Cryptic Species of a Cascade Frog from Southeast Asia: Taxonomic Revisions and Descriptions of Six New Species

PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3417, 60 pp., 14 ®gures, 15 tables October 29, 2003

RAOUL H. BAIN,1 AMY LATHROP,2 ROBERT W. MURPHY,3 NIKOLAI L. ORLOV,4 AND HO THU CUC5

ABSTRACT

The Southeast Asian cascade frog, Rana livida (Blyth, 1856), has long been suspected to be a complex of species. Several different forms are described from across its vast range. The loss of type material and disparate sampling efforts are challenges. Is variability in this species due to geographic variation or to the presence of multiple species? We use concordant evidence from morphology, morphometrics, cellular DNA content, and allozyme electrophoresis to in- vestigate diversity in R. livida from Vietnam. Three distinct species are recognized on the basis of morphology, as are four other suspect groups (morphotypes). Discriminant function analyses of morphometric data detect patterns of morphological variation among all seven groups. Pairwise comparison of cellular DNA content using t-tests shows signi®cant differences among sympatric morphotypes, suggesting they represent distinct species. This hypoth- esis is supported by an analysis of 14 allozymic loci, in which ®xed allelic differences are found among specimens in sympatry and allopatry. Examination of available type material of four junior synonyms of R. livida results in their recognition as species. One of these species,

1 Biodiversity Specialist, Center for Biodiversity and Conservation, and Division of Zoology (Herpetology), Amer- ican Museum of Natural History. e-mail: [email protected] 2 Herpetology Technician, Centre for Biodiversity and Conservation Biology, Royal Ontario Museum, 100 Queen's Park, Toronto, Ontario M5S 2C6, Canada. e-mail: [email protected] 3 Senior Curator, Herpetology, Centre for Biodiversity and Conservation Biology, Royal Ontario Museum, 100 Queen's Park, Toronto, Ontario M5S 2C6, Canada. e-mail: [email protected] 4 Research Scientist, Department of Herpetology, Zoological Institute, Russian Academy of Sciences, St. Petersburg, 199034, Russia. e-mail: [email protected] 5 Senior Herpetologist, Institute of Ecology and Biological Resources, Vietnam National Center for Natural Sciences and Technology, Nghia Do, Tu Liem, Hanoi, Socialist Republic of Vietnam.

R. chloronota, is a wide-ranging species erroneously referred to as R. livida. Seven species occur in Vietnam. We describe six new cryptic species belonging to the Rana chloronota complex, redescribe R. chloronota, R. livida, R. sinica, and R. graminea, and give comments on R. leporipes. Three of these new species (R. bacboensis, new species, R. hmongorum, new species, and R. daorum, new species) occur in montane forests in northern Vietnam, and two (R. banaorum, new species and R. morafkai, new species) are known only from the Tay Nguyen Plateau of Vi- etnam's Central Highlands. One species, R. megatympanum, new species, occurs in portions of both northern Vietnam and the Central Highlands. An identi®cation key for the Rana chloronota complex from Vietnam is provided. The ®nding of six cryptic species within a small portion of the geographic region of R. chloronota suggests that many more cascade ranids await discovery. This documentation has serious implications for conservation; each of the new species occurs in sympatry with at least one other member of the complex. Conse- quently, far more species are being affected by habitat loss than was previously thought.

INTRODUCTION (Inger, 1996). Rana livida has odoriferous, highly toxic skin secretions (Pope, 1931; Recent herpetological surveys in Vietnam Karsen et al., 1998). have resulted in the discovery of several pre- The taxonomy of R. livida is confusing viously undescribed species of frogs (e.g., and unstable. Variation within R. livida is Lathrop et al., 1998a, 1998b; Inger et al., documented. Taylor (1962) reported that 1999; Ohler et al., 2000; Orlov et al., 2001; male R. livida from Thailand are signi®cantly Ziegler and Kohler, 2001). Similarly, several larger than other conspeci®c males (Boulen- Southeast Asian species with vast ranges ger, 1882, 1920; Bourret, 1942). Inger and have been found to be complexes of cryptic Chanard (1997) also reported that male R. speciesÐmultiple species currently referred livida from northern Thailand differ in hav- to as a single taxon (Emerson, 1998; Inger, ing larger snout±vent lengths and colorless 1999; Fei et al., 2001). Disparate sampling spinules along the chin, throat, chest, and up- efforts contribute to this problem of uniden- per lip. Despite these observations, R. livida ti®ed cryptic species. One complex of cryp- has not been studied across its range. Major tic species likely involves an odoriferous, systematic works on Southeast Asian am- green cascade frog Rana livida (Werner, phibians continue to treat R. livida as a single 1930; Taylor, 1962; Murphy et al., 1997; In- taxon (e.g., Liu and Hu, 1961; Taylor, 1962; ger, 1999). Frost, 1985; Fei et al., 1990; Yang, 1991a; Rana livida (Blyth, 1856) occurs near fast- Zhao and Adler, 1993; Inger et al., 1999; moving montane rivers and streams of the Frost, 2002). Comparative study has been Southeast Asian mainland, from southern complicated by two factors. First, the type China, across Indochina, to the eastern Him- series of R. livida, and those of some of its alayas in India (Boulenger, 1920; Smith, currently recognized junior synonyms, can- 1921; Pope, 1931; Smith, 1931; Bourret, not be located because either the original de- 1942; Liu and Hu, 1961; Taylor, 1962; Frost, scriptions do not include specimen accession 1985; Zhao and Adler, 1993; Fellows and numbers or the types are lost. Second, the Hau, 1997; Fei, 1999; Stuart, 1999; Frost, type specimens are from disparate parts of 2002). Like all Southeast Asian cascade ra- the extensive distribution of R. livida. Thus, nids, it is dorsoventrally compressed and has it has been unclear whether the morpholog- long, powerful legs and fully webbed feet. ical variability of this species owes to geo- These attributes facilitate swimming in fast- graphic variation or the presence of multiple moving water. It has greatly expanded toe species. pads that allow it to cling to rocks in the Is R. livida a single species or a set of torrents and vegetation in the surrounding cryptic species? We used concordant evi- forest. The snout±vent length of female R. dence from morphology, morphometrics, cel- livida is nearly twice that of males. Females lular DNA content, and allozymic loci to in- have white eggs, but the larvae are unknown vestigate this question in a series of frogs 2003 BAIN ET AL.: CASCADE FROG FROM SOUTHEAST ASIA 3 from Vietnam. Because Vietnam represents to conclude that Polypedates smaragdinus only a small portion of the proposed range fails to conform to Article 12 of the Inter- of R. livida, this study asseses the potential national Zoological Code of Nomenclature variability that may be found across its large (1999). distribution. We also review the junior syn- Polypedates lividus: Blyth (1856) de- onyms and brie¯y discuss the speci®c, ge- scribed P. lividus in a report on specimens neric, and subgeneric taxonomic history in procured by Capt. Berdmore and W. Theo- order to establish a stable taxonomy. bold from Burma (now Myanmar). However, the type locality is uncertain. Although Blyth SYSTEMATIC REVIEW OF RANA LIVIDA (1856) reported that the specimens given to (BLYTH, 1856) him by Theobold were from the Tenasserim River Valley, Burma, Theobold (1860: 325) Rana livida (Blyth, 1856) reported that ``Mr. Blyth described a new ?Polypedates smaragdinus Blyth, 1852 Polypedates lividus Blyth, 1856 species of tree frog received from Major ?Rana nebulosa Hallowell, 1861 Berdmore at Mergui.'' Mergui (today Myeik) Polypedates chloronotus GuÈnther, 1875 is a coastal seaport, far from inland Tenas- Rana chloronota Boulenger, 1882 serim Valley. Rana livida Boulenger, 1887 The type series of P. lividus is not present Rana graminea Boulenger, 1899 in the Indian Museum Reptile Registry, any Rana (Hylorana) livida Boulenger, 1920 catalog or other early reports in the Indian Rana (Hylorana) sinica Ahl, 1925 Museum, and it cannot be found in the Zoo- Rana leporipes Werner, 1930 logical Survey of India collections (Chanda Odorrana livida Fei, Ye, and Huang, 1990 et al., 2000). The type specimens of P. livi- Rana (Eburana) livida Dubois, 1992 dus are neither in the British Museum (Nat- Polypedates smaragdinus: Blyth (1852) ural History) (Clarke, personal commun.), ambiguously described Polypedates smar- nor are they referred to in any other publi- agdinus in a three sentence description: `À cation other than the original description. No tree frog from the Naga hills, Assam (P. illustrations accompanied the description. smaragdinus, nobis). Length of the body 3¼ Presumably, the type series is lost. inches, hind limb 5¼ inches. Wholly green Polypedates chloronotus:GuÈnther (1875) above, changing in spirit to livid blue; undescribed another series of frogs collected by derparts white.'' No illustrations are associ- Jerdon from Darjeeling, India as Polypedates ated with the description and type specimens chloronotus, noting that it resembles a ``Hy- are not designated. Polypedates smaragdinus lorana''. He included specimens from the is not mentioned in the Indian Museum Rep- Khasi Hills, India, that Jerdon (1870) and tile Registry, in any catalog, or in other early Anderson (1871) referred to as P. smarag- reports in the Indian Museum (Chanda et al., dinus. Boulenger (1882) reevaluated the syn- 2000). There are no vouchers in the collec- types and assigned P. chloronotus to the ge- tions of the Zoological Survey of India col- nus Rana based on extensive webbing and T- lections (Chanda et al., 2000). The type ma- shaped distal phalanges. Reporting on the terial is not in the British Museum (Clarke, ``Fea series'' from northern Tenasserim Bur- personal commun.). Presumably, the types ma, Boulenger (1887) noted that some spec- are lost. Chanda et al. (2000) erroneously re- imens are ``beyond doubt to belong to the ported that Anderson (1871) redescribed the species named Polypedates lividus by Blyth, ``now lost syntypes'' of P. smaragdinus;in and also to be inseparable from GuÈnther's P. contrast, Anderson's redescription was based chloronotus''; he made Rana chloronota a on specimens already described by Jerdon junior synonym of P. lividus,asR. livida. (1870) from the Khasi Hills, India. GuÈnther Rana livida and R. chloronota differ sig- (1875) and Boulenger (1887) correctly stated ni®cantly: R. livida is `ùniformly dusky- that Blyth's original description was so vague plumbeous above'', but R. chloronota has a that it could be referred to a number of cas- green dorsum with dark sides that are cade ranids. This, coupled with the absence ``sharply de®ned'' and have ``dark cross of an illustration and type material, leads us bars'' on the limbs. This diagnosis of P. 4 AMERICAN MUSEUM NOVITATES NO. 3417 chloronotus agrees with the color plate that onymized R. graminea with R. chloronota accompanies the original description. More- (as R. livida) based on two male specimens over, whereas R. livida has smooth skin and from the Nakon Sritamarat Mountains of the is `ònly slightly granulose on the hind sur- Malay Peninsula. Nevertheless, some subse- face of the thighs'', R. chloronota has a quent works recognized R. graminea (e.g., smooth dorsum and granular ¯anks. These Pope, 1931; Boring, 1932; Bourret, 1942). characteristics, as well as the differences in Bourret (1942) considered R. graminea to be color patterns on the thighs and ¯anks, re- a ``northern variety'' of Indochinese R. chlo- main visible on the syntypes of R. chloronota ronota (as R. livida) because of its weak dor- and Fea's series of R. livida. Boulenger's solateral fold. Inger et al. (1999) did not ®nd (1890) description of R. livida in the Fauna evidence of R. graminea in their series of of British India is actually R. chloronota. All frogs from central Vietnam. Currently, R. subsequent works have used Boulenger's graminea is a junior synonym of R. chloron- (1890) taxonomy (e.g., Boulenger, 1920; ota (as R. livida) (Zhao and Adler, 1993; Smith, 1921; Pope, 1931; Smith, 1931; Bour- Frost, 2002), although this is unjusti®ed. ret, 1942; Liu and Hu, 1961; Taylor, 1962; Rana sinica: Ahl (1925) described Hylor- Fei et al., 1990; Yang, 1991a; Zhao and Ad- ana sinica, and Bourret (1942: 371) listed it ler, 1993; Inger and Chanard, 1997; Fellows as a possible synonym of R. chloronota (as and Hau, 1997; Karsen et al., 1998; Fei, R. livida): ``?Rana (Hylarana) sinica''. How- 1999; Inger, 1999; Inger et al., 1999; Stuart, ever, many differences separate the two spe- 1999; Ziegler, 2002; Orlov et al, 2002). cies: in R. sinica, ®nger I is shorter than ®n- Rana nebulosa: Hallowell (1861) de- ger II, and ®nger III is shorter than the snout; scribed this species from Hong Kong based in R. chloronota,IIϽ I Ͻ IV Ͻ III, and on a juvenile specimen. The ambiguous de- ®nger III is longer than snout. Rana sinica scription did not designate a type specimen. has large, round black spots on its ¯anks but The type is not included in the list of her- R. chloronota does not. Rana sinica lacks the petological type specimens of the National white lip-stripe that is present in R. chloron- Academy of Sciences, Philadelphia (Malna- ota. The tympanum of R. sinica is covered te, 1971), and it is not present in the collec- by a layer of skin, but in R. chloronota the tion (Gilmore, personal commun.). It is lost. distinct tympanum does not have a super®- Because the validity of the species can nei- cial layer of skin. It is still recognized as a ther be con®rmed nor denied, Boulenger (1882) considered R. nebulosa to be an un- junior synonym (Frost, 2002). certain species. Dubois and Ohler (2000) Rana leporipes: Werner (1930) described suggested that R. nebulosa is a possible ju- this species as a southern Chinese member nior synonym of R. chloronota (as R. livida) of the ``Rana livida group''. It differs from because its description does not contradict R. chloronota in having slender legs that lack that of R. chloronota. They advocated des- transverse bands, smaller disks, a white su- ignation of a lectotype from Hong Kong. We pratympanic fold, a weak dorsolateral fold, agree with Boulenger's assessment and con- and webbing that only reaches the base of clude that R. nebulosa fails to conform to the distal phalanx. Bourret (1942) synony- Article 12 of the International Zoological mized R. leporipes with R. livida var. gra- Code of Nomenclature (1999). minea, and Liu and Hu (1961) agreed with Rana graminea: Boulenger (1899) de- this synonymy. The type series of R. lepo- scribed Rana graminea from the Five Finger ripes is not in the Museum fuÈr Naturkunde Mountains on Hainan Island, China. Al- der Humboldt-UniversitaÈt, Berlin, Germany, though it closely resembles R. chloronota and is considered lost; all of Werner's col- (Boulenger, 1899, 1920), the two male syn- lections are deposited there. It is likely that types have a larger tympanum, oblique, near- the specimens were lost during World War II ly vertical (versus concave) loreal region, (Guenther, personal commun.). Despite its smaller hands, smaller digital disks, smooth current recognition as a junior synonym ¯anks (versus granulate), and weak dorsolat- (Frost, 2002), the diagnostic characters indi- eral folds (versus absent). Smith (1930) syn- cate that R. leporipes is a valid species. 2003 BAIN ET AL.: CASCADE FROG FROM SOUTHEAST ASIA 5

GENERIC AND SUBGENERIC PLACEMENTS OF and Ho, 1996; Karsen et al., 1998; Inger, RANA LIVIDA 1999; Inger et al., 1999). The status of Hy- larana remains controversial. Whereas Hy- Rana chloronota (as R. livida) is variously larana is usually considered to be a genus by placed in subgenera Eburana (Dubois, 1992; those working on the African fauna, those Duellman, 1993), Hylarana (Boulenger, working on the Asian fauna consider it to be 1920; Bourret, 1942), and Odorrana (Fei et a subgenus of Rana (Frost, 1985, 2002). al., 1990; Matsui, 1994; Zhao, 1994; Fei et Odorrana Fei, Ye, and Huang, 1990: Fei al., 1999). To add confusion, the latter two et al. (1990) placed 12 species of Chinese taxa are occasionally considered to be gen- odoriferous Hylarana in a new genus, Odor- era. Rana chloronota has diagnostic charac- rana. The genus is diagnosed primarily by ters that are unique to each of these taxa: external morphology and sternal elements, fully webbed feet are typical of Rana, en- including a very small and cartilaginous larged toe disks of Hylarana, odoriferous se- omosternum where the omosternal bone (ϭ cretions of Odorrana, and white eggs of style) is not forked; mesosternum long and Eburana. We review the generic and subge- slender and proximally much thicker; and neric taxonomy that includes R. chloronota xiphisternal cartilage much larger than omos- and discuss the validity of placement into ternum and deeply notched posteriorly (Fei each group. et al., 1990). The odoriferous properties of Rana Linneaus, 1758: The content and di- the skin are not included as a diagnostic agnosis of the genus Rana has changed since character, even though the generic name was its original description. Boulenger (1887) derived from this trait. Although Boulenger thought that Polypedates chloronotus be- (1920) placed R. chloronota (as R. livida)in longed in Rana because of its extensive web- section Ranae (Hylorana [ϭ Hylarana] chal- bing and T-shaped terminal phalanges; R. conotae), results from karyological studies chloronota remained in the genus Rana for suggest that it is a member of the Rana (Hy- almost a century. larana) andersonii group (Li and Wang, Hylarana Tschudi, 1838: Hylarana has 1985; Wei et al., 1993). Consequently, Fei et horseshoe-shaped circummarginal grooves al. (1990) placed the Rana andersonii group on the ventral surface of very large digital and R. chloronota (as R. livida) into Odor- disks. Boulenger (1882) originally thought rana. Neither Yang (1991a) nor Zhao and that Hylarana should not be recognized. Lat- Adler (1993) recognized Odorrana. Others er, he (Boulenger, 1920) revised its de®ni- have suggested that Odorrana be considered tion, making it a subgenus, but he errone- a subgenus because many species of Rana ously used the name Hylorana instead of Hy- share some of its ``diagnostic'' characters larana. (Dubois, 1992; Inger, 1996; Matsui, 1994; Rana chloronota conforms to the diagno- Zhao, 1994). sis of Hylarana in having dilated disks with Eburana Dubois, 1992: Dubois (1992) es- circummarginal grooves, T-shaped terminal tablished Subsection Hylarana with 10 sub- phalanges, and an unforked omosternal style. genera, including revisions of Hylarana and Boulenger (1920) placed R. chloronota (as R. Odorrana, and the description of a new sub- livida) in the subgenus Hylorana [ϭHylar- genus Eburana. Eburana is diagnosed by larana], section Ranae chalconotae, allying it val denticular formula (4±5/4), the absence with R. chalconota, R. hosii, and R. grami- of nuptial spines on the chest of the males, nea, among others. He recognized that the and the absence of pigmented eggs. Subsec- subgenus Hylorana [ϭHylarana] as an ``un- tion Hylarana is de®ned by the lack of hu- natural assemblage'' because it is formed on meral glands in the males. Dubois removed the basis of specialized ®ngers and toes. For R. chloronota (as R. livida) from Odorrana the next 70 years, R. chloronota was referred and placed it into Eburana (along with R. to as a member of the subgenus Hylarana swinhoana and R. narina). (e.g., Bourret, 1942), or its subgeneric place- Dubois' (1992) provisional classi®cation ment was ignored entirely (e.g., Pope, 1931; of the family Ranidae has received spirited Taylor, 1962; Zhao and Adler, 1993; Nguyen criticism due to philosophical and empirical 6 AMERICAN MUSEUM NOVITATES NO. 3417 issues. Criticisms focus on the phenetic basis IEBR Institute of Ecology and Biological Re- of the classi®cation, a selective treatment of sources, Hanoi characters inferred from the literature, and an ROM Royal Ontario Museum, Toronto inadequate analysis of variation of the de®n- ZMB Museum fuÈr Naturkunde der Hum- ing characters (Emerson and Berrigan, 1993; boldt-UniversitaÈt, Berlin Matsui et al., 1995; Inger, 1996, 1999). Data from karyological studies suggest that R. METHODOLOGICAL OUTLINE chloronota shares a closer af®nity with Specimens of R. chloronota were segre- Odorrana (sensu Fei et al., 1990) than with gated a priori into morphotypes. Consistently other Eburana (Wei et al., 1993; Matsui, detected morphological and molecular 1994; Matsui et al., 1995). Rana chloronota groups were considered to be distinct spe- is the only member of Eburana to have odor- cies. We analyzed morphometric data with iferous skin secretions. Furthermore, the lar- discriminant function analyses to distinguish vae of R. chloronota are unknown, making among forms and shapes. We also used t-test it impossible to evaluate this as a diagnostic evaluations of cellular DNA content data to character (Inger, 1996). differentiate among sympatric species, as- suming that conspeci®c individuals have TAXONOMIC CONCLUSIONS equal amounts of DNA. We further evaluated We adopt the collective recommendation species boundaries with allozyme electro- of Inger (1996), Matsui (1994), and Zhao phoresis. Fixed allelic differences provided (1994) in recognizing Odorrana as a subge- diagnostic characters for species. nus of Rana. The monophyly of Odorrana is supported by a phylogenetic analysis of more GENERAL COLLECTING PROCEDURES than 2000 base pairs of mitochondrial DNA Specimens of Rana chloronota from Hong sequence data (Chen and Murphy, unpubl.). Kong, and eight localities in Vietnam were We recognize P. smaragdinus and R. nebu- collected at various times of the year from losa as nomen dubium, since they fail to con- 1994 to 1999. Localities in Vietnam includ- form to Article 12 of the Code (1999). We ed: Hoang Lien Mountains near Sa Pa (Lao also remove R. leporipes, R. sinica, R. gra- Cai Province); Ba Be National Park (Bac minea, and R. chloronota from synonymy Kan Province); Tam Dao Mountain, Tam because each is de®ned by distinct, diagnos- Dao National Park (Vinh Phu Province); Na tic characters. Below we redescribe R. sinica, Hang Nature Reserve (Tuyen Quang Prov- R. graminea, R. chloronota, and R. livida, ince); Con Cuong region (Nghe An Prov- and comment on R. leporipes. We designate ince); and Tay Nguyen Plateau, An Khe Dis- a neotype of R. livida, since no name-bearing trict (Gia Lai Province) (®g. 1). Coordinates type exists and its designation quali®es under were recorded in geographic projection using the conditions of Article 75.3 of the Code handheld GPS units (see table 1). Although (1999). Because the voucher type specimens error and precision were carefully monitored, of R. leporipes have been lost, the type spec- position coordinates collected before 1 May imen of R. leporipes is now the photograph 2000 were subject to the effects of selective of the holotype (an iconotype), which is pub- availability (White House Press Secretary, lished as ®gures 1 and 2 in the original de- 2000). Because real-time and postdifferential scription (Werner, 1930: 49, pl. IV). corrections were not feasible or available in this region, positional accuracy should be MATERIALS AND METHODS judged accordingly. COLLECTION ABBREVIATIONS Most frogs were collected at night and all were euthanised using MS-222 or chlorobu- AMNH American Museum of Natural History, New York tanol within 24 hours of collection. Samples BMNH British Museum (Natural History), of liver, leg muscle, and heart were taken London from representative specimens. Frozen tis- FMNH Field Museum of Natural History, Chi- sues deposited in the ROM were ¯ash frozen cago in liquid nitrogen. All tissue samples in the 2003 BAIN ET AL.: CASCADE FROG FROM SOUTHEAST ASIA 7

Fig. 1. Map of Vietnam. Collecting localities of odoriferous frogs of the genus Rana are indicated by numbers, referable to table 1.

AMNH were preserved in 95% ethanol. quently stored at Ϫ80ЊC in the ROM. Frogs Blood samples obtained for ¯ow cytometry were ®xed in either 90% ethanol for 24 hours were suspended in a freezing solution (Vin- or in 10% formalin for several hours and delov et al., 1982; Murphy et al., 1997), ini- subsequently rinsed in water. All tissue col- tially frozen in liquid nitrogen and subse- lections are stored in the ROM and AMNH, 8 AMERICAN MUSEUM NOVITATES NO. 3417

TABLE 1 Locality and Seasonal data for Collections of Rana chloronota When known, exact coordinates and months are provided. Missing data suggest that the information was not available. Number refers to locations indicated on map in ®gure 1.

and all vouchers are maintained in 65% eth- of its synonyms were treated as separate anol in the AMNH, ROM, or IEBR. OTUs in subsequent analyses. MORPHOLOGY: The sex of each frog was LABORATORY PROCEDURES recorded and all measurements made with A total of 343 adult R. chloronota includ- digital calipers to the nearest 0.01 mm and ing recognized synonyms were examined rounded to the nearest 0.1 mm. External (appendix 1, ®g. 1, table 1). We compared measurements included snout±vent length all recently collected samples to specimens (SVL), head length (HDL) from tip of snout of R. livida from Fea's collection of R. livida to the articulation of the jaw, head width from Myanmar (see above), to currently rec- (HDW) between left and right articulations ognized junior synonyms of R. chloronota, of the quadratojugal and maxilla, snout as well as to series of R. chloronota from length (SNT), eye diameter (EYE), interor- Fujian Province, China, Lao PDR, and India. bital distance (IOD), tympanum diameter Frogs not identi®ed as R. chloronota or any (TMP), tympanum±eye distance (TEY), ®n- 2003 BAIN ET AL.: CASCADE FROG FROM SOUTHEAST ASIA 9 ger disk length (FPL) from the base of the from six localities within Vietnam and Hong pad of III to its tip, ®nger disk width (FPW) Kong using ¯ow cytometry (Murphy et al., at the widest part of the pad of III, toe disk 1997). DNA content analysis was carried out length (TPL), and toe disk width (TPL). Os- with a Becton Dickinson FACScan or a Bec- teological measurements taken from radio- ton Dickson FACScalibur ¯ow cytometer in graphs included the following: ®nger length the Faculty of Medicine, Department of Im- (FGR) from tip of the distal phalanx to the munology, University of Toronto, between base of the proximal phalanx of ®nger II, 1995 and 1997. Both systems used an argon hand length (HND) from distal end of radio- laser at a wavelength of 488 nm at 15 mW. ulna to tip of distal phalanx of II, foot length Fluorescence at wavelengths between 560 (FTL) from distal end of tibia to tip of distal and 640 nm was collected and digitally trans- phalanx of III, and tibial length (TIB). Pec- ferred to a histogram of mean DNA content toral girdle terminology follows Duellman concentration. Each sample was measured si- and Trueb (1986) and gular pouches are de- multaneously with a known standard, either ®ned using Inger (1956). Nerodia sipedon or Bufo woodhousi fowleri. MORPHOMETRICS: Morphometric analyses No sample was tested on both ¯ow cytome- were performed on 150 recently collected ters. Only samples measured on the same specimens from Vietnam, following the rec- machine at the same time were compared ommendations of Tabachnick and Fidell with each other to ensure accuracy; these in- (1989) and Hair et al. (1995). For this anal- cluded samples taken from a single locality. ysis, HDL (from posterior edge of otoccipital Cellular DNA content data of speci®c mor- to tip of premaxilla) and HDW (distance be- photypes were compared using t-tests to de- tween left and right quadratojugal-maxillary termine whether they exhibited statistically suture) were measured from radiographs; different levels of DNA. other measurements were taken as described PROTEIN ELECTROPHORESIS: Allozyme var- above. Raw morphometric data were log- iation was surveyed among 64 specimens of transformed and tested for normality using R. chloronota from six localities within Vi- categorized probability plots. Serial correla- etnam and six specimens from Hong Kong. tion plots were used to test for heterogeneity All morphotypes were used as a priori group- and detect redundancies among the variables. ings of prospective species. A population of A pooled within-group correlation-variance Huia nasica from Vietnam was included for matrix was used in a principal components comparison. All procedures follow protocols analysis (PCA). The ®rst principal compo- of Murphy et al. (1996). Electrophoretic con- nent (PC1) was used as a size component and ditions were optimized for 53 histochemical regressed against each log-transformed vari- stains on 11 buffer systems using a subset of able. Resulting residuals were used in a the samples (appendix 2). Locus nomencla- pooled-within group correlation-variance ture followed Murphy and Crabtree (1985) matrix for a discriminant functions analysis and Murphy et al. (1996). After optimal buff- (DFA). The effect of size on the measure- er systems were identi®ed, electrophoresis ments was factored out. All a priori classi®- was performed using the same lot of 11.5% cation error rates were made proportional to hydrolyzed potato starch (Connaught, lot no. group size. Robustness of the morphotype 514±3). Species boundaries were examined groupings was tested by reclassi®cation via population aggregation analysis modi®ed methods. When possible (i.e., where n Ͼ 15), from Davis and Nixon (1992). Genotypes of morphotypes were divided into subgroups, individuals within the same population (sym- with part of the group being used in the ini- patric morphotypes) were summarized in a tial DFA, and part being used in a reclassi- population pro®le, and groups of populations ®cation. Sexes were treated both separately were subsequently examined for ®xed allelic and together in the DFA. All analyses were differences. Populations with a sample size performed on STATISTICA ver. 5.1 (Stat- of one or two specimens were included with Soft, 1995). allopatric populations of their own morpho- CELLULAR DNA CONTENT: Cellular DNA types, regardless of their allelic pro®les. All content data were gathered for 73 specimens specimens were also evaluated using an un- 10 AMERICAN MUSEUM NOVITATES NO. 3417 biased minimum distance (Nei, 1978), and mm for females), a lip line, when present, genetic similarity (Rogers, 1972) using that is yellow (not white), the tympanum in BIOSYS ver. 1.7 (Swofford and Selander, males is enormous (120% of the eye diam- 1989). eter), the dorsal skin is shagreened with slight dorsolateral folds present in males, and RESULTS the dorsum is spotted brown-olive.

MORPHOLOGY MORPHOMETRICS Seven morphotypes occurred among specimens from Vietnam and Hong Kong. Mor- Morphometric analyses were performed photype 1, ``Typical'', conformed to the di- on female R. chloronota (no.1, ``Typical''), agnosis of R. chloronota and was assigned to ``Speckled'' (no. 3), ``Mottled'' (no. 5), and this species. None of the remaining six mor- ``Large'' (no. 7), as well as on male R. chlo- photypes was assignable to any synonym of ronota, ``Small'' (no. 4), ``Mottled'' (no. 5) R. chloronota. Specimens from Fujian Prov- and ``Southern Big Eye'' (no. 6). Small sam- ince, China, were identi®ed as Rana grami- ple sizes (n Ͻ 5) precluded the analysis of nea. the remaining morphotypes. Only R. chlo- Morphotypes 2, 3, and 4 are distinct spe- ronota included enough specimens to be di- cies based on morphological differences. vided into two subsets for reclassi®cation. Morphotype 2, or ``Black Egg'', differs from All other OTUs were included both in the all others in not having a white lip line, pos- classi®cation and reclassi®cation using dis- sessing a shagreened (not smooth) complete- criminant functions analysis (DFA). ly brown (green entirely absent) dorsum, and Categorical probability plots showed nor- in bearing black eggs. Morphotype 3, or mal frequency distribution in the log-trans- ``Speckled'', was unique in having large formed data. A high correlation between ®n- granules laterally and in males lacking gular ger length and hand length, and between toe pouches. Morphotype 4, or ``Small'', is dis- length and foot length was detected in serial tibguished by its small size, white dorsolat- correlation plots. Consequently, the ®nger eral glandules, white ¯ank spot, webbing to and toe length data were considered to be the distal subarticular tubercle but not to the redundant variables and were omitted from toe pad, a forked xiphisternum, and absence the analysis. The loadings of PC1 show that of vomerine teeth. Differences among the it represented a size factor (table 2). three remaining morphotypes were equivocal A DFA of females indicated strong, de- at the species level. Morphotype 5, or ``Mot- tectable patterns of morphological variation tled'', includes males that are relatively (F-statistic ϭ 6.86; p Ͻ 0.0001). Reclassi®- smaller than R. chloronota (SVL of 39±46 cation of all four OTUs was performed at a mm, vs. 41±53 mm) with a much larger tym- rate of 90% or higher (table 3). The discrim- panum (96% diameter of the eye) and web- inant loadings indicated that TMP had the bing that reached the distal tubercles. The greatest effect on factor 1, while FPW had dorsal skin of both males and females may the greatest effect on factor 2, and HDW and be either shagreened or smooth and their col- EYE each had the greatest load on factor 3 or varied from brown to green. Morphotype (table 4). A plot of factor 1 vs. factor 2 6, or ``Southern Big-Eye'', is very similar to showed clear differentiation among three ``Mottled''. The males are larger (42±55 OTUs: R. chloronota (``Typical'', no. 1), mm) with large tympani (89% diameter of ``Large'' (no.7), and ``Speckled'' (no. 3) (®g. the eye) and webbing to the proximal parts 2). ``Southern Big Eye'' had a centroid with- of the toe pads. The dorsal skin of males and in the 95% con®dence limit of both ``Large'' females is variable, being either shagreened and R. chloronota. A plot of factor 3 vs. fac- or smooth, spotted and/or uniform in color, tor 1 also showed separation of ``Speckled'' partially or wholly brown or green, with from the other three groups (®g. 3). slight dorsolateral folds. Morphotype 7, or A DFA of the four male OTUs resulted in ``Large'', has an SVL larger than that of R. an associated F-statistic of 9.64 (p Ͻ chloronota (48±55 mm for males, 93±105 0.0001). Rana chloronota was very distinct 2003 BAIN ET AL.: CASCADE FROG FROM SOUTHEAST ASIA 11

TABLE 2 TABLE 4 Coef®cients of the First Principal Component Factor Loadings of Female Morphogroups of the (PC1) for a Principal Components Analysis (PCA) Rana chloronota Complex from a DFA of 17 Variables Refer to text for variable abbreviations. Refer to text for variable abbreviations.

morphotypes from the south was re¯ected in the factor plot of the groups (®g. 4). Factor loadings indicated that the ®rst factor was most indicative of TMP, the second EYE, and the third FPW and HDL (table 6). A DFA of the combined male and female from ``Small'' (no. 4); the correct reclassi®- datasets resulted in an F-statistic of 6.78 (p 0.0001). All groupings were morphologi- cation scores were 97% for the former and Ͻ cally distinct, with the exception of male 100% for the latter (table 5). The two Indian ``Southern Big Eye'' (no. 6), which was re- specimens were reclassi®ed correctly as R. classi®ed as ``Mottled'' (no. 5) 54% of the chloronota. The reclassi®cation score of time (table 7). In the discriminant loadings, ``Mottled'' (no. 5) was also robust at 84%, TMP had the greatest effect on factor 1, FPW well above the a priori error rate. All incor- and EYE on factor 2, HDW and TIB on fac- rectly classi®ed ``Mottled'' were classi®ed as tor 3 (table 8). ``Southern Big Eye'' (no. 6). The 36% reclassi®cation of ``Southern Big Eye'' was CELLULAR DNA CONTENT well above the a priori expected error rate of 14%, but was still equivocal with ``Mottled''. t-tests indicated that cellular DNA content The similarity between the two sympatric for each pair of sympatric morphotypes was

TABLE 3 Reclassi®cation Matrix for Available Female Morphogroups of the Rana chloronota Complex from a Discriminant Function Analysis (DFA) 12 AMERICAN MUSEUM NOVITATES NO. 3417

Fig. 2. Results of a DFA using morphometric data from females referable to R. chloronota from Vietnam (factor 1 vs. factor 2). Morphotypes are represented by the following symbols: ⅜, no. 1 (Typical); Ⅺ, no. 6 (Southern Big Eye); Ð, no. 3 (Speckled); ⅷ, no. 7 (Large).

Fig. 3. Results of a DFA using morphometric data from females referable to R. chloronota from Vietnam (factor 3 vs. factor 1). Morphotypes are represented by the following symbols: ⅜, no. 1 (Typical); Ⅺ, no. 6 (Southern Big Eye); Ð, no. 3 (Speckled); ⅷ, no. 7 (Large). 2003 BAIN ET AL.: CASCADE FROG FROM SOUTHEAST ASIA 13

TABLE 5 Reclassi®cation Matrix for Male Morphogroups of the Rana chloronota Complex from a DFA

signi®cantly different (table 9). In Sa Pa, 12.10 pg of DNA/cell (Ϯ0.11), and ``South- ``Speckled'' (no. 3) had a mean DNA content ern Big-Eye'' (no. 6) 13.02 pg/cell (Ϯ0.10) level of 14.86 pg (Ϯ1.50), whereas ``Small'' (t ϭ 7.02, p Ͻ 0.0007). The congruence of (no. 4) had 13.26 pg (Ϯ0.42) (t ϭ 8.40, p Ͻ DNA content with the proposed morphoty- 0.0001). In Na Hang, R. chloronota had pes is indicative of their strength as a priori 13.05 pg (Ϯ0.54) of DNA/cell and ``Large'' groups used for electrophoretic studies. (no. 7) 14.43 pg (Ϯ0.06) (t ϭ 3.59, p Ͻ 0.007). In Con Cuong, R. chloronota had PROTEIN ELECTROPHORESIS 12.45 pg (Ϯ0.33) of DNA per cell, whereas sympatric ``Large'' (no. 7) had 13.98 pg Fourteen presumptive enzyme loci were (Ϯ0.14) (t ϭ 13.00, p Ͻ 0.0001). The two satisfactorily resolved on six buffer systems morphotypes from the Central Highlands (table 10). Two of these, AK-A and MDHP- also had distinct levels of cellular DNA from A, were monoallelic among all samples and each other, with ``Mottled'' (no. 5) exhibiting populations. The 12 remaining loci were po-

Fig. 4. Results of a DFA using morphometric data from males referable to R. chloronota from Vietnam (factor 1 vs. factor 2). Morphotypes are represented by the following symbols: ⅜, no. 1 (Typical); Ⅺ, no. 6 (Southern Big Eye); #, no. 5 (Mottled); ᭝, no. 4 (Small). 14 AMERICAN MUSEUM NOVITATES NO. 3417

TABLE 6 TABLE 8 Factor Loadings of Male Morphogroups of the Factor Loadings of Female and Male Rana chloronota Complex from a DFA Morphogroups of the Rana chloronota Refer to text for variable abbreviations. Complex from a DFA Refer to text for variable abbreviations.

lyallelic and they provided evidence of differentiation at the species level for all seven morphotypes (table 10). cause they did not exhibit any unique alleles WITHIN-GROUP GENOTYPES: Three of the that could have been interpreted as being OTUs were sampled from multiple popula- ®xed character differences. The all-male tions: R. chloronota was represented by ®ve ``Large'' series from Na Hang was distin- populations, ``Black Egg'' (no. 2) by four, guishable from the all-female sample at Con and ``Large'' (no. 7) by two. Neither R. chlo- Cuong by one ®xed allele at FUMH-A. ronota nor ``Black Egg'' exhibited ®xed dif- AMONG-GROUP GENOTYPES IN SYMPATRY: ferences among their populations. Sample Fixed allelic differences in eight loci sizes of the ``Black Egg'' populations from (mAAT-A, CK-C, GTDH-A, LDH-B, Na Hang and Tam Dao were too small (n ϭ mMDH-A, sMDH-A, PGM-A, FUMH-A) 1) to be used in the population aggregate diagnosed ®ve of the a priori OTUs (table analysis. They were combined with allopatric 10). ``Small'' (no. 4) had unique ®xed allelic populations of their own morphotypes be- differences at three loci, mAAt-A, mMDH-

TABLE 7 Reclassi®cation Matrix for Female and Male Morphogroups of the Rana chloronota Complex from a Combined DFA 2003 BAIN ET AL.: CASCADE FROG FROM SOUTHEAST ASIA 15

TABLE 9 t-test of Cellular DNA Contents of Sympatric Morphotypes Refer to table 1 for exact locality references.

A, and sMDH-A. ``Southern Big Eye'' (no. All allopatric groups exhibited at least one 6) had unique ®xed alleles at CK-C and ®xed allelic difference from each other (table Pgm-A. ``Speckled'' (no. 3) had one ®xed 10). Although the allozyme data provided unique allele at GTDH-A, as did ``Large'' substantial evidence for the recognition of (no. 7) from Con Cuong at Fumh-A. ``Black new species, the sample sizes were too low Egg'' (no. 2) had a unique ®xed allele at to make inferences about population sub- LDH-B. structure. Many ®xed differences were congruent with hypothesized species boundaries. The PHENETIC ANALYSES two species found in Sa Pa, ``Small'' (no. 4) A phenetic analysis of the allozyme data and ``Speckled'' (no. 3), exhibited ®xed al- summarized overall similarity for 14 loci. lelic differences from each other at 10 loci: Because all loci were not resolved in mAAT-A, CK-C, GPI-A, GTDH-A, mIHD- ``Small'' (no. 4), it was deleted from this A, LDH-A, LDH-B, mMDH-A, sMDH-A, analysis. Similarly, ``Black Egg'' (no. 2) and FUMH-A. ``Southern Big Eye'' and from Ba Be, Tam Dao, and Na Hang were ``Mottled'', both from the Central Highlands, also omitted in order to maximize the num- were also distinct from each other at seven ber of loci. A matrix of the two measures of loci with ®xed differences: CK-C, LDH-B, genetic distance, the unbiased minimum dis- sMDH-A, PEP-A, PGM-A, sSOD-A and tance (Nei, 1978) and Roger's genetic dis- FUMH-A. Huia nasica and R. chloronota tance (table 11), generally re¯ect the num- from Tam Dao were separated by six ®xed bers of ®xed allelic differences among differences: mAAT-A, CK-C, mIDH-A, groups. The largest genetic distance was LDH-A, sSOD-A, and FUMH-A. ``Large'' found between the ``Black Egg'' and ``Mot- (no. 7) and sympatric R. chloronota at Na tled'' morphotypes (Nei's D ϭ 0.595, Rog- Hang exhibited ®xed differences at ®ve loci: er's D ϭ 0.631). Allopatric populations of R. mAAT-A, CK-C, LDH-B, sSOD-A, and chloronota exhibited the smallest genetic dis- FUMH-A. tance. The smallest distance was found be- Three sympatric OTUs occurred in Con tween populations of R. chloronota from Cuong: R. chloronota (no. 1), ``Large'' (no. Tam Dao and Na Hang (Nei's D ϭ 0.000, 7), and ``Black Egg'' (no. 2). All three could Roger's D ϭ 0.003). be distinguished from each other with alter- nate ®xed alleles at FUMH-A. ``Large'' was PHYLOGENETIC ANALYSES OF also discernible with a ®xed difference at MORPHOLOGICAL AND MOLECULAR DATA sSOD-A, while ``Black Egg'' was distinct at three loci: LDH-B, CK-C, and mAATt-A. These morphological and allozyme data AMONG-GROUP GENOTYPES IN ALLOPATRY: were potentially amenable to phylogenetic TABLE 10 Allozyme Frequencies at all Resolved Presumptive Loci of the Rana chloronota Complex Locality data appear below species name/morphogroup number [refer to text] with sample sizes in parentheses. Alleles are indicated with lowercase letters. Missing data are indicated by ``nr''. Locality abbreviations: B, Ba Be: HK, Hong Kong; NH, Na Hang; TD, Tam Dao; CC, Con Cuong; AK, An Khe; S, Sa Pa. Refer to table 1 for locality references. Refer to appendix 2 for enzyme system abbreviations. 2003 BAIN ET AL.: CASCADE FROG FROM SOUTHEAST ASIA 17

analyses. Unfortunately, the morphological data gave a matrix of only plesiomorphies or . autapomorphies (i.e., all phylogenetically un- informative characters). The allozymic data had six potentially informative characters using mutation coding (Murphy, 1993; Murphy and Doyle, 1998). However, the putative syn- apomorphies exhibited no signi®cant character covariation (Faith and Cranston, 1991; Fu and Murphy, 1999). Although all of our data proved to be extremely valuable in di- agnosing species, they did not provide information on the phylogenetic relationships of these species.

TAXONOMIC CONCLUSIONS Complex We recognize each of the seven morphotypes as distinct species, based on concordant evidence of diagnostically ®xed character differences among sympatric samples. Mor- photype 1, ``Typical'', conforms to the di-

Rana chloronota agnosis of R. chloronota, and we assign it to this species. None of the remaining six species from Vietnam can be assigned to currently recognized synonyms of R. chloronota, and below all are described as new species. We include all of these species in a spe-

TABLE 11 cies complex as a matter of taxonomic convenience, while acknowledging the need for additional systematic work.

SPECIES ACCOUNTS Rana chloronota (GuÈnther, 1875) Figures 5, 6A, B, 7A, B, 11A, B, 12A±D, 13A Polypedates chloronotus GuÈnther, 1875 Rana chloronota: Boulenger, 1882 Rana livida: Boulenger, 1887 Genetic Distance Measures for Members of the Rana (Hylorana) livida: Boulenger, 1920 Odorrana livida: Fei, Ye, and Huang, 1990 Rana (Eburana) livida: Dubois, 1992

SYNTYPES STUDIED: BMNH 1947.2.28.4, 1947.2.28.6, and 1947.2.28.12 from Darjee- ling, India, on the hills collected by T.C. Jer- don. BMNH 1947.2.28.4 is an adult female; the other two specimens are adult males. The female has two incisions on its right side, one laterally, one ventrolaterally. The female also has had the shape of its head distorted during preservation. DIAGNOSIS: Rana chloronota, a member of Unbiased minimum genetic distance measures [Nei, 1978] are shown above the diagonal, and Rogers' [1972] genetic distance is shown below the diagonal Refer to text forPa. morphogroup Refer number. to Locality table abbreviations: 1 B, for Ba locality Be; references. HK, Hong Kong; NH, Na Hang; TD, Tam Dao; CC, Con Cuong; AK, An Khe; S, Sa the subgenus Odorrana (sensu Fei et al., 18 AMERICAN MUSEUM NOVITATES NO. 3417

Fig. 5. Syntypes of Rana chloronota: dorsal view of adult male (BMNH 1947.2.28.12) (left); adult female (BMNH 1947.2.28.4) (right). Not to scale.

1990), is characterized by a combination of (0.48); (7) supratympanic fold weak; (8) dor- the following attributes: (1) body dorsoven- sal skin smooth, ¯anks weakly granular, dor- trally compressed; (2) SVL means of males solateral folds absent, venter smooth; (9) dor- 46 mm (41±53 mm), females 92 mm (80± sum green, sometimes with black spots; fore- 100 mm); (3) vomerine teeth in rows oblique limbs and hindlimbs brown, with transverse to choanae; (4) lip-stripe white, extending bars; (10) median callous pad on ®nger III to across upper lip, terminating in glandule proximal tubercle; (11) disks on ®ngers and above insertion of arm; (5) head broad, snout toes greatly enlarged (Ͼ2ϫ base of phalan- rounded in dorsal view, bluntly rounded in ges); (12) feet fully webbed to disks, weak pro®le; (6) tympanum round, distinct, TMP: lateral fringes on I and V to terminal phalan- EYE of males (0.57) greater than females ges, webbing brown; (13) subarticular tuber-

Fig. 6. Hands and feet of (A, B) Rana chloronota, female syntype (BMNH 1947.2.28.4); (C, D) Rana graminea, male syntype (BMNH 1947.2.27.97); (E, F) Rana livida neotype (BMNH 1889.3.25.48). Scale equals 5 mm. 2003 BAIN ET AL.: CASCADE FROG FROM SOUTHEAST ASIA 19

Fig. 7. Heads in lateral and dorsal views: (A, B) Rana chloronota, female syntype (BMNH 1947.2.28.4); (C, D) Rana graminea, male syntype (BMNH 1947.2.27.97); (E, F) Rana livida, neotype female (BMNH 1889.3.25.48). Scale equals 5 mm. cles and inner metatarsal tubercle distinct, snout, and smaller females (SVL 67 mm, conical; (14) terminal phalanges T-shaped; mean of 92 mm in R. chloronota). Rana (15) xiphisternum large, deeply notched pos- hainanensis further differs from R. chloron- teriorly; (16) male with velvety nuptial pads ota in its size (mean female SVL of R. hain- on thumb, paired gular pouches, pectoral anensis 102.6 mm), relative lengths of ®n- spines absent; (17) eggs white. gers (II Ͻ IV Ͻ I Ͻ III for R. hainanensis, COMPARISONS: Rana chloronota super®- II Ͻ I Ͻ IV Ͻ III for R. chloronota), and cially resembles other Asian cascade ranids, absence of gular pouches in males. Rana hei- including: Huia nasica, Rana andersonii, R. jiangensis differs from R. chloronota in ®n- archotaphus, R. chalconota, R. grahami, R. ger size (II Ͻ I Ͻ III Ͻ IV) and smaller dig- graminea, R. hainanensis, R. hejiangensis, R. ital disks. Adult male R. jingdongensis (mean hosii, R. jingdongensis, R. junlianensis, R. SVL of 75 mm) are larger than those of R. kwangwuensis, R. leporipes, R. livida, R. chloronota (46 mm), lack subgular pouches, margaretae, R. schmackeri, R. sinica, and R. and possess pectoral spines (absent in R. tiannensis (table 12). Eggs of H. nasica, R. chloronota). Rana junlianensis has brown lip andersonii, R. chalconota, R. grahami, R. bands (absent in R. chlronota), and its larger junlianensis, R. margaretae and R. schmack- males (SVL 68±80 mm) possess pectoral eri are white with melanic poles, while those spines (absent in R. chlornota) and lack gular of R. chloronota are entirely white. The pouches. Unlike Rana chloronota, R. kwang- smooth green dorsum further distinguishes R. wuensis has small digital disks and males chloronota from H. nasica, R. chalconota, R. lack gular pouches. Rana andersonii has a graminea, R. hosii, R. jingdongensis, R. an- rough, olive-brown dorsum, small disks on dersonii, R. grahami, and R. tiannensis. Huia ®ngers, and males possess pectoral spines nasica differs from R. chloronota with its ol- and lack gular pouches. Rana grahami males ive-brown dorsum (R. chloronota green or are larger (SVL 66±84 mm), have pustules green with brown), longer and more pointed on the dorsum and ¯anks that can form dor- 20 AMERICAN MUSEUM NOVITATES NO. 3417

TABLE 12 Adult Frogs of the Rana chloronota Complex Compared with Other Southeast Asian Cascade Ranids 2003 BAIN ET AL.: CASCADE FROG FROM SOUTHEAST ASIA 21

TABLE 12 (Continued) 22 AMERICAN MUSEUM NOVITATES NO. 3417 solateral folds, and have no digital disks. mottling on the ¯anks (absent in R. chloron- Rana schmackeri has a smooth, heavily spot- ota). ted dorsum, and males possess pectoral DESCRIPTION: Head length greater than spines. Rana tiannensis has a brown dorsum head width (means are 132% for females, that is rough with large, prominent lateral 144% for males), head width 33% of SVL, granulations. Rana chalconota differs from length 47% of SVL; snout short, protruding R. chloronota by having distinct dorsolateral beyond margin of lower jaw, rounded in dor- folds, a pointed snout (vs. rounded) and sal view, bluntly rounded in pro®le; eye smaller SVL (males 32±44 mm, females 46± large, prominent, 70% snout length in fe- 59 mm), an outer metatarsal tubercle, and no males, 84% in males; eyelid broader than in- gular pouches in males. In R. archotaphus, terorbital distance. Top of head ¯at; canthus webbing on toe IV reaches base of distal su- rostralis rounded; loreal region concave; lip barticular tubercle (R. chloronota, webbing ¯ared just anterior to orbit; nostril about to base of disk), females are smaller (SVL three-fourths distance from eye to tip of 59±62 mm), TMP:EYE is more or less equal snout; supratympanic fold weak, reduced to in males and females (sexually dimorphic in a swollen rim of tissue dorsal to tympanum; R. chloronota), and the outer metatarsal tu- tympanum round, distinctly visible, in fe- bercle is present (absent in R. chloronota). males separated from eye by distance equal Rana sinica can be distinguished from R. to TMP, less than length in males; tympanum chloronota in lacking a lip-stripe, and in hav- 48% of EYE in females, 57% in males. Cho- ing smaller mature females (R. sinica holo- anae ovoid; vomerine dentigerous processes type of 66.6 mm), an indistinct tympanum prominent, oblique, posteromedial to choa- covered by a thin layer of skin (distinct, not nae, each bearing numerous teeth. Tongue obscured in R. chloronota), nares halfway cordiform, distinctly notched posteriorly, free between the eye and tip of the snout (nearer for approximately one-third its length. the snout in R. chloronota), and by relative Forearm robust; ®ngers moderately short, ®nger lengths (I Ͻ II Ͻ IV for R. sinica,II slender; hands 29% of SVL, relative lengths Ͻ I Ͻ IV for R. chloronota). Rana leporipes of ®ngers II Ͻ I Ͻ IV Ͻ III; ventromedial can be distinguished from R. chloronota by callous pad on III extends to proximal tuber- a white supratympanic fold (not colored in cle; disks greatly expanded (Ͼ2ϫ base of R. chloronota), a weak dorsolateral fold (ab- phalanges), relative pad size II Ͻ I Ͻ IV III, sent in R. chloronota), absence of transverse pad width (III) 50% pad length; circummar- bands on the arms and legs (present in R. ginal grooves present ventrally; terminal pha- chloronota), webbing that extends only to langes T-shaped; subarticular tubercles con- the distal phalanges (to base of disks in R. ical. Hindlimbs moderately robust; tibia chloronota), and smaller digital disks. Rana length 63% of SVL; FTL 68% of SVL in graminea, R. hosii, and R. livida bear the females, 80% in males; relative toe lengths I most striking resemblance to R. chloronota. Ͻ II Ͻ III Ͻ V Ͻ IV; inner tarsal fold absent; Rana graminea and R. hosii can be distin- feet fully webbed to base of toe disk, lateral guished by their dorsolateral folds. Rana fringes on I and V to terminal phalanges; graminea further differs from R. chloronota toes long, slender, with large, obliquely with its nearly vertical (versus concave) lo- rounded disks, relative pad size I ϭ II ϭ III real region, depressed head, larger tympa- Ͼ IV k V, pad width (IV) 82% of length in num, smaller hands, and smooth sides (gran- females, 90% in males; ventral circummar- ulate in R. chloronota), whereas R. hosii can ginal grooves present; subarticular tubercles be further distinguished by its lack of gular prominent, conical; inner metatarsal tubercle pouches and feeble tarsal folds. Rana livida ovoid, long; outer metatarsal tubercle absent. differs from R. chloronota in having a uni- Xiphisternum large, deeply notched pos- formly colored dorsum (green with dark teriorly. sides in R. chloronota), absence of transverse Skin on dorsum smooth, ¯anks with slight bars on the limbs, smooth skin with pustules granulations; dorsolateral fold absent; small dorsal to the cloaca (smooth dorsum, later- tubercles posteroventral to tympanum, some- ally granular in R. chloronota), and white times absent in males; granules on thighs and 2003 BAIN ET AL.: CASCADE FROG FROM SOUTHEAST ASIA 23 1 SD and Ranges ؎ Given as Means Rana chloronota TABLE 13 See Materials and Methods for abbreviations. Geographic Variation of Body Proportions in 24 AMERICAN MUSEUM NOVITATES NO. 3417 around cloaca; cloacal opening unmodi®ed, directed posteriorly, at upper level of thighs. COLOR IN LIFE (in preservative): Dorsum green, sometimes with brown spots (brown to livid blue), ¯anks gray (light brown) with yellow (white) marbling; lip-stripe white, loreal region brown (dark blackish brown); tympanum beige with dark brown central ring; dorsal limbs and digits brown with dark brown transverse bands; posterior surface of thighs yellow (white) with brown marbling; venter creamy white; ventral side of limbs creamy yellow sometimes with dark mottling; webbing marbled white on dark brown (brown). Iris golden, pupil outlined in a distinct yellow border. SECONDARY SEXUAL CHARACTERS: Gravid females with immaculate, white eggs, 2 mm in diameter (mature). Adult females have SVL approximately twice that of males. Fig. 8. Syntype of Rana graminea (BMNH Males have relatively larger tympanum, dis- 1947.2.27.97), dorsal view. Tag equals 29 mm. tance between tympanum and eye less than that of females, velvety nuptial pads on not ®nd the north-south partition of R. chlo- thumb, and paired gular pouches below jaw ronota described by Bourret (1942). The articulations. Nuptial spines on chest and bel- large Thai form of R. chloronota (Inger and ly absent. Chanard, 1997; Taylor, 1962) is not present GEOGRAPHIC VARIATION: Measurements for in collections from Vietnam, India, Laos, and the syntypes are given in Boulenger (1920) China. Male frogs from Khao Yai National as specimens 1±10 (types of P. chloronotus). Park and Loei Province, Thailand, are large Measurements for other series of R. chloron- (SVL 55±70 mm) and have white ventral spi- ota are given in table 13. Sexual size dimor- nules. None of our male specimens possess phism is most pronounced in specimens from pectoral spines or attain an SVL beyond 55 Tam Dao where mean SVL of males is 40% mm, and all possess the ®nger formula II Ͻ that of females. I. The Thai form is an undescribed species. DISTRIBUTION AND ECOLOGY: Rana chlo- Although male R. chloronota from Hong ronota occurs in forested, montane river sys- Kong have a TMP:EYE (0.76) that is more tems throughout Southeast Asia. These rivers similar to R. graminea (0.70) than it is to vary from shallow and slow moving to tor- Vietnamese populations of R. chloronota rential and deep. Specimens can be found on (0.57), they agree with R. chloronota in ev- boulders and logs, both in and around the ery other aspect, including allozymes, and water. Radiographs revealed that females we consider them as such. Rana chloronota feed on large invertebrates and small verte- occurs in China, northern and southern Vi- brates, including a megophryid frog. Rana etnam, India, and Lao PDR (FMNH chloronota calls from the tops of boulders 256493). This suggests that R. chloronota and overhanging branches in and around tor- occupies a large portion of its currently rec- rents and waterfalls (personal obs.). The au- ognized range of Assam, including Burma, dible part of the call is a very high squeak Thailand, Indochina, and southern China. but the roar of the cascades obscures the remainder of the vocalization, as also observed Rana graminea Boulenger, 1899 by Pope (1931). Figures 6C, D, 7C, D, 8, 11C OTHER SPECIMENS EXAMINED FOR VARIA- TION (see appendix 1, tables 13, 15). SYNTYPES: BMNH 1947.2.27.96, and REMARKS: Like Inger et al. (1999), we can- 1947.2.27.97, adult males from Five Finger 2003 BAIN ET AL.: CASCADE FROG FROM SOUTHEAST ASIA 25

Mountain, Hainan Island, China, collected by dorsum (R. graminea green or green with J. Whitehead. brown spots). Rana hainanensis further dif- DIAGNOSIS: Rana graminea, a member of fers from R. graminea in its relative lengths the subgenus Odorrana (sensu Fei et al., of ®ngers (II Ͻ IV Ͻ I Ͻ III for R. haina- 1990), is characterized by a combination of nensis,IIՅ I Ͻ IV Ͻ III for R. graminea), the following attributes: (1) body dorsoven- and absence of gular pouches in males. Rana trally compressed; (2) SVL means of males hejiangensis further differs from R. graminea 46 mm (42±53 mm), of females 94 mm (78± with its green ¯anks (brownish in R. grami- 100 mm); (3) vomerine teeth in rows oblique nea), relative ®nger length (II Ͻ I Ͻ III Ͻ to choanae; (4) lip-stripe white, extending IV for R. hejiangensis), and small disk size. across upper lip, terminating in glandule Adult males of R. jingdongensis are larger above insertion of arm; (5) head a little lon- than those of R. graminea (R. jingdongensis ger than broad, much depressed, snout blunt- SVL mean is 75 mm, R. graminea is 46 ly rounded in pro®le; (6) tympanum round, mm), lack subgular pouches, and possess distinct, transparent, TMP:EYE of males pectoral spines (absent in R. graminea). 0.77, females 0.56; (7) supratympanic fold Rana junlianensis has brown lip bands (ab- weak; (8) dorsal skin smooth, ¯anks weakly sent in R. graminea), small disks, and larger granular, weak dorsolateral folds present, males (SVL 68±80 mm) that possess pectoral venter smooth; (9) dorsum green, sometimes spines (absent in R. graminea) and lack gular with black spots; forelimbs and hindlimbs pouches. Rana kwangwuensis, unlike Rana brown, with transverse bars; (10) median cal- graminea, has small disks, and males lack lous pad on ®nger III to proximal tubercle; gular pouches. Rana andersonii has a rough, (11) disks on ®ngers and toes enlarged (Ͼ2ϫ olive-brown dorsum, small disks on ®ngers, base of phalanges); (12) feet fully webbed to and males possess pectoral spines and lack disks, but as a fringe from distal subarticular gular pouches. Male R. grahami are larger tubercle of IV in some specimens, weak lat- (SVL 66±84 mm), have pustules on the dor- eral fringes on I and V to terminal phalanges; sum, and have no digital disks. Rana (13) subarticular tubercles and inner metatar- schmackeri has a smooth, heavily spotted sal tubercle distinct; (14) terminal phalanges dorsum, and males possess pectoral spines. T-shaped; (16) male with strong forearms, Rana tiannensis has a brown dorsum that is slight hands, velvety nuptial pads, paired gu- rough with large, prominent lateral granula- lar pouches, pectoral spines absent; (17) fe- tions. Rana chalconota has a pointed snout males bear white eggs. (vs. rounded), smaller males (32±44 mm) COMPARISONS: Rana graminea super®cial- that lack gular pouches, and an outer meta- ly resembles other Asian cascade ranids, in- tarsal tubercle. In R. archotaphus, the web- cluding: Huia nasica, Rana andersonii, R. bing (toe IV) reaches the base of distal su- archotaphus, R. chalconota, R. chloronota, barticular tubercle (to base of disk in R. gra- R. grahami, R. hainanensis, R. hejiangensis, minea), and the outer metatarsal tubercle is R. hosii, R. jingdongensis, R. junlianensis, R. present (absent in R. graminea). Rana sinica kwangwuensis, R. leporipes, R. livida, R. lacks a lip-stripe (present in R. graminea), margaretae, R. schmackeri, R. sinica, and R. has an indistinct, skin-covered tympanum tiannensis (table 12). The presence of dor- (distinct and uncovered in R. graminea), na- solateral folds distinguishes R. graminea res halfway between the eye and tip of the from R. andersonii, R. chloronota, R. hain- snout (nearer the snout in R. graminea), and anensis, R. hejiangensis, R. jingdongensis, R. a different ®nger formula (I Ͻ II Ͻ IV for junlianensis, R. kwangwuensis, R. livida, R. R. sinica). Rana leporipes differs from R. margaretae, R. schmackeri, R. sinica and R. graminea in having a white supratympanic tiannensis. The presence of white eggs dif- fold, absence of transverse bands on the arms ferentiates R. graminea from H. nasica, R. and legs (present in R. graminea), and web- andersonii, R. chalconota, R. grahami, R. bing that only extends to the distal phalanges margaretae and R. schmackeri (white eggs (to base of disks in R. graminea). Rana chlo- with melanic poles). Huia nasica further dif- ronota, R. hosii, and R. livida bear the most fers from R. graminea with its olive-brown striking resemblance to R. graminea. Rana 26 AMERICAN MUSEUM NOVITATES NO. 3417 chloronota and R. livida can be distinguished present; subarticular tubercles moderately from R. graminea by the absence of dorso- prominent, rounded; inner metatarsal tuber- lateral folds. Rana graminea further differs cle ovoid, long; outer metatarsal tubercle ab- from R. chloronota with its nearly vertical sent. (versus concave) loreal region, depressed Skin on dorsum smooth, ¯anks with slight head, larger tympanum, transparent tympa- granulations; weak dorsolateral fold present; num, and smooth sides (granulate in R. chlo- white glandular tubercles posteroventral to ronota). Male R. graminea also possess tympanum; cloacal opening unmodi®ed, di- smaller, more slender hands than R. chloron- rected posteriorly, at upper level of thighs. ota. Rana hosii can be further distinguished COLOR IN LIFE (in preservative): Dorsum from R. graminea by an obtusely pointed green (brown to livid blue), ¯anks brown snout and the absence of gular pouches. (light brown) with yellow (white) marbling; Rana livida differs from R. graminea in hav- lip-stripe white, loreal region dark brown ing a uniformly colored dorsum (green with above (dark blackish brown), light brown be- dark sides in R. graminea), white mottling low; tympanum transparent, perimeter white on the ¯anks (absent in R. graminea), and (light brown); dorsal limbs and digits brown absence of transverse bars on the limbs. with dark brown transverse bands; posterior DESCRIPTION OF SYNTYPES: Head length surface of thighs yellow (white) with brown greater than head width (112%), head width marbling; venter creamy white; ventral side 35% SVL, length 39% of SVL; snout short, of limbs creamy yellow, sometimes with protruding beyond margin of lower jaw, dark mottling; webbing marbled white on bluntly rounded in dorsal view, much de- dark brown (brown). Iris golden, margin of pressed, bluntly rounded in pro®le; eye large, pupil outlined in a distinct yellow border. prominent, 117% of snout length; eyelid VARIATION IN OTHER SPECIMENS EXAMINED broader than interorbital distance. Top of (Fujian Province: AMNH A-28543±28545; head ¯at; canthus rostralis prominent; loreal A-28612; A-29973±29978; A-29980±29991) region feebly oblique or nearly vertical, (table 14): Faint dorsolateral folds are present weakly concave; nostril about three-fourths in some specimens of this series, although it distance from eye to tip of snout; supratym- is not clear whether their absence is an arti- panic fold weak; tympanum round, distinctly fact of preservation. Some specimens have visible, transparent, 75% of EYE. Choanae ®nger II Ͻ I, whereas others have ®nger I ϭ ovoid; vomerine dentigerous processes II. Male specimens from Fujian Province av- prominent, oblique, posteromedial to choa- erage 46.4 mm SVL (41.5±53.5), and fe- nae, each bearing numerous teeth. Tongue males average 94 mm (78.3±100.2) (table cordiform, distinctly notched posteriorly, free 14). for approximately one-third its length. SECONDARY SEXUAL CHARACTERS: Gravid Forearm robust; ®ngers very slender, rel- females bear immaculate white eggs. Adult ative lengths of ®ngers II Յ I Ͻ IV Ͻ III; females have SVL approximately twice that ventromedial callous pad on III to proximal of males and relatively larger digital disks tubercle; disks moderately expanded (Ͼ2ϫ than males. Males have strong arms, rela- base of phalanges), relative pad size II Ͻ I tively larger tympanum, smaller distance be- Ͻ IV Ͻ III; circummarginal grooves present tween tympanum and eye, velvety nuptial ventrally; terminal phalanges T-shaped; su- pads on thumb, and paired gular pouches be- barticular tubercles rounded. Hindlimbs long, low jaw articulations. Males lack nuptial slender; tibia length 65% of SVL; FTL 58% spines on chest and belly. of SVL; relative toe lengths I Ͻ II Ͻ III Ͻ DISTRIBUTION AND ECOLOGY: The species is V Ͻ IV; inner tarsal fold absent; feet fully known from Hainan Island (syntypes), the webbed to base of toe disk, but as a fringe Mao Son Mountains in Tonkin (northern Vi- from distal subarticular tubercle of IV, lateral etnam) (Boulenger, 1920), and Fujian Prov- fringes on I and V to terminal phalanges; ince, China (Pope, 1931). toes long, slender, with large, obliquely REMARKS: Boulenger (1920) gave mea- rounded disks, relative pad size I ϭ II ϭ III surements of the syntypes, as well as the Ͼ IV k V; ventral circummarginal grooves Mao Son specimen. Bourret (1942) consid- 2003 BAIN ET AL.: CASCADE FROG FROM SOUTHEAST ASIA 27

TABLE 14 Geographic Variation in Rana graminea from Fujian Province, China, Given as Means ؎ 1 SD and Ranges See Materials and Methods for abbreviations.

ered R. graminea to be a ``northern variety'' cluding Huia nasica, Rana andersonii, R. ar- of Indochinese R. chloronota because of its chotaphus, R. chalconota, R. chloronota, R. weak dorsolateral fold. Like Inger et al. grahami, R. graminea, R. hainanensis, R. he- (1999), we did not ®nd R. graminea in Vi- jiangensis, R. hosii, R. jingdongensis, R. jun- etnam. Our study of specimens treated as R. lianensis, R. kwangwuensis, R. livida, R. graminea by Pope (1935) agrees with the margaretae, R. schmackeri, R. sinica and R. original description. We did not study the tiannensis (table 12). Rana leporipes is Mao Son specimen. unique among all of the above species in having a white supratympanic fold, webbing Rana leporipes Werner, 1930 that only reaches the distal phalanx (not the disk), and oblong, somewhat rounded distal HOLOTYPE: Two ®gures (Werner, 1930: 49, pl. IV) of Mell no. 15660, an adult female phalanges (T-shaped in others, unknown for from Lung Tao Shan, North Kwangtung, R. hejiangensis, R. junlianensis, and R. China (700 m a.s.l.) collected on 4 July 1919 kwangwuensis). The absence of banding on by R. Mell. The voucher specimens of the the legs distinguishes R. leporipes from all type series have been lost and the photo- species listed here, except R. livida (present graphs of the holotype become the iconoty- or absent in R. chalconota and R. hosii). The pe. presence of a dorsolateral fold also distin- DIAGNOSIS (from text and plates of original guishes it from R. andersonii, R. chloronota, description): Rana leporipes is characterized R. hainanensis, R. hejiangensis, R. jingdon- by a combination of the following attributes: gensis, R. junlianensis, R. kwangwuensis, R. (1) body dorsoventrally compressed; (2) livida, R. margaretae, R. schmackeri, R. sin- SVL reported for females between 52 and ica, and R. tiannensis. Whereas Huia nasica 102 mm; (3) vomerine teeth in rows oblique has an olive-brown dorsum, R. leporipes is to choanae; (4) lip-stripe white; (6) tympa- dark green. Rana hainanensis further differs num very distinct, TMP:EYE is 0.5; (7) su- from R. leporipes in its size (SVL R. hain- pratympanic fold milky white; (8) dorsal skin anensis 103 mm) and relative lengths of ®n- smooth, ¯anks weakly granular, dorsolateral gers (II Ͻ IV Ͻ I for R. hainanensis,Iϭ II folds slightly distinguishable, venter smooth; for R. leporipes). Rana junlianensis has (9) dorsum dark green, ¯anks stony gray brown lip bands. Rana andersonii has a with white marbling, legs not banded; (11) rough, olive-brown dorsum. Rana grahami disks on ®ngers and toes only slightly en- has pustules on the dorsum and ¯anks and larged (Ͻ2ϫ base of phalanges); (12) feet no digital disks. Rana schmackeri has a fully webbed to base of distal phalanges; (13) smooth, heavily spotted dorsum. Rana tian- subarticular tubercles and inner metatarsal nensis has a rough, brown dorsum with large, tubercle large, projecting; (14) terminal pha- prominent lateral granulations. Both Rana langes oblong, somewhat pointed. chalconota and R. archotaphus possess an COMPARISONS: Rana leporipes super®cially outer metatarsal tubercle, but R. leporipes resembles other Asian cascade ranids, in- does not. Rana hosii can be distinguished 28 AMERICAN MUSEUM NOVITATES NO. 3417

Fig. 9. Dorsal view of Rana livida: adult female (BMNH 1889.3.25.47) (left); neotype, adult female (BMNH 1889.3.25.48) (right). Scale equals 5 mm.

from R. leporipes by its feeble tarsal folds. not be con®rmed, as the type series has been Rana chloronota differs from R. leporipes in lost. that it is larger (R. chloronota female SVL 80±100 mm) and it has a different digital for- Rana livida (Blyth, 1856) mula (II Ͻ I Ͻ IV Ͻ III for R. chloronota). Figures 6E, F, 7E, F, 9 Dorsum of R. livida is uniform (¯anks a different color for R. leporipes). Rana livida (Boulenger, 1887) REMARKS: The original description of R. NEOTYPE: BMNH 1889.3.25.48 an adult leporipes reported extensive variation in female from Thagata Juwa, Village on the snout±vent length (52±102 mm) and lacked hills southwest of Mt. Mooleyit, Dawna any description of secondary sex characters. Mountains, Myanmar (400±500 m), by M.L. The single male specimen of R. leporipes is Fea in early 1887. signi®cantly larger than males of R. chloron- DIAGNOSIS: Rana livida, a member of the ota (93 mm vs. a maximum of 53 mm). subgenus Odorrana (sensu Fei et al. 1990), However, it is unclear how the male was is characterized by a combination of the fol- sexed, as nuptial excrescences and gular lowing attributes: (1) body dorsoventrally pouches are not mentioned. Bourret (who did compressed; (2) SVL females 89.4, 97.1 mm; not indicate whether he had seen the type (3) vomerine teeth in rows oblique to choa- series) thought the male was an incorrectly nae; (4) lip-stripe white, extending across up- identi®ed female and that Werner's (1930) per lip, terminating in glandule above inser- frogs were all females spanning a range of tion of arm; (5) head broad, snout rounded varying degrees of maturity. We are also in dorsal view, bluntly rounded in pro®le; (6) skeptical about any males, although this can- tympanum round, distinct, TMP:EYE 0.41; 2003 BAIN ET AL.: CASCADE FROG FROM SOUTHEAST ASIA 29

(7) supratympanic fold weak; (8) dorsal skin chotaphus, webbing (toe IV) reaches base of smooth to ¯anks, pustules dorsal to cloaca, distal subarticular tubercle (R. livida, web- dorsolateral folds absent, venter smooth; (9) bing to base of disk), females are smaller dorsum uniform brown in preservative; fore- (SVL 59±62 mm), and it has an outer meta- limbs and hindlimbs lighter brown, without tarsal tubercle. Rana sinica differs from R. transverse bars; (10) median callous pad on livida in lacking a lip-stripe, and in having ®nger III to proximal tubercle; (11) disks on smaller mature females (R. sinica holotype ®ngers and toes enlarged (Ͼ2ϫ base of pha- 66.6 mm), an indistinct, skin-covered tym- langes); (12) feet fully webbed to disks, panum (distinct and uncovered in R. livida), weak lateral fringes on I and V to terminal nares halfway between the eye and tip of the phalanges; (13) subarticular tubercles and in- snout (nearer the snout in R. livida), and a ner metatarsal tubercle distinct, conical; (14) different ®nger formula (I Ͻ II Ͻ IV for R. terminal phalanges T-shaped; (15) xiphister- sinica,IIϽ I Ͻ IV for R. livida). Rana hosii num large, deeply notched posteriorly; (17) differs from R. livida by its obtusely pointed eggs white. head (rounded in R. livida), dorsolateral COMPARISONS: Rana livida super®cially re- folds, occasional bands on arms and legs, and sembles other Asian cascade ranids, includ- feeble tarsal folds (dorsolateral and tarsal ing Huia nasica, Rana andersonii, R. archo- folds absent in R. livida). Rana chloronota taphus, R. chalconota, R. chloronota, R. gra- differs from R. livida with its solid-colored hami, R. graminea, R. hainanensis, R. he- dorsum and dark sides (uniform in R. livida), jiangensis, R. hosii, R. jingdongensis, R. transverse bars on the limbs (absent in R. liv- junlianensis, R. kwangwuensis, R. leporipes, ida), smooth dorsum, ¯anks granular (slight- R. margaretae, R. schmackeri, R. sinica, and ly granulose on the posterior thighs in R. liv- R. tiannensis (table 12). The smooth, uni- ida), and absence of white mottling on the formly colored dorsum further distinguishes ¯anks (present in R. livida). Rana leporipes R. livida from all of the above species. The differs from R. livida in having smaller dig- unpigmented eggs of R. livida distinguish it ital disks, a white supratympanic fold, weak from H. nasica, Rana andersonii, R. chal- dorsolateral fold, and webbing that only conota, R. grahami, R. junlianensis, R. mar- reaches the distal phalanges (R. livida to base garetae, and R. schmackeri, which are white of the disk). Rana graminea further differs with melanic poles. Huia nasica differs from from R. livida with its nearly vertical (versus R. livida in its longer, more pointed head, and concave) loreal region and depressed head. smaller females (SVL 67 mm). Rana haina- DESCRIPTION: Head length greater than nensis further differs from R. livida in rela- head width (103%), head width 35% of SVL, tive lengths of ®ngers (II Ͻ IV Ͻ I Ͻ III for length 37% of SVL; snout short, protruding R. hainanensis,IIϽ I Ͻ IV Ͻ III for R. beyond margin of lower jaw, rounded in dor- livida). Rana hejiangensis differs from R. liv- sal view, bluntly rounded in pro®le; eye ida in its relative ®nger size (II Ͻ I Ͻ III Ͻ large, prominent, 72% snout length; eyelid IV) and smaller digital disks. Rana ander- broader than interorbital distance. Top of sonii has a rough, olive-brown dorsum and head ¯at; canthus rostralis rounded; loreal re- small disks on ®ngers. Rana grahami has gion concave; lip ¯ared just anterior to orbit; pustules on the dorsum and ¯anks (vs. nostril about three-fourths distance from eye smooth in R. livida) and slightly swollen dig- to tip of snout; supratympanic fold weak, ital tips. Rana junlianensis has brown lip swollen rim of tissue dorsal to tympanum; bands. Rana kwangwuensis has small digital tympanum round, distinctly visible, 44% of disks. Rana schmackeri has a smooth, heavi- EYE. Choanae ovoid; vomerine dentigerous ly spotted dorsum. Rana tiannensis has a processes prominent, oblique, posteromedial rough brown dorsum with large, prominent to choanae, each bearing numerous teeth. lateral granulations. Rana chalconota differs Tongue cordiform, distinctly notched poste- from R. livida by having coarsely shagreened riorly, free for approximately one-third its dorsum, distinct dorsolateral folds, pointed length. snout, smaller SVL (females 46±59 mm), Forearm robust; ®ngers moderately short, and an outer metatarsal tubercle. In R. ar- slender; hands 29% of SVL, relative lengths 30 AMERICAN MUSEUM NOVITATES NO. 3417 of ®ngers II Ͻ I Ͻ IV Ͻ III; ventromedial est villages to Mulayit Taung are Kyeik-ywa, callous pad on III to proximal tubercle; disks Daukkat-ywa, and Kyeik-don, to the west, greatly expanded (Ͼ2ϫ base of phalanges), and Mawkhi and Huthi, near the Thai border relative pad size II Ͻ I Ͻ IV Ͻ III; circum- to the east. marginal grooves present ventrally; terminal REMARKS: Male R. chloronota are substan- phalanges T-shaped; subarticular tubercles tially smaller than females and they posses prominent, rounded. Hindlimbs moderately paired subgular pouches and a larger tym- robust; tibia length 68% of SVL; FTL 85% panum. Presumably, male R. livida have the of SVL; relative toe lengths I Ͻ II Ͻ III Ͻ same attributes. Blyth (1852) appears to have V Ͻ IV; inner tarsal fold absent; feet fully included males in his original description; webbed to base of toe disk, lateral fringes on many of the specimens that Theobold as- I and V to terminal phalanges; toes long, sumed to be juveniles have whiter lip-stripes slender, with large, obliquely rounded disks, and a relatively large tympanum that is closer relative pad size I ϭ II ϭ III Ͼ IV Ͼ V; to the eye than in females. Although the col- ventral circummarginal grooves present; su- or is not recorded in life, Blyth (1856) re- barticular tubercles prominent, conical; inner ported that R. livida is ``uniform dusky- metatarsal tubercle ovoid, long; outer meta- plumbeous above, probably dull olive green tarsal tubercle absent. when alive.'' Boulenger (1920) recorded Xiphisternum large, deeply notched pos- measurements of two R. livida females from teriorly. Thagata, Tenasserim. It is not clear whether Skin on dorsum smooth to ¯anks; dorso- these two specimens are the same as those lateral fold absent; small tubercles posterov- described above. entral to tympanum; granules dorsal cloaca; cloacal opening unmodi®ed, directed poste- Rana sinica Ahl, 1925 riorly, at upper level of thighs. Figure 10 COLOR IN PRESERVATIVE: Dorsum uniform dark brown to ¯anks, some white spots and HOLOTYPE: ZMB 9785 an adult female mottling on ¯ank; prominent lip-stripe white; from China. tympanum brown; posterior surface of thighs DIAGNOSIS: Rana sinica is characterized by light brown with white spots, no transverse a combination of the following attributes: (1) banding; venter creamy white; ventral side of body dorsoventrally compressed; (2) SVL of limbs creamy white; webbing dark gray. single mature female is 66.6 mm, males un- SECONDARY SEX CHARACTERS: BMNH known; (3) vomerine teeth in rows oblique 1889.3.25.47, an adult female bears white to choanae; (4) lip-stripe absent; (5) head not eggs. No male specimens were studied. broad, snout rounded in dorsal view; (6) tym- MEASUREMENTS OF NEOTYPE (in mm): SVL panum round, indistinct, covered by a layer 89.4; SNT 14.2; HDL 32.8; HDW 31.6; EYE of skin, TMP:EYE 0.52; (7) supratympanic 10.2; IOD 8.0; TMP 4.5; TEY 2.8; HND fold weak; (8) dorsal skin smooth ¯anks 25.5; FPL 3.2; TIB 60.8; FTL 76.2; TPL 2.6. weakly granular, dorsolateral folds absent, MEASUREMENTS OF REFFERED SPECIMEN venter smooth; (9) dorsum bronze-green with (BMNH 1889.3.25.47): SVL 97.1; SNT irregular indistinct spots and blue-gray 16.7; HDL 36.1; HDW 33.4; EYE 11.9; IOD ¯anks, legs with black bands; cloacal region 9.2; TMP 4.6; TEY 4.1; HND 28.8; FPL 3.3; marbled black with white; (10) median cal- TIB 63.8; FTL 75.6; TPL 2.7. lous pad on ®nger III almost to proximal tu- DISTRIBUTION AND ECOLOGY: The range of bercle; (11) disks on ®ngers and toes slightly R. livida is unclear. The type material is lost enlarged (Ͻ2ϫ base of phalanges); (12) feet and the locality information has been poorly fully webbed on II, III, IV; I and V without recorded. Much confusion exists in its tax- lateral web fringes, webbing light gray in al- onomy. The only con®rmed locality is that cohol; (13) subarticular tubercles and inner of the neotype and maps do not show Tha- metatarsal tubercle distinct, conical; (14) ter- gata Juwa village, as indicated in by Feae. minal phalanges are slightly rounded; (15) Mount Mooleyit is currently found on maps xiphisternum not large, shallow notch pos- as Mulayit Taung (16ЊN, 98Њ30ЈE). The clos- teriorly; (16) male secondary sexual charac- 2003 BAIN ET AL.: CASCADE FROG FROM SOUTHEAST ASIA 31

Fig. 10. Holotype of Rana sinica (ZMB 9785), dorsal view (left), ventral view (right). Scale equals 5 mm.

teristics unknown; (17) eggs yellow (in al- brown dorsum (R. sinica green). Rana hain- cohol). anensis further differs from R. sinica in its COMPARISONS: Rana sinica super®cially re- size (mean female SVL R. hainanensis 103 sembles other Asian cascade ranids, includ- mm) and relative lengths of ®ngers (II Ͻ IV ing Huia nasica, Rana andersonii, R. archo- Ͻ I for R. hainanensis,IϽ II Ͻ IV for R. taphus, R. chalconota, R. chloronota, R. gra- sinica). Rana andersonii has a rough, olive- hami, R. graminea, R. hainanensis, R. he- brown dorsum. Rana hejiangensis has small- jiangensis, R. hosii, R. jingdongensis, R. er males (SVL 47 mm) and a different ®nger junlianensis, R. kwangwuensis, R. leporipes, formula (II Ͻ I Ͻ III) than R. sinica. Rana R. livida, R. margaretae, R. schmackeri and julianensis and R. kwangwuensis differ from R. tiannensis (table 12). Unlike the species R. sinica in their relative ®nger formula (II listed above, R. sinica has a layer of skin Ͻ I Ͻ IV). Rana jingdongensis possesses overlying its tympanum and its terminal pha- vertical lip bands, large toe disks, and is fully langes are slightly rounded (not T-shaped; R. webbed to all disks. Rana graminea differs leporipes has oblong, somewhat rounded dis- from R. sinica in that its nares is closer to tal phalanges). The absence of a lip-stripe in the tip of the snout than to the eye (halfway R. sinica further differentiates it from H. na- in R. sinica), and it has a different ®nger for- sica, R. archotaphus, R. chalconota, R. chlo- mula (II ϭ I Ͻ IV for R. graminea). Rana ronota, R. grahami, R. graminea, R. hejian- margaretae further differs from R. sinica in gensis, R. hosii, R. junlianensis, R. kwang- having larger females (SVL 78±88 mm). wuensis, R. leporipes, and R. livida. Eggs of Rana grahami has pustules on the dorsum H. nasica, Rana andersonii, R. chalconota, and ¯anks and no digital disks (only slightly R. grahami, R. junlianensis, R. margaretae, swollen tips). Rana schmackeri has a and R. schmackeri are white with melanic smooth, heavily spotted dorsum. Rana tian- poles, whereas those of R. sinica are immac- nensis has a rough, brown dorsum with large, ulate. The smooth green dorsum further dis- prominent lateral granulations. Rana chal- tinguishes R. sinica from H. nasica, R. an- conota differs from R. sinica by having dis- dersonii, R. jingdongensis, R. grahami, R. tinct dorsolateral folds, a pointed snout (vs. graminea, R. leporipes, R. schmackeri, and rounded), and an outer metatarsal tubercle. In R. tiannensis. Huia nasica has an olive- R. archotaphus the outer metatarsal tubercle 32 AMERICAN MUSEUM NOVITATES NO. 3417 is present. Rana hosii differs from R. sinica Xiphisternum small, with a shallow notch by its dorsolateral folds and feeble tarsal posteriorly. folds (absent in R. sinica). Rana chloronota Skin on dorsum smooth, ¯anks with slight differs from R. sinica in being larger (SVL granulations; dorsolateral fold absent; small for adult females 80±100 mm for R. chlo- tubercles posteroventral to tympanum; gran- ronota), having nostrils closer to the tip of ules on thighs and around cloaca; cloacal the snout than the eye (halfway for R. sini- opening unmodi®ed, directed posteriorly, at ca), and a different digital formula (II Ͻ I Ͻ upper level of thighs. IV Ͻ III for R. chloronota). Rana leporipes COLOR IN LIFE (in preservative): Dorsum also differs from R. sinica by having a white green (bronze-green to brown), ¯anks gray supratympanic fold (not colored in R. sinica). (blue-gray to light brown); loreal region Rana livida has a solid-colored dorsum with black (dark blackish brown), lip-stripe ab- white spots on its sides (¯anks a different sent; dorsal limbs and digits brown with dark color for R. sinica, lacking spots), and it brown transverse bands; posterior surface of lacks transverse bands on the arms and legs thighs black with white marbling; venter (present in R. sinica). creamy white; ventral side of limbs creamy DESCRIPTION OF HOLOTYPE: HDL greater yellow; webbing marbled white on dark than HDW (132%), HDW 26% of SVL, brown (brown). HDL 34% of SVL; snout short, rounded in SECONDARY SEXUAL CHARACTERS: The fe- dorsal view; protruding beyond margin of male holotype has large, immaculate yellow lower jaw, rounded in lateral view, EYE eggs (possibly white in life). prominent, smaller than SNT, IOD broader MEASUREMENTS OF HOLOTYPE (in mm): than EYE and upper eyelid. Top of head ¯at; SVL 66.6; HDL 22.5; HDW 17.6; EYE 8.2; canthus rostralis rounded; loreal region IOD 13.4; TMP 4.2; TIB 39.4; femur 33.7; steeply concave; nostril about one-half the FTL 33.7. distance from eye to tip of snout; supratym- REMARKS: The holotype is the only known specimen of R. sinica; all others are lost. This panic fold weak; tympanum round, indis- redescription is based on the original descrip- tinct, covered by layer of skin; TMP 52% of tion by Ahl (1925) with amendments and ad- EYE in females. Choanae ovoid; vomerine ditions based on our examination of the ho- dentigerous processes prominent, oblique, lotype. Ahl described the holotype as a male, posteromedial to choanae, each bearing nu- but it is clearly a gravid female. As well, Ahl merous teeth. Tongue cordiform, distinctly described a frog with large ®nger and toe notched posteriorly, free for part of its pos- pads. Although the condition of the specimen terior length. did not allow for accurate pad measurements Forearm robust; relative lengths of ®ngers to be made, the enlarged pads on the holo- I Ͻ II Ͻ IV, III Ͻ snout; ventromedial callous type are not as large as those on other species pad on III almost to proximal tubercle; disks in the Rana chloronota complex. This spe- slightly expanded (Ͻ2ϫ base of phalanges); cies differs signi®cantly from the other mem- circummarginal grooves present ventrally; bers of the Rana chloronota complex in pre- terminal phalanges rounded; subarticular tu- servative, but presumably has super®cial re- bercles conical. Hindlimbs moderately ro- semblances in life. Ahl (1925) believed that bust; extend 15 mm beyond snout when ad- R. sinica was closely related to Staurois na- pressed; TIB 59% of SVL; FTL 30% of tator. SVL; toe II is longer than all others, III ϭ IV; inner tarsal fold absent; feet fully webbed Rana bacboensis, new species to base of toe disk on II, III, and IV, I and V without external fringes; toes long, slender, (Previously referred to as species 2, ``Black with slight, rounded disks; ventral circum- Egg'') marginal grooves present; subarticular tuber- Figures 11D, 12E, F, 13B cles prominent, conical; inner metatarsal tu- HOLOTYPE: (ROM ®eld no. 13171) ROM bercle ovoid, long; outer metatarsal tubercle 29534 an adult female from the Khe Moi Riv- absent. er, approximately 24 km west of Con Cuong 2003 BAIN ET AL.: CASCADE FROG FROM SOUTHEAST ASIA 33

Fig. 11. Members of the Rana chloronota complex. (A) Male and female and (B) male R. chloronota, Nghe An Province, Vietnam. C. R. graminea, male, Hainan Island, China, (NLO ®eld series 26375). D. R. bacboensis, new species, holotype, a female from Nghe An Province, Vietnam. (E) Female holotype and male paratype in amplexus, and (F) juvenile paratype of R. daorum, new species, Lao Cai Province, Vietnam. Fig. 12. Heads of members of the Rana chloronota complex in lateral and dorsal views: (A, B) Rana chloronota, male (ROM 26402), and (C, D) female (ROM 26406); paratypes of (E, F) Rana bacboensis, new species (ROM 29529); (G, H) Rana daorum, new species (ROM 38538); (I, J) Rana hmongorum, new species (ROM 39891); (K, L) Rana morafkai, new species (ROM 39906); (M, N) Rana banaorum, new species (ROM39941); and (O, P) Rana megatympanum, new species (ROM 26398). Fig. 13. Hands and feet of members of the Rana chloronota complex: (A) Rana chloronota (ROM 26402); paratypes of (B) Rana bacboensis, new species (ROM 29529); (C) Rana daorum, new species (ROM 38538); (D) Rana hmongorum, new species (ROM 26379); (E) Rana morafkai, new species (ROM 39906); (F) Rana banaorum, new species (ROM 39941); and (G) Rana megatympanum, new species (ROM 26398). Scale equals 5 mm. 36 AMERICAN MUSEUM NOVITATES NO. 3417 village, Con Cuong District, Nghe An Prov- blotches; forelimbs and hindlimbs with trans- ince, Vietnam (18Њ56Ј30ЉN, 104Њ48Ј35ЉE) col- verse bands of distinct blotches to tips of dig- lected 24 October 1994 by I.S. Darevsky, its; (10) median callous pad on base of ®n- L.A. Lowcock, R.W. Murphy, and N.L. Orlov. gers II and III to proximal tubercle; (11) The holotype had leg and liver tissue removed disks on ®ngers and toes enlarged (Ͻ2ϫ base shortly after it was euthanised. of phalanges); (12) feet fully webbed to PARATYPES: ROM 29531±29533, all fe- disks, but as a fringe from distal subarticular males, collected with holotype on 24 October tubercle of IV, slight lateral fringes on toes I 1994 by I.S. Darevsky, L.A. Lowcock, R.W. and V to terminal phalanges, webbing mar- Murphy, and N.L. Orlov; ROM 26404, a fe- bled brown on white; (13) subarticular tu- male collected at the type locality by A. La- bercles distinct, conical; inner metatarsal tu- throp, R.W. Murphy, and N. Orlov on 5 June bercle distinct, ovoid; (14) terminal phalan- 1995; ROM 26357±26358, adult females col- ges T-shaped; (15) xiphisternum large, deep- lected from the type locality on 5 June 1995 ly notched posteriorly; (16) male with by A. Lathrop, R.W. Murphy, and N.L. Or- velvety nuptial pad on thumb, paired gular lov; ROM 29359, a female, from Ba Be pouches, no pectoral spines; (17) eggs black. Lake, Ba Be Lake National Park, Bac Kan COMPARISONS: Rana bacboensis is one of Province (formerly Cao Bang Province), Vi- the larger species of cascade ranids (SVL fe- etnam (22Њ25Ј05ЉN, 105Њ38Ј05ЉE), collected male 81±105 mm). It can be distinguished by R.H. Bain on 24 May 1995 at the out¯ow from all other cascade ranids of Southeast on the south side of Ba Be Lake; ROM Asia by its dark, pigmented eggs (immacu- 29526±29530, all females, from Na Hang late white or white with melanic pole in other Nature Reserve, Tuyen Quang Province, Vi- species) (table 12). Its black vertical lip-bars etnam (22Њ21Ј54ЉN, 105Њ25Ј40ЉE) approxi- distinguish it from R. archotaphus, R. chal- mately 15 km from Pac Ban village collected conota, R. chloronota, R. grahami, R. gra- by R.W. Murphy and A. Lathrop between 25 minea, R. hejiangensis, R. hosii, R. jingdon- and 30 May 1996; AMNH A-161248, a fe- gensis, R. kwangwuensis, R. leporipes, R. liv- male, from Hieng Stream, Chau Khe Com- ida, R. schmackeri, R. sinica, and Huia na- mune, Con Cuong District, Nghe An Prov- sica. The brown dorsum with dark spots ince, Vietnam (19Њ02Ј17ЉN, 104Њ42Ј06ЉE, el- distinguishes it from R. archotaphus, R. evation 300 m) collected on 29 April 1998 chalconota, R. chloronota, R. grahami, R. by N.Q. Truong; FMNH 255611 (adult fe- graminea, R. hejiangensis, R. hosii, R. jing- male), 255612 (adult male) along the Khe dongensis, R. junlianensis, R. kwangwuensis, Chat Stream, Pu Mat Nature Reserve, Con R. leporipes, R. margaretae, R. schmackeri, Cuong District, Nghe An Province, Vietnam and R. sinica, all of which have a predomi- (18Њ56ЈN, 104Њ45ЈE, elevation 300 m) on 8 nantly green dorsum. The presence of gular September 1998 by Bryan L. Stuart. pouches in males differentiates R. bacboensis DIAGNOSIS: Rana bacboensis, a member of from R. andersonii, R. chalconota, R. gra- the subgenus Odorrana (sensu Fei et al., hami, R. hainanensis, R. hosii, R. jingdon- 1990), is characterized by a combination of gensis, R. junlianensis, R. kwangwuensis, the following attributes: (1) body dorsoven- and R. margaretae. Huia nasica has a trally compressed; (2) SVL of males 54.9 smooth, olive-brown dorsum with lighter mm, means of females 96 mm (82±105 mm); ¯anks (R. bacboensis is shagreened and uni- (3) vomerine teeth present in rows oblique to formly brown with black spots both on the choanae; (4) vertical black stripes on upper dorsum and ¯anks), and adult females are lip (especially under eye), light colored glan- smaller than those of R. bacboensis (67 mm dule above insertion of arm; (5) head broad, vs. Ͼ80 mm). The absence of an outer meta- bluntly rounded in pro®le; (6) tympanum cir- tarsal tubercle and the large female SVL also cular, distinct, TMP:EYE 0.43 in females, distinguishes R. bacboensis from R. archo- 0.66 in the male; (7) supratympanic fold taphus and R. chalconota (female SVL 81± weak; (8) dorsal skin shagreened, becoming 105 mm for R. bacboensis, 59±62 mm and granular laterally, dorsolateral fold absent; 46±59 mm for R. archotaphus and R. chal- venter smooth; (9) dorsum brown with black conota, respectively). The absence of dorso- 2003 BAIN ET AL.: CASCADE FROG FROM SOUTHEAST ASIA 37 lateral folds distinguishes R. bacboensis from TMP:EYE 0.62. Choanae ovoid; vomerine R. chalconota, R. graminea, R. hosii, and R. dentigerous processes prominent, slightly leporipes (pustules on the dorsum of R. gra- oblique, posteromedial to choanae, each hami sometimes form a dorsolateral fold). bearing numerous teeth. Tongue cordiform, Rana bacboensis has webbing to the base of distinctly notched posteriorly, free for ap- the toe pad distinguishing it from R. lepori- proximately one-half its length. pes (webbing to distal phalanx). Rana sinica Forearms moderately robust; ®ngers mod- can further be distinguished from Rana bac- erately short, slender; hands 27% of SVL, boensis by its indistinct, skin-covered tym- relative lengths of ®ngers II Ͻ I Ͻ IV Ͻ III; panum, and different ®nger formula (I Ͻ II ventromedial callous ridge on ®ngers II and Ͻ IV for R. sinica,IIϽ I Ͻ IV for R. bac- III prominent, extending to proximal tuber- boensis). Rana bacboensis shares a super®- cle; disks expanded (Ͻ2ϫ base of phalan- cial resemblance to R. tiannensis, another ges), relative pad size II Ͻ I Ͻ IV Ͻ III, pad large brown cascade ranid, but differs in hav- length (III) 75% of pad width; ventral cir- ing shagreened dorsal skin with small lateral cummarginal grooves present; terminal pha- granulations (dorsum of R. tiannensis is langes T-shaped; subarticular tubercles con- rough with large, prominent lateral granula- ical. Hindlimbs moderately robust; tibia tions) and smaller toe disk than ®nger disk length 60% of SVL; foot length 62% of (the opposite condition of R. tiannensis). SVL; relative toe lengths I Ͻ II Ͻ III Ͻ V Rana bacboensis most closely resembles R. Ͻ IV; inner tarsal fold absent; feet fully hainanensis, R. jingdongensis, and R. ander- webbed to disks, but as a fringe from distal sonii. Rana bacboensis further differs from subarticular tubercle of IV, lateral fringes on R. hainanensis in its relative lengths of ®n- toes I and V to terminal phalanges; toes long, gers (II Ͻ IV Ͻ I Ͻ III for R. hainanensis) slender, with large, rounded triangular disks, and by its shagreened skin (smooth for R. relative pad size I ϭ II ϭ III Ͼ IV k V, pad hainanensis). Rana bacboensis also differs width (IV) 85% of pad length, circummar- from R. jingdongensis in pro®le of its snout ginal grooves present; subarticular tubercles shape (rounded or obtusely pointed in R. prominent, conical; inner metatarsal tubercle jingdongensis versus rounded in R. bacboen- ovoid, long; outer metatarsal tubercle absent. sis) and texture of skin (R. jingdongensis Xiphisternum large, deeply notched pos- dorsum scattered with tubercles and large teriorly. warts, lips and sides of heads with white Skin on dorsum shagreened with heavy spines, all absent in R. bacboensis). Rana granulations, leathery in alcohol preserva- bacboensis also differs from R. andersonii in tion; dorsolateral folds absent; small tuber- its head shape (obtusely pointed in R. ander- cles anterior and posterior to tympanum; sonii) and absence of ventral spines in the ¯anks with small granulations and large pus- males. tules; distinct granules on posterior thighs DESCRIPTION OF HOLOTYPE: A gravid fe- and around cloaca; cloacal opening unmod- male (ROM 29534), head length greater than i®ed, directed posteriorly, at upper level of width (127%), head width 34% of SVL, thighs. length 43% of SVL; snout short, protruding COLOR IN LIFE (in preservative): Dorsum, beyond margin of lower jaw, rounded in dor- ¯anks, and loreal region brown (brownish sal view, bluntly rounded in pro®le; eye gray) with small black spots, becoming larg- large, prominent, 81% of snout length; eyelid er on the ¯anks; upper and lower lips creamy broader than interorbital distance. Top of yellow with vertical black bars; dorsal limbs head ¯at; canthus rostralis rounded; loreal re- and digits brown with black transverse gion concave; lip ¯ared just anterior to orbit; bands; webbing on feet marbled white and nostril about three-fourths distance from eye dark brown (black); venter creamy white, to tip of snout; supratympanic fold barely ev- sometimes with light spotting on belly, chest, ident, curving posteroventrally from poste- and chin; iris golden, margin of pupil out- rior corner of eye to a level above insertion lined in a striking yellow and red border. of arm; tympanum round, distinctly visible, SECONDARY SEXUAL CHARACTERS: The ho- separated from eye by distance equal to lotype possesses large, black eggs (2 mm in 38 AMERICAN MUSEUM NOVITATES NO. 3417 diameter). The lone male paratype has gular posed to sunlight to promote development, a pouches, thickened forearms, and thick white requirement for some species with melanic nuptial pads. eggs (Duellman and Trueb, 1986). MEASUREMENTS OF HOLOTYPE (in mm): SVL 95.1; SNT 12.0; HDL 41.0; HDW 32.2; Rana daorum, new species EYE 9.7; IOD 6.4; TMP 6.0; TEY 5.0; HND (Previously referred to as species 4, 25.6; FGR 21.8; FPL 2.8; FPW 3.7; TIB ``Small'') 56.8; FTL 59.6; TPL 2.4; TPW 2.0. Figures 11E, F, 12G, H, 13C VARIATION OF PARATYPES: Variation in all type material is given in table 15. HOLOTYPE: (ROM ®eld no. 19047) ROM MEASUREMENTS OF FEMALE PARATYPES (in 26381 an adult female from approximately 5 mm, n ϭ 6, ROM 29359, 29526±29530): km NW of Sa Pa village, near O Qui Ho SVL 95.8 mm Ϯ 6.4 (81.8±105.1); SNT 13.8 Pass, Lao Cai Province, Vietnam Ϯ 1.4 (10.8±15.6) ; HDL 46.4 Ϯ 3.5 (43.5± (22Њ22Ј09ЉN, 103Њ50Ј14Љ E, elevation 1400 51.2); HDW 35.4 Ϯ 2.9 (34.1±39.6); EYE m) collected on 7 May 1995 along a water- 9.8 Ϯ 0.5 (9.4±10.8); IOD 7.9 Ϯ 1.7 (5.8± fall beside the road by A. Lathrop and B. 11.0); TMP 5.3 Ϯ 0.6 (4.0±6.0); TEY 4.9 Ϯ Hubley at approximately 1930 hours. 0.7 (3.6±6.0); HND 25.5 Ϯ 2.8 (18.6±30.3); PARATYPES: ROM 26382±26397, all males, FGR 20.8 Ϯ 2.3 (14.0±24.1); FPL 3.0 Ϯ 0.3 collected with the holotype, 7 May 1995 by (2.4±3.6); FPW 3.5 Ϯ 0.6 (2.6±4.3); TIB A. Lathrop and B. Hubley at 1030 hours; 58.6 Ϯ 3.7 (50.5±66.2); FTL 70.2 Ϯ 7.8 ROM 38500±38530, 38532±38540, 38542± (55.8±79.1). 43, 38546±38561 collected between 30 April MEASUREMENTS OF MALE PARATYPE (in and 15 May 1999 in the vicinity of the type mm, FMNH 255611): SVL 54.9, SNT 8.8, locality approximately 12 km northwest of HDL 28.0, HDW 18.1, EYE 6.6, IOD 6.1, Sa Pa Village near the O Qui Ho Pass TMP 4.4, TEY 1.8, HND 16.5, FPL 2.4, (22Њ20Ј58ЉN, 103Њ46Ј14ЉE, elevation 1900 FPW 1.8, TIB 32.4, FTL 45.4, TPL 2.4, m) by R.O. de SaÂ, C.T. Ho, A. Lathrop, T. TPW 1.6. Mason, R.W. Murphy, and N.L. Orlov. ROM ETYMOLOGY: The speci®c name, derived 38547 is a subadult; ROM 38503, 38507, from Bac Bo, the Vietnamese name for 38512, 38516, and 38530 are gravid females; northern Vietnam (often referred to as Ton- ROM 38500, 38517, 38526, and 38538 are kin), re¯ects this species' distribution. nongravid females; and the remaining spec- DISTRIBUTION AND ECOLOGY: Rana bac- imens are males with distended gular pouch- boensis occurs in forested montane river sys- es. tems across northern Vietnam. These rivers DIAGNOSIS: Rana daorum, a member of the vary from shallow and slow moving to tor- subgenus Odorrana (sensu Fei et al., 1990), rential and deep. Specimens may be found is characterized by a combination of the fol- on boulders and logs, both in and around the lowing attributes: (1) body dorsoventrally water and in the adjacent forest. Radiographs compressed; (2) SVL means of males 36 mm revealed that females feed on large inverte- (32±38 mm), females 55 mm (53±58 mm); brates, including small freshwater crabs. Fe- (3) vomerine teeth absent; (4) lip-stripe males were collected in April±May and Oc- white, extending across upper lip, terminat- tober. The holotype, collected in October, has ing in a glandule above insertion of arm; (5) fully developed ova, and two females (ROM head not broad, snout rounded in dorsal 26358, 29529) collected in the spring have view, rounded in pro®le; (6) tympanum undeveloped ova suggesting a fall breeding round, distinct, TMP:EYE of females (0.45) season. No calls are associated with this spe- greater than males (0.29); (7) supratympanic cies. The tadpoles are unknown. fold weak or absent; (8) dorsal skin smooth, REMARKS: Cascade ranids bearing white granular near cloaca and tympanum, dorso- eggs lay them under rocks, sheltered from lateral fold covered with small white gran- the sunlight (Pope, 1931; ROM ®eld notes, ules, ventral skin smooth; (9) dorsum green 1996). In contrast, the black eggs of R. bac- occasionally with black spots, ¯anks brown boensis might be found where they are ex- with at least one large white glandular spot; 2003 BAIN ET AL.: CASCADE FROG FROM SOUTHEAST ASIA 39 fore- and hindlimbs goldish brown, with brown, has vomerine teeth, and males have mottling or indistinct transverse bands; (10) a TMP:EYE ratio twice that of R. daorum. median callous pad on proximal two-thirds DESCRIPTION OF THE HOLOTYPE: An adult of ®ngers II and III; (11) disks on ®ngers and female (ROM 26381), head width 74% of toes greatly enlarged (Ͼ2ϫ base of phalanx), length, length 46% of SVL; snout short, ®nger pads larger than toe pads; (12) feet ful- rounded in dorsal view, rounded in pro®le, ly webbed to disks, but as a fringe from dis- protruding beyond margin of lower jaw; eye tal subarticular tubercle of IV, lateral fringes very large, prominent, 72% of snout; eyelid on I and V to terminal phalanges, webbing broader than interorbital distance. Top of mottled brown; (13) subarticular tubercles head ¯at; canthus rostralis rounded; loreal re- and inner metatarsal tubercle distinct, coni- gion vertical, concave; lip ¯ared just anterior cal; (14) terminal phalanges T-shaped; (15) to orbit; nostril about three-fourths distance xiphisternum narrow, forked posteriorly; (16) from eye to tip of snout; supratympanic fold males with nuptial pads on thumb, paired gu- indistinct, slight swelling above tympanum; lar pouches, pectoral spines absent; (17) eggs tympanum round, distinctly visible, separat- large, white. ed from eye by distance equal to that of TMP, COMPARISONS: Though bearing a super®- 41% of EYE. Choanae ovoid; vomerine den- cial resemblance to other cascade ranids of tigerous processes absent. Tongue cordiform, Southeast Asia, R. daorum is distinct (table distinctly notched posteriorly, free for ap- 12) by the absence of vomerine teeth, fe- proximately two-thirds its length. males with larger TMP:EYE than males, and Forearms robust; ®ngers moderately short, the presence of at least one large white spot slender, hand 28% of SVL, relative lengths on each ¯ank. Its noticeably smaller SVL, of ®ngers I Ͻ II Ͻ IV Ͻ III, ventromedial dorsolateral folds formed by white granules, callous pad on ®ngers II and III for two- and solid, bright green dorsum immediately thirds length of ®nger; disks greatly expand- distinguishes it from H. nasica, R. anderson- ed (Ͼ2ϫ base of phalanges), relative pad size ii, R. archotaphus, R. bacboensis, R. chlo- II Ͻ I Ͻ IV Ͻ III, pad length (®nger III) ronota, R. grahami (whose dorsal pustules equal to pad width, ventral circummarginal sometimes form a fold), R. graminea, R. grooves present; terminal phalanges T- hainanensis, R. hejiangensis, R. jingdongen- shaped; subarticular tubercles conical. Hind- sis, R. junlianensis, R. kwangwuensis, R. liv- limbs moderately robust; tibia length 60% of ida, R. margaretae, R. schmackeri, R. sinica, SVL; foot length 84% of SVL; relative toe and R. tiannensis. The entirely white eggs of lengths I Ͻ II Ͻ III Ͻ V Ͻ IV; inner tarsal R. daorum distinguish it from Huia nasica, fold absent; feet fully webbed to disks, but Rana andersonii, R. chalconota, R. grahami, as a fringe from distal subarticular tubercle R. junlianensis, R. margaretae, R. schmack- of IV, lateral fringe on toe V to terminal pha- eri (all with white eggs with a melanic pole), lanx; toes long, slender, with enlarged disks, and R. bacboensis (fully pigmented eggs). smaller than those on ®ngers, relative pad The presence of gular pouches in males dif- size I ϭ II ϭ III Ͼ IV k V, pad width (IV) ferentiates R. daorum from R. andersonii, R. 83% of length, each pad with ventral circum- chalconota, R. grahami, R. hainanensis, R. marginal grooves; subarticular tubercles hosii, R. jingdongensis, R. junlianensis, R. prominent and conical; inner metatarsal tu- kwangwuensis, and R. margaretae. A distinct bercle ovoid, long; outer metatarsal tubercle and uncovered tympanum also distinguishes absent. R. daorum from R. sinica. Rana daorum can Xiphisternum narrow, notched posteriorly. be further distinguished from R. livida, R. Skin on dorsum smooth, dorsolateral folds chloronota, R. maragaretae, and R. prominent in form of granules; small tuber- schmackeri by its ®nger formula (II Ͻ I Ͻ cles posteroventrally to tympanum, distinct IV for R. daorum,IϽ II Ͻ IV for others granules on ¯anks and dorsum to cloaca; clo- listed). Although R. daorum closely resem- acal opening unmodi®ed, directed posteriorly bles A. chunganenesis in size, the presence at upper level of thighs. of white granular dorsolateral folds, and di- COLOR IN LIFE (in preservative): Dorsum urnal behavior, A. chunganensis is red- green (livid blue), ¯anks brown (brown) and 40 AMERICAN MUSEUM NOVITATES NO. 3417 green (gray), each a prominent white spot; 0.6 (11.1±13.0); EYE 4.2 Ϯ 0.6 (3.2±5.2); granules of dorsolateral fold golden (white); IOD 3 Ϯ 0.4 (2.3±3.6); TMP 1.1 Ϯ 0.2 (1.0± lip-stripe white (silvery white) from nostril 1.9); TEY 0.6 Ϯ 0.2 (0.2±1.0); HND 10.1 Ϯ to above insertion of arm; tympanum dark 0.6 (9.0±11.1); FGR 9.0 Ϯ 0.3 (8.5±9.6); brown; loreal region dark brown (black); FPL 1.7 Ϯ 0.3 (1.2±2.0); FPW 2.0 Ϯ 0.2 dorsal surfaces of limbs mottled brown and (1.7±2.2); TIB 22.1 Ϯ 1.3 (19.1±23.5); FTL yellow with indistinct dark brown (black) 29.7 Ϯ 4.4 (27.1±40.4); TPL 1.6 Ϯ 0.2 (1.2± transverse bands; posterior surface of thighs 1.7); TPW 1.4 Ϯ 0.1 (1.2±1.6). yellow with brown (black) mottling; web- ETYMOLOGY: The speci®c name is a pa- bing marbled white (translucent) and dark tronym for the±D ao people (pronounced brown; venter creamy white; iris golden, pu- ``zao'') of northern Vietnam. pil outlined in a striking yellow and red bor- DISTRIBUTION AND ECOLOGY: This species der. is known from the vicinity of Sa Pa village, SECONDARY SEXUAL CHARACTERS: Gravid Lao Cai Province in northern Vietnam. The females have immaculate white eggs. They photograph of a froglet of R. livida (in Kar- are approximately 1.5 times larger than sen et al., 1998) also documents the occur- males. Males have a proportionally smaller rence of R. daorum in Hong Kong. The dis- tympanum than females (TMP:EYE for tinctive granular dorsolateral fold of R. dao- males 0.29, for females 0.45). The EYE:SNT rum is clearly visible in the misidenti®ed is also greater in females (0.72) than it is in frog. males (0.51). Males have velvety nuptial In early May, male Vietnamese R. daorum pads extending across thumb, and paired gu- are actively calling by 1000 hours on par- lar pouches located at the angles of the jaw. tially submerged rocks in cascades as well as Pectoral spines are absent. in vegetation adjacent to the streams. Fe- MEASUREMENTS OF HOLOTYPE (in mm): males, although not as common, can be SVL 55.7; SNT 7.8; HDL 25.4; HDW 18.8; found slightly farther away from streams in EYE 5.6; IOD 5.6; TMP 2.3; TEY 2.4; HND more dense vegetation. One male (ROM 15.5; FGR 13.9; FPL 2.8; FPW 2.8; TIB 26394) was found in amplexus with the ho- 33.6; FTL 47.0; TPL 2.3; TPW 1.9. lotype (®g. 6E). VARIATION OF PARATYPES: The loreal re- REMARKS: Rana daorum differs substan- gion on some specimens varies from dark tially from R. graminea (Boulenger, 1899; brown to green. The large white spot on the Bourret, 1942) despite having a dorsolateral ¯anks is sometimes accompanied by smaller fold. Its small, forked xiphisternum differs ones. Flanks also have varying degrees of from the large, deeply notched element of white mottling. The venter of some speci- other members in the Rana chloronota com- mens has light mottling on the chest and plex and the subgenus Odorrana (sensu Fei chin. Variation in all type material is given et al., 1990) and more closely resembles that in table 15. of male Huia nasica (Yang, 1991b). Huia na- MEASUREMENTS OF FEMALE PARATYPES (in sica shares range, habitat and morphological mm, n ϭ 8, ROM 38500, 38503, 38507, similarities with the Rana chloronota com- 38512, 38516, 38517, 38526, 38530): SVL plex: greatly expanded ®nger and toe disks, 55.0 Ϯ 1.2 (53.3±57.6); SNT 7.3 Ϯ 0.5 (6.8± tremendous sexual dimorphism in size, white 8.3); HDL 17.8 Ϯ 1.6 (16.7±19.4); HDW eggs, paired gular pouches, and a high chirp- 17.2 Ϯ 0.6 (15.6±17.6); EYE 5.8 Ϯ 0.4 (5.3± like call (Boulenger, 1920; Pope, 1931; 6.4); IOD 10.0 Ϯ 1.7 (10.0±11.3); TMP 2.5 Bourret, 1942). Ϯ 0.2 (2.3±3.0); FPW 2.3 Ϯ 0.3 (1.8±2.8); TIB 34.3 Ϯ 1.3 (32.7±36.4); TPW 2.3 Ϯ 0.3 Rana hmongorum, new species (1.8±2.8). (Previously referred to as species 3, MEASUREMENTS OF MALE PARATYPES (in ``Speckled'') mm, n ϭ 7, ROM 26383, 26386, 26387, Figures 14A, 12I, J, 13D 26389, 26390, 26392, 26394): SVL 36.2 Ϯ 1.2 (34.8±38.1); SNT 4.5 Ϯ 0.4 (4.0±4.9); HOLOTYPE: (ROM ®eld no. 19055) ROM HDL 18.4 Ϯ 1.7 (16.5±21.0); HDW 12.3 Ϯ 26376 an adult female approximately 5 km 2003 BAIN ET AL.: CASCADE FROG FROM SOUTHEAST ASIA 41

NW of Sa Pa village, near O Qui Ho Pass, phalanges T-shaped; (15) xiphisternum large, Lao Cai Province, Vietnam (22Њ20Ј09ЉN, deeply notched posteriorly; (16) males with 103Њ50Ј14ЉE, elevation 1400 m) collected 30 nuptial pad; gular pouches and pectoral April 1995 by A. Lathrop and B. Hubley be- spines absent; (17) eggs white. tween 1900 and 2100 hours. The holotype COMPARISONS: Rana hmongorum super®- had leg and liver tissue removed shortly after cially resembles other Asian cascade ranids, it was euthanised. including Huia nasica, Rana andersonii, R. PARATYPES: ROM 26370±26375, 26377± archotaphus, R. bacboensis, R. chalconota, 26380, 39235, and 39236 collected in the vi- R. chloronota, R. daorum, R. grahami, R. cinity of the holotype between 9 and 12 May graminea, R. hainanensis, R. hejiangensis, R. 1995 by A. Lathrop and B. Hubley. ROM hosii, R. jingdongensis, R. junlianensis, R. 26380, 39235, and 39236 are males and the kwangwuensis, R. leporipes, R. livida, R. remainder females. Additional specimens from margaretae, R. schmackeri, R. sinica, and R. the vicinity of the type locality include AMNH tiannensis (table 12). The white eggs of R. A-161480 collected on 27 August 1997 by hmongorum distinguish it from H. nasica, D.R. Frost and C.J. Raxworthy, and ROM Rana andersonii, R. chalconota, R. grahami, 39867±39896 (22Њ20Ј58ЉN, 103Њ46Ј14ЉE, ele- R. junlianensis, R. margaretae, R. schmack- vation 1900 m) collected between 30 April and eri (all with white eggs with a melanic ani- 2 May 1999 by R.O. de SaÂ, C.T. Ho, A. La- mal pole), and R. bacboensis (fully pigment- throp, T. Mason, R.W. Murphy, and N.L. Or- ed eggs). The dorsum and ¯anks with distinct lov. Male specimens are ROM 39874±39879, pustules distinguishes R. hmongorum from 39888, and 39890±39894; juveniles and sub- all of the above species, except R. jingdon- adults ROM 39867, 39868, 39880, 39895, and gensis, and R. grahami; R. daorum has two 39896; all others are females. distinct rows of granular dorsolateral folds. DIAGNOSIS: Rana hmongorum, a member Absence of gular pouches further differenti- of the subgenus Odorrana (sensu Fei et al., ates it from H. nasica, R. archotaphus, R. 1990), is characterized by a combination of bacboensis, R. chloronota, R. daorum, R. the following attributes: (1) body dorsoven- graminea, R. junlianensis, R. kwangwuensis, trally compressed; (2) SVL means of males R. schmackeri, and R. tiannensis. Rana 59 mm (54±65 mm), females 80 mm (74± hmongorum has an indistinct yellow lip- 87 mm); (3) vomerine teeth in rows postero- stripe distinguishing it from all other cascade medial to choanae; (4) lip-stripe yellow, in- ranids above, except R. chalconota, R. jing- distinct, extending across upper lip, termi- dongensis, R. junlianensis, and R. grahami. nating in a row of glandules; (5) head broad, The presence of webbing to the base of the bluntly rounded in pro®le; (6) tympanum toe pad distinguishes R. hmongorum from R. round, distinct, TMP:EYE in males (0.43) leporipes (webbing to distal phalanx) and R. greater than females (0.31); (7) supratympan- archotaphus. Larger females and the absence ic fold weak; (8) dorsal skin smooth with of an outer metatarsal tubercle further distin- pustules becoming more pronounced lateral- guish R. hmongorum from R. archotaphus ly, pustules may form two dorsolateral lines (female SVL 59±62 mm) and R. chalconota resembling a fold, ventrum smooth; (9) dor- (SVL 46±59 mm). Absence of ventral spines sum and forearms green with black spots, in males differentiates R. hmongorum from ¯anks reddish brown with yellowish pus- R. andersonii, R. grahami, R. jingdongensis, tules, transverse bars on fore± and hindlimbs; R. junlianensis, R. margaretae, and R. (10) median callous pad on ®ngers II and III schmackeri. Rana sinica is further differen- to proximal tubercle; (11) disks on ®ngers tiated from R. hmongorum by its indistinct, and toes moderately expanded (Ͻ2ϫ base of skin-covered tympanum, the relative length phalanges); (12) feet fully webbed to disks, of its digits (I Ͻ II Ͻ IV for R. sinica,IIϽ but as a fringe from distal subarticular tu- I Ͻ IV for R. hmongorum), and its rounded bercle of IV, lateral fringes on toes I and V distal phalanges (T-shaped in R. hmongo- to terminal phalanges, webbing brown-gray; rum). Huia nasica is different in that it has (13) subarticular tubercles distinct, conical; a smooth, olive-brown dorsum, a longer head inner metatarsal tubercle ovoid; (14) terminal and smaller size (67 mm vs. Ͼ75 mm for 42 AMERICAN MUSEUM NOVITATES NO. 3417

Fig. 14. Members of the Rana chloronota complex from Vietnam. A. Paratype of R. hmongorum, new species, male from Lao Cai Province. (B) Female, brown (night) phase, (C) female, green (day) 2003 BAIN ET AL.: CASCADE FROG FROM SOUTHEAST ASIA 43 females, 44±49 mm vs. 54±59 for males). ϭ III Ͼ IV k V, pad width (IV) 78% of pad The small male of R. hmongorum further dis- length, ventral circummarginal grooves pre- tinguishes it from R. jingdongensis, R. jun- sent; subarticular tubercles prominent and lianensis, R. andersonii, and R. grahami conical; inner metatarsal tubercle ovoid, (SVL 62±81 mm for R. jingdongensis,70± long; outer metatarsal tubercle absent. 80 mm for R. andersonii, 66±84 mm for R. Xiphisternum large, deeply notched pos- grahami). The relative length of ®ngers fur- teriorly. ther distinguishes it from R. hainanensis (II Skin on dorsum smooth with large pus- Ͻ IV Ͻ I Ͻ III) and R. hejiangensis (II Ͻ I tules, particularly on ¯anks, sacrum, and Ͻ III Ͻ IV). around cloaca; cloacal opening unmodi®ed, DESCRIPTION OF THE HOLOTYPE: ROM directed posteriorly at upper level of thighs. 26376, an adult female, head length greater COLOR IN LIFE (in preservative): Dorsum than width (136%), head width 36% of SVL, green (dark brown) and reddish brown (dark length 49% of SVL; snout short, protruding brown) with black spots. Sides reddish beyond margin of lower jaw, rounded in dor- brown (gray) and yellow (cream), with some sal view, bluntly rounded in pro®le; eye black spotting. Pustules on ¯anks yellowish large, prominent, 88% of snout length; eyelid white (creamy white), with black around the broader than interorbital distance. Top of bases. Cloacal region dark brown (black). head ¯at; canthus rostralis rounded; loreal re- Lip-stripe creamy white (gray); dorsal sur- gion concave; lip ¯ared just anterior to orbit; face of forearms with green (gray) patch; nostril about three-fourths distance from eye limbs mottled yellow (light brown) and to tip of snout; supratympanic fold indistinct, brown (dark brown) with black transverse slight swelling above the tympanum; tym- bands; anterior and posterior surfaces of panum round, distinctly visible, separated thighs with brown (dark brown) marbling on from eye by distance equal to TMP, 26% of yellow (cream); webbing uniformly brown; EYE. Choanae ovoid; vomerine dentigerous venter creamy white (creamy yellow). Iris processes prominent, posteromedial to cho- greenish yellow or brown, pupil outlined anae, each bearing numerous teeth. Tongue with a yellow border. cordiform, distinctly notched posteriorly, free SECONDARY SEXUAL CHARACTERISTICS: Ho- for approximately one-half its length. lotype is a gravid female with white eggs, 2 Forearms robust; ®ngers relatively short, mm diameter. Adult female SVL 135% that slender, hands 25% of SVL, relative lengths of males. Males have slightly larger tympa- of ®ngers II Ͻ I Ͻ IV Ͻ III; ventromedial num than do females, and velvety nuptial callous pad on ®ngers II and III to proximal pads on thumb. Paired gular pouches and tubercle, disks moderately expanded (Ͻ2ϫ pectoral spines absent. Venter of males varies base of phalanges), relative pad size III Ͼ IV from immaculate creamy white to darkly Ͼ I Ͼ II, pad length (III) 85% of pad width; mottled brown (gray in preservative). ventral circummarginal grooves on disks pre- MEASUREMENTS OF HOLOTYPE (in mm): sent; terminal phalanges T-shaped; subarti- SVL 86.8; SNT 11.1; HDL 42.3; HDW 31.1; cular tubercles conical. Hindlimbs moderate- EYE 9.7; IOD 6.4; TMP 2.5; TEY 3.9; HND ly robust; tibia length 57% of SVL; foot 22.3; FGR 16.5; FPL 2.3; FPW 2.7; TIB length 40% of SVL; relative toe lengths I Ͻ 49.2; FTL 35.0; TPL 3.0; TPW 2.3. II Ͻ III Ͻ V Ͻ IV; inner tarsal fold absent; VARIATION OF PARATYPES: Pustules can feet fully webbed to disks, but as a fringe sometimes form dorsolateral lines, which su- from distal subarticular tubercle of IV, lateral per®cially resemble folds. Pustules can be fringes on toes I and V to terminal phalan- white. Variation in all type material is given ges; toes long, slender, with large, triangu- in table 15. larly rounded disks, relative pad size I ϭ II MEASUREMENTS OF FEMALE PARATYPES (in

← phase, and (D) male of R. morafkai, new species, from Gia Lai Province. (E) Female and (F) male of R. banaorum, new species, from Gia Lai Province. (G, H) female of R. megatympanum new species. 44 AMERICAN MUSEUM NOVITATES NO. 3417 mm, n ϭ 9; ROM 26370±26375, 26377± tissue removed shortly after it was euthani- 26379): SVL 80.2 Ϯ 4.3 (74.3±86.8); SNT sed. 11.0 Ϯ 0.5 (10.1±11.7); HDL 38.8 Ϯ 1.9 PARATYPES: ROM 39904±39911, 39930, (37.0±41.0); HDW 29.8 Ϯ 1.0 (29.2±31.3); 39934, 39937, 39947, and 39949 collected EYE 9.3 Ϯ 1.1 (7.6±10.8); IOD 7.0 Ϯ 0.3 with the holotype between 15 and 19 June (6.4±7.3); TMP 2.9 Ϯ 0.4 (2.5±3.3); TEY 3.0 1996 by R.H. Bain, B. Hubley, A. Lathrop, Ϯ 0.6 (2.1±3.9); HND 23.2 Ϯ 1.3 (22.1± R.W. Murphy, and N.L. Orlov; ROM 39904± 25.6); FGR 18.6 Ϯ 1.1 (16.5±19.6); FPL 3.0 39011, and 39937 are males; the remainder Ϯ 0.4 (2.3±3.4); FPW 2.6 Ϯ 0.4 (2.0±3.1); are females. ROM 25094±25097, 25099, TIB 47.6 Ϯ 2.3 (43.1±49.2); FTL 39.6 Ϯ 5.0 25101, 25104±25106, 25108±25111, all (35.0±48.3) TPL 2.8 Ϯ 0.6 (2.0±3.5); TPW males collected in Buon Loi, An Khe Dis- 2.4 Ϯ 0.5 (1.9±3.3). trict, Gia Lai Province, Vietnam (elevation MEASUREMENTS OF MALE PARATYPES (in 700±750 m) by I.S. Darevsky and N.L. Or- mm, n ϭ 12; ROM 26380, 30876, 39235, lov on 11 November 1993. 39236, 39874, 39875, 39877, 39879, 39888, DIAGNOSIS: Rana morafkai, a member of 39891, 39892; AMNH A-161480): SVL 59.4 the subgenus Odorrana (sensu Fei et al., Ϯ 3.8 (54.7±65.3); SNT 9.4 Ϯ 0.9 (8.7± 1990), is characterized by a combination of 10.4); HDL 24.0 Ϯ 3.0 (20.7±30.4); HDW the following attributes: (1) body dorsoven- 20.0 Ϯ 1.2 (18.9±21.1); EYE 7.3 Ϯ 0.5 (6.7± trally compressed; (2) SVL means of males 8.3); IOD 5.3 Ϯ 0.7 (4.3±6.5); TMP 3.0 Ϯ 43 mm (39±46 mm), females 88 (80±100 0.4 (2.4±3.7); TEY 2.5 Ϯ 0.3 (2.2±3.0); mm); (3) vomerine teeth in rows oblique to HND 17.2 Ϯ 1.4 (14.1±19.4); FGR 13.3 Ϯ choanae; (4) yellow-white lip-stripe extend- 1.2 (12.8±14.8); FPL 2.5 Ϯ 0.5 (1.6±3.6); ing across upper lip, terminating in glandule FPW 2.1 Ϯ 0.6 (1.0±2.4); TIB 38.6 Ϯ 4.5 above insertion of arm; (5) head broad, (33.4±51.2); FTL 47.2 Ϯ 5.7 (30.4±52.1); bluntly rounded in pro®le; (6) tympanum TPL 2.3 Ϯ 0.4 (1.7±2.9); TPW 1.7 Ϯ 0.4 round, relatively large, distinct, approximate- (0.9±2.2). ly 96% of eye length in males, 71% in fe- ETYMOLOGY: The speci®c name is a pa- males; (7) supratympanic fold weak; (8) dor- tronym for the Hmong (pronounced ``huh- sal skin smooth or partly shagreened, becom- mung'') people, an ethnic group in the north- ing granular laterally, dorsolateral folds ab- ern montane regions of Vietnam. Their as- sent; (9) dorsum changing colors between sistance made it possible for us to document day and night, usually bright green in day- the fauna of Hoang Lien Mountains. light, brown at night sometimes with black DISTRIBUTION AND ECOLOGY: This species spots; forelimbs and hindlimbs with trans- is only known only from the vicinity of verse bars; (10) median callous pad on ®nger Mount Fan Si Pan at approximately 1900 m III to proximal tubercle; (11) disks on ®ngers or above. Adults were found along the cas- and toes greatly enlarged (Ͼ2ϫ base of pha- cades of a waterfall and along the mossy langes); (12) webbing on feet complete to slopes of a human-made culvert. base of disk in females, sometimes as a fringe on IV; in males to distal tubercle in Rana morafkai, new species males, lateral fringes on I and V to terminal phalanges; webbing brown-gray; (13) subar- (Previously referred to as species 5, ticular tubercles and inner metatarsal tubercle ``Mottled'') conical, indistinct; (14) terminal phalanges Figures 12K, L, 13E, 14 B±D T-shaped; (15) xiphisternum deeply notched posteriorly; (16) males with nuptial pads on HOLOTYPE: (ROM ®eld no. 7301) ROM thumb, paired gular pouches, pectoral spines 39932, a female from Tram Lap, An Khe absent; (17) eggs white. District, Gia Lai Province, Vietnam COMPARISONS: Rana morafkai resembles (14Њ26Ј39ЉN, 108Њ32Ј97ЉE, elevation ca. 900 other Asian cascade ranids, including Huia m) collected on 19 June 1996 by R.H. Bain, nasica, Rana andersonii, R. archotaphus, R. B. Hubley, A. Lathrop, R.W. Murphy, and bacboensis, R. chalconota, R. chloronota, R. N.L. Orlov. The holotype had leg and liver daorum, R. grahami, R. graminea, R. hain- 2003 BAIN ET AL.: CASCADE FROG FROM SOUTHEAST ASIA 45 anensis, R. hejiangensis, R. hmongorum, R. in R. morafkai), ®nger formula (I Ͻ II Ͻ IV hosii, R. jingdongensis, R. junlianensis, R. for R. sinica,IIϽ I Ͻ IV for R. morafkai), kwangwuensis, R. leporipes, R. livida, R. rounded distal phalanges (T-shaped in R. margaretae, R. schmackeri, R. sinica, and R. morafkai), and absence of a lip-stripe. Rana tiannensis (table 12). It differs from these leporipes has small disks and white supra- cascade ranids (except R. chalconota) in hav- tympanic folds, both distinguishing it from ing unique nocturnal and diurnal coloration, R. morafkai. The color pattern and skin tex- less distinct subarticular tubercles (all other ture of specimens of R. morafkai may be species bear distinct subarticular tubercles), identical to those of R. chloronota; however, and a webbing pattern that is unique to each the pronounced sexual dimorphism in the sex (females fully webbed to disk, males to tympanum size (TMP:EYE R. morafkai distal subarticular tubercles). Female R. mor- males 0.96; females 0.71) differentiates it afkai (SVL 80±100 mm) are considerably from all Vietnamese populations of Rana larger than female R. archotaphus (59±62 chloronota except those from Na Hang mm), R. chalconota (45±60 mm), R. daorum (males 0.60, females 0.43). (55±58 mm), and R. sinica (holotype 66.6 DESCRIPTION OF THE HOLOTYPE: An adult mm). The presence of a large tympanum in female (ROM 39932), head width 66% of the males (TMP:EYE 0.96) differentiates R. length, length 50% of SVL; snout short, morafkai from H. nasica (0.5), R. andersonii acutely rounded in dorsal view, bluntly (0.7), R. chloronota (0.57), R daorum (0.29), rounded in pro®le, protruding beyond margin R. grahami (0.53), R. graminea (0.77), R. of lower jaw; eye large, prominent, 38% of hainanensis (0.66), R. hmongorum (0.43), snout length; eyelid broader than interorbital and R. hejiangensis (0.5). Its broadly round- distance. Top of head ¯at; canthus rostralis ed snout distinguishes R. morafkai from H. rounded; loreal region concave; lip ¯ared just nasica, R. andersonii, R. chalconota, R. anterior to orbit; nostril about three-fourths schmackeri (obtusely pointed snouts in pro- distance from eye to tip of snout; supratym- ®le), R. graminea, and R. margaretae (dis- panic fold curving posteroventrally from tinctly depressed snouts). The presence of posterior corner of eye to a level above the gular pouches differentiates male R. moraf- insertion of the arm; tympanum round, dis- kai from those of R. andersonii, R. chalcon- tinctly visible, separated from eye by dis- ota, R. grahami, R. hainanensis, R. hmon- tance equal to TMP, 68% of EYE. Choanae gorum, R. jingdongensis, R. hosii, and R. ovoid; vomerine dentigerous processes margaretae. The absence of dorsolateral prominent, oblique, posteromedial to choa- folds distinguishes R. morafkai from R. chal- nae, each bearing numerous teeth. Tongue conota, R. daorum, R. graminea, R. hosii, R. cordiform, distinctly notched posteriorly, free junlianensis, R. kwangwuensis, R. leporipes, for approximately two-thirds its length. and R. margaretae (pustules on the dorsum Forearms moderately robust; ®ngers mod- of R. grahami and R. hmongorum sometimes erately short, slender, hands 21% of SVL, form a dorsolateral fold). The absence of an relative lengths of ®ngers II Ͻ I Ͻ IV Ͻ III, outer metatarsal tubercle distinguishes R. ventromedial callous pad on ®nger III to morafkai from R. archotaphus and R. chal- proximal tubercle, disks greatly expanded conota. The presence of completely white (Ͼ2ϫ base of phalanges), relative pad size II eggs differentiates it from R. bacboensis Ͻ I Ͻ IV Ͻ III, pad width (III) 90% of pad (black eggs), H. nasica, R. andersonii, R. length, ventral circummarginal grooves pre- chalconota, R. grahami, R. junlianensis, R. sent; terminal phalanges T-shaped; subarti- margaretae, and R. schmackeri (white eggs cular tubercles conical. Hindlimbs moderate- with a dark melanic pole). The absence of ly robust; tibia length 62% of SVL; foot spinules on the venter distinguishes R. mor- length 82% of SVL; relative toe lengths I Ͻ afkai from R. andersonii, R. grahami, R. II Ͻ III Ͻ V Ͻ IV; inner tarsal fold absent; jingdongensis, R. junlianensis, R. margare- feet fully webbed to toe disk, lateral fringe tae and R. schmackeri. Rana sinica differs on I and V to terminal phalanges, toes long, from R. morafkai by its indistinct and skin- slender, with large, rounded, triangular disks, covered tympanum (distinct and uncovered relative pad size I ϭ II ϭ III Ͼ IV k V, pad 46 AMERICAN MUSEUM NOVITATES NO. 3417 width (IV) 80% of pad length, each with EYE 6.0 Ϯ 0.9 (4.9±7.6); IOD 7.7 Ϯ 0.9 ventral circummarginal grooves; subarticular (6.1±9.0); TMP 4.1 Ϯ 0.7 (3.4±5.2); TEY 4.5 tubercles prominent and conical; inner meta- Ϯ 1.9 (3.2±8.3); HND 18.8 Ϯ 2.2 (16.7± tarsal tubercle ovoid, long; outer metatarsal 21.9); FGR 15.0 Ϯ 2.5 (12.2±18.1); FPL 3.5 tubercle absent. Ϯ 0.4 (2.8±3.9); FPW 3.0 Ϯ 0.3 (2.7±3.5); Xiphisternum large, deeply notched pos- TIB 53.5 Ϯ 4.7 (47.4±61.5); FTL 63.0 Ϯ teriorly. 11.3 (43.3±76.1) TPL 3.1 Ϯ 0.3 (2.5±3.5); Skin on dorsum shagreened, light granu- TPW 2.9 Ϯ 0.5 (2.3±3.7). lations laterally; dorsolateral folds absent; MEASUREMENTS OF MALE PARATYPES (in small tubercles posteroventrally to tympa- mm, n ϭ14, ROM 25094±25097, 25099, num; prominent granules on ¯anks and 25101, 25104±25106, 25108±25111, 39937): around cloaca; cloacal opening unmodi®ed, SVL 43.2 Ϯ 1.7 (39.2±45.9); SNT 6.8 Ϯ 0.4 directed posteriorly at upper level of thighs. (6.2±7.8); HDL 21.3 Ϯ 0.8 (20.5±22.4); COLOR IN LIFE (in preservative): Green HDW 13.8 Ϯ 1.0 (13.2±15.5); EYE 3.5 Ϯ (livid blue, with dark spots); ¯anks yellow 0.5 (2.5±4.5); IOD 3.9 Ϯ 0.4 (3.2±4.8); TMP and gray (cream and gray) with some white 3.3 Ϯ 0.3 (2.8±4.0); TEY 1.2 Ϯ 0.3 (0.8± mottling posteroventrally; lip-stripe yellow- 1.8); HND 11.8 Ϯ 1.4 (9.0±15.6); FGR 9.6 white (creamy white); loreal region brown Ϯ 0.8 (7.9±12.0); FPL 1.8 Ϯ 0.3 (1.16±2.3); (black); tympanum beige with dark brown FPW 1.6 Ϯ 0.2 (1.4±2.0); TIB 26.0 Ϯ 1.5 central ring; limbs above brown with black (22.8±28.4); FTL 27.1 Ϯ 7.7 (17.5±40.9); transverse bands, below creamy yellow with TPL 1.7 Ϯ 0.2 (1.3±2.0); TPW 1.3 Ϯ 0.1 black mottling; thighs marbled yellow and (1.1±1.7). brown; cloacal region black; venter creamy ETYMOLOGY: The speci®c name honors white; iris golden, margin of pupil outlined David Joseph Morafka, Research Associate in a striking yellow and red border. of the Royal Ontario Museum and California SECONDARY SEXUAL CHARACTERS: The ho- Academy of Sciences, in recognition of his lotype is a gravid female with creamy white unfailing friendship, his unsel®sh develop- eggs, 2 mm in diameter. Gravid females are ment and perpetuation of multiple, indepen- approximately twice the SVL of males. Tym- dent research programs, and for his catapult- panum in males (TMP:EYE 0.96) is larger ing the careers of many conservation biolo- than in females (0.71). Webbing on females gists. extends to disk, and only to distal subarti- DISTRIBUTION AND ECOLOGY: Rana moraf- cular tubercle in males. Males have velvety kai is known only from the Tay Nguyen Pla- nuptial pads extending across the thumb, teau of the Central Highlands, Gia Lai Prov- paired gular pouches located at the angle of ince, Vietnam. It inhabits forested montane the jaws, and pectoral spines absent. river systems. Specimens may be found on MEASUREMENTS OF HOLOTYPE (in mm): or near rapids or waterfalls. In at least May SVL 84.4; SNT 14.3; HDL 42.7; HDW 28.2; and June, male specimens may have distend- EYE 5.4; IOD 7.3; TMP 3.7; TEY 4.1; HND ed gular pouches, indicating that this species 18.0; FGR 14.0; FPL 3.0; FPW 2.7; TIB breeds in spring. 52.6; FTL 68.7; TPL 2.9; TPW 2.3. REMARKS: This species has been previous- VARIATION OF PARATYPES: The skin on the ly referred to as R. livida by Inger and Chan- dorsum can be smooth or partly shagreened. ard (1997) and Inger et al. (1999). These frogs are often brown at night, but di- urnally they can become green over their en- Rana banaorum, new species tirely body or only dorsally. The dorsum is (Previously referred to as species 6, occasionally colored with black spots. Vari- ``Southern Big Eye'') ation in all type material is given in table 15. Figures 12M, N, 13F, 14E, F MEASUREMENTS OF FEMALE PARATYPES (in mm, n ϭ 4, ROM 39930, 39934, 39947, HOLOTYPE: (ROM ®eld no. 7145) ROM 39949): SVL 87.6 Ϯ 6.9 (80.4±99.6); SNT 39944, an adult female from Tram Lap, An 13.4 Ϯ 1.7 (10.8±15.6); HDL 39.1 Ϯ 2.4 Khe District, Gia Lai Province, Vietnam (37.6±38.5); HDW 27.2 Ϯ 1.1 (27.7±28.2); (14Њ26Ј39ЉN, 108Њ32Ј97ЉE), elevation ca. 900 2003 BAIN ET AL.: CASCADE FROG FROM SOUTHEAST ASIA 47

TABLE 15 Variation in Body Proportions for Rana chloronota sensu stricto and Vietnamese Species of the Rana chloronota Complex Given as Means (in mm) ؎ 1 SD and ranges See Materials and Methods for abbreviations. 48 AMERICAN MUSEUM NOVITATES NO. 3417 m, collected on 15 June 1996 by R.H. Bain, tinct, conical; (14) terminal phalanges T- A. Lathrop, R.W. Murphy, and N.L. Orlov. shaped; (15) xiphisternum large, deeply The holotype had leg and liver tissue re- notched posteriorly; (16) males with nuptial moved shortly after it was euthanised. pads, paired gular pouches, pectoral spines PARATYPES: Four males (ROM 39912± absent; (17) eggs white. 39913, 39915±39916) and eight females COMPARISONS: Among Asian cascade ra- (ROM 39899±39901, 39928, 39929, 39931, nids, R. banaorum is most similar to sym- 39936, and 39941) collected with holotype patric R. morafkai and particularly R. chlo- between 15 and 29 June 1996 by R.H. Bain, ronota but it can be anatomically distin- A. Lathrop, R.W. Murphy, and N.L. Orlov. guished from both by its dorsolateral folds ROM 39716±39720, 39920±39922, and (table 12). Rana banaorum can further be 39924±39926 (males); 39942 (subadult) distinguished from R. morafkai by its larger from the Cha River, Buon Loi, An Khe Dis- males (SVL 42±55 mm, versus 39±45 mm). trict, Gia Lai Province, Vietnam (elevation Some Rana banaorum males have indistinct, ca. 900 m), collected on 27 June 1996 by microscopic spinules on the dorsal surface of R.H. Bain and N.L. Orlov. ROM 39700, and the leg extending to the feet, forming a 39702±39705 males from Krong Pa, An Khe ``saw-tooth'' formation along the lateral edge District, Gia Lai Province, Vietnam of toe V, which is present only to a slight (14Њ20Ј29Љ N, 108Њ28Ј46ЉE, elevation 850 degree or absent in R. chloronota. Its broad, m), collected on 13±25 August 1997 by C.T. rounded snout differs from the obtusely Ho, A. Lathrop, L.A. Lowcock, R.W. Mur- pointed snouts of H. nasica, R. andersonii, phy, and N.L. Orlov. ROM 25084±25086, R. chalconota, and R. schmackeri and from 25100, 25102, and 25103 (males) collected the depressed snouts of R. graminea and R. from Buon Loi, An Khe District, Gia Lai margaretae. A white lip-stripe differentiates Province, Vietnam (elevation ca. 700±750 R. banaorum from R. andersonii, R. bac- m), 5 November 1993 by I.S. Darevsky and boensis, R. grahami, R. hainanensis, R. N.L. Orlov. hmongorum, R. jingdongensis, R. junlianen- DIAGNOSIS: Rana banaorum, a member of sis, R. margaretae, R. sinica, R. schmackeri, the subgenus Odorrana (sensu Fei et al., and R. tiannensis. The gular pouches of R. 1990), is characterized by a combination of banaorum distinguish it from R. andersonii, the following attributes: (1) body dorsoven- R. chalconota, R. grahami, R. hainanensis, trally compressed; (2) SVL means of males R. hmongorum, R. hosii, R. jingdongensis, R. 50 mm (42±55 mm), females 93 mm (83±99 junlianensis, R. kwangwuensis, and R. mar- mm); (3) vomerine teeth in rows oblique to garetae. The dorsolateral folds of R. banao- choanae; (4) white lip-stripe extending across rum distinguish it from H. nasica, R. ander- upper lip, terminating in glandule above in- sonii, R. bacboensis, R. chloronota, R. hain- sertion of arm; (5) head broad, bluntly round- anensis, R. hejiangensis, R. jingdongensis, R. ed in pro®le; (6) tympanum round, distinct, junlianensis, R. kwangwuensis, R. morafkai, very large, TMP:EYE in males (0.89) greater R. livida, R. schmackeri, R. sinica, and R. than females (0.75); (7) supratympanic fold tiannensis; the dorsolateral folds of R. dao- weak; (8) dorsal skin shagreened, becoming rum are composed of distinct white granules, granular laterally, with slight dorsolateral and dorsolateral pustules sometimes form folds; (9) dorsum usually brown with yellow folds on R. grahami and R. hmongorum. The and black spots, sometimes green with black absence of an outer metatarsal tubercle im- spots; legs brown, lightly banded; (10) me- mediately differentiates R. banaorum from R. dian callous pad on ®ngers II, and III to archotaphus and R. chalconota. The pres- proximal tubercle; (11) disks on ®ngers and ence of white eggs differentiates R. banao- toes greatly enlarged (Ͼ2ϫ base of phalan- rum from R. bacboensis (black eggs), Huia ges); (12) feet fully webbed to disks, except nasica, R. andersonii, R. chalconota, R. gra- medial side of IV, where it reaches disk as a hami, R. junlianensis, R. margaretae and R. fringe, lateral fringes on I and V to terminal schmackeri (white eggs with a dark melanic phalanges, webbing brown; (13) subarticular pole). The absence of spinules on the venter tubercles and inner metatarsal tubercle dis- distinguishes R. banaorum from R. ander- 2003 BAIN ET AL.: CASCADE FROG FROM SOUTHEAST ASIA 49 sonii, R. grahami, R. jingdongensis, R. jun- reaches the disk as a fringe; lateral fringe on lianensis, R. margaretae, and R. schmackeri. toes I and V to terminal phalanges; toes long, Rana banaorum further differs from Huia slender, with large, rounded triangular disks; nasica by its larger females (83±99 mm vs. relative pad size I ϭ II ϭ III Ͼ IV Ͼ V, pad 67 mm in H. nasica). It is further differen- length (IV) equals pad width; each pad with tiated from R. sinica by its uncovered, dis- ventral circummarginal grooves; subarticular tinct tympanum (indistinct, covered by skin tubercles prominent and conical; inner meta- in R. sinica), large disks (small in R. sinica) tarsal tubercle ovoid, long; outer metatarsal and its relative ®nger lengths (I Ͻ II Ͻ IV tubercle absent. for R. sinica,IIϽ I Ͻ IV for R. banaorum) Xiphisternum large, deeply notched pos- and T-shaped distal phalanges (rounded in R. teriorly. sinica). Rana leporipes further differs from Skin on dorsum shagreened, with small R. banaorum with its white supratympanic pustules on ¯anks; dorsolateral folds weak; fold (not colored in R. banaorum), webbing small tubercles posteroventral to tympanum; to distal phalanges (webbing to disks in R. prominent granules on ¯anks and around clo- banaorum), and T-shaped distal phalanges aca; cloacal opening unmodi®ed, directed (oblong, somewhat rounded in R. leporipes). posteriorly, at upper level of thighs. DESCRIPTION OF HOLOTYPE: ROM 39944, COLOR IN LIFE (in preservative): Dark an adult female, head width 78% head brown (olive-brown, beige), ¯anks gray with length, length 46% SVL; snout short, acutely yellow spots (white spots); lip-stripe brown rounded in dorsal view, bluntly rounded in anteriorly and creamy white from level of the pro®le, protruding beyond margin of lower eye posteriorly (white); loreal dark brown jaw; eye large, prominent, 50% snout length; (black); tympanum beige with dark brown eyelid broader than interorbital distance. Top central ring; iris golden brown; dorsal limbs of head ¯at; canthus rostralis rounded; loreal grayish brown with black banding; cloacal region concave; lip ¯ared just anterior to or- region dark brown (black); webbing marbled bit; nostril about three-fourths distance from white on dark brown (uniformly brown); eye to tip of snout; supratympanic fold curv- venter creamy white, ventral surfaces of ing posteroventrally from posterior corner of limbs creamy yellow with black mottling. eye to a level above the insertion of arm; SECONDARY SEXUAL CHARACTERS: The ho- tympanum very large, round, distinctly visi- lotype is a gravid female with immaculate ble, separated from eye by distance equal to white eggs (2 mm in diameter). Gravid fe- TMP, 80% of eye length. Choanae ovoid; vo- males have an SVL nearly twice that of merine dentigerous processes prominent, males (mean SVL female, 93 mm; male 50 oblique, posteromedial to choanae, each mm). Male TMP:EYE (0.89) is larger than bearing numerous teeth. Tongue cordiform, females (0.75). Males have velvety nuptial distinctly notched posteriorly, free for ap- pads on the thumb and paired gular pouches proximately two-thirds its length. located at the angle of jaw. Pectoral spines Forearms moderately robust; ®ngers mod- are absent. erately short, slender, hands 23% SVL, rel- MEASUREMENTS OF HOLOTYPE (in mm): ative ®nger lengths II Ͻ I Ͻ IV Ͻ III, ven- SVL 98.0; SNT 13.5; HDL 45.4; HDW 35.6; tromedial callous pad on ®ngers II and III to EYE 6.8; IOD 7.4; TMP 5.4; TEY 4.7; HND proximal tubercle, disks greatly expanded 22.7; FGR 19.4; FPL 3.1; FPW 3.3; TIB (Ͼ2ϫ base of phalanges), relative pad size II 63.7; FTL 80.0; TPL 3.4; TPW 3.4. Ͻ I Ͻ IV Ͻ III, pad length (III) 94% of pad VARIATION OF PARATYPES: The skin on the width, ventral circummarginal grooves pre- dorsum varies from smooth to shagreened sent; terminal phalanges T-shaped; subarti- with small pustules on ¯anks. The lip-stripe cular tubercles conical. Hindlimbs moderate- varies from creamy yellow (white in alcohol) ly robust; tibia length 65% SVL; relative toe throughout its length to brown anteriorly and lengths I Ͻ II Ͻ III Ͻ V Ͻ IV; inner tarsal creamy white from the level of the eye pos- fold absent; feet very large, 82% of SVL, teriorly. The dorsal skin is variable from light fully webbed to terminal phalanges except green to dark brown (olive, brown, beige, or the inner side of IV where the webbing livid blue in alcohol) with or without large 50 AMERICAN MUSEUM NOVITATES NO. 3417 black spots. Variation in all type material is occurrence of three sympatric species of the given in table 15. R. chloronota complex from this region. MEASUREMENTS OF FEMALE PARATYPES (in Some male R. banaorum have indistinct, mi- mm, n ϭ 8, ROM 39899±39901, 39928, croscopic spinules on the dorsal surface of 39929, 39931, 39936, and 39941): SVL 92.7 the leg extending to the feet, forming a Ϯ 5.6 (83.4±98.7); SNT 13.8 Ϯ 1.6 (12.4± ``saw-tooth'' formation along the lateral edge 16.8); HDL 43.8 Ϯ 3.2 (37.5±48.8); HDW of toe (seen in FMNH specimens that are not 30.6 Ϯ 1.5 (26.7±33.4); EYE 6.1 Ϯ 0.7 (4.7± part of the type series). 6.9); IOD 8.2 Ϯ 0.9 (6.8±10.1); TMP 4.5 Ϯ 0.6 (3.5±5.6); TEY 4.5 Ϯ 0.6 (3.5±5.2); Rana megatympanum, new species HND 22.3 Ϯ 2.8 (17.5±25.0); FGR 18.2 Ϯ (Previously referred to as species 7, 2.8 (12.9±20.8); FPL 3.4 Ϯ 0.4 (2.8±4.3); ``Large'') FPW 3.1 Ϯ 0.2 (2.8±3.5); TIB 58.1 Ϯ 3.2 Figures 12O, P, 13G, 14G, H (52.7±63.7); FTL 68.0 Ϯ 7.0 (58.3±78.0) TPL 3.4 Ϯ 0.6 (2.3±4.3); TPW 2.8 Ϯ 0.4 HOLOTYPE: (ROM ®eld no. 12999) ROM (2.4±3.4). 39684, a gravid adult female from Khe Moi MEASUREMENTS OF MALE PARATYPES (in River, approximately 24 km west of Con mm, n ϭ 14, ROM 25084±25086, 25100, Cuong village (by road), Con Cuong District, 25102, 25103, 39912, 39913, 39915, 39916, Nghe An Province, Vietnam (18Њ56Ј30ЉN, 39920±39922, 39924): SVL 50.5 Ϯ 3.7 104Њ48Ј35ЉE) found between 24 and 29 Oc- (42.5±54.6); SNT 7.4 Ϯ 1.5 (3.1±8.7); HDL tober 1994 by I.S. Darevsky, L.A. Lowcock, 26.1 Ϯ 1.4 (24.6±28.1); HDW 17.4 Ϯ 0.6 R.W. Murphy, and N.L. Orlov. The holotype (17.8±18.3); EYE 4.0 Ϯ 0.5 (3.1±5.1); IOD had leg and liver tissue removed shortly after 3.8 Ϯ 0.4 (3.4±4.7); TMP 3.5 Ϯ 0.6 (2.0± it was euthanised. 4.7); TEY 1.7 Ϯ 0.6 (0.9±3.6); HND 14.0 Ϯ PARATYPES: Eight females (ROM 39263, 1.2 (11.8±16.0); FGR 11.1 Ϯ 1.1 (9.2±12.7); 39685±39691) collected with holotype be- FPL 1.9 Ϯ 0.3 (1.4±2.6); FPW 1.6 Ϯ 0.2 tween 24 and 29 October 1994 by I.S. Dar- (1.4±2.0); TIB 30.6 Ϯ 4.6 (24.1±43.6); FTL evsky, L.A. Lowcock, R.W. Murphy, and N.L. 32.6 Ϯ 6.8 (23.3±42.7); TPL 1.9 Ϯ 0.4 (1.3± Orlov. Four males (ROM 39237±39240) from 2.4); TPW 1.5 Ϯ 0.3 (1.1±2.1). Na Hang Nature Reserve, Tuyen Quang Prov- ETYMOLOGY: The speci®c name is a pa- ince, Vietnam (22Њ21Ј54ЉN, 105Њ25Ј40ЉE) tronym for the Ba Na people, an ethnic group found along waterfalls between 26 and 27 living on the Tay Nguyen Plateau (Central May 1996 by A. Lathrop and R.W. Murphy Highlands) of south-central Vietnam where between 193 and 0200 hours. ROM 26398± this species occurs. 26400 Con Cuong District, Nghe An Prov- DISTRIBUTION AND ECOLOGY: Rana banao- ince, Vietnam (18Њ56Ј30ЉN, 104Њ48Ј35ЉE), rum is known only from the Tay Nguyen Pla- collected 5 June 1995 by B. Hubley, A. La- teau of the Central Highlands, Gia Lai Prov- throp, R.W. Murphy, and N.L. Orlov. ince, Vietnam. It inhabits forested montane DIAGNOSIS: Rana megatympanum, a mem- river systems on or near rapids or waterfalls ber of the subgenus Odorrana (sensu Fei et of primary and disturbed second growth. In al., 1990), is characterized by a combination May and June males have tight, leathery gu- of the following attributes: (1) body dorso- lar pouches, suggesting that they are not call- ventrally compressed; (2) SVL means of ing and that the breeding season occurs dur- males 52 mm (48±55 mm), females 100 mm ing some other time. (93±105 mm); (3) vomerine teeth in rows REMARKS: Inger and Chanard (1997) and oblique to choanae; (4) yellow lip-stripe pre- Inger et al. (1999) noted that specimens of sent in males, absent or indistinct in females; Rana chloronota (as R. livida) from An Khe (5) head broad, bluntly rounded in pro®le; have more pronounced dorsolateral folds (6) tympanum round, distinct, TMP:EYE in than elsewhere in Vietnam. These frogs do males enormous (1.20), greater than in fe- not ®t Bourret's (1942) interpretation of R. males (0.51); (7) supratympanic fold weak; graminea as a ``northern variety'' of R. chlo- (8) dorsal skin shagreened, dorsolateral folds ronota. These differences likely re¯ect the present only in males; (9) dorsum olive to 2003 BAIN ET AL.: CASCADE FROG FROM SOUTHEAST ASIA 51 brown sometimes with black spots; ¯anks ferentiate male R. megatympanum from R. marbled yellow and brown-gray; forelimbs andersonii, R. bacboensis, R. chloronota, R. and hindlimbs barred; (10) ventromedial cal- hainanensis, R. hejiangensis, R. morafkai, R. lous pad on ®ngers II, III, and IV to proximal jingdongensis, R. junlianensis, R. kwang- tubercle, fringes on ®ngers II, III, and IV; wuensis, R. livida, R. schmackeri, R. sinica, (11) disks on ®ngers and toes greatly en- and R. tiannensis. The dorsolateral fold of larged (Ͼ2ϫ base of phalanges); (12) feet male R. megatympanum differs from that of fully webbed to toe disk, lateral fringes on I R. daorum, which is composed of minute and V to terminal phalanges, webbing white granules, R. hmongorum and R. gra- brown; (13) subarticular tubercles and an in- hami, whose dorsolateral pustules sometimes ternal metatarsal tubercle distinct, conical; form a dorsolateral fold, and R. banaorum (14) terminal phalanges T-shaped; (15) xi- and R. chalconota, which have folds that are phisternum large, deeply notched posteriorly; distinct and continuous, extending to the (16) males with nuptial pads, paired gular groin. The olive brown coloration with black pouches, pectoral spines absent; (17) eggs spots of R. megatympanum differentiates it white. from H. nasica (olive-brown dorsum, lighter COMPARISONS: Rana megatympanum su- brown laterally) and from R. archotaphus, R. per®cially resembles other Asian cascade ra- chalconota, R. chloronota, R. daorum, R. nids, including Huia nasica, Rana anderson- graminea, R. hejiangensis, R. hmongorum, ii, R. archotaphus, R. bacboensis, R. ban- R. hosii, R. jingdongensis, R. junlianensis, R. aorum, R. chalconota, R. chloronota, R. dao- kwangwuensis, R. leporipes, R. margaretae, rum, R. grahami, R. graminea, R. R. schmackeri, R. sinica (all with green). hainanensis, R. hejiangensis, R. hmongorum, Rana megatympanum lacks an external meta- R. hosii, R. morafkai, R. jingdongensis, R. tarsal tubercle present in R. archotaphus and junlianensis, R. kwangwuensis, R. leporipes, R. chalconota. Absence of ventral spines in R. livida, R. margaretae, R. schmackeri, R. male R. megatympanum separates them from sinica, and R. tiannensis (table 12). It can be R. andersonii, R. grahami, R. jingdongensis, differentiated from all other Odorrana by the R. junlianensis, R. margaretae, and R. enormous sexual dimorphism in tympanum schmackeri. The presence of white eggs dif- size (TMP:EYE 1.20 in males, 0.51 in fe- ferentiates R. megatympanum from H. nasi- males). Male R. megatympanum have a yel- ca, R. andersonii, R. chalconota, R. grahami, low lip-stripe and females have an indistinct R. junlianensis, R. margaretae, R. schmack- or absent yellow lip-stripe, differentiating it eri (white eggs with melanic poles), and R. from H. nasica, R. archotaphus, R. banao- bacboensis (eggs completely melanic). Rana rum, R. chalconota, R. chloronota, R. dao- megatympanum differs from R. sinica by its rum, R. graminea, R. hejiangensis, R. hosii, distinct, uncovered tympanum (indistinct and R. morafkai, R. leporipes, and R. livida (all covered with a layer of skin in R. sinica), its with white lip-stripes); R. andersonii, R. bac- disk size (small in R. sinica), its relative ®n- boensis, R. hainanensis, R. jingdongensis, R. ger lengths (I Ͻ II Ͻ IV for R. sinica,IIϽ margaretae, and R. tiannensis have vertical I Ͻ IV for R. megatympanum), and its T- lip-bars; R. junlianensis has a yellow lip- shaped distal phalanges (rounded in R. sini- stripe with brown lip-bars; R. schmackeri has ca). Rana megatympanum can be distin- no lip-stripe or vertical lip-bars. The broad, guished from R. leporipes by its webbing rounded snout differentiates R. megatympan- (only to the basal end of the distal phalanx um from H. nasica, R. andersonii, R. chal- in R. leporipes, to the disk in R. megatym- conota, and R. schmackeri (obtusely pointed) panum), supratympanic fold (colored white and from R. graminea and R. margaretae in R. leporipes), and its T-shaped distal pha- (depressed). Its gular pouches distinguish R. langes (oblong, somewhat rounded in R. leo- megatympanum from R. andersonii, R. chal- pripes). Rana megatypmanum also differs conota, R. grahami, R. hainanensis, R. from R. hejiangensis by its ®nger formula (II hmongorum, R. hosii, R. jingdongensis, R. Ͻ I Ͻ III Ͻ IV for R. hejiangensis), and by junlianensis, R. kwangwuensis, and R. mar- its large disks (small for R. hejiangensis). garetae. Dorsolateral folds immediately dif- Rana megatympanum closely resembles R. 52 AMERICAN MUSEUM NOVITATES NO. 3417 andersonii, R. hainanensis, R. jingdongensis, disks greatly expanded (Ͼ2ϫ base of phalan- and R. tiannensis. In addition, R. megatym- ges), relative pad size II Ͻ I Ͻ IV Ͻ III, pad panum differs from R. andersonii in that its width (III) 87% of pad length, ventral cir- males are smaller (53±78 mm for males of cummarginal grooves present; terminal pha- R. andersonii), and its ®nger and toe-disks langes T-shaped; subarticular tubercles con- are relatively larger. Rana megatympanum ical. Hindlimbs moderately robust; tibia differs from R. hainanensis in its smaller length 69% of SVL; foot length 53% of SVL (R. hainanensis males 49±62 mm, fe- SVL; relative toe lengths I Ͻ II Ͻ III Ͻ V males 75±122 mm, R. megatympanum males Ͻ IV; inner tarsal fold absent; feet fully 48±55 mm, females 93±105 mm) and its rel- webbed to base of toe pads, lateral fringes on ative ®nger lengths (II Ͻ IV Ͻ I Ͻ III for R. I and V to terminal phalanges; toes long, hainanensis,IIϽ I Ͻ IV Ͻ III for R. me- slender, with large, rounded triangular disks, gatympanum). Rana megatympanum further relative pad size I ϭ II ϭ III Ͼ IV k V, pad differs from R. jingdongensis by its smaller width (IV) 75% of pad length, each with males (R. jingdongensis SVL 62±81 mm) ventral circummarginal grooves; subarticular and skin (R. jingdongensis dorsum scattered tubercles prominent and conical; inner meta- with tubercles and large warts, lips and sides tarsal tubercle ovoid, long; outer metatarsal of heads with white spines, all absent in R. tubercle absent. megatympanum). Rana megatympanum most Xiphisternum large, deeply notched pos- closely resembles R. tiannensis, another large teriorly. brown cascade ranid, but differs from it by Skin on dorsum shagreened, becoming in- having shagreened dorsal skin with small lat- creasingly granular laterally; dorsolateral eral granulations (dorsum of R. tiannensis is folds absent; small tubercles posteroventral rough with large, prominent lateral granula- to tympanum; prominent granules on ¯anks tions), and toe disks are smaller than those and around cloaca; cloacal opening unmod- on ®ngers (the opposite condition of R. tian- i®ed, directed posteriorly, at upper level of nensis). thighs. DESCRIPTION OF HOLOTYPE: ROM 39684, a COLOR IN LIFE (in preservative): Dorsum gravid female, head width 77% of head olive-brown, ¯anks yellow and brown-gray length, length 50% of SVL; snout short, (gray to olive); lip-stripe absent (tympanum acutely rounded in dorsal view, bluntly beige with dark brown center ring); loreal rounded in pro®le, protruding beyond margin brown (black); iris gold; top one-third red, of lower jaw; eye very large, prominent, 73% dorsal limbs brown with black banding of snout length; eyelid broader than interor- (brown); posterior surface of thighs brown bital distance. Top of head ¯at; canthus ros- with black marbling (cloacal region black, tralis rounded; loreal region concave; lip thighs gray with white mottling); webbing ¯ared just anterior to orbit; nostril about marbled white on dark brown (brown on three-fourths distance from eye to tip of white); venter creamy white (creamy yellow snout; supratympanic fold curving posterov- with black mottling). entrally from posterior corner of eye to a lev- SECONDARY SEXUAL CHARACTERS: The el above the insertion of arm; tympanum eggs of the holotype are creamy white and 2 round, distinctly visible, separated from eye mm in diameter. Adult females have SVL ap- by distance equal to TMP, 52% of EYE. Cho- proximately twice that of males. Males have anae ovoid; vomerine dentigerous processes a yellow lip-stripe, and females either lack prominent, oblique, posteromedial to choa- one or have an indistinctly yellow lip. Males nae, each bearing numerous teeth. Tongue also possess a weak dorsolateral fold, where- cordiform, distinctly notched posteriorly, free as females do not. Males have a larger tym- for approximately two-thirds its length. panum than females, velvety nuptial pads ex- Forearms moderately robust; ®ngers mod- tending across the thumb, paired gular erately short, slender, hands 27% of SVL, pouches located at the angle of the jaw, and relative lengths of ®ngers II Ͻ I Ͻ IV Ͻ III, no pectoral spines. lateral fringes on ®nger II, III, and IV, with MEASUREMENTS OF HOLOTYPE (in mm): median callous pads to proximal tubercle; SVL 93.6; SNT 15.0; HDL 46.5; HDW 35.7; 2003 BAIN ET AL.: CASCADE FROG FROM SOUTHEAST ASIA 53

EYE 10.8; IOD 6.8; TMP 5.6; TEY 4.1; KEY TO VIETNAMESE SPECIES IN THE RANA HND 25.7; FGR 19.6; FPL 3.0; FPW 2.6; CHLORONOTA COMPLEX TIB 65.0; FTL 49.7; TPL 3.2; TPW 2.4. The Rana chloronota complex in Vietnam VARIATION OF PARATYPES: Variation in all includes species that are dorsoventrally com- type material is given in table 15. pressed with long legs. Snout±vent length of MEASUREMENTS OF FEMALE PARATYPES (in females is approximately twice that of males mm, n ϭ 10, ROM 26398±26400, 39685± (table 14). The dorsum may be green, brown, 39691): SVL 100.3 Ϯ 4.2 (93.6±105.3); SNT or a combination thereof; dorsum usually 14.8 Ϯ 0.8 (13.8±16.5); HDL 45.1 Ϯ 3.3 smooth, sometimes shagreened, often be- (41.3±47.6); HDW 35.2 Ϯ 0.7 (34.1±35.7); coming more rugose laterally. Tympanum is EYE 10.2 Ϯ 0.7 (9.3±11.6); IOD 8.8 Ϯ 1.0 distinct. Venter is entirely smooth. Legs with (6.8±10.0); TMP 5.3 Ϯ 0.4 (4.6±5.9); TEY transverse bands or mottled with indistinct 4.7 Ϯ 0.3 (4.1±5.0); HND 25.2 Ϯ 2.5 (20.6± bands, posterior surface of thighs marbled 29.7); FGR 21.0 Ϯ 1.0 (19.6±22.6); FPL 3.5 yellow and black. An outer metatarsal tuber- Ϯ 0.6 (2.8±4.4); FPW 3.0 Ϯ 0.5 (2.5±3.8); cle is absent. Disks on ®ngers and toes well TIB 63.0 Ϯ 3.3 (55.8±67.7); FTL 72.0 Ϯ dilated, with ventral circummarginal 12.1 (49.7±88.0) TPL 3.4 Ϯ 1.0 (1.9±5.2); grooves. Webbing is complete or nearly so, TPW 2.8 Ϯ 0.4 (2.2±3.4). extending to most distal tubercle or to base MEASUREMENTS OF MALE PARATYPES (in of toe pad (as a fringe in some species). mm, n ϭ 4, ROM 39237±39240): SVL 52.3 Males have velvety nuptial pads. Ϯ 3.4 (48.6±55.2); SNT 8.7 Ϯ 0.8 (8.3±9.6); VARIABLE CHARACTERS: Lip-stripe is pres- HDL 28.0 Ϯ 1.4 (24.6±27.1); HDW 18.5 Ϯ ent in most species and is usually white. Vo- 0.8 (18.0±19.1); EYE 4.1 Ϯ 0.9 (3.17±4.6); merine teeth, if present, are in rows oblique IOD 5.0 Ϯ 0.5 (4.7±5.6); TMP 4.7 Ϯ 0.3 to the internal choanae. Males usually have (4.3±5.1); TEY 1.7 Ϯ 0.3 (1.5±2.0); HND paired gular pouches. Eggs completely pig- 15.2 Ϯ 0.4 (14.8±15.5); FGR 12.5 Ϯ 0.4 mented or unpigmented. (12.2±13.0); FPL 2.0 Ϯ 0.5 (1.6±2.5); FPW HABITAT: Ubiquitous throughout highland 1.9 Ϯ 0.4 (1.4±2.3); TIB 32.7 Ϯ 0.4 (32.3± waterways of Southeast Asia. Found in and 33.1); FTL 38.3 Ϯ 7.3 (29.9±42.8) TPL 1.8 around water, including the forest ¯oor and Ϯ 0.6 (1.3±2.5); TPW 1.6 Ϯ 0.1 (1.5±1.6). canopy of surrounding forests. Commonly ETYMOLOGY: The speci®c name is a noun found perched on rocks beside or among the in opposition, derived from the Latin pre®x cascades. ``mega'' (meaning very large) and ``tympa- 1. Dorsolateral fold distinct, composed of small num'', in reference to the relatively large gold (white in preservative) granules; no- tympanum of this species. ticeably smaller than other cascade ranids DISTRIBUTION AND ECOLOGY: Rana mega- (males 35 mm, females 55 mm), large white tympanum is known from northern and lateral spot, and vomerine teeth absent .... north-central Vietnam. It occurs in montane ...... R. daorum rivers that vary from shallow and slow mov- ± Dorsolateral fold not as above or granules ab- ing to torrential and deep. It may be found sent ...... 2 2. Indistinct glandular dorsolateral folds present on boulders and logs, both in and around the ...... 3 water as well as in the adjacent forest. Ra- ± Dorsolateral folds absent ...... 4 diographs revealed that large invertebrates 3A. Lip-stripe indistinct yellow or absent in fe- (both insects and crustaceans) form part of males, yellow in males; male tympanum the diet. Vocalizations and tadpoles are un- 120% of eye length, 51% for females; skin known. completely shagreened, brown, and snout± REMARKS: The dorsolateral fold and very vent length notably large (males 52 mm, large tympanum of male R. megatympanum females 100 mm) . . . R. megatympanum potentially make it Bourret's (1942) ``north- 3B. White lip-stripe present in both males and females; dorsum green to olive, shagree- ern form'' of R. chloronota (ϭ R. graminea). ned or incompletely smooth, ¯ank gray; However, R. graminea is bright green above male tympanum 89% eye, females 75%; (Boulenger, 1899) in contrast to the olive- SVL males 50 mm, females 93 mm .... brown dorsum of R. megatympanum...... R. banaorum 54 AMERICAN MUSEUM NOVITATES NO. 3417

3C. Dorsum bright green with or without black patrically with R. chloronota: R. bacboensis, spots, completely smooth; ¯ank brownish; R. hmongorum, R. daorum, and R. megatym- male tympanum 77% eye, females 56%; panum. Two species from the Central High- SVL males 46 mm, females 94 mm .... lands of Vietnam also occur sympatrically ...... R. graminea with R. chloronota: R. banaorum and R. 4. Dorsum completely smooth; dorsum bright green, vomerine teeth present, white lip- morafkai. stripe present, eggs immaculate, and SVL The amphibian diversity in Southeast Asia females 93 mm, males 46 mm ...... is underestimated. Given that three of the ...... R. chloronota new species (R. bacboensis, R. daorum, and ± Dorsum in part smooth and shagreened, or R. hmongorum) are distinct in their gross with heavy granulations ...... 5 morphology and that many areas remain un- 5. Dorsum with heavy granulations; skin heavi- surveyed or poorly explored, we predict that ly granulated laterally and on pelvis, dor- the amphibian diversity in Indochina is sum bright green with black spots, males grossly underestimated; since the review of without gular pouches, SVL females 80 Vietnamese amphibians by Inger et al. mm, males 60 mm ...... R. hmongorum (1999), the number of recorded species has ± Dorsum shagreened or in part smooth . . . 6 increased by more than 50% to 154 species, 6. Black vertical bar on lip; dorsum shagreened, including those described herein (Orlov et brown with some black spots, lip bands al., 2002). Museum specimens have not al- black, extend vertically across mandible, ways been studied with close attention to (white lip-stripe absent), webbing marbled morphological detail. This raises the likeli- white on dark brown to toes, eggs pig- hood that previously undescribed species in mented black, SVL 95 females, 55 mm the R. chloronota complex will be found, males) ...... R. bacboensis both in the wild and in current museum col- ± No black vertical bar on lip; dorsum sha- lections. greened or incompletely smooth, color variable green to brown in daylight, changing At every surveyed location in Vietnam at to brown at night, white lip-stripe present, least two, and sometimes three, species of the webbing brown, eggs immaculate white, Rana chloronota complex occur sympatri- SVL females 88 mm, males 43 mm .... cally. Fixed diagnostic characters differenti- ...... R. morafkai ate these sympatric species. If these criteria are extended to allopatric populations across the vast range of R. chloronota, additional DISCUSSION species are recognizable. This concordant ap- The discovery of a high degree of diver- proach would be bene®cial to the elucidation sity in the R. chloronota complex in a small of species within other problematic species area is astonishing given that the species complexes such as Polypedates leucomystax group has been studied for more than 145 (e.g., Narins et al., 1998; Inger, 1999), P. du- years. The historical confusion regarding the gritei (e.g. Orlov et al., 2001), Limnonectes identi®cation of Rana chloronota is a result blythii (e.g., Emerson, 1998), and L. kuhlii of several factors: the loss of type material, (e.g., Inger, 1999). Furthermore, the com- the disparate localities of sampling areas, and plexity of the alpha taxonomy is exacerbated the presence of multiple cryptic species (both by the absence of phylogenetic data of sympatric and allopatric). Rana chloronota Southeast Asian ranids. inhabits a large portion of its historically recognized range, although the ranges of other IMPLICATIONS FOR CONSERVATION species in the complex appear to be more restricted than R. chloronota. Interestingly, The occurrence of multiple species in the multiple species of the complex are usually Rana chloronota complex has important im- found in sympatry in Vietnam (and presum- plications for conservation. Montane forests ably elsewhere). As many as three species across Southeast Asia are increasingly threat- can be found in the same stream on a given ened by human pressure (Wege et al., 1999). evening. One or two of the following four Rana chloronota was presumed to be rela- species from northern Vietnam occur sym- tively secure from threat when it was consid- 2003 BAIN ET AL.: CASCADE FROG FROM SOUTHEAST ASIA 55 ered to be a single, widespread species. How- acquired under permits to Michael M. Lau. ever, as a complex of species with unknown, Laotian specimens were acquired under per- but presumably smaller ranges, the complex mits to Bryan Stuart by the Ministry of Ag- may be much more susceptible to habitat de- riculture and Forestry, Vientiane. Agriculture gredation and pressures. Other similar spe- Canada issued import permits for frozen tis- cies complexes are equally at risk and they sues and preserved specimens deposited in require urgent study and international collab- the ROM, whereas the United States Fish oration to ensure that they can be identi®ed and Wildlife Service issued permits for fro- in a timely manner. zen tissues and preserved specimens depos- The Rana chloronota complex is poten- ited in the AMNH and FMNH. All collecting tially useful as a biomonitor of forest health. and euthanasia of specimens were performed These frogs are harvested for food by some under approved animal use protocols. ethnic groups in Vietnam (and by Chinese This study is based on work supported by farmers), but are avoided by other groups the Natural Sciences and Engineering Re- (Nguyen, 2000). Local people are aware of search Council (NSERC) of Canada grant their toxic skin; in our experience, the Viet- A3148 to R.W.M.; the National Science namese do not prefer these frogs as food giv- Foundation under grant no. 98±70232 to the en a choice. In contrast, R. chloronota is re- Center for Biodiversity and Conservation at ferred to as ``green chicken'' in southeastern the American Museum of Natural History; Yunnan Province, China. These frogs must the generous assistance of the ROM Sciences be skinned, thoroughly cleaned, and some- Fieldwork Fund, the ROM Future Fund, the times decapitated before being consumed. ROM Foundation, the Department of ROM Cognizant of conservation, Chinese farmers Volunteers, Department of Zoology of the only eat ``adult'' frogs (females) and leave University of Toronto, and Cathay Paci®c ``juveniles'' (males) to sustain the resource Airlines to R.W.M.; a University of Toronto (Murphy and Orlov, personal obs.). Open Masters Fellowship and American Mu- The skin secretions likely make Odorrana seum of Natural History Collections Study unpalatable and possibly even dangerous to Grant to R.H.B.; Deutsche Wissenschaftliche other would-be predators. Snake stomach Gesellshaft Grant (Zoological Museum of contents across Southeast Asia have not in- Hamburg University) for N.L.O. Bryan Stu- cluded R. chloronota and yet they have in- art's collections were made possible by sup- cluded other large ranids (Bourret, 1936; port from the National Geographic Society Pope, 1935). Consequently, if the R. chlo- grant no. 6247±98 to Harold Heatwole. ronota complex experiences low hunting and We thank Eleanor J. Sterling for her con- harvesting pressure in an area where resource tinued support of this project. We are grateful use by humans is otherwise intense, their to Nguyen Van Sang, Pham Duc Tien, Khuat population status could be a useful indicator Dang Long, Nguyen Tien Hiep, and the late of the health of the montane forest. That said, Cao Van Sung of the IEBR for their invalu- almost nothing is known about the natural able assistance with on-site arrangements. history of the complex. Their breeding habits We are further indebted to Nguyen Quang have not been reported. Their feeding habits Truong, Leslie A. Lowcock, Tom Mason, Ra- are poorly known. The composition and fael O. de SaÂ, and Brad Hubley for their source of their skin toxins is unknown. Even valuable assistance in the ®eld. Chen Liqiao their distributions are uncertain. provided access to unpublished data, and Ned Gilmore provided information regarding ACKNOWLEDGMENTS specimen holdings at the Academy of Natu- ral Sciences, Philadelphia. For permission to Collecting and export permits were made study specimens under their care, we thank available through Vietnam's Institute of Harold K. Voris and Robert F. Inger Ecology and Biological Resources, Hanoi, (FMNH), Charles W. Myers and Darrel R. and the Forestry Protection Department, Frost (AMNH), Mark Wilkinson and Barry Ministry of Agriculture and Rural Develop- Clarke (BMNH), and Rainer Guenther ment, Vietnam. Hong Kong specimens were (ZMB). We also thank the AMNH library 56 AMERICAN MUSEUM NOVITATES NO. 3417 staff, particularly Mary DeJong and Amanda Whitehead in the interior of Hainan. Proceed- Bielskas. For invaluable assistance during ings of the Zoological Society of London 1899: museum visits, we thank Linda S. Ford and 956±962, 4 pls. Margaret Arnold (AMNH), Alan Resetar and Boulenger, G.A. 1918. Remarks on the batrachian James B. Ladonski (FMNH), and Frank Til- genera Cornufer, Tschudi, Platymantis, Gthr., Simonantis, g.n., and Staurois, Cope. Annals lack (ZMB). We thank Roman Khalikov for and Magazine of Natural History, ser. 9(1): imaging assistance. We thank Bryan Stuart 372±375. for granting us permission to study and in- Boulenger, G.A. 1920. A monograph of the south clude recent collections from Laos in our Asian, Papuan, Melanesian, and Australasian study. frogs of the genus Rana. Records of the Indian Chunmei Huang, Xiao-Chun Wu, Ho-Ling Museum 20: 1±126. Poon, and Jinzhong Fu provided translations Bourret, R. 1936. Les serpentes de l'Indochine. of Chinese descriptions, and Nasreen Rah- Toulouse: Henri Basuyau & Companie, 2 vols., man and Johan Lindell did the same for Ger- 646 pp. man publications. For reading and comment- Bourret, R. 1942. Les batrachians de l'Indochine. 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APPENDIX 1

SPECIMENS EXAMINED Pac Ban, Tuyen Quang Province: (females) ROM 29526±29530; Con Cuong Region, Nghe An Rana chloronota: Vietnam; Tam Dao, Vinh Phu Province: (females) ROM 26404, 29531±29534, Province: (females) ROM 26411±264416, 39248± 26357, 26358, AMNH A-161248, FMNH 39249, 39313±39315, 39338; (males) ROM 255611; (male) FMNH 255612; Rana daorum:Sa 26433, 39348±39350, 39353±39354, 39357± Pa and vicinity, Lao Cai Province: (females) 39358, 39360, 39361, 39364; Ba Be National ROM 26381, 38500, 38503, 38507, 38512, Park, Bac Kan Province (formerly Cao Bang 38516, 38517, 38526, 38530, 38538; (males) Province): (females) ROM 26348±26356, 26367; ROM 26382±26397, 38501, 38502, 38504± (males) ROM 26360±26366, 26368, 26369, 38506, 38508±38511, 38513±38515, 38518± 39362; Na Hang Nature Reserve, Pac Ban, Tuyen 38525, 38527±38529, 38532±38537, 38539, Quang Province: (females) ROM 39278±39284, 38540, 38542±38543, 38546, 38548±38561; (ju- 39286, 39287; (males) ROM 39755, 39757, veniles and subadults) ROM 38547; Rana hmon- 39759, 39760, 39767, 39774, 39775; Con Cuong gorum: Sa Pa and vicinity, Lao Cai Province: (fe- Region, Nghe An Province: (females) ROM males) ROM 26370±26379, 39867±39873; 26405, 39260±39262, 39264±39271, 39276, AMNH A-161480; (males) ROM 26380, 39235, 39277; (males) ROM 26401±26403, 39272± 39236, 39874±39879, 39890±39894; (juveniles 39275; Gia Lai Province: (females) ROM 26422, and subadults) ROM 39868, 39880±39889, 26424, 26425, 26427, 26428; (males) ROM 39895, 39896; Rana morafkai: Gia Lai Province: 26420; Lang Bien Plateau (female) FMNH 83223; (females) ROM 39930, 39932, 39934, 39947, China; Hong Kong: (female) ROM 39243; 39949; (males) ROM 25094±25097, 25099, (males) ROM 39241, 39242, 39244, 39245; Lao 25101, 25104±25106, 25108±25111, 39904± PDR: FMNH 256493; India; Darjeeling: ROM 39911, 39937; Rana banaorum: Gia Lai Province: 14057, 14058, BMNH 1947.2.28.4, 1947.2.28.6, (females) ROM 39899±39901, 39928, 39929, 1947.2.28.12; Assam: FMNH 72416, 74158; 39931, 39936, 39941, 39944; (males) ROM Rana graminea: China; Fujian Province: AMNH 25084±25086, 25100, 25102, 25103, 39700, A29973±29991, A28543±28545, 28612; Hainan 39702±39705, 39716±39720, 39912, 39913, Island: BMNH 1947.2.27.96, 1947.2.27.97; NLO 39915, 39916, 39920±39922, 39924±39926; (sub- Field Series 26375; Rana livida: Myanmar; Ten- adult) ROM 39942; Rana megatympanum:Na asserim: BMNH 1889.3.25.47, 1889.3.25.48; Hang Nature Reserve, Pac Ban, Tuyen Quang Rana sinica: China; undisclosed: ZMB 9785; Province: (males) ROM 39237±39240; Con Rana bacboensis: Vietnam; Ba Be National Park, Cuong Region, Nghe An Province: (females) Bac Kan Province (formerly Cao Bang Province): ROM 26398±26400, 39263, 39684±39689, (female) ROM 29359; Na Hang Nature Reserve, 39691; (males) ROM 39690. 60 AMERICAN MUSEUM NOVITATES NO. 3417

APPENDIX 2 ISOZYME LOCI ASSAYED AND BUFFER SYSTEMS USED TO RESOLVE THEM Buffer abbreviations are as follows: tris-citrate II, TC 8; tris-citrate III, WTC; amine-citrate (Morpholine), CT 6.3, CT 6.5, CT 6.7; tris-citrate/borate, Plk 8.7; lithium borate, LiOH; tris-borate-EDTA, EBT.

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