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PCR tI a Liner ar PEE UDR a hea Ai a PPP ELLA Kale plage veers er APIO gt . ries sae ay see is Maines orale Perey’ sD PTR et Ck ae eet 1 8 ss ce as WERT OORT pues SAKA SNS A bas AA Pty) PREECE CEN A NG SOURIS ese ¢ Pada va in's eC wee res ‘ v4 : ‘ Slee dv bine eset $9 ; wre ‘3 3 SELECT Ae ERE ROR TOT Fey ¢ ROO ae eee PAP ener Ye TONE KA acne ooh erate iy : Car tiees deg es oa fafa a eee : dev eevts nae aN Va ya PROP nOCeOC OOO O8 1) ANY): aye 4¥ f es we’ RECORDS of the INDIAN MUSEUM Vol. XX. A MONOGRAPH OF THE SOUTH ASIAN, PAPUAN, MELANESIAN AND AUSTRALIAN FROGS OF THE GENUS RANA. By G. A. BOULENGER, LL.D., D.Sc., F.R.S. Ww Calcutta : PUBLISHED BY THE DIRECTOR, ZOOLOGICAL SURVEY OF INDIA. PRINTED AT THE BAPTIST MISSION PRESS, JUNE, 1920. ae i {OS Seas a M MONOGRAPH OF THE SOUTH ASIAN, PAPUAN, MELANESIAN AND AUSTRA- ELAN PROGS OF, THE GENUS TRANS At.) By G. A. BouLENGER, LL.D., D.Sc., F.R.S., etc. (Published by permission of the Trustees of the British Museum). The species of the large genus Rana, now about 195 in num- ber,? have long been in need of a thorough revision. In the following pages I propose to deal with those inhabiting the Indo- Malay Region, or rather Southern Asia, to which I have added the few which extend the range of the genus to the Papuan and Melanesian islands and the extreme north of Australia (Cape York). I have taken as the northern and western limit of the area dealt with a line drawn from Palestine through Northern Persia, the Hindoo-Coosh (Hindu Kush) and Karakorum mountains to and including the Yang-tse-Kiang, a limit answering better the requirements of a survey of the distribution of Tailless Batrachians than those usually adopted for other groups of animals. The collection of the British Museum has greatly increased of iate and affords ample material for the study I have undertaken ; in addition to this I have received much help from Dr. Annandale, both in material and in discussing some knotty points in con- nexion with Indian species. My grateful thanks are due to him. I therefore hope and trust the detailed descriptions I have drawn up, making use of several characters hitherto neglected, will prove of lasting value for the better understanding of the species, and of their inter-relationships. The description of Frogs is much more difficult than that of Reptiles, the naked skin depriv- ing us of the aid derived from the proportions and disposition of the shields and scales and their numerical characters; measure- ments of the different parts of the body are of first importance, and I have therefore prepared tables of very numerous adult or half-grown specimens, in order to convey exact information on this point, and also to show the amount of individual variation, a matter of great importance which is so often passed over. t At Dr. Boulenger's request I have added notes on the habits, geographical range, etc., of species with which I am personally acquainted. These notes are enclosed in square brackets and initialled. NW. Annandale. 2 Nearly twice the number given in my Catalogue of 1882, after elimination of the Madagascar species which are now placed in the genera Mantidactylus, Blgr. and Aglyptodactylus, Blgr. 2 Records of the Indian Museum. [VoL. XX, The time has come to rid ourselves of the empirical methods which have necessarily prevailed so long in zootaxy, and to en- deavour to group species, as far as possible, according to their probable phylogenetic relationships, and for this purpose it is desir- able to fix one’s ideas by establishing a theoretical prototype from which phyletic lines may be drawn up. This I have re- cently attempted to do in a paper! of which an abstract is here given. The Firmisteynia are certainly derived from the Arcifera, as shown by their ontogeny, and among the latter the Cystignathidae agree in so many points with the Ranide that we may legiti- mately assume a direct genetic relationship between them. Dealing first with external characters, I consider the com- plete separation of the outer metatarsals by web as the most primi- tive character, which is not correlated with the degree of develop- ment of the web between the toes. In all the series of allied forms in which the phyletic tendencies are clearly indicated, we can follow the gradual reduction of the membrane; the species with fully webbed toes are, therefore, the least specialized, so far as this character is concerned. The dilatation of the tips of the digits is a specialization culminating in the large scansorial discs of the species adapted to an arboreal life. ; The primitive frog must have had a well-developed tympanum covered by a very thin skin. The reduction of the organ, which may be completely concealed under the thick skin, leads to its suppression (R. plesket). The prominent glands on the back have often a tendency to run together in longitudinal ridges, irregular at first and ulti- mately localized, such as the dorso-lateral fold which characterizes a great number of species. However, this fold may again disap- pear, as some species of Hyloyana seem to show; it is therefore necessary to distinguish, from the other characters taken in con- junction, whether the absence of the fold is primary or secondary. The skeleton affords important indications for judging of the phylogenetic relationships. I have limited myself to the use of characters which may be ascertained without difficulty on spirit specimens. Such are those derived from the skull, which can be examined by lifting up the skin, usually feebly adherent, and the pectoral arch, which can be exposed by making an incision along the breast and removing the muscles. The shape of the terminal phalanges is easily revealed by the dissection of the tip of one of the fingers or toes. I regard as the most typical a skull such as that of R. hexa- dactyla, in which the nasals are large, and in contact with each other and with the frontoparietals, the upper surface of the eth- moid being covered over. From sucha type we may derive on the one hand those forms in which the cranial roof is expanded, and ' Bull. Soc. Zool. France, 1918, p. 111. 1920. ] G. A. BOULENGER: Frogs of the Genus Rana. 3 the bones rugose and more or less confluent with the skin, on the other those in which the nasals and the frontoparietals are reduced. Most species agree with the type of the genus, Rana tem- poraria, in having a strong, horizontal clavicle (so-called precora- coid), and a simple omosternum (episternum). But the omo- sternum may be forked at the base, A-shaped, and this I regard as a specialization; whilst the oblique direction of the clavicle, diverging from the coracoid towards the median line, as in a few African species, is certainly an approximation to the original condition in the Cystignathidae. The African frogs of the subgenera Hildebrandtia and Ptychadena show a high specialization in the reduction of the clavicles. The species with simply pointed terminal phalanges I regard as the most primitive ; a transverse distal expansion, which may be carried so far as to give the bone the shape of a T or a Y, in con- nexion with the scansorial discs, is a departure from the primitive type, and so is the claw-shaped phalanx which is characteristic of Ptvchadena.! Basing our considerations on these theoretical conceptions, we find that Rana hexadactyla is, among Asiatic species, the nearest approach to the ideal prototype: Outer metatarsals separated, fully webbed toes, pointed fingers and toes, distinct tympanum, absence of dorso-lateral glandular fold, condition of the skull. In the shoulder-girdle, with horizontal clavicles and forked omosternum, it is, however, more advanced than some of its African congeners. I therefore start with R. hexadactyla, from which the series headed by R. tigrina and R. grunntens can be directly derived. How other series, corresponding to the subgenera into which the genus is divided, may be derived from Rana, sensu striclo, is expressed in the synopsis given further on. Having explained the principles by which I have been guided in the classification of the species, I need hardly add that their application is not without many difficulties, which I am still unable to overcome, I trust, however, that the present arrangement con- stitutes a marked progress on past attempts. A few words of explanation are necessary concerning the method of taking measurements. The length of the head and of the snout are taken along the axis, and the posterior extremity of the head corresponds to the articulation of the skull with the vertebral column, which can be approximately ascertained, in the flesh, by feeling with the points of the compasses. The width of the head is the greatest width, at the commissures of the jaws. The tympanum,’ of which the greatest diameter is given, is compared with the eye, measured along the side of the head. The limbs are measured fully stretched out, the hind limb from the ! Cf. Boulenger, C.R. Ac. Sci. Paris, CLXV, 1917, p- 987. 2 In a recent paper published in the Records of the Indian Museum (XV, 1918, p. 97), Mr. Baini Prashad proposes to substitute the name ‘tympanic area ’ 4 Records of the Indian Museum. [VoL. XX, middle line of the thigh where it joins the body. By foot is meant the foot without the tarsus; it is measured from the tarso-meta- tarsal articulation. Each finger or toe is measured from the point at which it joins its fellow, and if this is not the same on both sides, as in the case of the fourth toe, from the point most remote from the tip. The length of the first toe is reckoned from the distal extremity of the base of the inner metatarsal tubercle. Genus Rana, L. Tongue extensively free and more or less deeply notched and bifid behind. Vomerine teeth (exceptionally absent). Pupil roundish-subtriangular or horizontal, or rhombic and very contrac- tile.’ Fingers free, toes more or less extensively webbed (excep- tionally nearly free). Omosternum and sternum with a bony style. No intercalary bone between the penultimate and distal phalanges. Divided into 9 subgenera :— I. Clavicles strong, straight or feebly curved, horizontal or directed forward, narrowly separated on the median line. A. Digits not dilated at the end, or with dilatations devoid of horizontal groove. 1. Tympanum present ; squamosal not forming a suture with the maxillary, a. Outer metatarsals separated by web, at least in the distal half. Rana, sensu stricto. Type: R. temporaria, L. 14 Eurasian species, 22 American, 11 African, 45 Indo-Malayan, 1 Papuan. 6. Outer metatarsals united or separated only in their distal extremity ; omosternum usually forked at the base.? Tomopterna, D. et B. Type :. T. delalandii, D. et B. 4 Indian species, 4 African. Burrowing forms, connected with Rana (R. tigrina-limnocharis group) through the Indian R. rufescens, Jerd. 2. Tympanum present ; squamosal forming a suture with the maxillary. a. Outer metatarsals completely united; omosternum entire. Pyxicephalus, Tsch. Monotype: P. adspersus, Tsch., Africa. Burrowing form like the preceding, probably derived from another group of Rana. 6. Outer metatarsals separated in the distal third or fourth; omosternum forked at the base. Aubria, Blgr. Monotype: R. subsigillata, A. Dum., Africa. Probably derived from the &. tigrina group, like Tomopterna, but in a different direction. to ‘tympanum’ as universally used. When we say the tympanum is distinct, we do not convey the idea that it is exposed ; just as we are fully justified in stating that the eye of a frog is distinct even when the transparent lower lid is drawn over it. Mr. Baini Prashad has quoted various authors to show that ‘a great deal of confusion exists regarding the tympanum being a structure distinct from the skin or otherwise.’’ He should have referred to the introduction to my book, ‘The Tailless Batrachians of Europe’ (Ray Society, 1897), in which (p- 11), dealing with the external characters, I have thus described the state of things :— ‘The tympanum, or drum of the ear, is absent in Bombinator and Pelobates. When present it may be concealed under the skin, as in some specimens of Discoglossus, Pelodytes, and Bufo vulgaris, or appear on the temple behind the eye as a round or oval disk covered with thin skin.” ! As in R. corrugata, kuhlii and spinosa. * The character suffers exceptions, as Miss Procter has pointed out to me that the omosternum may be simple in specimens of R.delalandii. Is this to be regarded as a case of reversion to the original condition? It shows at any rate that great systematic importance cannot be attached to the shape of the omo- sternum. 1920.] G. A. BouLENGER: Frogs of the Genus Rana. 5 3. No tympanum ; outer metatarsals separated in the distal third or fourth ; omosternum entire. yoy ~---Nanorana, Gthr. Monotype: N. pleskei, Gthr., Kashmir and Thibet. Con- nected with the R. liebigii group through R. blanfordiz, Blgr. B. Toes, often also fingers, dilated at the end, the dilatation, or disc, bearing a crescentic or horseshoe-shaped horizontal groove. 1. Outer metatarsals united or separated only in the distal third ; omosternum forked at the base. { /09---Discodeles, Blgr. Type: Rk. guppyi, Blgr. 6 Indo-Malayan species, 3 Mela- nesian. Connected with Rana (R. grunniens group); leads to the Indo-Malayan, Papuan, and Melanesian genera Cornufer, Tsch., and Platymantis, Gthr. 2. Outer metatarsals separated at least in the distal half ; omosternum entire. 23---Hylorana, Tsch. Type: R. erythraea, Schleg. 58 South Asian species, 1 African, 5 Papuan and Melanesian. Completely connected with Rana (Ranae typicae) ; leads to the Indo- Malayan genera Staurois, Cope, and Simomantis, Blgr.! II. Clavicles very slender, curved or directed obliquely backwards, widely sepa- rated on the median line. A. Outer metatarsals united ; omosternum entire. Hildebrandtia, Nieden. Type: &. ornata, Peters. 7 species from Africa. Burrowing forms. — B. Outer metatarsals separated ; omosternum forked at the base ; distal pha- langes claw-shaped. a Ptychadena, Blgr. Type: RX. mascareniensis, D.et B. 16 species from Africa. These two last subgenera are probably derived, as divergent series, from a common ancestral type connected with the subgenus Rana. The following diagram expresses the relations between these subgenera, as I conceive them. Ptychadena. Aubria. Hylorana. Discodeles. Nanorana. rey) Vis i Hildebrandtia, —~_ Ue eee 2X ype ae Tomopterna, ——— —— Rana. ————— Pyxicephalus. Subgenus Rana, s. str. SYNOPSIS OF THE SPECIES. I. Dorso-lateral glandular fold absent or, if present, not connected with the supratemporal fold; outer metatarsals separated at least in the distal half. A. Tympanum very distinct, smooth; nasal bones in contact with each other, with two exceptions?; omosternal style forked at the base. 1. Inner metatarsal tubercle small but very prominent, pointed, digitiform ; fingers pointed ; toes webbed to the tips ; male with external vocal sacs. Vomerine teeth in strong oblique series in the adult ; first finger longer than second; toes pointed ; tibia 2 to 24 times as long as broad; throat and belly with granules and small warts... ~ .. | R. hexadactyla, Less. L Cf. Boulenger, Ann.and Mag. N. 7. (9) 1, 1918, p. 372. 2 R, microdisca and palavanensis. 6 Records of the Indian Museum. [Vor, XX Vomerine teeth in small groups; first and second fingers equal; tips of toes swollen or dilated into very small discs; tibia 23 to 33 times as m8 as broad ; throat and belly usually smooth ... Rk. cyanophlyctis, Schn. 2. Inner metatarsal tubercle small or ‘large, blunt or compressed, not digiti- form ; toes 4 to entirely webbed. a. Toes with obtuse or slightly swollen tips; vomerine teeth between the choanae, often extending posteriorly beyond them; male with a more or less developed pad on the inner finger. a. No outer metatarsal tubercle; toes ~ to entirely webbed ; first finger longer than second ; tibia 2} to 3 times as long as broad ; male with external vocal sacs. Toes completely webbed; tibio-tarsal articulation reaching tip of snout ... 5 R. verruculosa, Roux. Toes completely or nearly completely webbed ; tibio- i tarsal articulation not reaching tip of snout ..? R. tigrina, Daud. — Toes incompletely webbed, or with the web deeply 2 notched ; tibio-tarsal articulation not reaching tip of snout * R. cancrivora, Gravh. | A. Usually < a more or less distinct outer metatarsal tubercle. * Toes 3 to 2 webbed; tibia 33 to 4 times as long as broad ; head and back rough with warts of unequal size; male with internal vocal sacs R. verrucosa, Gthr., ** Toes usually not more than 5 webbed, rarely 3; male with ex- ternal vocal sacs. Toes } to 2 webbed); tibia 2} to 3} times as long as broad ; tibio- tarsal articulation not reaching poate tip of snout ; first finger longer than second . 7 R. limnocharis, Wiegm. Toes not quite } webbed ; tibia 3 to 4 times as long a as broad ; tibio- tarsal articulation not reaching tip of snout ; first finger not or but very slightly longer than second me se ac .. R. greenit, Blgr. Toes barely 4 webbed; tibia 4 to 43 times as long as : broad ; tibio-tarsal articulation reaching tip of snout or beyond; first finger longer than second R. brevipalmata, Peters. j. Toes with the tips dilated into distinct discs ; ; ses without or with inter- nal vocal sacs, without enlargement of the fore limb or inner finger, often with tooth-like bony processes in front of lower jaw. a, Vomerine teeth extending behind level of choanae, or entirely behind them. * Tibia 3 to 4 times as long as broad ; nasal bones large and in contact with each other and with the frontoparietals. + Toes $ to entirely webbed ; interorbital region flat. { First finger much longer than second. Tympanum } to % diameter of eye; tibia more than twice in length from snout to vent ; heels not over- lapping ; male without vocal sacs, without tooth-like processes in lower jaw _ OR. grunniens, Daud. Tympanum 3 to § diameter of ¢ eye; tibia 13 to 21 times in length from snout to vent; heels overlapping ; discs of toes very small, male without vocal sacs, with tooth- like precesses in lower jaw... ./R. macrodon, D. et B. Tympanum j to 2 diameter of eye; tibia 2 to 2} times : in length from snout to vent; heels not overlapping ; , Bax discs of toes rather large ; male with vocal sacs, with eS tooth-like processes in lower jaw “RR. magna, Stejn. Tympanum to $ diameter of eye; tibia nearly 2 to 21 times in length from snout to vent; heels ov erlapping ; ; discs of toes very small; male with vocal sacs, with tooth-like processes in. lower jaw .. 2 R. modesta, Blgr. §§ First finger not much longer than second. Tympanum 4% size of eye; first finger as long as or a [v. Kamp. little shorter than second ,,, ae ... AR. microtympanum, 1920. ] G. A. BOULENGER: Frogs of the Genus Rana. 7 Tympanum } to 3 diameter of eye; first finger a little longer than second; tibia 12 to 2} times in length from snout to vent ; heels overlapping ; male without vocal sacs, without tooth-like processes in lower jaw“ R. doriae, Blgr. +4 Toes 3 to nearly entirely webbed; male with the interorbital region very convex, swollen behind, with tooth-like processes in the lower jaw, no vocal sacs, and the tympanum as large as or a little larger than the eye. Tibia at least twice in length from snout to vent ; inner metatarsal tubercle } to 3 length of inner toe; inter- orbital swelling of male rounded behind ... R. macrognathus, Blgr. Tibia at least twice in length from snout to vent ; inner metatarsal tubercle 3 to } length of inner toe; inter- orbital swelling of male followed by a dermal flap .../7 R. pileata, Blgr. Tibia 14 to 2 times in length from snout to vent ; inner metatarsal tubercle about 3 length of inner toe ; back with 8 or 10 glandular longitudinal folds ; interorbital swelling of male foliowed by a knob-like prominence “R. plicatella, Stol. TH Toes 4 to 3 webbed ; tibia 13 to 2 times in length from snout to vent. Tympanum 4} to 2 diameter of eye; inner metatarsal tubercle feebly prominent, 2 to * length of inner toe ; male without vocal sacs, without tooth-like promi- nences in lower jaw ie Ae ... YR, hascheana, Stol. Tympanum & to $ diameter of eye; inner metatarsal tubercle very prominent, 3 to $ length of inner toe; male with vocal sacs and tooth-like prominences in lower jaw ae ae bes .. JOR. limborgii, W. Scl. *“* Tibia 4 to 5 times as long as broad, less than twice in length from snout to vent; toes 3 to 3 webbed; first finger as long as or a little longer than second; nasal bones moderately large and separated from each other. No dorso-lateral fold; male without vocal sacs, with tather small tooth-like prominences in lower jaw ...°/ R. microdisca, Boettg. A dorso-lateral fold; male with vocal sacs, without tooth-like prominences in lower jaw : 8. Vomerine teeth in nearly transverse series on a level with the posterior borders of the choanae ; nostrils close to the tip of the snout ; tympanum 3 diameter of eye ; first finger longer than second; toes nearly entirely webbed ... ae ..27R. narina, Stejn. B. Tympanum hidden or but feebly distinct ; tips of toes dilated into more or less distinct discs; male without vocal sacs, without enlargement of the fore limb, with large tooth-like prominences in front of lower jaw; nasal bones in contact with each other ; omosternal style forked at the base. Tibio-tarsal articulation reaching temple or posterior border of eye; heels not meeting ; tibia 2 to 24 times as long as broad, 24 to 2} times in length from snout to vent; toes entirely webbed; first finger usually a little shorter than second aie en 2) R. corrugata, Peters. Tibio-tarsal articulation reaching temple or eye ; heels meeting or not ; tibia 24 to 3 times.as long as broad, 2 to 23 times in length from snout to vent; toes entirely or nearly entirely webbed; first finger as long as or a little longer than second _... <. Tibio-tarsal articulation reaching beyond eye; heels overlapping ; tibia 24 to 3} times as long as broad, 14 to 2 times in length from snout to vent ; toes 3 to nearly entirely webbed ; first finger as long as or a little longer than second te aS ... oR. laticeps, Blgr./ © C. Tympanum hidden or only moderately distinct ; no tooth-like processes in front of lower jaw ; omosternal style not forked at the base. Z. Tibio-tarsal articulation not reaching beyond eye; no dorso-lateral fold ; R. palavanensis, Blgr. R. kuhlit, D. et B. 8 Records of the Indian Museum. [VoL. XX, toes entirely or nearly entirely webbed ; breeding male with much thickened fore limb and horny spinules on the inner fingers and on the breast. a. Tympanum distinct; a tarsal fold; male with internal vocal sacs. a. Tibio-tarsal articulation reaching eye; tibia 23 to 3} times as long as broad, twice in length from snout to vent. First finger longer than second ; tips of toes not swollen ; ) inner metatarsal tubercle 2 length of inner toe —_...“7 R. feae, Blgr. ). oe ORs First and second fingers equal; tips of toes slightly dilated; inner metatarsal tubercle compressed, sharp- edged ss aa ss . oR. yunnanensis, And. First finger slightly longer than second ; tips of toes swollen ; inner metatarsal tubercle = length of inner toe os oh Mi ...27R. tibetana, Blgr./, 70 B. Tibio-tarsal articulation reaching temple or posterior border of eye; tibia 3 to 3% times as long as broad, 2} to 24 times in length from snout to vent ; first finger as long as or a little shorter than second ; tips of toes obtuse or slightly swollen ; inner metatarsal tubercle 2 to} length of inner toe .. OR. sternosignata, Murr. /\. // b. Tympanum hidden or very indistinct ; first finger longer than second. ‘ Tibio-tarsal articulation reaching shoulder or tympa- num ; tibia 2} timesas long as broad, 2} to 25 times in length from snout to vent: heels not overlapping ; tips of toes not swollen; inner metatarsal tubercle 4 length of inner toe; male with internal vocal sacs ../R. phrynoides, Blgr. / Tibio-tarsal articulation reaching eye, tibia 23 to 3} times as long as broad, about twice in length from snout to vent ; heels overlapping ; tips of toes swollen ; inner metatarsal tubercle 3 length of inner toe; male with internal vocal sacs ee Be .. 32 R. spinosa, David. Tibio-tarsal articulation reaching eye; tibia 13 to nearly 2 times in length from snout to vent; heels overlapping ; tips of toes swollen; inner metatarsal tubercle } length of inner toe; male without vocal SacS, cs a Ba ye 22 R. gammiei, And. 5 ZI. Tibio-tarsal articulation reaching nostril, tip of snout, or beyond¢ heels overlapping; tibia 3 to 4 times as long as broad, 1% to 2 times in length from snout to vent ; first finger as long as or a little shorter than second ; tips of toes swollen. a. Male with much thickened fore limb, with horny spinules on the inner finger and on the breast. Head a little broader than long: tympanum feebly dis- tinct ; toes $ webbed ; dorso-lateral fold distinct only in front ; male without vocal sacs C3 ..~ 1 R. annandalii, Blgr. Head much broader than long; tympanum hidden or very indistinct ; toes entirely webbed ; a dorso-lateral | 6. Male with fore limb not much thickened, without spinules, without vocal sacs; head a little broader than long. Tympanum distinct ; toes entirely webbed; a feeble dorso-lateral fold a ee .. JR. assamensis, W. Sch 20 Tympanum hidden ; toes entirely webbed ; no dorso- lateral fold ee. BS aa .o7 R. vicina, Stol. Tympanum very indistinct; toes 3 to 2 webbed; no dorso-lateral fold wR. blanfordit, Blgr./ II. A glandular dorso-lateral fold,! connected with the supratemporal fold ; tympanum very distinct, smooth. A. Omosternal style not forked at the base. I. Outer metatarsals separated nearly to the base. a Canthus rostralis very obtuse or indistinct; interorbital space much ! Absent or indistinct in some specimens of R. grahami and R. malabarica. fold usually distinct ; male with internal vocal sacs STR. liebigit, Gthr. 7a baw / 1920.] G. A. BOULENGER: Frogs of the Genus Rana. 9 narrower oe the upper eyelid ; tibia 3 to 4 times as long as broad ; nasal bones large and in contact with each other or narrowly separated. Fingers pointed ; toes entirely webbed ; male with in- ternal vocal sacs . er er ../ R. plancyi, Lataste. Fingers obtuse or obtusely pointed ; toes } to entirely webbed ; male with external vocal sacs ... .. YOR. esculenta, L., Fingers obtusely pointed ; toes } webbed; male with external vocal sacs and a very large flat gland on the side of the body ... wee ore ...0 R. pleuraden, Blgr. 6. Canthus rostralis distinct ; interorbital space at least 3 the width of the upper eyelid ; nasal bones small and widely separated from each other ; male with internal vocal sacs. Head as long as broad or a little broader ; tympanum 2 to § diameter of eye ; tibio-tarsal articulation reach- ing tip of snout or a little beyond; tibia 3 to 4 times as long as broad ; toes entirely webbed . ER. grahami, Blgr. Head as long as broad or a little broader ; tympanum 8 6 * diaméter of eye; tibio-tarsal articulation reach- ing nostril, tip of snout, or a little beyond; tibia 4 to 5} times as long as broad; toes } to 4 webbed... R. japonica, Gthr. / Head longer than broad; tympanum % to § diameter of eye; tibio-tarsal articulation reaching beyond tip of snout; tibia 54 to 6 times as long as broad ; toes 3 webbed oe An $3 ... 77R. longicrus, Stejn. If. Outer metatarsals bound together in their basal third orhalf ; tibia 4 to 43 ae times as long as broad ; tibio-tarsal articulation reaching anterior border of eye or between eye and tip of snout ; toes }to webbed ; tympanum 3 to ? diameter of eye. Heels strongly overlapping ; inner metatarsal tubercle compressed, very prominent, 2 to 3 length of inner foeene ros : .. 5 R. lateralis, Blgr./ Heels not overlapping ; inner metatarsal tubercle less than } length of inner toe... as .. 4 R. okinavana, Boettg. B. Omosternal style forked at the base ; outer meta- tarsals united in their basal half; tibia 3} to 4 times as long as broad ; fibio-tarsal articulation teaching tympanum or eye; toes } to } webbed ; tympanum % to once diameter of eye ; male with external vocal sacs and a humeral gland ...47R. malabarica, Tsch. The synopsis is not an artificial key merely for the ready identification of the species; although I trust it will answer that most important purpose, it also aims at conveying the expression of what I conceive to be the true relationships from a phyletic standpoint. These relationships are best expressed by the follow- ing attempt at grouping the species into natural sections :— A. RANAEK HEXADACTYLAE. The most generalized type, from which the three following sections, as well as the genus Oxyglossus. may be conceived to have been derived. R. hexadactyla, R. cyanophlyctis. B. RANAE TIGRINAE. Lead to Tomopterna. R. verruculosa, R. cancrivora, R. tigrina, R. verrucosa, R. lam nocharis, R. greenti, R. brevipalmata. C. RANAE GRUNNIENTES. Lead to Discodeles. R. grunniens, R. macrodon, R. magna, R. modesta, R. micro- tympanum, R. doriae, R. macrognathus, R. pileata, R. plicatella, R. hascheana, R. limborgii, R. microdisca, R. palavanensis, R. narina. 10 Records of the Indian Museum. [VOL. XX, D. RANAE KUHLIANAE. Lead to the genus Nyctibatrachus. R. corrugata, R. kuhlit, R. laticeps. E. RANAE LIEBIGIANAE. Derived from some primitive type, unknown.'! Leads to Nanorana. R. feae, R. yunnanensis, R. tibetana, R. sternosignata, R. spi- nosa, R. phrynoides, R. gammiet, R. liebigit, R. annandalit, R assamensis, R. vicina, R. blanjordit. F. RANAE TYPICAE. The group characteristic of the Eurasian and North American regions, of which the Ranae catesbianae prob- ably represent the most primitive type. Lead to Hylorana. R. plancyt, R. esculenta, R. pleuraden, R. grahami, R. japonica. R. longicrus, R. lateralis, R. okinavana. G. RANAE MALABARICAE. Probably derived from the preced- ing ; a second species (2. galamensis) in Africa. R. malabarica. 1. Rana hexadactyla. Rana grunniens, part., Daud, Hist. Rain. Gren. Crap. p. 65 (1803), and Hist. Rept. VII, p. 127 (1803). Rana hexadactyla, Lesson, in Bélang., Voy. Ind. Or., Rept. p. 331 (1834) ; Tschudi, Class. Batr. pp. 40, 80 (1838) ; Ginth., Cat. hae. Sal. p. 11 (1858), and Rept. Brit. Ind. p. 405 (1864) ; Steind., Novara, Amph. p- 19 (1867); Gunth., Proc. Zool. Soc. 1875, p 568; Bouleng., Cat. Baty. Ecaud. p. 17 (1882), and Faun. I[nd., Rope. Pp. 441 (nie0), Ferguson, Fourn. Bomb. N.H. Soc. XV, i904, p. 500, pl. A, fig. 2 Annand. and Rao, Rec. Ind. Mus. XV, 1918, p. 31, pl. ii, fig, 2. Dactylethra bengalensis, Lesson, /ll. Zool. pl. xlvii (1834). Rana cutipora, Dum. et Bibr., Erp. Gen. VIII, p. 339 (1841). Rana robusta, Blyth, Fourn. As. Soc. Beng. XXIII, 1854, p. 208. 2? Rana neweraellia, Kelaart, Prodr. Faun. Zeyl. 1, p. 292 (1852). Vomerine teeth in strong oblique series from the inner anterior corners of the choanae to a little beyond the level of their pos- terior borders; less developed in the young, in which they form short oblique series or small groups just behind the level of the posterior borders of the choanae. The cornua at the back of the tongue longer and more pointed than usual in the genus, often with a median process between them. Form robust, body not at all constricted at the waist. Head as long as broad or broader than long, much depressed ; snout rounded or obtusely pointed, scarcely projecting beyond the mouth, longer than the eye in the adult; canthus rostralis indis- tinct ;_loreal region very oblique, feebly concave; nostril nearer the end of the snout than the eye; distance between the nostrils equal to or a little greater than the interorbital width, which is much less than that of the upper eyelid; tympanum distinct, 2 to once the diameter of the eye, 1} to 4 times as long as its distance from the latter. ! Of which the Erithrean-Abyssinian R. beccarit, Blgr. is perhaps the nearest surviving representative. 1920. ] G. A. BOULENGER : Frogs of the Genus Rana. 1 Fingers slender, pointed, with a more or less distinct dermal border, first longer than second, third longer than the snout ; sub- articular tubercles small, feebly prominent. Hind limb thick, moderately long, the tibio-tarsal articula- tion reaching the tympanum or the eye, the heels meeting or not when the limbs are folded at right angles to the body; tibia 2 to 2} times as long as broad, 2} to 2} times in length from snout to vent, shorter than the fore limb or than the foot. Toes pointed, with very broad web reaching the tips, the free border almost rectilinear; the fourth toe not extending very much beyond the third and fifth; outer metatarsals separated nearly to the base; subarticular tubercles very small or indistinct; a strong dermal fringe on the outer toe; tarsal fold slightly distinct or absent; a small but very prominent, pointed, digitiform inner metatarsal tubercle,’ its base about } the length of the inner toe; no outer tubercle. Skin smooth above, with small porous warts on the sides; a strong fold from the eye to the shoulder, more or less dis- tinctly connected with its fellow across the head just behind the eyes. Lower parts more or less distinctly granulate, with enlarged porous warts on the throat and under the thighs, and with a curved series of porous warts (sensory organs) along each side of the belly, and another near the flanks. Adult olive or dark brown above, sometimes with a yellow vertebral streak, or with a yellowish lateral band from behind the shoulder to the groin. Young with two or three yellowish bands along the back, in addition to the lateral band. Lower parts white, the thighs usually striped black and white, the black stripes two or three in number. Males with a white external vocal sac on each side, projecting through a slit close to the posterior third of the mandibular ramus, the slit as long as or a little longer than the eye; no other secondary sexual characters.” Nasal bones large, in contact with each other and with the frontoparietals, which are narrow and grooved along the sagittal suture; ethmoid entirely covered over in the adult; zygomatic branch of squamosal very long. Omosternal style forked at the base. Terminal phalanges obtusely pointed. The tadpole has been described and figured by Ferguson and by Annandale and Rao, and I have also examined specimens from Cochin. ‘Tail 2 to 22 times as long as body, acutely pointed, with the dorsal crest extending on the body. Beak white, broadly edged with black; lip forming two lobes on each side; only one series of horny teeth in the upper lip, marginal; two or three on the lower lip, the outer marginal, all, or only the outer, uninter- rupted. 1 Containing 3 or 4 ossicles on end. 2 As observed by Ferguson in &. cyanophlyctis, the male does not clasp the female in the manner our European frogs do, but digs the hands into the axils. 12 Records of the Indian Museum. [Vo.. XX, Measurements in millimetres. T 2 3 4. 5. 6 7 8 a Go Here g ¢ g $ From snout to vent.. 93 130 48 65 115 104 59 60 Head... at 30) 39 18 23 35 33 20 20 Width of head a5 SG 50 18 26 42 41 21 20 Snout .. dou un 14 6 8 12 II 7] 7 Eye 30 ae ts) 10 6 7 10 9 7 7 Interorbital width .. 3 5 2 Basie wea! coy 2 2 Tympanum sit nO Wf. 4 6 9 7 4 4 Fore limb 56. 9B jo 27 37 60 57 32 30 First finger oe litt 16 7 fe) 14 14 7 vi Second finger at LO 13 6 8 II II 6 6 Third finger Boe 3} 17 8 II 16 15 8 9 Fourth finger Bo 6) 13 5 8 II II 6 6 Hind limb Ber ye aieye) 72 98 153 148 84 84 Tibia ant oy) ue 55 21 28 45 43 25 25 Foot BG 0) 46 6L 26 34 SI 2 30 30 Third toe ou. 78) 38 17 20 32 30 18 18 Fourth toe we Ae 53 23 29 45 45 2 26 Fifth toe ac 35 46 19 2 37 37 21 22 I—2. Madras. 3. Malabar. 4—7. Ceylon. 8. S. Ceylon. Habitat.—Southern India and Ceylon. [In life the adult of this species has the back of a bright leaf- green colour, with the mid-dorsal streak, if present, primrose- yellow. In half-grown individual the colours are paler and the conspicuous spots of the young frequently persist. The locality ‘‘Puri’’ given by Mr. Narayan Rao and myself (Rec. Ind. Mus. XV, p. 31) is probably incorrect, being based on somewhat faded specimens of tadpoles from running water (see my note on R. cyanophivetis). On a recent tour through the eastern districts of the Madras Presidency I failed to find R. hexadactyla north of Nellore. This frog usually lives amongst dense aquatic vegetation, in which its green colour conceals it admirably. In Madras it is eaten by Frenchmen, being captured by means of a hook baited with a red rag and pulled on a line through the weeds it frequents. ING Zlclp 2. Rana cyanophlyctis. Rana cyanophlyctis, Schneid., Hist. Amph. I, p. 137 (1799); Peters, Mon. Berl. Ac. 1863, p. 78; Ginth., Rept. Brit. Ind. p. 406 (1864); Steind., Novara, Amph. p. 20 (1867); Stoliczka, Proc. As. Soc. Beng. 1872, p. 102; Blanf., Zool. E. Persia, p. 433 (1876); Bouleng., Cat. Batr. Ecaud. p. 17 (1828), and Faun. Ind., Rept. p. 442 (1890); Murray, Zool. Sind., p. 398 (1884); Anders., Proc. Zool. Soc. 1895, p- 660, pl. xxxviil. fig. 2; Ferguson, Fourn. Bomb. N.H. Soc. XV, 1904, p. 500; Bouleng., Faun. Mal. Pen., Kept. p. 228 (1912) ; Annand. and Rao, Rec. Ind. Mus. XV, 1918, p. 30, fig. Rana bengalensis, Gray, Ind. Zool. I, pl. Ixxxii, fig. 2 (1834); Kelaart, Prodr. Faun. Zeyl. 1, p. 192 (1852). Rana leschenaultit, Dum. et Bibr., Erp. Gen. VIII, p. 342 (1841) ; Cantor, Cat. Mal. Rept. p. 138 (1847); Gtnth., Cat. Bat. Sal. p. 11 (1858). Dicroglossus adolf, Ginth., Proc. Zool. Soc. 1860, p. i158, pl. xxviii, fig. B, and Rept. Brit. Ind. p. 402 (1864). Rana ehrenbergii, Peters, Mon. Berl, Ac. 1863, p.70; Bouleng., Cat. Batr. Ecaud., p. 18. 1920. | G. A. BOULENGER : Frogs of the Genus Rana, 13 Vomerine teeth in small, round or oblique groups on a level with the posterior borders of the choanae or just behind them, equally distinct from each other and from the latter, or closer together. ‘Tongue as in the preceding species. Form less robust than in the preceding, with more distinct waist. Head broader than long, rarely as long as broad, much depressed ; snout rounded or very obtusely pointed, scarcely pro- jecting beyond the mouth, as long as or longer than the eye; canthus rostralis indistinct; loreal region very oblique, feebly con- cave ; nostril equally distant from the eye and from the end of the snout, or a little nearer the former; distance between the nostrils equal to or a little greater or a little less than the inter- orbital width, which is usually much narrower than (not more than half) that of the upper eyelid; tympanum } to nearly once the diameter of the eye, 13 to 5 times as long as its distance from the latter. Fingers more or less pointed, with a more or less distinct dermal border, first and second equal, third longer than the snout; subarticular tubercles small, feebly prominent. Hind limb rather thick, moderately long, the tibio-tarsal articulation reaching the tympanum, the eye, or between the eye and the nostril, the heels meeting or not when the limbs are folded at right angles to the body ; tibia 24 to 3} times as long as broad, 2 to 24 times in length from snout to vent, shorter than the fore limb or than the foot. Toes with the tips swollen or dilated into very small discs, with very broad web reaching the tips, the free border very feebly emarginate; the fourth toe not extending very much beyond the third and fifth; outer metatar- sals separated nearly to the base; subarticular tubercles very small, feebly prominent ; a strong dermal fringe on the outer toe ; tarsal fold feeble or absent ; a small but very prominent, pointed, digitiform inner metatarsal tubercle, its base 1 to 2 the length of the inner toe; no outer tubercle. Skin of upper parts with large pores, smooth or granular, often with small warts which may be tipped with minute asperities ; a strong fold from the eye to the shoulder, often connected with its fellow across the head just behind the eyes. Lower parts smooth. Greyish or olive above, with dark olive round spots or mar- blings; a more or less distinct dark, light-edged band along each flank, and on the front and back of the thighs, often disappearing in the adult ; spots not forming complete cross-bands on the limbs. Lower parts white, marbled spotted, dotted or vermiculated with blackish. Males with a grey or blackish external vocal sac on each side, projecting through a slit close to the posterior half of the mandi- bular ramus, the slit as long as or a little longer than the eye; no other secondary sexual characters. 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