian posterior probabilities (≥ 0.90) and numbers after slashes indicate ML bootstrap supports (≥ 90).

Hylarana bannanica (Rao & Yang, 1997)

Figs 3, 4, 5A

Rana bannanica Rao & Yang, 1997.

Type material

Holotype. KIZ 94001, adult male. Paratypes. KIZ 94002–KIZ 94008, seven adult males.

Type locality

Mohan Town, Mengla County, Xishuangbanna Prefecture, Yunnan Province, China.

Specimen examined

KIZ2019BN, adult male, collected by Shuo Liu from Manzhuang Village, Mohan Town, Mengla County, Xishuangbanna Prefecture, Yunnan Province, China (21°21'51"N, 101°37'17"E, 760 m elevation), on 7 May 2019.

Figure 3. 

The holotype (KIZ 9401) of Hylarana bannanica in preservative. A. Dorsal view; B. Ventral view; C. Close-up view of the right side of the head; D Close-up view of the left side of the head. Photos by Dingqi Rao.

Description of the newly-collected topotypic specimen

SVL 40.4 mm; head width less than head length (HW/HL 0.83); snout pointed, projecting beyond lower jaw; nostril slightly closer to snout than to eye; canthus rostralis distinct; loreal region vertical, slightly concave; snout length greater than eye diameter; pineal body indistinct; tympanum relatively large (TD/ED 0.82), rounded; vomerine teeth distinct; choanae close to vomerine teeth; tongue oval, posterior notch U-shaped; vocal sac openings small, on floor of each corner of mouth; vocal sacs present.

Figure 4. 

The topotypic specimen (KIZ2019BN) of Hylarana bannanica in preservative. A Dorsal view; B Ventral view; C Close-up view of the right side of the head; D Close-up view of the left side of the head; E Close-up view of the left palm; F Close-up view of the left sole. Photos by Shuo Liu.

Forelimbs relatively slender; relative lengths of fingers III > I ≥ IV > II; all finger tips expanded into small discs; webbing on fingers absent; subarticular tubercles distinct, oval and prominent; supernumerary tubercles below base of fingers II–IV present, small and rounded; thenar tubercle oval, large; median metacarpal tubercle oval, prominent; outer metacarpal tubercle elongated, indistinct; grey nuptial pad on finger I indistinct.

Hind limbs medium long; relative lengths of toes IV > V > III > II > I; all toe tips expanded into small discs, discs on toes slightly larger than on fingers; webbing on toes undeveloped; subarticular tubercles distinct, oval, prominent; inner metatarsal tubercle oval; outer metatarsal tubercle rounded; tarsal fold present.

Dorsal skin slightly rough, upper lateral skin granular, lower lateral and ventral skin smooth; dorsolateral folds distinct, supratympanic fold absent; one white gland above base of forelimb behind tympanum on each side; some tiny tubercles form indistinct skin ridges on dorsal tibia.

In preservative, dorsum light brown, lateral side greyish-brown, ventral side of head and limbs yellowish-white, belly greyish-white, iris black, pupil white.


Currently, Hylarana bannanica is known to be distributed in Xishuangbanna Prefecture, Yunnan Province, China, as well as northern Laos (Phongsaly Province), Thailand (Chanthaburi, Ubon Ratchathani, Amnat Charoen, Sa Kaeo, Prachin Buri, Nakhon Ratchasima and Chachoengsao provinces), southwest Cambodia (Cardamon Mountains) and peninsular Myanmar (Taninthary Region), based on data from Rao and Yang (1997), Chuaynkern et al. (2004), Stuart and Emmett (2006), Yang and Rao (2008), Poyarkov et al. (2021), Zug and Mulcahy (2020), Zug (2022) and this study.

Conservation status

Further research is required to clarify the extent of the distribution, population size, trends and conservation status of the species. Hylarana bannanica is distributed over a large area including many protected areas. Across its range, the species seems not to be rare. Thus, we tentatively suggest Hylarana bannanica be considered a Least Concern (LC) species, following the IUCN’s Red List Categories (IUCN Standards and Petitions Committee 2019).


Dubois (1992) partitioned the genus Rana into many sections, one of which is the section Hylarana, which contains two subsections, namely subsection Hydrophylax and subsection Hylarana. He treated Hydrophylax as a subgenus and put it together with the newly-erected subgenera Amnirana, Humerana, Papurana, Pulchrana and Sylvirana in the subsection Hydrophylax and treated Clinotarsus, Glandirana, Hylarana, Odorrana and Pterorana as subgenera and put them together with the newly-erected subgenera Chalcorana, Eburana, Nasirana, Sanguirana and Tylerana in the subsection Hylarana and explicitly stated that these arrangements were preliminary hypotheses (Dubois 1992). Oliver et al. (2015) revised Hylarana sensu lato by utilising an inclusive molecular phylogeny, elevated Amnirana, Chalcorana, Hydrophylax, Hylarana, Humerana, Papurana, Pulchrana and Sylvirana to genus rank and erected two new genera, namely Abavorana and Indosylvirana. Chan et al. (2020) performed a comprehensive analysis of Hylarana sensu lato and also recognised the ten genera Abavorana, Amnirana, Chalcorana, Hydrophylax, Hylarana, Humerana, Indosylvirana, Papurana, Pulchrana and Sylvirana in it. Later, Dubois et al. (2021) found Chalcorana to be paraphyletic with Pulchrana and Amnirana and Sylvirana, Papurana, Hydrophylax, Hylarana and Indosylvirana to be polyphyletic; therefore, they placed all of the genera within Hylarana sensu lato, except for Abavorana, into the synonymy of Hylarana. However, we consider that these synonymous genera may be valid and need to be resurrected, at least for some of them. For example, Dubois et al. (2021) recovered Sylvirana to be polyphyletic due to Sylvirana guentheri as the sister taxon of Humerana miopus. In our phylogenetic analysis, Hylarana guentheri was also nested in the subgenus Humerana with strong support. Nevertheless, we cannot say directly that Sylvirana or Humerana is invalid, unless Hylarana guentheri is the type species of Sylvirana. Hylarana guentheri is a very common species in southern China and northern-central Vietnam and we have collected specimens of this species multiple times. This species has a large body size, which is different from most species of the subgenus Sylvirana and consistent with most species of the subgenus Humerana. Since Hylarana guentheri is not the type species of Sylvirana, therefore, we consider that Hylarana guentheri should be moved to Humerana from Sylvirana, and Sylvirana and Humerana are still valid, instead of synonymising them. Anyway, we still adopt the current arrangement (Dubois et al. 2021; Frost 2023) of Hylarana sensu lato for the time being; further morphological and molecular studies are required to elucidate the division of genera within Hylarana sensu lato.

Hylarana bannanica was described from Xishuangbanna, Yunnan, China, in 1997 and no new specimens from its type locality have been reported since then. The specimen (KIZ2019BN) we collected is the first rediscovery from the type locality of this species since it was described more than 20 years ago. Unfortunately, this specimen was misidentified as a subadult of the local common species H. cubitalis (Smith, 1917) and was not taken seriously when it was collected, so that the habitat information at the collection site and photos of this specimen in life were not recorded. In the subsequent process of specimen sorting, we discovered that this specimen is an adult male due to the presence of nuptial pad and vocal sacs, but without a large oval gland on the inner side of each elbow, which shows it is obviously not H. cubitalis, so we conducted a detailed study on this specimen. This experience reminds us that it is necessary to have sufficient taxonomic competence when doing fieldwork and that those who visit different locations should be aware of what species are distributed or what species may be distributed there.

In previous studies (e.g. Frost et al. (2006); Oliver et al. (2015); Chan et al. (2020); Dubois et al. (2021); Reilly et al. (2022)), Hylarana milleti (Fig. 5B) was once placed in the genera or subgenera Sylvirana, Indosylvirana and Papurana, respectively. In our phylogenetic analysis, it is evident that H. milleti and H. bannanica are not conspecific, but their phylogenetic positions within the genus have not been resolved. At present, we cannot determine which subgenus these two species belong to respectively. The systematic relationship within this genus needs to be further resolved.

Figure 5. 

Comparison between Hylarana bannanica (A from Chanthaburi, Thailand) and H. milleti (B from Bidoup-Nui Ba, Lam Dong, Vietnam) in life. Photos by Parinya Pawangkhanant (A) and Nikolay A. Poyarkov (B).


We thank Parinya Pawangkhanant (AUP) for providing the photo of Hylarana bannanica from Thailand. We thank George Zug (USNM) for his help in this study. Thanks to the forest rangers of Xishuangbanna National Nature Reserve for their help in the field. We also thank the editors and reviewers for their working on the manuscript. This work was supported by the Science-Technology Basic Condition Platform from the Ministry of Science and Technology of the People’s Republic of China (Grant No. 2005DKA21402), the project of the second comprehensive scientific investigation of Xishuangbanna National Nature Reserve, the project of Ministry of Ecology and Environment of China: investigation and assessment of amphibians and reptiles in Jinghong City, Menghai County and Mengla County, in part the Russian Science Foundation to N.A. Poyarkov (Grant No. 22-14-00037, molecular analyses) and in part the Rufford Foundation to T.V. Nguyen (Grant No. 39897-1; data analysis).


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