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Cant ates is a maT (aac aageepenell etry stil Cae et Brahe aetna asian Sa eee Sprain eee Duets ares ss | us: RE ates Pes ceriatte Site eadel toes eiecratta ratecot ee Keaeatie eee see ats ae sbeedhtoe tate ff moe sate Sater ayiattorais ai i car Morte itesn iy i ae se ie ay bande hyd ead sy} a _ rly? + i : t; Fp ee ash aT se i ui aie Meal 1 ae Are ea A ec Bc eee RU NUSTAC A ent Sa Mec : Nan eae Oe ei es gh a es nor fag ‘ ‘Y rats tyes ; sah Drea Bates Miah gre oY faa se shh Ginte oy Fara da dive | MAb eg ' 5 po eae i vies Be Ranta Ringe bg skin nat) fae) : ie Site 3 Picyay! ; fat b ir i Seis g ae ; t hasaecaes pees oye foealitet gies : ‘i pee pet pes ie hs aS : eeentie: pat wh eS ip Fatgrtentettony wa Sate ery stented innit tle reyes eyes? aan Desi Hi ‘ iy wo Se SA epee, MEW psiay ev ! ts ait eee) ypu Ny * =n ue $roeecertetas Sree hits: a aaa E Ririsaraatrt eae seat inert mei lereseh s erie sso tacstyaphoibandegs teesps ne tare Sesptltae or : saigset = Littittregsoprtee ee bes vere: ths Mise ent ecen ee ess cae a Reet restate ‘ is Sy et ramon bS6) / | vf it BIOLOGY AMPHIBIANS OF WESTERN CHINA Ch’eng-chao Liu “SUL 47 4051) Mati! { is} “ FIELDIANA: ZOOLOGY MEMOIRS VOLUME 2 Published by CHICAGO NATURAL HISTORY MUSEUM JUNE 15, 1950 FIELDIANA: ZOOLOGY MEMOIRS VOLUME 2 cr \CA Go NATURAL HISTORY MUSEUM ESTAR FOUNDED ey Mag iter’ ere il CHICAGO NATURAL HISTORY MUSEUM CHICAGO, U.S.A. 1950 THE LIBRARY OF THE JUN 2 11950 UNIVERSITY OF H.LINOIS Editors KARL P. SCHMIDT Chief Curator, Department of Zoology LILLIAN A. ROSS Associate Editor of Scientific Publications AMPHIBIANS OF WESTERN CHINA Fieldiana: Zoology Memoirs, Volume 2 Frontispiece — Courtesy of American Geographical Society and Dr. Herold J. Wiens, Yale University THE RUGGED TIBETAN BORDER COUNTRY OF NORTHWESTERN SZECHWAN AMPHIBIANS OF WESTERN CHINA CH’ENG-CHAO LIU PROFESSOR OF BIOLOGY, WEST CHINA UNION UNIVERSITY RESEARCH ASSOCIATE, DIVISION OF AMPHIBIANS AND REPTILES CHICAGO NATURAL HISTORY MUSEUM FIELDIANA: ZOOLOGY MEMOIRS VOLUME 2 Published by CHICAGO NATURAL HISTORY MUSEUM JUNE 15, 1950 PRINTED IN THE UNITED STATES OF AMERICA BY CHICAGO NATURAL HISTORY MUSEUM PRESS on —Ned KY-¢ Ale ie PREFACE I have the pleasant duty of expressing my profound gratitude to the num- erous institutions and to the host of individuals who have aided me in the studies from which the present volume results. At the head of the list must stand West China Union University, where, under President L. K. Chang, I found a new university sponsor and freedom to engage in research, and where Professor M. C. Chang, Head of the Department of Biology, gave me sympathetic and effective support. Second must come my American scientific haven, Chicago Natural History Museum, where I was made welcome by Colonel Clifford C. Gregg, the Director, and by my colleagues in herpetological studies, Messrs. Karl P. Schmidt, Clifford H. Pope, D. Dwight Davis, and Robert F. Inger. I must, of course, thank the United States of America, through whose Division of Cultural Rela- tions in the State Department I received the opportunity to spend a year in the United States in order to bring together my accumulated data into publish- able form. Messrs. Francis J. Colligan and Earl A. Dennis, of the State Depart- ment, smoothed my way throughout my stay in America. I have examined nearly every specimen of Chinese amphibians preserved in American museums and must thank the authorities and my colleagues at the United States National Museum, the Museum of Comparative Zoology, the American Museum of Natural History, the Carnegie Museum, Cornell University, and the Museum of Zoology of the University of Michigan. The notes and manuscripts of the late Leonhard Stejneger, the great pioneer in the critical study of the herpetology of Japan and China, were courteously placed at my disposal by Dr. Doris M. Cochran, Curator of Herpetology at the National Museum. I was received everywhere as a welcome friend and colleague, and I realized that there is to be found in scientific circles the true spirit of the One World of the future. In the course of visits to various museums and universities I have consulted with Dr. E. R. Dunn, Mr. Charles M. Bogert, Mr. M. Graham Netting, and Mr. Arthur Loveridge. I owe them a large share of my debt of gratitude. My former teacher at Yenching University, Dr. Alice M. Boring, has my grateful remembrance for her continued encouragement and help during the war years. In China itself, the Institute of Zoology of the Academia Sinica lent me valued support, and especial thanks are due to Dr. H. W. Chang for bibliographic aid. In western China, my repeated expeditions would have been impossible with- out the faithful help and encouragement of my wife, who aided me in the study 5 6 PREFACE of collected specimens, made measurements and calculations, and typed the manuscripts of all of my preliminary papers. In various years I was assisted in field exploration by H. W. Chang, Y. W. Kao, H. C. Chang, P. L. Luh, C. K. Liu, C. T. Chin, K. F. Cheng, and T. T. Liu. But for Dr. T. L. Yui, of Sichang Hospital, who saved my life when I had typhus, these words could not have been written. I was materially aided by Mr. Y. C. Chiao, of the Department of Communications of the Central Government, in travels in Sikang, Kansu, and Shensi. Mr. L. C. Liu, Director of the Sikang-Kantze Highway Construction, aided me in reaching objectives in Sikang and Kansu. Many salamanders and frogs are intrinsically beautiful creatures, with pleasing colors and harmonious patterns. Their coloration is of great interest for the distinction of the species and still more for its relations to the environ- ment in which the species are found. I was extremely fortunate in finding a capable artist in Chengtu, in the person of Mr. I. S. Wang, young and mostly self-trained, who became my invaluable aid in the field and at home. The illustration of this volume, with the exception of plates 1 and 2 and figures 10 and 82, is his work. I am therefore very proud to present a series of forty-seven of Mr. Wang’s water colors, assembled on eight plates, the publication of which was made possible by a special contribution from Mr. Albert H. Wetten, Trustee of Chicago Natural History Museum. I am profoundly grateful to Mr. Wetten for this colored illustration of my work, and feel certain that its users will be equally indebted to him. The manuscript, as it grew in the laboratory of the Division of Amphibians and Reptiles at Chicago Museum, was a product of co-operative labors. Mr. Karl P. Schmidt, Chief Curator of the Department of Zoology, had aided me in studies on Chinese amphibians in 1934 and 1935, my reception in Chicago being no doubt the more cordial for our relation as successive students of Dr. A. H. Wright of Cornell University. On my return in September, 1946, we found ourselves old friends. Mr. Schmidt’s aid in the organization of the work and in clarifying my English has been generous beyond my power to thank him. Mr. Pope, Curator of the Division of Amphibians and Reptiles, joined Mr. Schmidt in reading the manuscript through successive corrections. Much of the subject matter of the work was discussed jointly by Messrs. Schmidt and Pope and myself, and I had the valuable aid of Mr. Pope’s notes on type specimens preserved in European museums. Mr. Robert F. Inger, Assistant in the Division, was ever helpful, and continued his interest and aid through the preparation of the finally voluminous manuscript for the press. Mr. Davis, Curator of the Division of Vertebrate Anatomy, aided me with translations from the French, as Mr. Schmidt did from the German. Miss Laura Brodie rendered an invaluable service to the manu- script by her careful mounting of the illustrations. Miss Margaret J. Bauer, Mrs. Helen G. Moyer, and Mrs. Howard K. Gloyd shared the tedious work of typing from an extraordinarily interlined and corrected longhand original manuscript, taking a personal interest in the project. PREFACE ff I wish to mention the fact that the library facilities at Chicago Natural History Museum are especially notable for herpetological material, combining the personal libraries of Messrs. Schmidt and Pope with the general library of the Museum. Mr. Carl W. Hintz, the Librarian, and all the members of his staff, were extremely cordial and helpful to me in the borrowing of books. CH’ENG-CHAO LIU November 1, 1948 CONTENTS PAGE ISTO LEEUSTRATIONS 2 or baht hd BAe ee pt oo beer ak et ted ice ene yore tL PMT RODOCTIONG fe 5a iets a0 rte cata ws etal wien nie ai.ch ad en eae et TL iPreservationrands Measurements occ. tes os in oe ty tee nS ewe Ist Orsbocalities. 5< eis eo he ae toe ere ee ey a ee ee ee 0) ZOOLOGICAL EXPLORATIONS IN WESTERN CHINA, 1988-46 ............. 28 Emigration of Teachers and Students to Western China ........... «28 Herpetological Exploration of Mount-Omel 37:02 (32 eee es ee Hsp loravionstateyViuping Sikang-lObOr scr 2e reps en Cae te Coral ee bane eee (| JourneytoNorthwestern:szechwWan, 5 + tase sins bo) Ce co ne ee ee JOurNey: LOMNINgshus Sikan gel GAD. atte Vt ate Sed ees em See Ra ee cee OL Explorations ine mangshusesikang O43 0c a tele er vt ae eerie De Gee 9 OO! DIscoveries*ine Western GRIN a. caw iy ee eee oa a a DISTRIBUTION AND SPECIATION OF AMPHIBIA OF WESTERN CHINA. ........ . 48 INtHOGUCHONG 54 Ween cee eal Raa ate eee aNY Cao) eee: OR Pa tena General Relations between Amphibia of Western China and Other Parts of the Ao) (6 es rea ik a i petites al ME he tral eM As oe eae ee ee Gee Ae General Relations between Amphibia of Western China and Other Parts of China. 45 Distribution of the Amphibia of Western China ............... 46 Discnssionvofebndemic’ Generac 5k ae ee ee Re ne ee ea aS Habitatelisolation: 21.48 port ee see erg Fa oe cia knot Woe etek tee wre eo Provincial Lists of the Species of Amphibia of Western China ......... 58 Adaptations of the Amphibia of Western China ............... 54 Adaptationiin Adults ands bad pol esa Sar tay ca cpa ei ce BP is ee tent ney Oe. EISTIORN SPECIES) Pewee ee ee a ene ge ee ate See Reis ea ee Cee ee OO Cad ata ice tee a eee Os Be co A Wee ee teh ect a Gem A eae OD Salieri tigi rc she oe ay eine Beets etd uve ye mn een hee Ane eae ae eae Lee bl) IVE PERE NOH Ss fogs getaway We Myre ede ct eet! A Sea Wp ee hel alma ene AER hoe ut OO INDE Xie tras ck anes es Re ry ett we ee at ie he Wena N SR SOS | ak oye mt eee: ie = 1 on _ ee, LIST OF ILLUSTRATIONS PLATES Frontispiece. A view of the rugged Tibetan border country of northwestern Szechwan. 4; 2. 3. 10. Courtesy of American Geographical Society and Dr. Herold J. Wiens, Yale University. Salamanders of western China. Figs. 1-3. Batrachuperus tibetanus. Figs. 4,5. Batra- chuperus pinchonit. Salamanders of western China. Figs. 1-3. Tylototriton kweichowensis. Salamanders of western China. Fig. 1. Batrachuperus pinchonii. Fig. 2. Triturus wolterstorfi. Fig. 3. Tylototriton taliangensis. Pelobatid frogs of western China. Fig. 1. Scutiger pingit. Fig. 2. Megophrys omei- montis. Fig. 8. Vuibrissaphora boringii. Fig. 4. Aelurophryne tainingensis. Fig. 5. Megophrys minor. Fig. 6. Aelurophryne glandulata. Pelobatid frogs of western China. Figs. 1, 2. Scutiger schmidti. Fig. 3. Megophrys shapingensis. Fig. 4. Megophrys boulengeri. Fig. 5. Scutiger popei. Frogs of western China. Fig. 1. Rana nigromaculata. Fig. 2. Rana chaochiaoensis. Fig. 3. Rana andersonii. Fig. 4. Rana shuchinae. Fig. 5. Nanorana pleskei. Fig. 6. Rana margaretae. Cascade frogs of western China. Fig. 1. Staurois loloensis. Figs. 2, 6. Staurois mantzorum. Fig. 3. Staurois chunganensis. Fig. 4. Staurois kangtingensis. Fig. 5. Staurois lifanensis. Tree frogs of western China. Figs. 1, 2. Rhacophorus chenfui. Fig. 3. Rhacophorus omeimontis. Figs. 4,5. Rhacophorus bambusicola. Aerial egg-mass of tree frog, Rhacophorus omeimontis. Tadpoles of salientians of western China. Figs. 1, 2. Megophrys minor. Fig. 3. Scutiger rugosa. Fig. 4. Megophrys oshanensis. Fig. 5. Microhyla ornata. Fig. 6. Rana boulengeri. Fig. 7. Aelurophryne glandulata. Fig. 8. Bombina maxima. Fig. 9. Rana adenopleura. Fig. 10. Hyla annectans. Fig. 11. Kaloula macroptica. Fig. 12. Staurois lifanensis. Fig. 18. Scutiger schmidti. Fig. 14. Rhacophorus leucomystax. TEXT FIGURES PAGE Map of western China, showing routes of eleven expeditions ......... 16 Illustrations of measurements used in this paper. A. Adult. B. Foot of adult. Cre Tad Poles ae ett eer ne he ce NE ere Code ee TeO On My eta os 19 Map of western Szechwan and eastern Sikang, te jocalities: mountain ranges, andeal tijudesite ache werk tne eh a wr Medes Mt CE tree agit aay Map of Mount Omei, showing routes, temples, and altitudes. ........ 26 Map of Paohsinghsien (Muping), Sikang, showing main rivers and mountain ranges, and routes and localities of collecting trip made in 1989. ...... 28 11 12 LIST OF ILLUSTRATIONS PAGE Map of northwestern Szechwan, showing collecting routes, localities, and ELA [nh 61 - Ngee aR MIN ee ei at ies Ie pul a rine Se aoe oeiia SMe PS en dN earth 30 Map of Ningshu, Sikang, showing collecting routes, localities, and altitudes. . . 33 Map of Kangshu, Sikang, from Kangting to Kangtze, showing collecting routes, localities:-and:altitudes oq. ee Ge oe a ee ar ee 37 Megalobatrachus davidianus. A. Dorsal view of head. B. Ventral view of head. 73 Megalobatrachus japonicus. A. Dorsal view of head. B. Ventral view of head . 73 . Hynobius shihi. A. Adult. B. Oral cavity. C. Ventral view of hand. D. Finger, showing cornified tip. E. Ventral view of foot .............. 78 ty ED GUACHUDETUS KATISChNIA AOUIC: seit casi sey ne ars ge ha ee cee 88 . Batrachuperus karlschmidti. A. Egg-case. B. Larva just after hatching. C. Fully developedilarvart pags k corte Oe ek tae ot SO eae ed Pome era? 93 . Batrachuperus yenyuanensis. A. Adult female. B. Oral cavity. C. Ventral view or.nand. 1). Ventral:view: of foot nce. Bars eee sea Sues bene nee Ps 99 VE BOMUING: NATIMGs MOAIO 2 Pony Sie, ec ik cess ene CE sees Re ae ee oe 111 . Bombina maxima; male. Ventral view of foots 244.5. ie Sk Sa. 125 PAA CULTODRTUNE: OT CUIDES NAIC se. us one Cede ee ee on eee 126 . Aelurophryne brevipes; tadpole. A. Lateral view. B. Mouth ........ 130 . Aelurophryne tainingensis; male. A. Ventral view of head and thoracic region. B. Ventral view of hand. C. Ventral view of foot ........2.2.2.. 133 Aelurophryne maculata. A. Dorsal view. B. Ventral view of hand. C. Ventral VIC WLOLOOUwe rs 3 ty Sete mae een eerie eth Bye Rages Ae a teem Mae eh ees ee ein 136 Aelurophryne glandulata; male. A. Ventral view of hand. B. Ventral view of foot . 138 Aelurophryne glandulata; male. Ventral view of head and thoracic region. . . . 139 Aelurophryne glandulata; tadpole. A. Ventral view. B. Mouth ....... 142 . Scutiger pingii; male. A. Ventral view of head and thoracic region. B. Oral cavity. C. Ventral view of hand. D. Ventral view of foot ........ 147 . Scutiger pingit; tadpole. A. Lateral view. B. Mouth ........... 149 . Scutiger rugosa. A. Young adult. B. Oral cavity. C. Ventral view of hand. DE" Ventral view Or footacces, dese: oct ee on oh ee a pee re 152 WS CUséser 7 NHOSa MOUtH) OL CAGDOIC =. 5 6.04. Shien ee eke wr dC te ec css 153 Scutiger popei. A. Oral cavity. B. Ventral view of hand. C. Ventral view of LOOM Soin aie, Meanie tae niday oe Sean ile aslo i Reese etn ee ee 155 . Scutiger popei; tadpole. A. Lateral view. B. Mouth ........... 159 Scutiger schmidti. A. Ventral view of head and thoracic region of male. B. Oral cavity. C. Ventral view of hand of male. D. Ventral view of foot... . . 163° Scutiger schmidti. A. Egg-mass. B. Mouthoftadpole .......... 167 Scutiger sp. from Mount Omei; tadpole. A. Lateral view. B. Mouth ... . 172 . Scutiger sp. from Lungtung; tadpole. A. Lateral view. B. Mouth. ..... 176 Vibrissaphora boringiit. A. Oral cavity. B. Ventral view of hand. C. Ventral VICWOL LOOU ss erry arr inc iad delet nnd Siete cy oc Sou ae Steaee, ain Sette ge 178 . Megophrys boulengeri. A. Oral cavity. B. Ventral view of hand. C. Ventral VIEW OL TOOU ws coca whee Sp cA cn Ee eta neat A eee elicit Om he Beg td or 182 LIST OF ILLUSTRATIONS 13 PAGE . Megophrys minor; tadpole. A. Tadpole with lips folded. B. Expanded ‘funnel.’ COBO uti ay tae a aeetoc eRe goat er: Miler nce ec en Wa Wad nang AOR ee ae 188 . Megophrys omeimontis. A. Oral cavity. B. Ventral view of hand. C. Ventral view:or 1oots» «ls ateraleview: of head'h=s 2k he etsceteey B i ce 192 . Megophrys shapingensis. A. Oral cavity. B. Ventral view of hand. C. Ventral VIeWwLOMooty ~)aAbateralcview Ob head. Ae tet et We ents eee eee 195 . Megophrys oshanensis. A. Adult. B. Oral cavity. C. Ventral view of hand. Da Ventral: views 01 1OOGS,s0 a oom hu eeet) oe eee ee ee. ba iar hoes BAe wh 198 Megophrys oshanensts-moutnOl tadpole 4) ss ne ae aes te ee 199 . Bufo raddei. A. Ventral view of hand. B. Ventral view of foot ....... 204 . Bufo raddei. A. Segment of a strand of eggs. B. Tadpole. C. Mouth of AC DOLE Sesser rt et eA ROA cay ae cdi San ern Panne Mee tole ae nt nec oe 205 Bufo tibetanus; adultmale ..... . Sete tae ah eae er ete ee 208 . Bufo tibetanus. A. Ventral view of hata B. Ventral view of foot ..... 209 . Bufo tibetanus; tadpole. A. Lateral view. B. Mouth. ........... yal IBuforousowregnit; ad ultiemalew, ta og Gee ee, ek he Tees ee 215 . Bufo bufo wrighti. A. Ventral view of hand. B. Ventral view of foot. ... . 216 . Bufo bufo wrighti. A. Segment of a strand of eggs. B. Tadpole. C. Mouth of Lad POley ewan br ee een eee Wee ar tO ge ee a ORT PC NOT Ee eee 218 . Bufo bufo gargarizans. A. Ventral view of hand of male. B. Ventral view of LOOT Rae ee ee ee A) a ae iat, BEE Re ot eS es LN Oa 221 . Bufo bufo gargarizans; tadpole. A. Lateral view. B. Mouth ........ 223 . Hyla annectans. A. Adult male. B. Ventral view of hand. C. Ventral view of FOOD ey eee AES eran OS ated Ko hee aes Oars Moers COA oe: ot ees hee P| Hylavannectans=Moutheor tadpoles eawetare tn ecets C8 1 ed eh ees 231 Calluella ocellata. A. Dorsal view. B. Ventral view ............ 233 . Calluella ocellata. A. Ventral view of hand. B. Ventral view of foot .... . 234 Kaloula rugifera. A. Adult male. B. Ventral view of hand. C. Ventral view of foot of males). “Topiottingertipcs ~.- 4 sn ie ; WNT get Pn FPO TSUNG » eae gles 4, 2 = FF MW 2 : Z 2 = > = = TTS 2 Ba = % se Ss A Z iss =: 3 3 % PS Siti, NW, YE. ¥ 1% 3, My XS NG, : Sy gre = : 7 E> S Maui, ys os? - s ay Gi z lid, = Na = 8 3 Ei S K ¥ 2 ® * Si B SS oe oe © ponSING=CiTy 3 Ans = ee AS, aa why ZB, S Amma = way wn Tee 7 SS Wi er" 3, . Al SUNG -NAN-PING 2 Ya ras. et wes ull” oe 3 “yy iT Ss = "7 ® aes Wy Ail gs MILES YY iat We dale 102°s0° Fic. 5. Map of Paohsinghsien (Muping), Sikang, showing main rivers and mountain ranges, and routes and localities of collecting trip made in 1939. 28 LIU: AMPHIBIANS OF WESTERN CHINA 29 After five days on foot we arrived at Muping, where we made our headquarters in the primary school. After four days of collecting around this city, we started for Tupakou. It was only about fifty miles from the city to Tupakou, but it took us three days to get there. This was partly because we made collections on the way, but more because our path led us across swift mountain streams and deep gorges. Sometimes we had to use a tree trunk bridge to cross a river. Our path some- times led over the flimsiest of platforms made of branches of trees, skirting a sheer cliff with rapids more than 100 feet below. Tupakou, selected as our head- quarters, is a small village at the upper end of a valley (altitude 8,000 feet). Our collections from Muping are remarkable especially for primitive sala- manders of the genus Batrachuperus, frogs of the rare Himalayan genus Scutiger, and the life history of the poisonous snake Trimeresurus jerdonii, remarkable for its brilliant coloration and viviparous reproduction. Journey to Northwestern Szechwan Northwestern Szechwan (fig. 6) is another interesting geographical region, with a very complicated topography. There is a high plateau of grass land cut by mountain ranges that run in a north-south direction and are separated by river valleys. The water for the irrigation of the Chengtu plain comes mainly from the high mountains. I made five trips for field studies to these mountains, which were accessible from Chengtu. The fauna and flora are rich, and there are characteristic plants and animals. The people in this part of the country are mostly Chinese, with a small area occupied by another non-Chinese tribe known as the Ch’iang people. The Ch’iang Physically, the Ch’iang are of the yellow race, possibly of the Tibeto-Burman group, with dark eyes, black and straight hair, and a brown complexion. The nose is a trifle narrower than that of the Chinese, and the epicanthic fold is often lacking. These people were originally a nomadic tribe who depended for a livelihood on their flocks of sheep and goats. The Chinese character for Ch’iang consists of the character for sheep or goat and the character for man; this, literally, means ‘‘a people who raise sheep and goats.” The Ch’iang figure considerably in early Chinese writings and go back to the Yin Dynasty (1765-1122 B.c.). The oracle bones contain numerous references to them. They lived to the west of the Yin, who made war upon them and took them captive. In the course of Chinese history, they were a constant menace to the northwestern borders of the empire. Today the Ch’iang inhabit a region along the T’o and the Min rivers, near the Chinese districts of Wenchwanhsien, Weichowhsien, and Lifanhsien. The 103° & A900 Ao- chun-kow r E -MEN-KWAN ; mins HSIN-TIEN-TZE % A660 TUNG-HUA, A 400 fens Pea WS A 4300 ie <0 SALIFAN ws B A ASi00 WEI-CHOW € A 4500 ‘TSA- KU-NAD >> A 6000. Tt. A 4300 te, PAN-CH’IAD A A200 ©) WENCHWAN A 3900 S0-CHA0 A380 ES 3 | QD TA0-Kwan ‘ A370 + z. - SHA-PIN- KWAN / PD HSING-WEN-PING Quand ZE-uNe i Mind A 2900 KWANHSIEN 31° aS > p/o3s0 i] A2x00 * 3 TSING-CHENG-SHANA Ss Fic. 6. Map of northwestern Szechwan, showing collecting routes, localities, and altitudes. 30 LIU: AMPHIBIANS OF WESTERN CHINA 31 rivers are from four to ten thousand feet above the sea level. The surround- ing mountains rise in some places to an altitude of about 15,000 feet. There are extensive loess deposits that make good farm land. Ethnologically, this region is a marginal area of the Tibetan culture, but the people have had constant contacts with the Chinese, who have influenced their lives more than any of the neighboring groups. Most of the Ch’iang live on farms or in fortified villages. They are all farmers. Houses, usually having two or three stories, are built of stones mortared together with clay; the roofs are flat. The first floor is generally used for raising pigs and sheep, goats and cattle, the second floor is for living quarters, and the top floor is the place for worship. The kitchen on the second floor is used not only for cooking and as the dining room but also as a social room. There is some buying and selling in the villages, but the principal market places are in the Chinese cities. Here the Ch’iang bring their agricultural products for sale, and purchase salt, cotton cloth, and other commodities. They lead a very simple life. There are no theatrical performances, few dances, and feasts only during the first moon of the year. Marriage, birth, sickness, death and burial are attended with complicated ceremonies. There are elaborate rituals with sacrifices for worship in the sacred groves and on the housetops, where a sacred white stone is enthroned. This is a triangular quartz stone, symbolic of the purity of the gods, and near it communion with the gods is natural and easy. It is also believed to possess superhuman potency and is sometimes used as a means of protecting the homes from demons. The Ch’iang have a supreme god whom they call Mu Bya Sei, or God of the Sky. He resembles closely the Tien or Ti of early Chinese history. How- ever, like the Chinese, they are not monotheists, for they have numerous gods whom they have probably worshipped for many years, in addition to the Chinese, the Buddhist, and the Christian gods, whom they have learned to worship in the sacred groves. The Ch’iang have no written language, and the chants and rituals have been handed down by word of mouth. Their language, which is dying out, belongs to the broad Tibeto-Burman group, and the preservation of primitive forms, such as case distinction, seems to indicate that the language represents a very old phase of this group. Journey to Ningshu, Sikang, 1942 The southeastern corner of Sikang, originally a part of Szechwan, is known as Ningshu (fig. 7) and is a distinct political region. It is inhabited by the Lolo. The topography of the Lolo country is very characteristic, with high mountain ranges running in a north-south direction, cut by deep valleys. The Lolo permit few strangers to cross their borders, and the flora and fauna are consequently very little known. Through the help of the Minister of Educa- 32 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 tion, and the local government of Sichang, four members of the staff of West China Union University went to the Lolo country, starting in mid-April, 1942. Sichang, the center for our work, is the largest city in Ningshu, and is the political, economic, and educational center of that part of the country. Through the arrangements made by the local government and the kind assistance of friends in a hospital in Sichang, my assistant and I were able to start from Sichang for Chaochiao, in the heart of the Lolo section, on May 1, 1942. Dr. Yii (a medical doctor) and two nurses from the hospital in Sichang accompanied us. We had horses to carry our supplies of food, baggage, and drugs, together with gifts, such as salt, cloth, needles, and thread, for the Lolo people. The distance from Sichang to Chaochiao is only about 120 miles, but it took us six days to reach the city. On the way, we got a good collection of amphibians. The third evening we stayed in a headman’s house at Lolokou. The old man intended to be very kind to us. He served us a chicken dinner, but we could not appreciate his kindness. The chicken was hammered to death and part of its feathers were removed; it was then drawn, and the intestines were crudely cleaned; thereupon it was chopped into pieces and put in a big cauldron of cold water, which was then brought to a boil. When the chicken was half done, some salt was added and it was ready to be served to the guest. Out of courtesy, we had to eat our dinner with smiling faces, but once is enough in my life. At the same time, we prepared some of our own rice to serve them. We spent two evenings in Lolo houses. After six days of walking and insufficient food all of us were very anxious to get to the city, where we expected to get a rest and have some good meals. As we approached, we saw the beautiful wall and gates. We hurried forward, but we were greatly disappointed and greatly surprised when we reached the south gate. There were only three houses in the town, one for the magistrate and his family, one for the army, and one for the district National party. The only tree in the city stands in front of the magistrate’s house. The magistrate, the only inhabitant of the city, welcomed us, and gave us a dinner of rice, cooked dried meat, and fried soy beans, which seemed extremely good to starving people. The empty city was the result of fighting between the Chinese and the Lolo in 1936. All of the Chinese in the city were either killed or taken as slaves by the Lolo, and the houses were burned. After that time, the magistrate could not come to Chaochiao, so he kept his office in Sichang, about 120 miles away. The present magistrate had fought his way through with his soldiers the previous winter and the Lolo near the city had run away, leaving empty houses on the hillsides. The next day, we started our work around the city, and collected specimens of a new species of wood frog. In the afternoon I felt very ill, and had a high temperature. The next day I was unconscious, and my friends decided to take me back to Sichang immediately. Dr. Yii and the magistrate had two poles cut from small trees, and ropes were put between the poles so that two soldiers Fic. 7. Map of Ningshu, Sikang, showing collecting routes, localities, and altitudes. 33 34 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 could carry me back to the hospital in Sichang. It took five days, during which time I was for the most part unconscious. At Sichang Hospital, Dr. Yii found that I had typhus. It took me about fifty days to recover, and I was fortunate that I did not lose my life. The later investigations in this part of the country were carried out by my assistants, Mr. Chang and Mr. Kao, as I could not go to the field. In their brief collecting, eight new species of amphibians were discovered, and the life histories of two rare frogs were completed. The Lolo country is really virgin territory for the studies on its flora and fauna, but it is still much too dangerous for scientists. The Lolo usually take other people for slaves, and once you have been caught by them there is little chance of escape. I may give a very brief account of this little-known people. The Lolo The Lolo dwell in the mountainous regions of southeastern Sikang and southwestern Szechwan. This section of territory is the Taliangshan, which lies east of the Yalung River and extends south of Tatsienlu (Kangting). The land covers an area of nine or ten thousand square miles, with a population of about 500,000 people. During the Chou Dynasty (1122-221 B.c.) this people was known as the Lu. According to the Shuching, they helped Emperor Wuwang in his expedition against the Shangs in eastern China. Today the Chinese do use the term “‘Yichia”’ but the great majority use the term ‘‘Manchia,” when referring to the people of this tribe. The name Lolo is most frequently used by foreign investigators, and by a very small minority of Chinese and No Su themselves. When referring to themselves, they generally prefer the terms “No Su,” “Ijia” or ‘‘Heikut’ou.” In the course of centuries, some Lolo have emigrated to Yunnan and have acquired different names. Along the Red River they are known as the Woli; along the Salween, the Liso; between the Salween and the Mekong, the Lohei; and on the southernmost border of the province, the Ak’a. Linguistically they belong to the Tibeto-Burman family. They have a written language, which is used mainly by the priests or magicians for the purpose of performing rituals and divination. Villages are perched on the mountain sides in what seem to be precarious positions. The house of a family is always a unit by itself, but sometimes several houses are built side by side to form a hamlet. The house is built of wood, sup- ported by poles, walled with stamped earth or boulders, and roofed with planks or the bark of trees. Stones are placed in rows on the roof so that the structure will not be shaken by the wind. Around the house is a square enclosure sur- rounded by earthen walls or a bamboo or wood fence. In one corner of the enclosure stands a watch tower of two or three stories, in which are guns and other weapons that are used for the protection of the house from the raids of other families. LIU: AMPHIBIANS OF WESTERN CHINA 35 Inside the house the head of the family lives in the left room, which is furnished with a small wooden bed. His wife and children all sleep on the wooden or earth floor. The fireplace is situated in the middle room, where the people gather to have their meals. The room on the right is for animals and for the shelter of the slaves. The men wear a rather picturesque armor, made of untanned leather deco- rated with designs in red, black and yellow. The upper half provides protection for chest and back, and to this is attached a skirt made of plaited strips of leather. Worn with the armor there is a large leather gauntlet for the sword arm, and a small leather wristlet that protects the wrist when drawing the bow. The women favor bodices and long, pleated skirts that come to the heels and swing with the rhythm of the wearer as she marches along. Neither men nor women wear shoes or sandals. The men wear long caps, which are referred to as their “heavenly horns,” or the hair may be drawn into a forwardly directed “horn.” The women wear a cloth on the head. The Lolo are mainly divided into two classes, the Black Lolo and the White Lolo. The Black Lolo are also called the Black Bones, and rule over the White Bones, who are composed of five classes of slaves. The children of the Black Bones do not learn any occupation except hunting for recreation. Their daughters are often trained to manage the household affairs, and hold a high position in the family. The children of the White Bones are born to be slaves and are the property of their father’s overlord. They do all the manual labor and work in the fields. The Lolo grow corn, their staple food, on the mountain slopes. Be- sides corn, the principal crops are wheat, rice, potatoes, beans, and turnips. Their principal domestic animals are chickens and ducks, and pigs, sheep, cattle, and ponies. The household unit consists of the overlord’s family and his slaves, whom he considers as a form of property to be bought and sold at will. His family is patrilineal and related to other families to form a clan, living in a definite area of the country with one or more headmen. Marriage is arranged by the parents and the middlemen. The “‘bridal money,” in the form of silver, cattle, weapons, and slaves, is paid in installments. On the wedding day the ceremonies are elaborate. Gifts are exchanged, and there are contests between parties of both families. The marriage system is clan exogamy but class endogamy. Polygamy is known to be practiced. The husband must ask his wife’s permission to take concubines. When the husband dies the wife must marry a brother or nephew, or even the father or the uncle, which is a form of levirate. The class system cuts across the organization of the family and the clan. The relations of one family with another are usually very unfriendly, and feuds are common among them. The No Su are proud of raiding and plundering in their feuds, which are carried on indefinitely. It is these feuds that keep up the warlike spirit among the No Su. 36 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 Explorations in Kangshu, Sikang, 1943 Kangshu is the vast western section of Sikang Province, originally known as East Tibet. My exploration was limited to the northeastern part of Kangshu, from Kangting, the capital of Sikang, to Kantze (fig. 8). Transportation is difficult, and indeed any collecting trip in Sikang is not without hardship. On my travels in 1948 only a collector and one assistant went with me. Early in the morning of June 27, we started our long journey from Chengtu. For six days we sat on top of a loaded truck, passed the dangerous Erlong Mountain, and, with eight days of additional stopovers at various places, arrived at Kang- ting on July 8. Kangting, well known as Tatsienlu to the Chinese as well as to foreigners, was the center for our preparations for a trip to the high plateau and high mountain country inhabited by the Tibetans, who are quite different from the Chinese in language and customs. It took us fifteen days to prepare ourselves for the adventure to a new country. The most essential things to take with us were food, including rice, wheat flour, salt, sugar, smoked meat, and lard; heavy bedding; warm clothes; and heavy shoes. As we could not hire a local interpreter we tried to learn a few necessary phrases and sentences of Tibetan so that we could get along with them. I wrote down the sounds of phrases of the Tibetan language, in place of the actual Tibetan words, in my notebook. With these words and phrases we managed to make our expedition without an interpreter. Another problem we had to solve for our round trip was the means of transportation. Through the | help of the local government in Kangting, we got three horses and six yaks for carrying our food stuffs, baggage, and collecting equipment; but we had to change our animals at different stations. We started our journey on the morning of July 23. Four Tibetans looked after the horses and yaks and served as our guides. We traveled very slowly in order to stop at different places to collect specimens and make observations. The round trip from Kangting to Kantze, a distance of about 600 miles, took us 388 days. We stopped overnight in mon- asteries, the houses of peasants, and the tents of nomads, as well as in the houses of magistrates of districts, and in schoolhouses. In this way we learned much about conditions among the Tibetans in Sikang. As I am not an ethnologist, and our purpose in going there was especially for the study of amphibians, my account of the Tibetans is necessarily very general, and mostly limited to our own experiences. The Tibetans The Tibetans are a mixture of many races, with Mongol stock predominating. Their origin is not known. Their language is classified as of the Tibeto-Burman family. Very little is known about their early history. Originally there were three basic factors in their social organization: the tribe, the kingdom, and the lamasery. Any one of these may form a social unit; the nomads, for example, are organized into tribes; the greater part of the country is divided into upmys ~ U74? - UL a SUNG-LIN-KDU-PASS A 15000 YAKIANG HSIN =TU-CHIAQ! TAUCH'ENG Fic. 8. Map of Kangshu, Sikang, from Kangting to Kangtze, showing collecting routes, localities, and altitudes. 37 38 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 kingdoms; and the lamasery may, like the western parish, be the dominating factor in a certain area. The lamasery, in fact, plays much the same part in Tibetan life as did the monastery in mediaeval Europe, often forming the chief center of social, economic, religious, and cultural life. It is also a political center. As the Tibetans in Sikang Province are now under the control of the provincial government, the kingdom has been disappearing, but the lamaseries and the tribes remain unchanged. The lamasery retains its social and political influence in Sikang. In addition to the lamas in the lamaseries of Sikang, the Tibetans or Kang can be classified into three categories: the farmers, the nomads, and the traders. The great majority of the Tibetans are engaged in farming and in cattle- and sheep-raising, but the herdsmen live a life so different from that of the agricultural peasants that they form a separate group. The Kangshu region of Sikang is one of the poorest countries in the world; the great elevation, the rigorous climate, the brief summer and the lack of men to till the ground make the peasant’s life a hard one. The land cannot be plowed till May, when the staple crops of barley and peas are sown. Irish potatoes, turnips and a few other vegetables can be grown at the lower levels in Sikang, at altitudes ranging from nine to ten thousand feet. On the first day of plowing, a religious service is held to invoke the blessing of the gods and their protection of the land from hail, the farmers’ worst enemy. The Tibetans practice crop rotation, according to which the land must lie fallow approximately one year in every three. The harvest is gathered in September, though at the higher altitudes the crops may never ripen because of the early frosts; crops harvested before they ripen can only be used as fodder. Generally the men do the plowing and the women sow the seed, milk the cattle and make butter, while both sexes do the weeding, irrigation, reaping, threshing, and winnowing. During harvest time, the peasants work eighteen hours a day, and the women bring their food to the fields. Dotted all over the plains or hillsides, wherever there is grazing, may be seen the black hair tents of the nomad herdsmen, who may be considered the original inhabitants of Tibet and are probably the purest specimens of the race. The natural grazing grounds (there are no other), in which the wild pasture is coarse and sparse, is spread over the vast area of Kangshu, and consequently the herds must be continually on the move as the grass is consumed. For this reason, different tribes have their own areas to feed their herds, and may come into conflict. The herds consist mainly of yaks and sheep with a few horses and a few goats. No matter where they go, they always take their dogs to watch the herds, especially during the evening. These dogs are known as “Mango,” which means ‘‘wild people’s dog.”” They are very large, with especially large heads and big feet. Each dog has the value of several cattle. A nomad values his dog as much as his own life. LIU: AMPHIBIANS OF WESTERN CHINA 39 The yak (Poéphagus grunniens) is a heavily built ox with long coarse hair that is usually black, but may be gray or even white; it is hardy and slow, and is the chief means of transportation in Kangshu. Economically it is the nomad’s chief asset, for it carries his home on its back, its hair is used to make tents and coarse cloth, its hide is exported for leather, and its meat forms the chief diet of the tribe. The female gives excellent milk, which produces cheese and the white butter that plays such a large part in Tibetan life. The rectangular tent, which the nomad carries with him in his endless search for fresh pasture, is made of yak hair. When he pitches his tent, he stacks blocks of turf all round the inside, to protect him from the piercing wind and to form a shelf on which to put his meager possessions; these consist of the barest es- sentials—a churn for making butter and buttered tea, a teapot, a few cooking vessels of iron or clay, and two or three large pans used in boiling milk to make cheese. In the center of the tent is a large stove made of mud and stones; dry yak-dung is the main fuel, as the plateau is treeless. Stacked around the walls are the rugs, saddles and leather bags containing food, so arranged that the family can lie down between them to sleep. The nomads may carry a little “tsamba”’ (barley powder) with them, but it is an expensive food for them as they grow no crops of their own, and must buy from the agricultural peasants; their diet consists almost entirely of meat, butter, and cheese. Some of the nomads may plunder their fellow tribesmen, or they may prey upon travellers, even attacking large caravans. The traders form a small middle class that is now increasing greatly in importance, as foreign trade is taking a more important part in Tibetan life. These traders may be independent, or they may be the agents of the big mon- asteries or of government officials, who do not hesitate to use their official position for trade purposes. The Tibetans are born traders, willing at any time to bargain for their neighbors’ possessions in order to sell them again at a profit. Monasteries as well as the city markets are the places for trading. Once a year, the herdsmen come down from their pastures and trade their wool and skins for household articles; they seldom use money, preferring the system of barter. Sometimes the traders may go to the pastures to exchange goods with the nomads. The main diet of the Tibetans consists of yak meat, mutton, barley flour, butter, cheese, and tea. The wealthy, or those who live at lower altitudes, may supplement this with green vegetables and fruit. The cattle are slaughtered at the end of the summer, after they have had the benefit of the green pasture, and the carcasses are stored, without any curing, throughout the winter. Sometimes the meat is cut into strips and dried. The chief grain used for food is barley. This is eaten usually in the form of ‘‘tsamba,” which is prepared in the following way: first the grain is washed, then it is roasted in an iron pan, then ground in a water-mill or a small hand mill, to make it into flour. The flour is then mixed with butter and tea, or just water, in a wooden bowl, kneaded into a roll with the fingers, and then eaten. 40 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 The Tibetans drink even more tea than the Chinese or the English. The tea leaves consist of old leaves of various kinds of tea shrubs, together with twigs and dirt. These leaves and twigs are pressed together to form “‘tea-bricks,”’ which are made solely for the Tibetans. Their way of preparing the tea is unique. They take from the tea-brick a handful of leaves and put them into a cauldron of cold water, adding a small quantity of soda; the mixture is then boiled. When traveling, they drink the tea from a wooden bowl, adding a little butter. In their tents or homes, the boiled tea is poured through a strainer into a wooden churn, in which it is vigorously churned, usually with the addition of butter and a little salt. It is then poured into an earthen-ware or copper teapot, from which it is served. This tea is really more like soup than tea, but it makes a warm and nourishing drink. Huge quantities of it are consumed in the mon- asteries, especially during festivals. Besides tea, the Tibetans drink milk, barley beer, and spirits. The uses of butter are manifold; in the household it is used in cooking and is rubbed on the skin of children to keep out the cold; it is mixed with “‘tsamba”’ and taken in tea; it is used as hair-dressing by the ladies; and it provides the fuel for lamps. The Tibetans also use butter as money to pay their taxes to the landlords and monasteries. Monasteries consume large amounts of butter for the rows upon rows of little copper lamps that are kept burning constantly on their altars. Butter is also made into images and decorative forms at the time of their festival, which is held at the end of February each year. Basically, the dress for Tibetan men and women of all classes is the same, consisting of a long, wide gown; there are, however, strict rules governing the material, quality, and color of the robes worn by different classes. The poor people have but one robe, which is made of coarse homespun and is gathered at the waist with a girdle, to give greater freedom of movement. In the pocket formed in front in the fold of the robe, the peasant will store away food, meat, butter, snuff, gunpowder, his wooden bowl, and anything else he may need during the daytime. The herdsmen wear a sheepskin robe all the year round, with the hair on the inside, and a skin or fur cap. The peasant wears rough felt boots with rawhide soles. Only the nobles may wear silk in public. The middle-class Tibetans may wear silk, but are forbidden to appear in public in such dress. The lamas wear red woollen robes. The women wear a great variety of ornaments—gold and silver rings in the hair, ear-rings of different sorts, and necklaces of beads or even jade. Men always carry a sword in a beautiful scabbard that is given by the father at the time of birth. Both sexes wear a charm-box, a small case of chased gold or silver, often studded with precious stones and varying in shape from a small cylinder to a box about five inches square. The charm-box contains a talisman, usually consisting of an image and an inscription, which must be written by a lama. The image may be the “Lord of Mercy” or more often the ‘“‘Lady Deliverer,’’ who is believed to cure the sick. The charm-box is worn for protection against accident, various kinds of sickness, and evil spirits. To the Tibetan his charm- LIU: AMPHIBIANS OF WESTERN CHINA 41 box is more than an ornament; it is regarded as essential to his safety and health. “Bon” (commonly known as Buddhism) is the chief religion of the Tibetans, and one third of the men are lamas. The monastic system has attained such huge proportions probably because the monasteries are the centers of culture and upon them has fallen the task of providing for the people’s education. At the present time there are five sections of lamas, each section wearing a different color of robe—yellow, red, white, black and flowered. Yellow Bon is the most influential one. Tibetans have no less than five different ways to dispose of their dead. The commonest is called ‘‘chagoppo” and consists in feeding the corpse to the vultures. At the time of death, a lama magician administers the last rites, which include the reading of passages from the book of the dead, as a kind of guide to the disembodied spirit in its wanderings. When the dying man has ceased to breathe, the lama, sitting at the head of the body, swiftly plucks a few hairs from the scalp, uttering the magic word ‘“‘Hik”’ to release the soul from the body through the tiny hole that is believed to have been made in the skull. The corpse is then placed in a sitting position in a corner of the room and the relatives are invited to a feast, at which a good deal of wine is drunk, a little of the food and drink being offered to the corpse. After some days—the time is determined by the sorcerer—the body is carried out to a hillside where, to the sound of the chanting of lamas, it is cut into pieces. The flesh is separated from the bones and is fed to the vultures and magpies that sit patiently waiting a few yards away. The bones are crushed with a heavy stone and are also given to the birds. Meanwhile, the service continues at the home of the deceased. If the vultures do not eat the flesh of the corpse, or do not finish it, the dead person is thought to have been in sin, and his soul to be unfit for heaven. The next most popular form of disposal of the dead is by cremation, but on account of the difficulty of obtaining fuel this method is confined mostly to the religious dignitaries. Water burial is a third method, the corpse being thrown into a river; but this is rarely done, except in the case of convicted criminals. Burial in a grave is limited to those who have died of an infectious disease such as smallpox. A fifth method is reserved for high lamas and living buddhas. The body is placed in a large box in a sitting position, salt is packed around it, and the box is closed and sealed. For three months the box rests in the temple, until the corpse has become completely dehydrated and the skin is hard and shrunken. It is then taken out, built up with clay mixed with wood fibers to represent the features of the lama, and gilded. Then a shrine is built and the mummy takes its place among the sacred relics of the monastery. The salt in which it was preserved is sold as a medicine to the people. Discoveries in Western China The amphibians form a small group of vertebrates. In China, only 88 species were recognized by Pope and Boring in 1939, and they list only 25 species 42 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 for western China. After careful study of my amphibian collections, in addition to six species newly recorded, I found that I had some twenty new species of amphibians, one of which represents a remarkable new genus. The amphibian fauna of Szechwan and Sikang is decidedly rich in unusual forms. The high plateau of central Sikang and the mountains of eastern Sikang and western Szechwan are the centers of speciation for the primitive pelobatid frogs (Aelurophryne, Scutiger, Megophrys, and Vibrissaphora). The giant sala- mander, Megalobatrachus, of western China is closely related to the large hell- bender, Cryptobranchus, of eastern North America; the fire-bellied toad, Bombina, of Sikang is related to European species; and the tree frog, Rhacophorus leuco- mystax, is a common tropical form of southeastern Asia. The Siberian toad, Bufo raddei, is a Palearctic species that reaches northern Szechwan and Sikang. Therefore, Szechwan and Sikang are in a transitional zone from the Palearctic in the north to the Oriental in the south; and ecologically their range is from the warm temperate lowland of the east to the high cold plateau of the west. The mountain ranges running in a north-south direction between these two provinces afford the greatest variety of topographic and ecological conditions. An outstanding element of the amphibian fauna of western China is Aeluro- phryne, which is found only on the high plateau; originally there was only one species known, A. mammata, but now we know no less than five species. Frogs adapted to mountain streams are another special group of western Chinese Amphibia. Tadpoles of the genus Stawrois have large ventral suckers by which they adhere to the surface of slippery stones under cascades, and the adults have well-developed digital disks. Tree frogs of the genus Rhacophorus are not un- common in western China. They live in trees, and their eggs are laid in nests of leaves on trees, not in the water as are those of common frogs. DISTRIBUTION AND SPECIATION OF AMPHIBIA OF WESTERN CHINA Introduction The faunistic distribution of Amphibia is often simpler than that of other vertebrates, because amphibians are bound almost absolutely to the land and to fresh water. The relatively small number of species is another factor simplify- ing the study of their distribution. The range of morphological specialization is relatively small, for caecilians, salamanders, and frogs form such compact orders that each can be recognized at once, even by the non-specialist. The Amphibia, in the process of evolution from the aquatic to the terrestrial environ- ment, are the smallest and simplest of the classes of higher vertebrates. The factors influencing amphibian distribution are obvious and great. Com- mon salt, even in weak solution, is an almost insuperable barrier to their dis- tribution, for one per cent salt solution kills the eggs and the larvae of most Amphibia. They are, in general, unable to cross seas, salt lakes, or saline plains. It has been thought that occasionally they might cross bodies of salt water on floating vegetation, but this must be an extremely rare event. Many terrestrial amphibians tolerate relatively high temperatures if moisture is available, but not cold and drought. Permanently frozen regions can have no Amphibia, as is true also of absolutely dry regions, not only because of the low temperature or humidity but also because of the lack of other animals, especially insects, as a food supply. Ranges of mountains are less effective barriers than the oceans, but they are more effective in the distribution of amphibians than in that of birds or mammals. The physical features may limit certain groups, as cascade frogs are limited to mountain streams, and vegetational features may limit other groups to grassy plains or to forests. The great centers of the tree frog evolution are the tropical forests of Malaysia, Africa, and tropical America. The geographic location and the physical features of Szechwan and Sikang are favorable to differentiation of amphibians and to certain types of adaptation. The forms found on the high plateau are adapted to cold; the inhabitants of the high mountains may be torrent breeders, and tree frogs (not necessarily found in forests), burrowing forms, and dry land and swamp types are represented. The geographic location of Szechwan and Sikang places them mostly in the Palearctic region, but at the south they exhibit a few relations with tropical southeastern Asia, that is, with the Oriental region. The physical features of 43 44 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 these two provinces, especially on the boundary between Szechwan and Sikang, are very remarkable. The great amount of topographic differentiation affords the greatest diversity of ecological conditions and thus favors speciation. This seems to be the chief factor producing the richness of the amphibian fauna of western China. General Relations between Amphibia of Western China and Other Parts of the World There is a definite pattern of distribution of world Amphibia. Species of Cryptobranchidae are found only in the Eastern Palearctic and Southeastern Nearctic; those of the Discoglossidae are limited to the Palearctic and are absent in the Nearctic. Various species of true toads (Bufo) are widely distributed over the world, being absent only in Malaysia, Australia, and New Zealand. The amphibian fauna of western China is very characteristic, with numerous genera developed chiefly on the border of the Tibetan Plateau, but with other species, genera, or families exhibiting varied relations with amphibian faunas of the other parts of the world. The broadest geographic relations of the Amphibia of western China are demonstrated at the generic level by Bufo and Rana. Toads of the genus Bufo in Szechwan and Sikang include four species. There are sixteen species of the frogs of the genus Rana in western China. Rhacophorus, with five species, repre- sents a tropical element in the fauna, the genus being very widespread in the East Indies and southeastern Asia. Bombina is a Eurasian genus with two species in Europe and two in eastern Asia. These fire-bellied toads are forms having a red belly variegated with black and yellow or orange. The distribution of the species of this genus is curious; orientalis is found only in Shantung and Manchuria and certain adjacent areas, while the West Chinese species, maxima, is found only in southeastern Sikang, the southwest corner of Szechwan, Yunnan, and adjacent Tongking. Stejneger, in 1905, before Bombina maxima was discovered, suspected that Bombina arose in southeastern Asia and emigrated to the north and west, that is, to northeastern Asia and to Europe. The dispersal of Bombina throughout northern Asia offers no great difficulty, there being no significant barriers. The present restriction of the ranges of the several species appears to indicate relict distribution of a primitive type. Megalobatrachus davidianus of the family Cryptobranchidae is widely dis- tributed in southwestern China, ranging as far north as the middle region of Shensi. Another more primitive member of the family, Cryptobranchus, is found in eastern North America. The original headquarters of Cryptobranchidae was presumably in Asia. It must be assumed that the North American stock came from Asia by crossing the land bridge that once connected Asia and North America via Alaska. All of the living members of the family are aquatic forms living in rivers or small streams. During the latter part of the Tertiary, they were widely distributed. Fossil eryptobranchids are known from the Miocene LIU: AMPHIBIANS OF WESTERN CHINA 45 of Europe and the lower Pliocene of Nebraska, where today the family is extinct. There are numerous other east Asiatic-southeast North American faunal relations, presumably of age equivalent with that of the Cryptobranchidae, for example, the Microhylidae and Pelobatidae. The amphibian fauna of Szechwan and Sikang, especially in the south of these provinces, exhibits certain close relations with the amphibian fauna of tropical Asia. The best examples are the various species of Megophrys of the otherwise Holarctic family Pelobatidae. The pelobatids appear to be Holarctic in origin, but they invaded the Oriental region in giving rise to the genus Mego- phrys, which is extremely characteristic of the East Indies. The presence of this genus in temperate western China thus seems to be a secondary northward dispersal of the genus. Megophrys boettgeri, found in Shansi and Kansu, is the northernmost species of the genus. General Relations between Amphibia of Western China and Other Parts of China The mountain ranges between Szechwan and Sikang, especially toward the southern parts of these two provinces, afford favorable topographic and geo- graphic conditions for the differentiation of the amphibian fauna. The transition from the Palearctic to the Oriental region gives this area a mixed amphibian fauna, with Palearctic forms at the north and Oriental species in the south, somewhat confusing the evident endemic and more characteristic special West Chinese fauna. The distribution of the Oriental and Palearctic forms overlaps to a certain extent, but each fauna still has its own distribution pattern. The following lists show the distribution and relations of the Amphibia of western China. I. Szechwan and Sikang species extending into other parts of China. 1. Batrachuperus pinchonii 3. Bufo bufo wrighti 5. Bufo tibetanus 2. Batrachuperus tibetanus 4. Bufo bufo minshanicus 6. Rana boulengeri Bufo bufo minshanicus was first known from Kansu, but the center of its range is evidently northwestern Szechwan. II. Northern and northeastern species found in Szechwan or Sikang. 1. Bufo bufo gargarizans 2. Rana nigromaculata 3. Rana temporaria chensin- ensts Rana temporaria chensinensis is a northern species widely distributed in Siberia and northern China. Rana nigromaculata and Bufo bufo gargarizans, representative of the European Rana esculenta and Bufo bufo bufo, are essentially northern types, but with a wide range in western China. III. Southern and southeastern species found in Szechwan or Sikang. 1. Megalobatrachus davidi- 4. Hyla annectans 8. Rana japonica nbs : 5. Staurois chunganensis 9. Rana adenopleura 2. Bombina maxima 6. R h ‘d 10. R theri 3. Hyla chinensis . Rana phrynoides . Rana guen ia (Nomaterialcollected) 7. Rana pleuraden 11. Rana andersonii 46 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 This list includes forms that are well represented in Fukien, and some, like Megalobatrachus davidianus and Bombina maxima, with their centers of dispersal only little to the south and east of Szechwan. IV. Tropical forms extending into Szechwan and Sikang. 1. Microhyla ornata 3. Rana limnocharis 2. Microhyla butleri 4. Rhacophorus leucomystax V. Endemic species of Szechwan and Sikang. A. ENDEMIC SPECIES OF SZECHWAN 1. Megophrys minor 5. Scutiger sp. 8. Aelurophryne glandulata 2. Megophrys omeimontis (Mount Omei) 9. Kaloula rugifera 3M h ne , 6. Scutiger sp. 10. Staurois lifanensis ee oe as me ee (Panlungshan) 11. Staurois ricketti minor a oN Qcrmeaten 7. Vibrissaphora boringit 12. Rhacophorus chenfui B. ENDEMIC SPECIES OF SIKANG 1. Batrachuperus yenyuan- 6. Scutiger sp. : 11. Calluella ocellata ensis (Lungtung, Paohsing) 12. Staurois mantzorum 2. Batrachuperus cochranae 7. Aelurophryne mammata 13. Staurois kangtingensis 3. Tylototriton taliangensis 8. Aelurophryne brevipes 14. Staurois loloensis 4. Scutiger pingti 9. Aelurophryne maculata 15. Rana chaochiaoensis 5. Scutiger rugosa 10. Kaloula macroptica 16. Rana shuchinae C. SPECIES FOUND IN BOTH SZECHWAN AND SIKANG, AND CONFINED TO THOSE Two PROVINCES 1. Batrachuperus karl- 3. Scutiger popei 6. Staurois mantzorum schmidti 4. Aelurophryne tainingensis 7. Rhacophorus omeimontis 2. Megophrys boulengeri 5. Nanorana pleskei 8. Rhacophorus dugritei The first two lists (A and B) may be expected to yield various species to the third (C) when their ranges become better known. Many forms appear to have a very sharply localized range, which contributes to the richness of the fauna of western China. DISTRIBUTION OF THE AMPHIBIA OF WESTERN CHINA The amphibian fauna of Szechwan and Sikang is decidedly rich in distinctive forms. The high plateau of Sikang and the mountains of eastern Sikang and western Szechwan are the centers of dispersal and probably of the origin of the genera Batrachuperus, Scutiger, Vibrissaphora, and Aelurophryne; of a series of species of Megophrys, and of the ranids Altirana and Nanorana; and of Bufo bufo minshanicus and Bufo bufo wrighti. The amphibian fauna of Szechwan and Sikang is much more closely related to that of the southern and southeastern parts of China than to that of northern and northeastern China. The ecological conditions in northern China are not favorable for Amphibia, so that only a few species that are adjusted to low temperatures and low humidity exist there, and most Palearctic forms are absent. LIU: AMPHIBIANS OF WESTERN CHINA 47 There are only six species of amphibians, with no salamander, in the Province of Hopei. Although the number of northern species found in Szechwan and Sikang is apparently small, no less than four of these six species are found in western China; but this is associated with very wide range of these forms. The amphibian fauna of southern and southeastern China is much richer than that of northern China, and various species and genera are common to southeastern China and western China. The forms of northern China are found in northern Szechwan and Sikang, and the southern and southeastern forms are scattered in southern Szechwan and the southeast corner of Sikang. Overlapping of the distribution of the northern and southern species occurs, especially for the northern Bufo bufo gargarizans and Rana nigromaculata and the southern Megalobatrachus davidianus and Microhyla ornata. According to zoogeographical relations, Szechwan and Sikang are in a tran- sitional zone from the Palearctic in the north to the Oriental in the south; eco- logically they range from the lowlands of the east to the high plateau of the west. The mountain ranges running in a north-south direction between these two provinces afford great variety of topographic and ecological conditions in which the habitat niches have been filled by various adaptive groups. Mount Omei presents the most varied and interesting amphibian fauna of any of the collect- ing stations. This mountain is the southeastern end of the Hungshan range with the Tatuho at the west and south. The amphibian fauna of Mount Omei consists mainly of forms found in the northern parts of the Hungshan, whereas the amphibians of the Taliangshan south of the lower arm of the Tatuho are mostly quite distinct species. The high plateau of Sikang and northwestern Szechwan, separated from Mount Omei by the Tatuho, has a different amphibian fauna. For the present discussion, western Szechwan and Sikang are arbitrarily divided into five zones: Mount Omei; Taliang Mountains and Ningshu in Sikang; the Kang- shu region in Sikang from Kangting to Kantze; northwestern Szechwan and the adjacent region in Sikang; and the Chengtu plain. The amphibian fauna of Mount Omei is characteristic in having more species of Scutiger, Megophrys, and Rhacophorus. The following lists of species found in each region indicate the similarities and the differences between the amphibian fauna of those regions. I. Mount Omei. 1. Megalobatrachus davidi- 10. Vibrissaphora boringii 20. Rana limnocharis anus 11. Bufo bufo gargarizans 21. Rana nigromaculata 3. Megophrys minor 13. Kaloula rugifera 23. Rhacophorus leuco- 4. Megophrys omeimontis 14. Microhyla ornata mystax 5. Megophrys boulengert 15. Staurois chunganensis 24. Rhacophorus omeimontis 6. Megophrys oshanensis 16. Rana margaretae 25. Rhacophorus chenfui iain it oo 17. Rana bowlengeri 26. Rhacophorus bambusi- 8. Scutiger schmidti 18. Rana adenopleura cola 9. Scutiger sp. 19. Rana japonica 48 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 II. Taliang Mountains and Ningshu, Sikang. 1. Megalobatrachus davidi- 7. Hyla annectans 14. Rana andersonii anus 8. Kaloula macroptica 15. Rana phrynoides 2. Batrachuperus yenyuan- 9. Calluella ocellata 16. Rana pleuraden ade 3 . 10. Staurois loloensis 17. Rana adenopleura re A aeons taliangensis 11. Staurois ricketti minor 18. Rhacophorusbambusicola 5. Sei para 12. Rana chaochiaoensis 19. Rhacophorus omeimontis Gaede tances 13. Rana shuchinae 20. Rhacophorus dugritei III. Kangshu, Kangting to Kantze, Sikang. 1. Batrachuperus karl- 4. Aelurophryne taining- 8. Altirana parkeri schmidti ensis 9. Nanorana pleskei 2. Aelurophryne mammata 5. Aelurophryne maculata 10. Staurois kangtingensis 3. Aelurophryne brevipes 6. Bufo tibetanus 11. Rana temporaria chensin- 7. Bufo bufo wrighti ensis IV. Northwestern Szechwan and adjacent Sikang. 1. Megalobatrachus davi- 10. Scutiger sp. 20. Staurois lifanensis dianus Paronestat: Kwan- 21. Staurois loloensis 2. Batrachuperus pinchonii sien) 22. Staurois chunganensis 3. Batrachuperus karl- 11. Aelurophryne glandulata 93 Pana margaretae schmidti 12. Aelurophryne taining- 24. Rana boulengeri 4. Batrachuperus cochranae ensis ep 25. Rana japonica 5. Batrachuperus tibetanus 18. Bufo bufo wrighti . . ; : 26. Rana temporaria chensin- 6. Megophrys minor 14. Bufo bufo gargarizans ensia 7. Scutiger poper 15. Hyla annectans 27. Rana limnocharis 8. Scutiger schmidti 16. Kaloula rugifera 28. Rana nigromaculata 9. Scutiger sp. ? 17. Microhyla ornata 29. Rhacophorus leucomystax (Lungtung, Paohsing) 18. Nanorana pleskei 30. Rhacophorus omeimontis 19. Staurois mantzorum 31. Rhacophorus dugritei V. Chengtu Plain. 1. Bufo bufo gargarizans 3. Microhyla ornata 5. Rana nigromaculata 2. Kaloula rugifera 4. Rana limnocharis 6. Rana guentheri The vertical distributions of the Amphibia of western China are discussed under the species headings. Discussion of Endemic Genera The factors involved in the rise of a new species are numerous and varied. The interaction of the various types of isolation—topographic, geographical or spatial, chronological or seasonal, ecological or physiological—in combination with genetic change of the populations and natural selection, produces speciation. The amphibian fauna of western China affords an interesting example of specia- tion, with numerous closely related species of endemic generic groups. Topo- graphic isolation, favored by the rugged mountainous terrain with deep per- manent streams and their valleys, ecological and physiological isolations, favored by varied climatic and vegetational environmental niches, and the pressure of LIU: AMPHIBIANS OF WESTERN CHINA 49 natural selection have been the chief factors in the evolutionary specialization of the amphibians of Szechwan and Sikang. The topography of western China is determined by the relation of the region to the Tibetan Plateau and by the north-south mountain ranges between Sze- chwan and Sikang. Altitudes in western China range from 1,750 feet at Chengtu to 25,000 feet on Koonga Mountain in Sikang. The low and mild temperate Chengtu Plain passes into high, cold plateaus at 14,000 feet. The mingling of the northern and southern faunal elements in a zone transitional between the Palearctic and the Oriental clearly has been favorable to species formation. The pelobatid genus Aelurophryne and the ranid Nanorana (with its close relative Altirana) flourish only on the high plateau of Sikang and in the north- western corner of Szechwan, and are completely absent from the warmer lowlands. These two genera, both adapted to cold, are found under different ecological conditions. The various species of Aelurophryne always inhabit stony mountain streams, especially near their upper reaches, where the water is shallow and very cold. Nanorana pleskei is found only in the quiet water of the marshes on the high plateaus, where there is usually a rich vegetation. A third high plateau species, Bufo tibetanus, endemic in western China, ranges north to Tsinghai. This toad is found only under stones or in holes on hillsides. None of these three quite differently adapted Salientia of the high plateau range into the equally cold lowlands of the Palearctic region, and they are found nowhere else in the world. The salamander genus Batrachuperus of western China also illustrates topo- graphic as well as physiological and geological isolation in relation to speciation. There is no doubt that the center of differentiation of Batrachuperus is in north- western Szechwan and the northeastern corner of Sikang, where four of the five species are found. B. pinchonii is found together with Aelurophryne brevipes on the high plateau of Sikang, and with A. glandulata in northwestern Szechwan. It occurs with Megophrys boulengeri on the top of Mount Omei. Curiously enough, it is never found in association with frogs of the genus Scutiger, or indeed with any other amphibian. Batrachuperus and Aelurophryne thus charac- terize an ecological group, the former being somewhat more a mountain form, and the latter more an inhabitant of the high plateau. The Himalayan pelobatids of the genus Scutiger have long been rarities in museum collections. They are really not rare, but because of their characteristic isolation in very small and inaccessible streams at high altitudes, they have not often been collected. Scutiger sikkimensis is recorded from the Burmo-Sikang border, and S. alticola was discovered in the southern part of Tibet. As a result of my collecting trips in western China, I can record four new species of Scutiger— pingit, rugosa, schmidti and popei—and it appears that there are three more undescribed species in western China, known only from tadpoles and specimens just metamorphosed. It now appears that the center of differentiation of Scutiger is unquestionably in the mountains of the western border of Szechwan 50 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 and in the eastern and the southeastern parts of Sikang. The species of Scutiger are all cold-adapted animals found in upper reaches of mountain streams; but they are not adapted to the extremes of the high plateau or high mountain zones like Aelurophryne and Batrachuperus. All the species of Scutiger are definitely limited to a topographic zone and a special habitat—small mountain streams from 3,000 to 8,000 feet altitude. Aelurophryne, closely related to Scutiger, is sharply limited to the high plateau and mountain slopes from 8,000 to 12,000 feet altitude. Geographic isolation is also well illustrated by various species of the am- phibian fauna of western China. Bufo raddei and Rana temporaria chensinensis, which are northern Chinese forms, are limited to the northern part of Szechwan and Sikang. The high plateau species of Aeluwrophryne, Bufo tibetanus, and Nanorana pleskei never extend southeastward to the high mountains south of Wenchwanhsien in Szechwan. The western Chinese species of Megophrys are found mainly south of Lat. 30° N., and especially in the Mount Omei region. The results of my investigations in western China indicate that the am- phibian fauna of Mount Omei is more closely related to the amphibians of the mountains north of Mount Omei than to those of the Taliang Mountains to the south. This is especially clear in the genus Scutiger. Scutiger schmidti and S. poper are found on Mount Omei and also in the mountain streams of Panlung- shan, west of Kwanhsien and in the vicinity of Paohsinghsien (Muping); but not one specimen of these two forms has been taken on the Taliang Mountains. In the streams on the west side of the Taliang Mountains there are two other quite different species, Scutiger pingit and S. rugosa, which are not found on Mount Omei or anywhere else. The Tatu River, south of Mount Omei and north of the Taliang Mountains, marks the boundary of two different faunal areas that are characterized by distinct species but related by their genera. Mount Omei, at the south end of Hungshan range, has an altitude of 11,000 feet, much higher than the mountains just to the west and northwest. Above the level of 5,000 feet it is really an ecological island, isolated like the islands in the sea. This may be the chief factor in its amphibian speciation. The extent of this speciation is well illustrated by the distinctness of the genus Vibrissaphora, which I discovered on the mountain in the zone between 3,000 and 4,000 feet, and which is now known to have a representative also in the mountains of Fukien. Megophrys presents other good examples of Mount Omei speciation, M. boulengeri being found only at the top of the peak. Another endemic species, M. omeimontis, is found at about 3,000 feet altitude. Habitat Isolation Certain closely related species of the amphibian fauna of western China are found breeding in the same place. The northern Chinese woodfrogs, Rana temporaria chensinensis, and the southern Chinese woodfrogs, R. japonica, lay their egg-masses in side pools of Kialing River near Peip’ei, north of Chungking. LIU: AMPHIBIANS OF WESTERN CHINA 51 Although they are closely related species and have the same breeding site, they are still easily distinguished; their breeding seasons at Chungking are also dif- ferent. According to H. W. Chang, R. japonica lays its eggs from the end of November to the end of December, whereas, for the closely related species R. temporaria chensinensis, the breeding season is as late as January. There are very good illustrations to show the physiological isolation of the amphibian fauna of Mount Omei. Four species of Rhacophorus found on Mount Omei differ from each other morphologically as well as physiologically. The common tree frogs of the tropical species R. lewcomystax have a different voice from that of three Rhacophorus of western China. R. chenfui is an endemic form of Mount Omei; R. omeimontis and R. bambusicola are endemic to Szechwan and Sikang and are found with chenfuz on Mount Omei. In their habits, and thus in their physiological reactions, these three species are more closely related to each other than to R. leucomystax. The females of leucomystax lay their egg- foam masses on the walls of small pools or ponds or on vegetation in flooded rice fields. The other three species never choose this kind of site for egg-laying. This is one of the factors that separates them from the widespread leucomystax. The breeding season of leucomystax is also later than that of the other three tree frogs. The habitat of leucomystax is characteristic. I rarely found it on trees or bushes; it is a terrestrial form. The vertical distribution of this species is from 1,000 to 4,000 feet altitude. In other words, in western China it is a charac- teristic lowland frog, though absent from the Chengtu plain. The other three closely related species found on Mount Omei are distinctly different species, rather than subspecies. The development of three species of tree frogs in one region has been effected by adjustment both to specific habitat and to physiological requirements of the habitat zones. R. bambusicola is found from 8,000 feet upward to the very top of the mountain. Frogs of this species are physiologically adjusted to cold temperatures. In connection with adapta- tion to cold they are more terrestrial than arboreal, with stouter bodies, smaller adhesive disks, shorter legs, and much better developed inner metatarsal tubercles. They lay their egg-foam masses under the roots of vegetation, under stones near pools, ponds, or ditches, or even in holes in the ground. They never lay eggs on trees as does R. omeimontis. This terrestrial breeding habit has a positive correlation with the zonal vegetation, as there are no broad-leaved trees above 8,000 feet altitude, but only pine trees, dwarf bamboos, and numerous low bushes. R. omeimontis is found from 3,000 to 5,000 feet altitude, but never asso- ciated with bambusicola. This species is the most typical tree frog, and its habitat is presumably that of the ancestral type. R. omeimontis has the longest legs and the largest adhesive disks of the three endemic forms, and it lays its eggs in nests made by pulling together the leaves of broad-leaved trees or other vegetation near the water. This species is found at lower altitudes, with the trees appro- priate for its breeding habit. : wie’ ‘ \ oe i. 52 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 R. chenfui is found on Mount Omei together with omeimontis. R. chenfui is limited to two places on the mountain and is certainly not widely distributed like omeimontis, bambusicola, or lewcomystax. This species still has arboreal adaptations, but has shorter legs and smaller adhesive disks than omeimontis. Unfortunately, the breeding habits of chenfui are still unknown, although my assistants and I tried very hard throughout three summers to learn more about them, after finding the tadpoles during our first season. The adaptive complex of bambusicola, omeimontis, and chenfui is distinct in each species, but they are nevertheless closely allied forms. Their croaking is very similar; an inexperienced person never recognizes the differences between their voices. The morphological characters of their tadpoles are similar. I failed to distinguish them during my first investigations; but once one knows the differences between them, one never fails to recognize them in the field. Physiological isolation (including behavior characters) as well as habitat isola- tion, are the factors that make possible the presence of three closely related species of frogs on the same mountain. Rana adenopleura, also found on Mount Omei, is another good example of speciation through habitat and behavioral isolation. The breeding habits of this frog are absolutely different from those of other Chinese species of Rana. Eggs are laid in nests near the margin of the water of small pools but never in the water. (Pope [1931] stated that this frog lays its eggs in flooded rice fields.) Through this habit, Rana adenopleura is excluded from any possibility of in- breeding with other species. Again, this frog can only perpetuate its race in places where rains are frequent, so that the hatched tadpoles are washed into the pool for further development. The vertical distribution of the vegetation of the Mount Omei region ex- hibits four zones. These are the warm-temperate, up to 2,000 feet altitude; the temperate, from 2,000 to 5,000 feet; the cool-temperate, from 5,000 to 10,000 feet; and the sub-Alpine, from 10,000 feet upward. In the lowest zone, at the foot of Mount Omei, there are seven species of frogs and toads but only Rana guentheri is limited to this area, the remaining six extending into the next zone. The temperate zone has the richest amphibian fauna, consisting of nineteen species. Six of these are more commonly found in the warm-temperate zone below and three range upward to the cool-temperate, leaving ten species confined to the 2,000 to 5,000 foot zone. In the cool-temperate zone, there are eight species of frogs and toads but only three of these are limited to this area. The pelobatid frogs of the genus Scutiger are characteristically cool-temperate. The ecological conditions on the top of Mount Omei are absolutely different from those in the zones below and only a few species of cold-adapted Amphibia exist there. Batrachuperus pinchonii and Megophrys boulengeri occur only in this zone and breed there freely. They are not found below 10,000 feet altitude. The third species found in the uppermost zone is Rhacophorus bambusicola, which ranges downward into the cool-temperate, to about 8,000 feet. LIU: AMPHIBIANS OF WESTERN CHINA 53 PROVINCIAL LISTS OF THE SPECIES OF AMPHIBIA Megalobatrachus davidianus Batrachuperus pinchonii Batrachuperus tibetanus Batrachuperus karlschmidti Hynobius shihi Megophrys oshanensis Megophrys minor Megophrys boulengeri Megophrys omeimontis Megophrys shapingensis Scutiger popet Scutiger schmidti Scutiger sp. (Mount Omei) Scutiger sp. (Panlungshan) Megalobatrachus davidianus Batrachuperus pinchonii Batrachuperus tibetanus Batrachuperus karlschmidti Batrachuperus yenyuanensis Batrachuperus cochranae Tylototriton taliangensis Bombina maxima Megophrys boulengeri Megophrys omeimontis Scutiger popet Scutiger pingit Scutiger rugosa Scutiger sp. (Lungtung) Tylototriton verrucosus Triturus wolterstor ffi Bombina maxima Megophrys lateralis Megophrys carinensis Bufo bufo wrighti Bufo melanostictus Hyla annectans Bufo tibetanus OF WESTERN CHINA I. SZECHWAN Vibrissaphora boringii Aelurophryne glandulata Aelurophryne tainingensis Bufo bufo gargarizans Bufo bufo wrighti Bufo bufo minshanicus Hyla annectans Microhyla ornata Microhyla butleri Kaloula rugifera Nanorana pleskei Staurois chunganensis Staurois lifanensis Staurois rickettt minor II. SIKANG Aelurophryne mammata Aelurophryne brevipes Aelurophryne tainingensis Aelurophryne maculata Bufo bufo gargarizans Bufo bufo wrighti Bufo tibetanus Hyla annectans Microhyla ornata Kaloula macroptica Calluella ocellata Altirana parkeri Nanorana pleskei Staurois mantzorum III. YUNNAN Microhyla ornata Kaloula verrucosa Calluella yunnanensis Kalophrynus pleurostigma in- terlineatus Staurois afghanus Rana limnocharis Rana tigerina rugulosa IV. TsSINGHAI Bufo raddei Rana limnocharis Rana nigromaculata Rana guentheri Rana boulengeri Rana adenopleura Rana margaretae Rana andersonii Rana temporaria chensinensis Rana japonica Rhacophorus dugritet Rhacophorus omeimontis Rhacophorus bambusicola Rhacophorus leucomystax Rhacophorus chenfui Staurots chunganensis Staurois kangtingensis Staurois loloensis Rana limnocharis Rana nigromaculata Rana boulengeri Rana phrynoides Rana pleuraden Rana andersonit Rana temporaria chensinensis Rana japonica Rana chaochiaoensis Rana shuchinae Rhacophorus bambusicola Rana kuhlit Rana guentheri Rana phrynoides Rana feae Rana pleuraden Rana andersonir Rana chaochiaoensis Rhacophorus leucomystax Rana temporaria chensinensis 54 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 V. KANSU Batrachuperus tibetanus Bufo bufo minshanicus Rana boulengeri Megophrys boettgeri Bufo raddei Rana temporaria chensinensis - Bufo bufo gargarizans Rana nigromaculata ADAPTATIONS OF THE AMPHIBIA OF WESTERN CHINA Adaptive radiation in the Amphibia is limited in comparison with this mode of evolution in other groups of vertebrates. The external modifications for protection and adjustment to new ecological conditions are poorly developed or lacking in the adult form of modern amphibians. They are accordingly very easily controlled and influenced by environmental factors, especially temperature and moisture. Although various ways of protecting the eggs and the tadpoles have been developed among the Amphibia, most amphibians in these stages are exposed to biological, physical, and chemical hazards. Furthermore, the eggs and the tadpoles themselves have a narrow range of adaptive structure and are very poorly equipped for protection against the environment. This limited adaptability helps to account for the small amphibian fauna in the world in general. The Chinese fauna and flora are by no means well studied, and this is true for the Amphibia. Little has been written on the habits and habitats of Chinese salamanders, or on the toads and frogs, except for the paper by Pope (1931), who made intensive field studies of the amphibians of Fukien and Hainan, and for my own series of papers (1930-31), examining the habits and life histories of the Amphibia of northern China. Although many people have collected in western China, no studies in nature have been made. The following account of the adaptation of amphibians of western China is mainly the result of my own observations. Discussion of the habitat preferences of the Amphibia requires great caution, as it is difficult to draw a definite line between the different kinds of habitats and the various species may overlap to some extent from one habitat to another. When I record a frog from a certain habitat, I mean that it was found there by me, but this does not mean that this species is invariably to be found only in such an environment. This is well illustrated by Rana adenopleura, which Pope found in flooded fields on mountains in Fukien, whereas I invariably found it in pools, and very rarely in ponds, on Mount Omei. Amphibians, like other animals, may make short distinct migrations at the breeding season and in pursuit of food. The same species of frog may be found in different ecological niches during different seasons of the year and different stages of its life history. A good example of this is Stawrois chunganensis, found only in large mountain streams during its breeding activities. Before and after the breeding season these frogs are rarely seen, and it is probable that they live in the vegetation on hillsides. I obtained two specimens in such a habitat at the time of rain. Finally, the vertical distribution of amphibians varies from species to species, in correla- LIU: AMPHIBIANS OF WESTERN CHINA 55 tion with differences of habitat, but often with large overlap. Bufo bufo wrighti has a vertical range extending from 3,000 to 10,000 feet. This toad may breed in pools and ponds, in stationary bodies of water along mountain streams or in mountain streams themselves. The tadpoles of a given species may be found in different types of water; they are subject to conditions of current and may be washed down from pond to stream or from running water to swift water. Such displacement is well illustrated by the tadpoles of Scutiger popei. It is difficult to draw sharp lines between different types of water habitats, as explained below, and it may therefore be difficult to distinguish the frog types from one habitat to the next. Examples of this difficulty are the indefinite boundaries between swift and running water; or between ponds and pools. After a heavy rain, the water in a small mountain stream, normally with “running water,” changes into “swift water”; and a pool may temporarily become a pond. It is also true that after a heavy rain the quiet water of a ditch, pond, or pool may become “‘slow-running water.’’ Thus, bodies of water may change from one current condition to another during the year. When we say ‘‘swift water’? we mean that of permanent streams with cascades. Adaptation in Adults and Tadpoles The high mountains of western China afford various kinds of interesting environments that are good laboratories for studies of the fauna and flora in nature and excellent experimental ground for ecological and taxonomic studies. The amphibian fauna in western China is rich and characteristic, especially in montane forms such as the different species of the family Pelobatidae, the cascade frogs of the genus Staurots, and the different tree frogs of the genus Rhacophorus. The habitats of the adult amphibians fall naturally into three categories, the aquatic, the terrestrial, and the arboreal. AQUATIC ADAPTATION.—The aquatic forms of amphibians can be divided into three subordinate categories, the swift-water or cascade group, the running- water group, and the quiet-water group. In the cascade group, we shall take up only two examples, Rana margaretae and Staurois lifanensis. Rana margaretae is commonly seen adhering closely to the surface of the rocks that form the steep banks of swift mountain streams, or sitting on mats of green vegetation on such rocks. This frog usually directs its head toward the current, into which it dives at the slightest disturbance. These frogs usually sit from one to four feet above the water level. They are green in color, matching the vegetation. Staurois lifanensis, as well as all the other species of Staurois except chunganensis, is a true cascade frog, usually seen adhering closely to the surface of a rock by the side of a shaded cascade, either on bare rock or on matted vegetation reached by the spray. The body of this frog is slender and dorso-ventrally depressed, the legs are long, and the toes are fully webbed and provided with adhesive disks at the digital extremities. The depressed body and the adhesive disks enable the animal to maintain its hold 56 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 on the slippery stones within the spray zone. The webs and the long legs enable it to swim vigorously in the swift water. The coloration, varying greatly and changing under different conditions, is of great survival value in the effective protection of the species. Only a trained eye can detect one of these frogs before it moves. The running-water habitat, in about the same altitude zone as the cascading larger streams, consists mainly of the smaller headwater brooks. There are only a few species of salamanders and frogs in western China that inhabit this niche. Rana boulengeri, the large mountain frog, and the allied species, R. phry- noides, are adapted to mountain streams. The body is stout and heavy but the legs are long and provided with very strong muscles and fully developed webs. These frogs are good swimmers in running water. Their coloration is very similar to that of the stones in the water or near the margin of the mountain streams, where they usually sit. The primitive pelobatid frogs, like Aelurophryne glandulata and Scutiger popei, may be found under stones in the water of mountain streams. However, they are not truly aquatic forms and they hide normally in crevices between stones and in similar situations on hillsides along or near the mountain streams. The high mountain salamanders of the species Batrachuperus pinchonii hide themselves under stones in small mountain brooks from 6,000 feet upward, but are never found in larger and swifter streams. They are occasionally seen under decayed logs of pine on hillsides. The strong tail of this species and the undulating movements of the body form the powerful means needed for locomotion in its brook habitat. The quiet-water group of Amphibia exhibits great variety of size and form. It is more heterogeneous than any of the other ecological groups of amphibians and its species are widely distributed vertically and horizontally. Mountain forms and plains types can be recognized. Nanorana pleskei is a swamp-dweller in grassland on high plateaus and mountains; it is found under stones or roots of vegetation in quiet water from 9,000 to 14,000 feet altitude. The depressed body resembles that of the cascade frogs; this type of body is not found in other frogs living in quiet water. Body form in Nanorana pleskei seems to be associated with the habits of the species. These frogs hide them- selves under flat stones in the water during the summer and at the time of hibernation they may go deep in the ground in the crevices between stones. Their legs are weak and short, but their feet are provided with well-developed webs for swimming in the temporary pools and ponds and open water of the high mountain marshes. Bombina maxima, the most primitive salientian in China, is more aquatic than terrestrial. This frog is generally found under stones in or at the edge of the water of small ponds or pools of high mountains. The legs are moderately developed and the toes are fully webbed, especially in the males. The beautiful musical frog, Rana adenopleura, especially famous on Mount Omei, is another good example of adaptation to quiet water. It inhabits pools and is occasionally found in ponds and marshes. This species is a typical mountain quiet-water form with much stouter body, rather short and weak legs, and LIU: AMPHIBIANS OF WESTERN CHINA 57 moderately developed webs between the toes. There are only a few species of frogs inhabiting the quiet water of the lower plains. The common Chinese pond frogs, Rana nigromaculata and R. guentheri, are the best illustrations of the quiet-water type of low altitudes (from 1,000 feet above sea level at Chungking to 1,750 feet at Chengtu). These frogs are well equipped for a pond life and at the same time are good swimmers, with well-developed legs and webs. R. nigro- maculata may be found in ponds on the lower slopes of the mountains up to about 3,000 feet altitude. TERRESTRIAL ADAPTATION.—Many species of various genera of salamanders and frogs visit the water only during their breeding season, and live on land, often far from water, at other seasons. Such forms should be classified as terrestrial, and they may be distinguished as a special ecological group. Ter- restrial amphibians may have different habitats from time to time and under different conditions. The burrowing, narrow-mouthed toads (Kaloula rugifera) remain in their hiding places during the daytime, but during or after a heavy rain they may come out onto open ground, or hide themselves among vegetation. They feed at twilight. This case clearly illustrates the difficulties of classifying the amphibians into definite groups, according to habitat. However, in general, they can be divided into two adaptive groups: the grassland inhabitants and the burrowing forms. The grassland group includes quite a few species of Amphibia, both montane and plains forms. The common members of this group are the wood frogs, Rana japonica and R. temporaria chensinensis; the rice-field frog, R. limnocharis; the pond frog, R. nigromaculata; the small narrow-mouthed frog, Microhyla ornata; and the pelobatid frog, Megophrys minor. M. minor is the only species found in grass or among bushes of mountain sides; all the rest inhabit the hills as well as the Chengtu Plain. Sometimes one may find Rana limnocharis and Microhyla ornata in cracks of the earth or in crevices in stones. There is no definite struc- tural differentiation for such habits. Some of the terrestrial forms of amphibians hide during the daytime in cracks of stones and earth or in holes, dug by themselves. They are burrowers in varying degree, and are usually provided with shorter legs and strong inner metatarsal tubercles for digging. They are protected from desiccation by the thicker skin with its large number of mucous glands, as in Kaloula rugifera and K. macroptica. The toads of the genera Kaloula and Bufo have sometimes been regarded as forming, ecologically, a stone pile and old-building group, because they are so generally found near old houses. These species obviously meet more enemies than do other terrestrial forms, and are much more exposed to predation than the cascade and running-water species. In amphibians there are no structures for active offense, the defensive weapons of toads and frogs being mainly chemical in nature. Little is known of the biochemistry of their glandular secretions. Bufo bufo gargarizans has an ugly appearance with large numbers of warts of varying size; there is a super- 58 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 stition that by touching them one may acquire warts. That the chemical secre- tions of the skin of toads provide an effective defense is shown by the fact that cats and dogs, especially the former, will eat frogs but not toads. It is likely that these animals learn by experience in each generation to avoid toads; one experience, perhaps, will be remembered for life. Histological study demon- strates the many poison glands found in the skin of Bufo bufo gargarizans and Kaloula rugifera. Other toads, Bufo raddei, B. tibetanus, and B. bufo wrighti, are protected in the same way. The pelobatid frogs of the genera Scutiger and Aelurophryne also belong to the digging and burrowing group, hiding in holes in the earth or in crevices in stones on hillsides except at the breeding season. After this season one may find some Aelurophryne glandulata under stones in the water, but only rarely can specimens of the species of Scutiger be found in this habitat. The dull coloration in these amphibians, very much like that of the earth and stones under which they hide themselves during the day, seems to be correlated with their habitat. When they come out at twilight to search for food, their coloration may be protective to a degree. Tree frogs of the Rhacophorus group (Rhacophoridae) and the tree toad Hyla annectans (Hylidae) are mostly arboreal in habits and sometimes have terrestrial adaptations. They are never truly aquatic. The chief structural specializations for arboreal habits are the large adhesive disks at the tips of the fingers and toes, the long limbs and the dorso-ventrally depressed body. Rha- cophorus omeimontis is the best example of arboreal adaptation among West Chinese frogs, with the most highly developed adhesive disks and the most striking resemblance to its environment in coloration. This tree frog is usually found on trees or on the roofs or walls of houses, only rarely on bushes. The color of the back of R. omeimontis is very similar to that of the lichens of its usual habitat. Without learning how to look for it, and without keen eyes, very few people will be able to find this frog. R. bambusicola is generally found among bushes or in grass, and its adhesive disks are much smaller than those of R. omei- montis. Disk size is also correlated with breeding habits, as the females of bambusicola lay their eggs in holes in the ground or under stones out of water, and omeimontis lay theirs in leaves at the tips of the branches of trees over- hanging water. The small green tree toad, Hyla annectans, may be found on bushes or in grass by the side of streams or in marshes; the legs and adhesive disks are only moderately developed. In this species there is very good color protection. BREEDING ADAPTATION.—Most of the frogs breeding in swift mountain streams of western China belong to the genus Staurois; only one species of Rana breeds in such streams. In the genus Staurois, sex dimorphism is distinctly marked. Males are much smaller in size than the females; for example, the average body length of fifty males of Stawrois chunganensis is 36.5 mm., and of fifty females 49.8 mm. This proportion of male to female size holds roughly for all other species of Stawrois studied, and for Rana margaretae as well. Means of clasping and holding fast to the female are moderately developed. Nuptial LIU: AMPHIBIANS OF WESTERN CHINA 59 pads of dark or creamy color are moderately developed on the inner dorsal side of the first finger of the males of the several species. The arms in the males of Staurois are moderately enlarged as compared with the arms of the females, except in S. chunganensis, in which the males have only slightly enlarged arms. Enlargement of the arms of the male is also to be seen in Rana margaretae, which probably belongs to the cascade-frog group, for both adults and tadpoles are found with the species of Staurois, along large mountain streams. Dimorphism in size of the sexes is distinctly an adult adaptation to breeding in swift water. The smaller size of the male is correlated with comparatively slender and dorso-ventrally depressed body; this is interpreted as facilitating breeding activities by decreasing the total area of the breeding pair exposed to the swift current in the mountain torrents. The method of egg-laying, the nature of the egg-mass, and the type of egg are especially adapted to breeding in swift water. The process of egg-laying was observed only in Staurois chunganensis. The attachment of the eggs to the under side of the stones in the water is functional in keeping the eggs from being washed downstream. The small amount of jelly also reduces the resistance to the current. The white color of the eggs of S. chunganensis is not clearly adaptive; the egg color is unknown for other species of Staurois. Tadpoles inhabiting the swift water of mountain streams of western China belong chiefly to the genus Staurois, but some tadpoles of various species of Rana are also found in relatively quiet pools back of the large stones of such mountain streams. Staurois tadpoles adhere tightly to the surface of slippery rock in the water by means of a ventral sucker formed by the mouth and the adjacent region of the body. The flat body of the tadpole decreases the likeli- hood of its being washed down from the stone to which it adheres, and also diminishes the effect of the swift current. The strong tail muscle and the low thick crest of the tail enable the tadpole to swim even against a swift current. Tadpoles of Rana margaretae are not equipped to live in such swift water, but are found in rather quiet pools behind large stones in swift mountain streams. Sometimes the tadpoles of Scutiger popei, Aelurophryne glandulata, Megophrys minor, and Rana boulengeri may also be found in such mountain streams. These tadpoles are much more abundant in the smaller streams where the adults as well as eggs and young tadpoles of those species are found. Floods and mere accident account for the carrying down of some of the tadpoles to larger, swifter parts of the stream below. Among the Amphibia of western China most of the frogs of the family Pelobatidae breed in small mountain streams, especially in their upper reaches. A very few species of Rana also lay their eggs in this type of stream. The frogs of the genus Scutiger afford the best examples of breeding adaptations to this habitat. The adults of various species of Scutiger, Aelurophryne, and Megophrys inhabit running water only in the breeding season, when they repair to the smaller mountain streams. Sex dimorphism in size in these forms is slight, and the males are in fact found to be smaller only in Scutiger schmidti. 60 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 All the males of the species of Scutiger have two patches of fine spines on the chest and nuptial spines on the inner dorsal sides of the first and second fingers. The arms of the males are much stronger than those of the females. Field observation proves that the spines on the chest of the male have nothing to do with maintaining the firm grip of the male, the amplexus being inguinal. The mating occurs in small mountain streams, under stones or among roots, and there is little evident need for difference in size between the sexes. The secondary sex characters correlated with amplexus are nevertheless moderately developed. The frogs of the genus Megophrys also lay their eggs under stones in small mountain streams. The secondary sex characters in this genus are poorly developed, the males being generally a little smaller than the females. The males do not have strongly developed arms, and have only a very moderate gray nuptial pad developed on the inner dorsal side of the first finger, instead of spines, as in Megophrys minor. Rana boulengert, also characteristically a running- water form, is an example of very different adaptations in its secondary sex characters. Males of R. boulengeri are distinctly larger than the females, and are equipped with strong spines on the throat, the chest, the belly, and the inner dorsal side of the first, the second and sometimes the third fingers. The arm of the male is enormously enlarged, and with the spines this assures a very strong hold on the female. In sex dimorphism another pelobatid frog, Aeluro- phryne mammata, resembles Rana boulengeri rather than Megophrys. It is obvious that frogs breeding in the running water of small mountain streams differ ecologically, as a group, from those that breed in swift water. Frogs of running water have much less sex dimorphism in size, and may have little development of nuptial spines. However, when the males are distinctly larger than the females, the other secondary sex characters, especially the nuptial spines and the enlargement of the arm, are very strongly developed. In the running-water forms the mode of egg-laying among the frogs of the genus Scutiger has been observed only for S. schmidti. It may be presumed to be similar in other frogs of this habitat, since the egg-masses of other species of Scutiger are likewise known to be attached to the under side of rocks or roots. This mode of egg-laying is similar to that of Staurois chunganensis. As far as is known, the color of the egg is white in all of the frogs breeding in running water, except in Rana boulengeri, in which the egg is gray at the animal pole and yellowish white at the vegetal pole. Thus the mode of egg- laying, the color of the eggs, their attachment, and the amount of jelly are very similar in the running-water and the swift-water species. Rana boulengeri affords a variety of exceptions (see p. 267). Tadpoles inhabiting running water of mountain streams illustrate two types of adaptation, one for surface feeding and the other for bottom feeding. Tadpoles of Megophrys minor swim at the surface of water when feeding, in quieter places back of large stones, with their funnel-like mouth parts expanded at the surface film. In such clear waters even the surface plankton must have LIU: AMPHIBIANS OF WESTERN CHINA 61 an impoverished micro-fauna and micro-flora. The bottom-feeding forms have very strong mandibles and labial teeth, as in the tadpoles of different species of Scutiger, Aelurophryne, Megophrys pelodytoides, and Rana boulengeri. Both surface and bottom feeders are good swimmers in the running water of their habitat. Their bodies are elongated and cylindrical, with strong, long tails provided with low thick crests. It is difficult to find a clear demarcation between “running water’ and “‘slow- running water.” In slow-running water, distinguished as a breeding habitat of frogs, the flow of the water is so much reduced that it does not carry away the unattached eggs laid there by certain species of frogs. As set forth above, all the eggs laid in running water are of the attached type, and no free-floating or submerged eggs are to be found. It is difficult to find a species that lays its eggs in slow-running water and not in completely quiet water, because there is no effective difference between these two habitats. Comparatively speaking, how- ever, there are certain species that generally lay their eggs more commonly in slow-running water than in quiet pools or ponds. The West Chinese mountain toad, Bufo bufo wrighti, is often found laying its egg-strings in slow-running water in mountain streams, and this is true also for some Chinese wood frogs, such as Rana chaochiaoensis. In the frogs with this breeding habit, the female is a little larger than the male and other secondary sex characters associated with breeding habits are only moderately developed and never as strongly so as in the running-water frogs, which include striking forms like Rana boulengeri and Aelurophryne mammata. The eggs are laid in masses in Rana chaochiaoensis and in strings in Bufo bufo wrighti but always with an amount of jelly much greater in quantity than the jelly of the eggs of the swift-water and running-water frogs. The eggs of Rana boulengeri are exceptional (see p. 267). The eggs of the frogs of slow-— running water are strongly pigmented and black or dark gray in color; the size of these eggs is generally less than that of the eggs found in the cascade and running-water habitats. Tadpoles of Bufo bufo wrighti are of the slow-running-water type, differing from the tadpoles of the very nearly allied species, B. b. gargarizans, in having a rather depressed body and a stronger tail with low thick tail crest. They are only rarely found in quiet water. There are two small depressions on the throat, which is flattened in the tadpole of wrighti. This specialization is closely as- sociated with the adaptation of this tadpole, since it can be used to adhere to the substratum in slow-running water. The tadpoles of Chinese wood frogs can adjust themselves to slow-running as well as to quiet water. The form of the body and the shape of the tail are really intermediate between those of the tadpoles of running water and those of the tadpoles of quiet water. Other quiet-water tadpoles may be found in slow-running water when their normal pools are invaded after rains by flood water that drives the tadpoles into the nearest stream. 62 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 Numerous species of frogs and toads lay their eggs in pools, ponds, ditches, or flooded rice fields. In the frogs breeding in this habitat the males are usually smaller than the females, and the other secondary sex characters are moderately developed. However, sex dimorphism in size is not as great as in the swift- water group. The secondary sex characters correlated with amplexus are less strongly developed than in frogs of the running-water group. The general phenomena connected with breeding adaptations are similar to those found in the slow-running-water group. Males are provided with nuptial asperities, as in Bufo bufo gargarizans, or nuptial pads, as in Rana nigromaculata, and usually the arms of the males are enlarged for amplexus in most species of this group of frogs. The egg-laying adaptations of the different species of the quiet-water frog vary greatly, but the general principle is the same for all, with adjustments essential for the preservation of the race. Species of Kaloula lay their eggs in temporary pools, usually near old houses or in roadside ditches. The eggs float singly on the surface of the water like drops of oil, and are very easily overlooked. The external layer of jelly has a winglike projection near the animal pole. This functions as a floating apparatus. In this surface situation the eggs get the optimum amount of light (or heat) and oxygen for rapid development, which is of great importance in shallow pools that are easily dried up in a short time. The eggs are small and dark in color, and thus have high capacity for absorption of heat. The early developmental stages—cleavage, blastulation, and gastrula- tion—are much more rapid than in any other amphibians. The species of Microhyla seem to form another ecological group adapted to breeding in quiet water. M. ornata lays its eggs in rice fields, and more rarely in small ponds or pools. The jelly surrounding the small, light brown eggs forms a film floating on the surface of the water. Without special attention, one usually fails to see the egg-film. The process of development is much more rapid than in other pond frogs or toads. This is presumably because the water bodies, in which the eggs are laid, are not permanent. The pool may dry up, or the farmer may drain the rice field for a season. The process of development, under such hazards, must be rapid in order to perpetuate the species. This condition is similar to that found in Kaloula. Many species of Rana lay their eggs in the various types of quiet water. R. nigromaculata generally goes to ponds to breed, and limnocharis and guentheri lay their eggs mostly in flooded rice fields. The eggs of the various species of Rana are mostly medium in size and dark brown or nearly black at the animal pole, with a large amount of jelly surrounding and connecting the egg envelopes to form the egg-mass. Toads, such as Bufo bufo gargarizans, lay their eggs in strings entangled with aquatic plants in ponds and ditches. The eggs are black and are protected by their jelly strings, within which each egg is enclosed by a thin layer of jelly. The species of tadpoles inhabiting the quiet water form a heterogeneous group, both in phylogenetic and in morphological features. These tadpoles vary LIU: AMPHIBIANS OF WESTERN CHINA 63 greatly as to the form of the body, the type of the mouth parts, and the nature of the tail, being derived from different families of Salientia. The convergent characters that adapt them specifically to the ecological environment of quiet water will be described below. Arbitrarily, the quiet-water tadpoles could be divided into three categories, the pond group, the pool group, and the flooded rice-field group. But the tadpoles of a single species may be found in all of these habitats. Presence in the several types of quiet water is best illustrated by the tadpoles of Rana nigromaculata; Bombina maxima inhabits ponds and pools in the tadpole stage; and the tadpoles of Kaloula rugifera are found only in pools, as distinguished from the more permanent ponds. The typical tadpole inhabiting quiet water is not exposed to strong currents and does not require a capacity for powerful swimming. These tadpoles have a stout body, moderately developed mouth parts, and a weak tail of characteristic form, provided with a high delicate crest and with a pointed or attenuate and delicate tip. The best examples of this type are the tadpoles of Microhyla ornata, Rana nigromaculata, and Rhacophorus leucomystax. Some of these tadpoles may have their body form approaching the condition suitable for running water, as in the tadpoles of Bufo bufo gar- garizans in the lowland, and Rhacophorus bambusicola at higher altitudes. The mouth parts of the quiet-water tadpoles in general are provided with moderately developed mandibles and labial teeth, but never have such strong mouth parts as those of Scutiger or Aelurophryne. The quiet-water tadpoles of Kaloula and Microhyla have no mandibles and no labial teeth in adjustment to their open- water feeding habits. Most amphibians lay their eggs in water, and this element is necessary for their development, unless they have evolved specialized protection against drying. There are, however, many salamanders and frogs that lay their eggs on the ground, in hollows in the ground, or on vegetation. The eggs of frogs in this ecological group do not develop into normal living tadpoles if they are placed in water, as I have shown by some simple experiments with the eggs of Rhacophorus omeimontis. In western China there are five such species, one of Rana and four of Rhacophorus, that belong to this ecological group. Rana adenopleura, the famous musical frog of Mount Omei, lays its eggs in small nests excavated in the banks of small pools from six to twelve inches above the water’s edge. In this species the males make the nest; two males found in new nests had bleeding snouts when they were collected. The nests are rounded or oval with a smooth inner surface and a comparatively narrow opening. The eggs, which are always laid in the nests, are small and are scattered in a large amount of jelly that is much more watery than the jelly of other amphib- ians of western China. The function of the enormous amount of jelly for this breeding habit is obvious. If there is no rain to moisten the eggs in the nests, or to wash the newly hatched tadpoles from the nests to the pool, then the jelly, which soon becomes liquefied, is the medium in which development takes place. This type of jelly seems to protect the embryos and young tadpoles while waiting 64 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 for the rain to wash them into the pool nearby. Rana adenopleura is found only in regions where rains are usually frequent, as on Mount Omei, where this frog is abundant. Its breeding season is from June to September, which is the rainy season of this sacred mountain. The breeding adaptations seem to be the chief factors in the distribution of this species. The secondary sex characters (including dimorphism in size) are poorly developed in association with the habit of laying eggs in nests. The longer snout of the male may have something to do with the construction of the nest. This character recalls the sharp spade- like snouts of males of various species of the American Leptodactylus. The tree frogs of the genus Rhacophorus present further interesting develop- ments of terrestrial breeding habits. Some species, among them bambusicola, lay their eggs in holes under stones or roots of vegetation; some lay on walls of pools (such as those of unused manure pools) or on vegetation in flooded rice fields, as does lewcomystax; and others place their eggs on the leaves of low plants, or most strikingly in leaf-nests made in trees overhanging pools or ponds, as does omeimontis. Rhacophorus bambusicola, a tree frog discovered in the Lolo country of Chaochiaohsien, Sikang, June, 1942, is characterized by short legs. Males are much smaller than females, and have other secondary sex characters poorly developed. The disks are much smaller than in other species of the same genus found in western China. The body is stout for a tree frog. All these characters have something to do with the breeding habits. The eggs are laid in holes near pools or ditches or sometimes just under wood, stones, or even among the roots of plants, wherever a space can be found, the egg jelly being whipped up into foam by the female. Since bambusicola is a ground breeder, it is not necessary for the frogs of this species to have the long legs, the large disks, or the somewhat flattened bodies adapted to the arboreal breeding habit, the extreme of which is illustrated by omeimontis. The eggs of bambusicola are rather small (2.1 mm.) and white in color; they are buried in a foam mass, the fluid secreted with the eggs being beaten by the legs of the female into foam, a little of this fluid being secreted before the eggs are laid. Eggs are scattered singly or in groups in the foam and each egg is enclosed by a thin capsule. The foam protects the eggs from drying and the liquefaction of the foam on the inside corresponds func- tionally more or less to a small pool of water. This serves to protect the embryos or newly hatched tadpoles when there is no rain to wash them into the adjacent pond or ditch, where they can metamorphose. Rhacophorus leucomystax, which has a wide distribution in southeastern Asia, is another interesting tree frog that enters western China. It could be considered as a form intermediate in the evolution of breeding habit between the short- legged bambusicola, which is a ground breeder, and the arboreal breeder, omei- montis. Males of lewcomystax are smaller than females. The average diameter of the disks of males and females is 4.2 per cent of the body length. The male has a nuptial pad developed on the first and second fingers as in bambusicola. The eggs are laid in foam masses, which may be hung on vegetation in flooded LIU: AMPHIBIANS OF WESTERN CHINA 65 rice fields, or, commonly, are attached to the walls of vertical-sided pools. If there is no chance to reach such a site for egg-laying, the egg foam may be placed on the ground anywhere near a pool. The eggs are small (1.8-2 mm.) and white in color, scattered singly or in groups inside of the foam. Each egg is enclosed by a capsule firmly connected with the foam substance. At a later stage, a yellowish layer of liquid forms the egg capsule. This is the beginning of liquefaction. The foam is essential for the development of the embryos. It also protects the young tadpoles from drying up when there is no rain. Ata certain stage of development the whole foam mass may drop into the water be- cause of liquefaction of the foam by which it is attached; or the young tadpoles may drop into the water through an opening in the foam mass. When the foam drops into the water, it spreads out on the surface as a mass of small bubbles and the tadpoles free themselves from it and swim vigorously. Arboreal breeding habits are the most extraordinarily interesting phenomena in Amphibia of western China. The beautiful tree frog, Rhacophorus omeimontis, has a bird-like voice and whistles on trees at twilight during the breeding season (April-June) to attract the females. With a flashlight one may see him standing on the small branches of trees just above a pool, and singing without distended throat. Males are much smaller than females, and both have a dorso-ventrally depressed body. The legs are long and have very large disks at the tips of the digits; the average diameter of the disks in this species is 3.2 mm., 5.4 per cent of the body length. Nuptial pads are developed on the inner dorsal sides of the first two fingers of the male. According to my observations, these frogs usually lay their eggs on leaves of trees that hang over a body of water. If there are no trees on the edge of the ponds, the frogs lay their eggs on low vegetation on sloping _ banks of small stationary bodies of water. The nests exhibit much variation, being formed by a varying number of leaves. The eggs are partly exposed and partly concealed by the leaves. There is no foam mass. The exposed portion of the egg mass is covered by a sticky secretion, by which the leaves of the nest are glued together. The eggs are large (8.5 mm.) and light green in color, lighter than the leaves of the nest; such a color may offer some protection for the eggs. The eggs are well protected by their three capsules, with two layers of liquid, the first between the egg and the inner capsule, the second between the first and second capsules. The outer layer is a white opaque elastic mem- brane. This is a porous egg coat that connects egg to egg and attaches the egg mass to the leaves. Although the jelly is not so abundant as in Rhacophorus bambusicola and leucomystax, the eggs of omeimontis are better protected by leaves, by the sticky secretion just inside of the leaves, and by the elastic outer coat of each egg. Furthermore, inside of the outer coat there are two capsules and two layers of liquid (see pl. 9). Tadpoles of different species of Rhacophorus in western China and of Rana adenopleura are found in pools and ponds on the hillsides, between 1,500 and 11,000 feet above sea level. According to the form of the body and the shape of the tail, the tadpoles of Rhacophorus leucomystax and Rana adenopleura are 66 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 interpreted as typical quiet-water forms. Their bodies are stout, with high delicate tail crests. The tadpoles of other species of Rhacophorus tend toward the running-water type, as they are very much like the tadpoles of wood frogs and toads. However, they are still found mostly in pools and ponds, rarely in ditches. On the basis of structural adaptations for breeding, Amphibia of western China might be classified into three categories. The cascade forms, Stauwrois, and the tree frogs, Rhacophorus, constitute the first category, with certain ex- ternal similarities, especially in breeding adaptations. These adjust them on one hand for cascade life, or on the other for arboreal life. The general body form is somewhat dorso-ventrally depressed in both Staurois and Rhacophorus. It is unquestionably true that the depressed body of Stawrois is associated with its life in the crevices of rocks by the sides of the mountain streams, and the depressed body may assist it in getting beneath stones for egg-laying. The flattened body of Rhacophorus omeimontis is evidently associated with arboreal life, but it also bears some relation to the arboreal breeding habits. The males of Staurois and Rhacophorus are much smaller than the females, in correlation with breeding behavior. A big male on the back of a female must offer much resistance in swift water and in getting beneath stones for egg-laying. It may be even harder for a female Rhacophorus to bear a large male at the breeding site on the tree or on the walls of pools. Furthermore, if the male of a Rhacophorus is larger than the female it is difficult or even impossible for the female to move her legs freely to beat the secretion into foam. The disks at the tips of the digits are characteristic specializations for Staurois and Rhacophorus. They are an essential adaptation for cascade and arboreal life, but they also aid the female to maintain herself while laying her eggs in swift water and on trees. The disks on the digits of bambusicola are much smaller than those of other species of Rhacophorus, possibly because they are less needed in the egg-laying process in or on the ground. In leucomystax the disks are much larger than in bambusicola but smaller than in omeimontis: these disks come into use in climbing on stone walls or among vegetation. R. omeimontis is best adapted to arboreal life, and lays its eggs on leaves of trees. The disks in this species are better developed than those of any other tree frog of western China. : The second category consists of the frogs that breed in running water. Ac- cording to the relative size of the male and female, this category may be sub- divided into two groups, in one of which the males are slightly smaller than the females, while in the other the males are distinctly larger. If the male is some- what smaller than the female, the secondary sex characters may be well developed, as in Scutiger schmidti, or may be poorly developed, as in Megophrys minor. The more interesting condition is found in the second group, in which the males are larger than females, as is the case with Rana boulengeri and Aelurophryne mammata. Secondary sex characters correlated with breeding adaptations are extraordinarily developed, as may be seen in the greatly enlarged arm, the strong LIU: AMPHIBIANS OF WESTERN CHINA 67 spines on the fingers, the spines over the entire belly of Rana boulengeri, and the two patches of spines on the chest of Aelurophryne mammata. It is not easy to explain these relations, or to see what adaptive advantage the remarkable sex differences in the above two species of Amphibia of western China may have. It is often difficult to distinguish slow-running water from quiet water. Moreover, a quiet body of water after a heavy rain may become slow-running water. In any case, the frogs of the quiet-water category have stout bodies and moderate secondary sex characters, quiet water affording suitable sites for them to breed and lay their eggs. Eggs of frogs in all the categories vary greatly in color, size, and amount of jelly. It is hard to draw any conclusion about the significance of such varia- tion. Itis true, however, that eggs attached to the under sides of stones, protected by foam or covered by leaves of vegetation, are white in color. The exposed floating eggs and the eggs in unconcealed submerged egg masses are more or less pigmented; furthermore, eggs of these types laid in early spring are noticeably pigmented and even black; and those eggs laid later in the season are usually much lighter in color. There is no positive correlation between the size of the eggs and the size of the adults. It is true that the size of the egg has a positive correlation with the nature of the body of water in which the eggs are laid. The narrow-mouthed toads lay their small eggs in temporary pools. The developmental process of these eggs is so extraordinarily rapid that they can complete their metamorphosis before the pools dry up. This is an ecological adaptation. The jelly of the eggs of Amphibia provides both biological and physical protection. From the study of Amphibia of western China, it is evident that jelly is much more important for protection against the physical environment than for protection against biotic elements of the environment as a whole. The eggs of the cascade and running-water forms have very little jelly, as they are not in danger of evaporation and their smaller amount of jelly may be thought to offer an advantage in lessened resistance to current. Although the eggs of Rana boulengeri are an attached running-water type, each egg appears to hang down individually. These eggs are protected by a large amount of jelly. This is easily explained, as part of the jelly is used for attachment and for the jelly cables that suspend the eggs. Again, as these eggs are laid in small mountain streams, usually just after rains, the water level may go down quickly and leave the eggs exposed and in need of greater protection from the jelly. The large amount of jelly in the eggs of the nest-breeding frog, Rana adenopleura, is more obviously and definitely for the physical protection of the eggs, as there is no water in the nests. In the foam masses of tree frogs the liquefying foam takes on the function of the water and jelly of the aquatic breeding forms. The foam protects the eggs, which are only enclosed by a thin capsule. The jelly may be specialized for the formation of a floating apparatus composed of cap-like pro- jections, as in Kaloula. 68 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 The most remarkable specialization in the tadpoles of western Chinese Am- phibia is the large ventral adhesive suction disk of the cascade species of Staurois. It is used as a vacuum adhesive apparatus to attach to and move back and forth on stones under cascades. The tadpoles inhabiting running water always have a cylindrical body, a strong tail, and a low and thick tail crest. All these struc- tures are necessary for effective swimming. On the other hand, tadpoles found in quiet water mostly have a stout body and a weak tail, with a high and delicate crest drawn out to a narrow pointed tip. Feeding adaptations afford other in- teresting problems in the natural history of amphibian tadpoles. The surface- feeding form has its lips expanded into a funnel to collect its plankton food; the bottom-feeding forms naturally require strong mouth parts in order to scrape off the food stuffs from the surfaces of stones, or to eat other organisms. The mouth parts of quiet-water tadpoles are moderately developed and without strong specialization. LIST OF SPECIES ~CAUDATA Megalobatrachus davidianus' Blanchard Steboldia davidiana Blanchard, 1871, Compt. Rend. Acad. Sci., Paris, 73: 79—Thebet oriental; idem, 1871, Ann. Mag. Nat. Hist., (4), 8: 212; David, 1871, Nouv. Arch. Mus. Hist. Nat. Paris, 7, Bull.: 95—Tchongpa [Chungpa], Szechwan; Gray, 1878, Ann. Mag. Nat. Hist., (4), 12: 188. Sieboldia davidi David, 1875, Jour. Trois. Voy. Chinois, 1: 326; idem, 2: 20-22, 216— Ouang-Kia-Ouan [Huang-kia-wan], southwestern Shensi. Hoplobatrachus davidi Mollendorff, 1877, Jour. N. China Asiat. Soc., 11: 105. Megalobatrachus maximus Boulenger, 1882, Cat. Batr. Grad., Brit. Mus., p. 80 (part, China; not of Schlegel 1837); Boettger, 1885, Offenb. Ver. Naturk., 24-25: 166. Megalobatrachus japonicus Barbour, 1912, Mem. Mus. Comp. Zool., 40: 125; Stej- neger, 1925, Proc. U. S. Nat. Mus., 66, Art. 25: 3. Megalobatrachus sp. Despax, 1913, Bull. Soe. Zool. France, 38: 134. Cryptobatrachus maximus Stanley, 1915, Jour. N. China Asiat. Soc., 44: 14. Megalobatrachus sligoi Boulenger, 1924, Proc. Zool. Soc. London, 1924: 173—Hongkong. Megalobatrachus japonicus davidi Chang, 1935, Bull. Soc. Zool. France, 60: 347; Chang, 1936, Contr. Etude Morph. Biol. Syst. Amph. Urodéles Chine, p. 82. Megalobatrachus japonicus davidianus Pope and Boring, 1940, Peking Nat. Hist. Bull., 15, pt. 1: 18; Freytag, 1943, Wochenschr. Aquar.-Terrarienk., 4: 88, figs. History of species.—Megalobatrachus davidianus was regarded as a lizard in the books of the pupils of Confucius. In these books, ‘‘salamander,” “lizard,” “skink” and “‘gekko” are regarded as four names for the same kind of animal, and applicable reciprocally to one another. One can also find there the word “Nei-yu,”’ the Chinese name still used to designate this giant salamander. It is described as follows: “In the streams of Ille-tzu, Shansi Province, there are found numerous Nei-yu; their form is similar to that of a serpent; the largest one is called Sha-yu.” This passage is repeated in Shan-hat-ching (‘“Documents of the sea and mountains”), which was written by an anonymous author about 600 B.c. About 100 B.c., Tze Ma-chien, the most celebrated author of the Han Dynasty, remarked that the giant salamander also existed in the province of Shensi, where it was called Jen-yu (‘‘man-fish”’ or ‘‘a fish like a man’). Koo Po (A.D. 300) illustrated the salamander in Er-ya, and described it in the follow- 1 For further references, see Chang, 1936, p. 82. Throughout the paper, synonymies are in abbreviated form, giving only the essential references. 69 70 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 ing words: ‘‘Nei-yu shows nearly the form of the catfish with four feet. The anterior feet resemble those of the monkey, the posterior ones those of the dog. It emits cries similar to those of a small infant; the ones a size of 8-9 feet are called Sha-yu.”’ About A.D. 840 Tau-haun-tsin, a naturalist, noted the presence of this salamander in the provinces of Hunan, Hupei, and Anhwei. Five hundred years later, Yan-fu noted it from Kwangtung Province in his Documents on rare ob- jects, from which Li-fang and his collaborators cited the following passage in compiling their encyclopedia of the Sung Dynasty, “Tai-ping-yu-lan: Nei-yu has four feet, the form of a fish, walks slowly by means of its feet, approaches the catfish in size, lives and grows in the mountain streams.” About A.D. 1120 Koen-tzeun-pi dissected this creature for the first time. He describes it as follows: ‘‘Nei-yu, the giant salamander, has a form similar to that of the otter, four feet, a belly hanging like a heavy sack, without scales like the catfish; its body has a purple color; the stomach contains small fishes, small crabs and small stones. I have opened it and determined the food.”’ But he placed it in the same class as the fishes, and he did not distinguish it from the lizard. Tsai-yuen-dou (1070) described it more correctly. In Mau-tze-mine-fu-chia (‘Interpretation of the objects mentioned in Shih-ching’’) he writes: “The salamander, although it closely resembles the lizard in its general form, may be distinguished from the latter by its habitat. Those that live on land are lizards; those that live in the water are salamanders, belonging to the Pisces.”’ In 1569, Li Chi-chen mentioned the existence of the giant salamander in the provinces of Szechwan, Honan and Shensi. It was not until 1837 that Europeans suspected that the giant salamander might be found in China. In the Fauna Japonica, Temminck and Schlegel (p. 135) state that de Guignes (1813) mentions the ‘‘Nei-yu” in his Chinese Diction- ary, page 895. This supposition was confirmed when Abbé Armand David dis- covered this animal in western China in 1869. The exact type locality of Megalobatrachus davidianus has never been de- finitely recorded. In Emile Blanchard’s original description of the species, which, by the way, properly lists the difference in the tuberculation of the head and the length of the digits among the specific characters, the type locality is given only as western China, qualified as follows: “La gigantesque Salamandre vit sur les frontiers du Celeste-Empire dans des eaux claires et limpides qui descendent des montagnes du Khou-khou-noor.” It may, however, be possible to ascertain more exactly the place where Abbé David obtained his specimens. The giant salamander was collected during David’s second voyage, which lasted from May, 1868, to June, 1870. His epochal stay at the Catholic mission- ary station in the then independent principality of Muping occupied most of the year 1869. He left Chengtu on February 22 and arrived at the station, now referred to as Paohsing, six days later. With the exception of a period from August 30 to September 27, when he was obliged to go to Chengtu on account LIU: AMPHIBIANS OF WESTERN CHINA (et of ill health, the rest of the year was spent at Muping, where he made the famous collections that startled the scientific world. By the end of November, 1869, he was back in Chengtu. He has recorded in considerable detail his doings from day to day while at Muping (Nouv. Arch. Mus. Hist. Nat. Paris, 10), but, although he mentions the various novelties as they were collected, nowhere is there any mention of the giant salamander. It may therefore be safely concluded that it was not obtained at Muping, for he would certainly not have failed to record such a startling find. Moreover, we have his positive statement (Nouv. Arch. Mus. Hist. Nat. Paris, 7, Bull.: 97) that it was obtained during his trip to eastern Kokonor (Tsinghai). He says: ‘‘Mon excursion du Kokonoor oriental m’a fourni quelques nouveautés. J’en ai rapporté la grande salamandre (laquelle vit aussi au Setchuan, au Tibet et au Yunnan, d’aprés mes informations).” Of this excursion to eastern Kokonor I have been unable to find any detailed account, although he promised to publish one. We only know that after a short rest at Chengtu he left on a trip to the northern mountains on December 26, 1869, arriving at the “eastern corner of Kokonor’”’ on January 7, 1870. Shortly after his arrival he was taken ill with the cholera, which stopped his further progress. He returned by way of Lunganfu to Chengtu, where he arrived at the end of March, 1870. He must have obtained the giant salamander during this return voyage. During his third voyage to China he again obtained specimens, this time in the province of Shensi. In relating this fact (Jour. Trois. Voy. Chinois, 2, 1875, p. 20) he says that they were “bien les mémes que le Sieboldia Davidi, de Tchong-pa.”’ This is the first and only reference known to the type locality of M. davidianus. On the road travelled by David from Lunganfu, halfway between that city and Chengtu, the maps show a district city, Chungpa, on the river known as Fowho, and there can be little doubt that this is David’s Tchong-pa and the specific type locality of the Chinese giant salamander. Various authors have concluded that the Chinese and Japanese giant sala- manders are identical, while others have considered the Chinese form to be a subspecies of the Japanese Megalobatrachus japonicus. I have studied living specimens of the Chinese form in their natural condition near Yaan, in Sikang Province, and preserved specimens of both Chinese and Japanese giant sala- manders in Chicago Natural History Museum and the United States National Museum. The manuscript notes of Stejneger have been available to me, thanks to Dr. Doris M. Cochran, and Mr. Clifford H. Pope has turned over to me his own notes as well. His comparative study indicates that the Chinese and Japanese forms are specifically distinct. The tail of M. davidianus is proportionately longer than the tail of M. japonicus. But the most distinct character that separates the two species of Asiatic giant salamanders is the nature of the tuber- cles on the head. The tubercles of M. davidianus (fig. 9) are mostly in pairs, and are much smaller and fewer than those of M. japonicus (fig. 10). The tubercles on the throat differ even more and are characteristic for each species. In M. davidianus, the very small paired tubercles are arranged in rows parallel with 72 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 the lower jaw; in M. japonicus they are mostly single and large and irregularly scattered. The central position of the throat is free from tubercles in both species. Although Chang (1935) mentioned these differences, he intended to keep the Chinese form as a subspecies Megalobatrachus japonicus davidi. All the Japanese specimens have the top of the head densely tuberculated, especially around the eyes; only a narrow space down the middle of the inter- orbital region is smooth. Even in the youngest specimen (No. 6163), in spite of its soft condition, the tubercles are quite perceptible. In the largest Chinese specimen (No. 69456) the whole median part is smooth, as is also a considerable space around the eyes; Nos. 52409 and 16801 from Yaan are smaller, but the tubercles are similar. In No. 69347, which is only slightly smaller than the largest, the whole top of the head in front of the eyes is perfectly smooth and there are only a few scattered tubercles on the sides of the postorbital region. Finally, in No. 65454, also from Yaan, a somewhat younger individual than the smallest Japanese specimen, but in an excellent state of preservation, the skin of the head is absolutely smooth, without trace of tubercles. The most conspicuous differences between the two forms are well shown in figures 9 and 10. Mr. E. G. Boulenger (Proc. Zool. Soc. Lond., 1924, pt. 1, p. 173) has de- scribed a living specimen of uncertain type locality, probably Chinese and possibly from the mountain regions of northern Kwangtung or Kwansi, as Megalobatrachus sligot. It has the characters that distinguish M. davidianus from M. japonicus and is evidently related to the former, not to the latter. Boulenger differentiates M. sligoi from maximus (under which he includes both japonicus and davidianus) by its longer, flatter, and smoother head, and by the supposed lesser distance between eye and labial border. Neither the length of the head (which is difficult to define, and which was not defined by Boulenger) nor the relative distance of the eye from the labial border are valid characters. Boulenger’s type specimen, however, agrees in every essential, especially in the nature of the tubercles, with specimens from the upper Yangtze Valley. I regard the Hongkong locality as certainly representing a transported specimen. This animal is eaten, particularly by the Cantonese, and is known to be transported for this purpose. Stejneger’s manuscript notes show that he had reversed his decision of 1925 and reached the conclusion that the two giant salamanders are distinct species. - He wished to retain M. sligot as possibly representing a third form with a more eastern range in China than that of davidianus. Pope and Boring, following Chang, have placed sligoz definitely in the synonymy of davidianus. They have used the trinomial forms M. j. davidianus and M. j. japonicus, because the two forms can be regarded as geographically representative. There is of course no possibility of intergradation between forms separated by more than a thousand ‘In No. 15098, from Yaan, there are only a few large paired tubercles on the postero- dorsal a of the head, a few very small ones on the snout, and a large smooth area on top of the head. 2.0. Ex a5 hak °o se G .- 8S om os y : See Rees ge Af Fic. 9. Megalobatrachus davidianus (X 34). view of head. A. Dorsal view of head. B. Ventral Fic. 10. Megalobatrachus japonicus (X 34). view of head. Drawn by Margaret Bradbury. 73 A. Dorsal view of head. B. Ventral 74 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 miles. It seems preferable to me to recognize their long separation in both time and space by referring the two forms to distinct species. Original description.—‘‘Among the objects collected by David after his departure from eastern Tibet we had the skin of the great batrachian. It was important to compare the salamander from China with that from Japan and to ascertain whether the two animals were the same species or different species. Comparison leaves no doubt: The salamander reported by David, very close to the salamander discovered by Siebold, is distinguished by several very apparent characters. It has, on the head and anterior part of the body, tubercles less confluent and regularly arranged so as to form lines and very definite patterns. In this way, the eye is as if enclosed by a double row of tubercles which, on the internal side, become angular like a widely open V. In the Japanese species the tubercles present, on the contrary, only a confused arrangement. The Chinese species also appears to us to have the digits and the four limbs a little longer, and we believe that the general color of the body is more black, but the imperfect state of preservation of our single specimen prevents us from insisting on several details. We give to the great salamander of western China the name of Sieboldia Davidiana, which will recall once again the memory of the admirable explorer of China, Mongolia, and Tibet. The gigantic salamander lives on the frontiers of the Celestial Empire in the clear and limpid waters that descend from the mountains of Khou-Kou-Noor. It appears that it reaches an enormous size; David reports that he took specimens of which the weight is 25 to 30 kilo- grams. One understands that such animals are a valuable food resource for the inhabitants of the country.” Description.—A specimen collected at Fuhsienkwan, Yaan, Sikang, by Floyd Tangier Smith, Chicago Natural History Museum No. 16801, serves well for redescription. Form very robust; head strongly depressed, snout obtusely truncate; nostrils small, rounded, close to the edge of the upper lip and at the corners of the truncated snout, the internasal space less than half of the interorbital space; eye small, rounded, dorso-lateral in position, and without eyelid; top of head from interorbital region to snout more or less flat, with a rounded temporal protuberance above and behind each eye; angles of the mouth about at the level of the middle of the temporal protuberance; vomerine teeth in an arched series starting between the choanae, parallel to the maxillary and premaxillary series; a thin lower labial fold starting about midway between nostril and eye to the angle of the mouth. Trunk less depressed than head, with about fifteen costal grooves (in most cases not very distinct), a strong vertebral groove, and strong lateral dermal folds. Foreleg short, dorso-ventrally flattened, with dermal fold at the posterior side, fingers four, 3-2-4—1 in order of length, the outer one flattened and widened with a dermal fold continuous with the dermal fold of the arm, the other three LIU: AMPHIBIANS OF WESTERN CHINA 75 Megalobatrachus davidianus: Measurements U.S.N.M. U.S.N.M. C.N.H.M. C.N.H.M. U.S.N.M. No. 69456 No. 52409 No. 15098 No. 16801 No. 65454 Totallength* esas eee eS 1056 125 695 660 240 Body: length. stein eee cis 621 436 385 415 150 Head width at angle of jaw... 115 95 96 84 30 Ratio to body length (%)..... 10.8 10.8 13.8 18.7 12.5 Internasal’ space sin. sci. ote 23 20 18 19 7.5 Ratio to head width (%)..... 20.0 21.0 18.7 22.6 25.0 Nostril to edge of lip......... 5 4.5 5 455 255 Ratio to head width (%)..... 4.7 4.8 5.2 5.3 8.3 Interorbital space............ 76 51 46 44 16 Ratio to head width (%)...... 66.0 52.6 47.5 5223 53.3 Eye to edge of lip............ 18 10.5 13 10 5 Ratio to head width (%)..... 15.6 11.0 13.5 11.9 16.6 Tip of snout to eye.......... 56 38 37 35 13 Ratio to head width (%)..... 48.6 40.0 38.5 41.6 43.3 Length of foreleg............ 102 74 75 64 29 Ratio to body length (%)..... 16.4 17.0 «(19.4 15.4 19.3 Length of longest finger....... 31 22, 21 15 8 Ratio to body length (%)..... 4.9 5.0 5.4 3.6 bes Length of hind leg........... 140 93 88 84 33 Ratio to body length (%)..... 22.5 21.3 2228 20.2 222, Length of longest toe......... 36 29 yAL 21 8 Ratio to body length (%)..... bad 6.6 5.4 5.0 5.3 Length from axilla to groin... 300 205 DA 230 73 Ratio to body length (%)..... 48.0 47.1 60.0 55.4 48.6 . Eength or taller ae 435 290 310 245 90 Ratio to body length (%)..... 70.4 66.6 80.0 59.2 60.0 fingers less flattened and with rounded tips; no webs between fingers; tips of fingers light brown, cornified; and a large rounded cornified pad at the bases of the first and second fingers. Hind leg short and flattened, with a strong dermal fold at the back and very loose skin folded on the ventral side except for the distal part of the leg; toes flattened with rudimentary webs, 3-4—-2-5-1 in order of length, the outer toe with the strongest dermal fold continuous with the dermal fold of the leg; the second and third toes fringed on their outer edges, the fourth toe more strongly fringed; fringes developed on the inner sides of the two outer toes; tips of toes rounded, with light brown cornified epidermis; a round, brown cornified pad at the base of the first and the second toes. Tail short, 59 per cent of the body length, compressed, nearly rounded at the base and thin on last one-fourth; dorsal tail fin extending to the trunk, the first half thick with a dull edge and the distal half thin and sharp-edged; vent slit-like, just posterior to the base of the leg. Skin rough and porous, with numerous wrinkles, folds, and tubercles. The largest and most prominent tubercles with characteristic arrangement in the Sieg 0°0S aa 9ST G°3g €°0S raat LOT eg Tey, IT &I G° 3s 3°61 8h 09 9°F 9°F OT S°FT L’8T 9°9T OF eg GLY TOV 61 LZ Sor S°Sr g 8 0's T'sg 63 vé 39 0° G'S oF 6 1Z 9°81 g°8 rai € OI 9°ST OF v9 &1Z alg Gos 897 €9T9 LIGVE WNSQ WNSND 067 6PETT 667 €vg9 Ets LLY 8°&¢ €°3S 008 802 G06 186 062 v'Lv | g°3g 6°97 006 £06 SES 082 092 rs ey COM] o°g vg 02 LT 6S 83 0€ €°06 6°12 T'6T 6°8T 6L §8 &6 c0T SOT vv UP ov 9°7 8b Seo LT 8T GZ LZ o ST LT €°9T L°8t v9 69 &L 18 vor &°&P 0°07 STV 8°98 v& 68 8& 9v 6S €°&T G°Or 0°0T ama IT él él IT 9T 9.99 9°&S 6 LS €°0¢ cv 0g Tg 69 TL LL t'9 LS 8°9 OL 9 > a GL OT | 9°9T 6°91 T'vT cT ST 9T 8T 06 6°91 iS vst SLT LL 06 G6 OIT TT OLE LOV &6P €&¢ vg OLS gT9 G39 068 As) 9TZvE STSVE VISE 61869 69L9¢ sjUsWeInsea fT :snavwodnl snyrn.synqojnba yy 1°08 CRO en ewee ee (%) y4Bue] Apoq 07 oney Oke ccc nes [req Jo yySuery 6°9S ba ia ee atari (0/4 yqsue, Apog 03 o1ey og oc ulo 13 0} BlIxe Woy yysuey Oce per haete sey: (%) y43u9] Apoq 0} oryey 12 aAaces cha laictuatavetene Gat iat 30} 4yseBuo] jo yysue'T Wg) ti Amin acre Sg ek (%) y3u9] Apoq 04 oryey 221 © Sy ayo las0: 0) Ae, BL SUSE aL eee 08's 39] pury jo yysue'T Le $0.6 So sy al. svete © Lois (%) yy sue] Apoq 04 oney be ee eS SD Lata Jo3uy yso3uo] jo yysue'T CNA di ah teh acer (%) y43u9] Apoq 0} o1rjyey FIT o ates Wiel mye sl aite a Vel ee) Wiese tae 89210] jo yqsue'T FiO tes aneey Tt (%) YUypIA peey 0} orjyey 6S re Tk Tat at ea aa SW EN ou Para eee afo 04 qynous jo diy, A | GD eg cca (%) Up pray 07 o1ey mat neeeMsnalehelits cut rote tere sr ahetthe diy jo a3pe 0} CYNG | 8° 9F 61. Rpg easigl meus ls 5.2 (%) pia peoy 0} oryey 28 ara 0.6, 0" 0.6 eT al Te 16a hee cane) ores goeds [@}1q40.10} UT 1: ee aR (%) YPM peasy 0} Oey 6 so ie Oxe, @ erases. 458 ps dy jo aspe 0} [L1qSON PST So) -priia ie 1's) tea tn eta eatiel ve (%) pia peoy 0} o1yey 12 Ct ie at A Se Ot Cnt ee Me ee goeds [eseue}UT Got Geeta aa a eae (%) y4Bue] Apoq 04 o1ryey CLT elone te eneh Roe teta te wel jo a]sue ye Yapia peo Gag eee? (quoa 0} ynous) Yq 3ue] Apog 086 SE MOR Ds te es Da ee ee ee te Ve hae ee yysue] [?70.L 9ESTE WNS WNSO WNSf WNSO WNSO WNSO WNSN WHNO 76 LIU: AMPHIBIANS OF WESTERN CHINA AG, head; large tubercles scattered mostly in pairs on the postero-dorsal side of the head, with a central median long smooth area from the top of the snout to the neck region; smaller tubercles arranged more or less distinctly in rows and in pairs on the dorso-lateral sides of the head, the smallest tubercles near the margin of the upper jaw. Throat provided with even smaller tubercles, arranged in regular pairs and in rows parallel to the lower jaw, and a very regular row of small paired tubercles near the edge of the lower jaw from one corner of the mouth to the other; the central large portion of the throat free from tubercles. Paired tubercles scattered on the body, especially along the lateral dermal fold, but not on the limbs and the belly; a few paired tubercles scattered on the sides of the tail, with a row of paired tubercles in the lateral groove of the tail. The prominent lateral dermal folds originate at the side of and below the temporal protuberances, con- tinuous with the lateral parts of the upper jaw, and extend caudally, ending on the postero-dorsal sides of the hind legs. Color (in alcohol) Saccardo Umber, irregularly blotched and marbled with dusky spots, limbs similarly spotted, tips of digits light brown. Hynobius shihi sp. nov. Type.—No. 49384 Chicago Natural History Museum, from Chihsinling, Tachangsze, eastern Szechwan, China. Adult female, collected July, 1938, by Pei-nan Shih. Diagnosis.—A distinct Hynobius with vomerine teeth in two arched series convex antero-laterally and not meeting; black cornified claw-like digital tips with sharp edges curved downward; tail with thin dorsal and ventral tail fins, thin and very pointed at tip; light brown cornified epidermis on palms and soles. Apparently reaches a large size before transforming (fig. 11). Geographically nearest Hynobius chinensis, from which it is sharply distinguished by the characters above. Description of type-——Body rather stout, somewhat depressed, with ten costal grooves. Head length 27.5 per cent of the length of the body from snout to vent, its width 19 per cent of the body length, and its height 10 per cent of the body length; snout very much depressed and somewhat squarish, with nostrils near the antero-lateral corners; internasal space greater than the length of the eye and about twice as wide as the interorbital space; angle of the mouth just back of the posterior corner of the eye; eyes dorso-lateral in position; sides of jaws flattened out laterally, with poorly developed dorsal and ventral labial folds; skin on the throat very thin and colorless with many longitudinal folds, gular fold distinguishable; small black gill stumps forming a knob covered with skin just back of the sides of the head. Limbs well developed, the digits over- lapping when the arms and legs are appressed, fingers 3-2-4—1 and toes 3-4-2-5-1 in order of length; palms and soles covered with light brown cornified epidermis; digital tips covered with black cornified claw-like epidermal structures with sharp edges curved ventrally, fingers and toes flattened, with thick lateral fringes. Tail 78 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 shorter than the body, posterior half laterally flattened and distally with very thin dorsal and ventral fins, and with a very thin pointed tip. Vent a lon- gitudinal slit with swollen lips, much pigmented at the antero-lateral mar- gin. Vomerine teeth in two arched series convex forward and outward, eight teeth in the left series and seven teeth in the right, beginning just inside the choanae, curving forward and then toward the mid-line, but not meeting. Premaxillary teeth and the teeth at the anterior region of the maxillae much Fic. 11. Hynobius shihi. A. Adult (x 1). B. Oral cavity (x 3). C. Ventral view of hand (x 4). D. Finger, showing cornified tip. E. Ventral view of foot (x 4). larger than the post-maxillary teeth. Color on the back gray, densely and un- evenly stippled with dark brown, belly lighter, throat colorless. Paratypes.—Three males, two females, and three larvae (Liu coll.) were collected at the same time and place by Mr. Shih. No. 371, a female, resembles the type except in having a tubercle at the base of the fourth finger, a large pad at the base of the first toe, and a small tubercle at the base of the fifth toe; the vomerine teeth are like those of the type except that the number in each series is nine. No. 397, adult female, has light brown cornified claw-like digital tips, seven with the black cornified part lost; two tubercles on the left palm, one at the base of the first and the other at the base of the fourth finger; tubercle at the base of the fifth toe distinct; vomerine tooth series as in type but with teeth 8-10. In No. 382, adult female, the tubercles and cornified condition on the palms and soles are as in No. 371; vomerine tooth series shorter, more widely separated, teeth 6-7. In two adult males collected from the same locality the tubercles and cornification of the palms and digits are as in No. 371; in No. 387 the two vomerine tooth series are narrowly separated, teeth 9-9; in No. 383, light brown cornified epidermis covers the palms and soles, all the digital tips are capped by the black claw-like structure of the type, and each palm and sole is provided with two distinct tubercles; vomerine teeth 10-10. LIU: AMPHIBIANS OF WESTERN CHINA to Larvae.—Three larvae, Nos. 344, 377, and 380, ready to transform, range in length from 81 to 82mm. The description of the larva is based upon specimen No. 344. Most of the external characters are similar to those of the adults, except as to size, presence of gills, and condition of eye and mouth. The propor- tionate length of the tail of the young is much less than in adults, about 78 per cent of the body length, whereas in the adult it is about 88 per cent. The eye of the larva is covered by a transparent membrane connecting with the developed eyelids. The angle of the mouth reaches a point opposite the middle of the eye in the larva and to a line at the posterior corner of the eye in adults. There are three pairs of external gills developed on the sides of the neck. The gill filaments are in a stage of absorption, being black and shrivelled and more or less entangled on their thin low rami. There are four gill slits: the first is the largest, with the antero-median edge of the slit at the level of the angle of the jaw, covered externally by the gular fold; and the last is the smallest, with a slightly developed transparent membrane at its caudal edge. The size of the second and the third gill slits is about two-thirds of the first, and each is pro- tected by a colorless membrane on the latero-ventral side of the second and third gill arches. On the postero-inner margin of the first gill arch there is a row of five gill filaments; the second gill arch has two rows of filaments, five in each row; the third gill arch has two rows, four in each row; and the fourth gill arch has four filaments in a single row on the antero-inner margin. Discussion.—Hynobius is the most generalized genus of salamanders and is widely distributed in Asia, especially in the northeastern region. The four species in China are Hynobius leachii, in southern Manchuria and Korea; H. key- serlingit, in Outer Mongolia and northern Manchuria; H. sonani in Formosa; and H. chinensis in southeastern China, in Hupeh, Chekiang, and Fukien. The type locality of chinensis is Ichang, Hupeh. Mell (1929) states that Pratt, who sent specimens labeled Ichang to the British Museum, collected his material in the mountains south of the Yangtze River near Changyang. The type locality of the new species in eastern Szechwan is not very far from the type locality of chinensis, but the new species does not appear to be directly related to chinensis. Hynobius shihi can be distinguished from all other species of the genus in China by the characters named in the diagnosis. The claw-like digital tips are very similar to those of Onychodactylus, but shihi differs in having well-developed lungs and a short compressed tail with delicate tail fins and slender tip. The larva of Hynobius shihi has a maximum size of 82 mm., while in H. chinensis the maximum size is 62 mm. (Pope, 1931). The larva of chinensis has a very large head, short body, and high tail, while in the larva of the new species the head is not so conspicuously large, the body is long, and the tail is low. There is a black, fine, horny line on the inner margin of the lower jaw in larval chinensis but not in shihi. The general form of the larva of shzhi is very much like the larva of Batrachuperus pinchonii or B. karlschmidti. This new species of Hynobius is an interesting addition to the amphibian fauna of Asia, especially on account of its morphological retardation. Dunn 80 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 (1923) described Hynobius retardatus, in which some specimens have the vomerine tooth series not meeting in the median line, which he regards as a larval condition. In H. shihi, all the adults in hand have separated vomerine tooth series. Still more interesting is the fact that the gill stumps are still found in the adults, covered by skin. In order to check sexual maturity, two females were opened and both were found to have matured ova. Besides the matured ova as the check of sexual maturity, the four female specimens have the antero-lateral lips of the vent greatly pigmented and swollen, while the female larvae have slit-like vents without pigmentation. H. shihi represents a further change in the direction of neoteny and eventually may prove to be truly neotenic, producing eggs in the larval stage. Hynobius shihi: Measurements of Type and Paratypes No. 49384 (type) No. 371 No. 383 No. 387 Female Female Male Male Ratio Ratio Ratio Ratio to body to body to body to body Measurements length length length length mm (%) mm (%) mm. (%) mm. (%) Bodyslength 3.4 osrece sca 69 tere Ul eee ee) seers kOO PS Headilength. 2235S .052 cn. 19 Dhcor 220 2120) LE 29.3) 219 29.2 Head: widthia.=.... 2 en 13 1828 S15 20:2 ~12 20.7 138 20.0 Trunk: Jength:< 26532534: 31 44.9 38 50.1. 29+ 50.0 27 41.5 Length of foreleg........... 1.5. 2b.0) 19 25.6: 16 20.5, 17 26.1 Length of hind leg......... 22 3128) 6 222 30.9 20 34.4 21 32.3 Parliength ieee, eye eta 62 89.8 65 87.8 62 89°6...50 87.8 aU Width see ss fae eae tan 1058 20, 101 6:5. 4L1.2 1.6 GINS batheight=200 ee 8.0. iE fos ela 8.0 18.7 920) 21328 Genus BATRACHUPERUS Boulenger Batrachuperus Boulenger, 1878, Bull. Soc. Zool. France, 3: 71 (monotype, Salamandrella sinensis Sauvage); Barbour, 1912, Mem. Mus. Comp. Zool., 40: 126. Batrachyperus Boulenger, 1882, Cat. Batr. Grad., Brit. Mus., p. 37. KEY FOR IDENTIFICATION OF SPECIES OF BATRACHUPERUS I. Well-defined tubercles on palms and soles. A. Tail longer than body, without brown horny epidermis covering its tip; palms and SOLES. WIGHOUT COPNINCALIONS 1: cree ioe ride esi eae rae Calera ae tee yenyuanensis. AA. Tail equal to or shorter than body, with brown horny epidermis covering its tip; palms:and!solesicornified’).2 s5-20 Siok see ra ne eee ees pinchonit. II. No distinct tubercles on palms or soles. A: No labial; fold; skin' of the throat pitted? «2.2.2: Yo ceca kata cochr anae. AA. Labial fold present, skin of throat not pitted. B. Coloration uniform, without distinct spots or marbling; size large. ; karlschmidti. BB. Body finely mottled or with distinct dark spots.................. tibetanus. General Account of Batrachuperus Several species of Batrachuperus, found in the high mountains of Szechwan, Sikang, Tsinghai, Kweichow, and Kansu, can be demonstrated as a result of LIU: AMPHIBIANS OF WESTERN CHINA 81 my collections and studies. The first species of the genus was described as Dermodactylus pinchonti by David in 1871 from specimens collected by himself in 1870 at Muping, Sikang. In 1878 Boulenger studied the same material and named the genus Batrachuperus, using the name sinensis, of Sauvage, who had renamed the same Muping specimens as Salamandrella sinensis in 1877 (Dermo- dactylus, type scutatum, is a synonym of Hemidactylium). In 1925 Schmidt described Batrachuperus tibetanus from the “Tibetan border of Kansu, at about 33° N. Lat.”” Chang (1936) and Pope and Boring (1940) placed B. tibetanus in the synonymy of B. pinchonti, which name had been shown by Stejneger (1925) to be earlier than sinensis. In 1945 I published the life history of B. pinchonii, as the result of my field studies during eleven collecting trips in the mountains and high plateaus of western China, including Szechwan, Sikang, Kansu, Tsing- hai, and Shensi. In this paper I suggested that there are at least two more species besides B. pinchonii, one the form collected from Yenyuanhsien, Sikang, and the other the smaller spotted one from Lifanhsien, Szechwan, and from Muping, Sikang. Since studying my collections with the library facilities of Chicago Natural History Museum, and discussing them with Dunn, Schmidt, Pope, and Netting, and after examining the additional material in Chicago, Washington, New York, Cambridge, and Pittsburgh, I find that there are no less than five species of Batrachuperus. The smaller spotted form from Lifan, Muping, and Mount Omei fits the original description and in fact all of the descriptions of the types of B. pinchonii. Boulenger (1882) figured the type. The larger and uniformly colored form that I had identified as B. pinchonii in 1945 differs from pinchonii in lacking (1) conspicuous spots or marblings; (2) light brown horny epidermis covering the palms, soles, digits, and ventral lower parts of the arms and legs; and (3) tubercles on palms and soles. Also, it has shorter legs. This new species I name B. karlschmidti after Mr. Karl P. Schmidt. The long and weak-tailed form taken in Yenyuanhsien is a distinct species, as I suspected, and is here named B. yenyuanensis; a specimen collected from Muping, together with pin- choni and karlschmidti, lacks the labial fold, has much stronger jaws and stronger vomerine teeth, and is named B. cochranae after Dr. Doris M. Cochran. B. tibetanus is different from the other species, and may be regarded as a further distinct form, known only from the types. The West China amphibian fauna is complicated by the complexity of the ecological and topographic conditions. It seems at first unbelievable that five species of salamanders of the genus Batrachuperus should occur in the mountains and the plateaus of western China. This wealth of species is similar to that of the amphibian fauna of the Appalachian Mountains of the eastern United States. In the Appalachians one may find two or three species of Desmognathus in the same mountain stream. In our experience it is difficult to distinguish these species, especially from a few preserved specimens. Much more material of Batrachuperus from western China is desirable, and further studies in the field are required to clarify the systematics and ecology of these closely related forms. 82 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 Complicated speciation may be discerned in five groups of amphibians in western China: the aquatic salamanders of the genus Batrachuperus (Hynobiidae) ; the cascade frogs of the genus Stawrois (Ranidae); the high plateau frogs of the genus Aelurophryne (Pelobatidae); the montane Scutiger (Pelobatidae) ; and the tree frogs of the genus Rhacophorus (Rhacophoridae). Among these five groups it is easier to distinguish the species of the arboreal or more terrestrial types than those of the aquatic forms. Thus, it is difficult to identify the species of Batrachuperus and Staurois, easier to separate the forms of Aelurophryne, and easiest of all to distinguish the species of Scutiger and Rhacophorus. The center of speciation of Batrachuperus is in the mountains around Muping, Sikang, and around Wenchwan, Weichow, and Lifan of western Szechwan. This is also the center for the species of Stawrois. Batrachuperus pinchonii and B. karlschmidti are widely distributed and more abundant than other species of the same genus. Batrachuperus cochranae is a more specialized form, with much bet- ter developed vomerine and maxillary teeth and also with stronger mandibles. In Batrachuperus cochranae the labial folds are lacking in old adults but are indicated in somewhat younger specimens. This fact clearly indicates that B. cochranae is the most specialized form in the genus. The long tail and very thin and high tail fin of B. yenyuanensis represent a special direction of evolution, with adaptation to quiet water. The other species of the genus are found in mountain streams. The rich ecological and complex topographic conditions are the chief factors influencing the speciation of such a group of cold-adapted salamanders. Further investigation may reveal still more forms, or may indicate that one or more of the forms here described do not warrant recognition. The specimens of Batrachuperus in Fan Memorial Institute of Biology, Pei- ping, collected from Maha, Kweichow, and referred to pinchonii by Pope and Boring, may be supposed to represent still another undescribed species. Batrachuperus pinchonii David Dermodactylus pinchonit David, 1871, Nouv. Arch. Mus. Hist. Nat. Paris, 7: 95— Muping. Salamandrella sinensis Sauvage, 1877, Bull. Soc. Philom. Paris, (7), 1: 117—Muping (same types). Batrachuperus sinensis Boulenger, 1878, Bull. Soc. Zool. France, 3: 72; Dunn, 1923, Proc. Amer. Acad. Arts Sci., 58: 520. Salamandrella keyserlingii Bedriaga, 1898, Wiss. Res. Przewalski Central Asien Reisen, Zool., 3, (1): 3. Batrachuperus pinchonii Stejneger, 1925, Proc. U. S. Nat. Mus., 66: 5; Chang, 1932, Contr. Biol. Lab. Sci. Soc. China, Zool. Ser., 8: 142, figs. 3-4; Chang, 1936, Contr. Etude Morph. Biol. Syst. Amph. Chine, Paris, p. 77; Pope and Boring, 1940, Peking Nat. Hist. Bull., 15, pt. 1: 17; Liu, 1945, Jour. West China Border Res. Soc., 15, (B): 44, figs. 1-11. History of species.—Batrachuperus pinchonti was long a poorly defined species because it was known only from the types and only from preserved material. The complicated history of its synonymy has been set forth above. LIU: AMPHIBIANS OF WESTERN CHINA 83 David (1871) described the color of Batrachuperus pinchonii as “ranging from mottled brown to gray-yellow mottled with brown, with numerous blackish spots. Belly lighter.”” Sauvage (1877) stated that the body is gray, slightly pinkish, with brown spots more or less in rows, varying from specimen to speci- men. Boulenger (1882a) gave a much better description of the types as follows: “greyish brown above, more or less variegated with blackish; lower surface lighter, mottled with brown; the limbs being adpressed along the body, the median fingers and toes meet.”’ He did not mention the light brown horny epidermis that covers the tips of the fingers and toes and extends on the ventral surfaces of the digits and on the palms and soles to the base of the arm and leg. His figures, however, show this character very clearly. Pope noted this light brown horny epidermis when he examined the cotypes in London. Chang (1936) states that the color of the types in alcohol is generally grayish or olive brown, mottled above with blackish spots and marked with denser specks below; the horny palms and soles are intense brown, deeper at the tips of the digits; and the crest of the terminal quarter of the tail is brown. In 1945, in my paper on the life history of Batrachuperus pinchonii, I suspected the existence of additional species, but had not reached definite conclusions. Distribution and collection data.—Batrachuperus pinchonii is a widely dis- tributed species in streams of the mountain ranges along the boundary between Szechwan and Sikang. Its northern limits of distribution may be at Sung- pan, Szechwan, and its southern limits in the northern Taliangshan at the southwestern corner of Szechwan and the southeastern corner of Sikang. The known vertical distribution is from 5,000 to 13,000 feet. I collected this species from the top of Mount Omei during the summers of 1938, 1940, and 1945; from Lianghokou of Paohsinghsien during August, 1939; and from Tsakunao and Tsingshan of Lifanhsien during the summer of 1941. Thirty-one specimens were collected from Yenmenkwan, near Weichow, Szechwan, at about 5,000 feet altitude, September 16, 1934, by F. T. Smith. Comparison with allied species.—Batrachuperus pinchonwi (pl. 1, figs. 4 and 5; pl. 3, fig. 1) can be distinguished from all the other species of the genus by the light brown epidermis covering its palms and soles and extending to the lower parts of the arms and legs, covering the ventral sides of all digits and even ex- tending to the dorsal sides of the first or second segments of the fingers and toes. The “brown crest” of the tip of the tail mentioned by Chang is a cornified covering like that of the feet; it is present in all of the specimens referred to pinchonti. There are usually two small tubercles on each palm and sole in pinchonii and in yenyuwanensis, but such tubercles are not present in the other species of the genus. B. pinchonii and B. yenyuanensis are easily distinguished: the former has a shorter and stronger tail, longer and stronger limbs, and large dark spots on the back; yenywanensis has a much weaker and longer tail, more slender and shorter limbs, and a finely marbled dorsal coloration. Original description.—The original description, in French, is very brief. It may be translated as follows: ‘“‘Another salamander, of small size, is of special 84 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 interest in that it is not related to any known European genus, but to the American genus Dermodactylus. It is described as follows: Size like that of our common [European] salamander. Color ranging from mottled brown to gray- yellow mottled with brown, with numerous blackish spots. Belly lighter. Iris golden gray. Lives in the cold streams of the high forest-covered mountains, and often retires into holes in pine trees called ‘Chamou’; whence it is called ‘Chamouyre’ (fish of the pines). I have dedicated this remarkable amphibian to Monseigneur Pinchon, apostolic vicar of upper Szechwan, under the name Dermodactylus pinchonii, in memory of the service he has rendered to my expedi- tion, and because he gave me the first information about this species.”’ Chang’s amplification of this inadequate description, based on the four cotypes in the Paris Museum, is as follows (translation) : “Head depressed, longer than wide. Snout very short, rounded, somewhat elevated. Loreal region oblique; canthus rostralis indistinct. Naris very small, closer to eye than to that of opposite side; internasal space greater than interorbital space, which is somewhat depressed. Eye large, rather prominent, projecting laterally, larger than the distance that separates it from the end of the snout. Eyelids well developed, upper a little less than the interorbital space, joined to the lower by a dermal fold behind the eye. Mouth wide, buccal angle situated below eye; labial fold well developed partly covering lower jaw, which also has a fold but less well developed. Vomerine-palatine teeth in two small transverse rows, situated at the middle and on the line of the choanae, separated from each other by a distance equaling half of the length of each row. Neck rather long, provided with a fold curved anteriorly. Body cylindrical, of medium length, measuring three times length of head, with twelve costal grooves. Tail long, shorter than head and body combined, cylindrical at base and gradually diminishing toward the end, where it is very compressed, especially the last quarter. It shows a straight superior keel that becomes a low crest terminating in a rather obtuse point. Feet well developed, flattened as far as the ends of the digits, over- lapping or meeting if placed along the body. Four fingers, in order of length: 2-3-4-1; four toes: 3-2-4-1. A soft, rather indistinct cornified sheath under the carpus and tarsus, extending to beyond the phalanges like a nail. Anus a cruciform slit, the short transverse slit situated at the anterior part of the longi- tudinal slit; edge in front of anus slightly elevated. “Skin fairly smooth, with minute pores everywhere except on the throat and under the palms and soles. A vertebral groove; another groove extending from eye to gular fold marks the parotoid; throat provided with very distinct longitudinal folds in front of gular fold. “Color in alcohol: generally grayish or olive brown, mottled with blackish spots above and marked with denser specks below. Cornified skin intense brown, deeper at tips of fingers. Crest of terminal quarter of tail brown.” Measurements of types (from Chang).—Total lengths of types 148, 187.5, 152.7, and 137 mm., tail lengths 64, 64, 71.6, and 61.4 mm., respectively. LIU: AMPHIBIANS OF WESTERN CHINA 85 Variation.—On September 16, 1934, thirty-one specimens were collected by F. T. Smith from Yenmenkwan, near Weichow, Min Valley, Szechwan, at about 5,000 feet altitude. Of the twenty-eight specimens of this Batrachuperus in Chicago Museum, fifteen are adult males and thirteen adult females. Variation is not great. All the specimens have more or less cornification of the digits, palms, and soles, the lower parts of the lower limbs, and the tip of the tail. Thirteen males have two tubercles on each palm or sole and two males have no tubercles on the feet; ten females have the two tubercles on both palm and sole, one specimen has them only on the hands, one has them only on the feet, and they are indistinct in one. The coloration is rather uniform, with light brown background color on the back and dorsal sides of the limbs and on the dorsal and dorso-lateral aspects of the tail; all have irregularly scattered dark spots and markings on the sides of the tail, especially near its tip. Some specimens have the dark spots more crowded, and arranged more or less in dorso-lateral rows. I collected many specimens of B. pinchonii at Mengtunkou, Lifanhsien, 50 miles from Weichow, where the Chicago Museum specimens were obtained. Both series agree closely and also agree with the characters of the types. Two specimens in the Museum of Comparative Zoology (Nos. 17427 and 17428) from Ulongkong, in western Szechwan, are in close agreement with other pinchonii examined. Twenty specimens, five of each sex from two localities, were measured to examine relative proportions (see table). Sexual dimorphism is shown in this species. Males are smaller than females and the length of the trunk is slightly longer in the female than in the male. The number of costal grooves is normally twelve, but this character is of little systematic importance in Batrachuperus. The vomerine tooth patches vary in shape, in position and in number of teeth. Among seven specimens from Weichow, chosen at random, one has two oblique patches of vomerine teeth between the choanae with postero-lateral extensions slightly posterior to the choanae, and seven teeth in each patch. These tooth patches are narrowly separated by a space about one-fourth the length of each patch. Another specimen has slightly oblique vomerine tooth patches be- tween the choanae, with six teeth in each patch, the space between the patches being about one-fifth of the length of each patch. Five specimens have two more or less straight tooth patches, with teeth 4-4, 4-5, 5-4, 4-5, and 5-6, between the choanae, separated by a space equal to about one-half the length of one of the patches. It is clear that the vomerine teeth are not good taxonomic characters. In the male, the vent is a transverse crescentic opening, convex anteriorly, with a light-colored papilla in the middle of the anterior lip. A shallow longi- tudinal groove extends backward from the middle of the transverse opening. The vent of the female is a longitudinal opening, with or without lateral grooves. The vent is more prominently swollen in the female than in the male. Differences in preservation produce variation in the appearance of the vent in the female, but the sexes are nevertheless readily distinguishable. 86 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 Batrachuperus pinchonii: Measurements of Adult Yenmenkwan, Weichow Mengtunkou, Lifan Measurements Five Males Five Females Five Males Five Females Body engin ac. ccer ee ey 65-75 71-77 60-75 66-76 IAVETAQCE Scherr te cee (69.6) (74.6) (67.4) (72.2) Heads width oe tyccisy 13-16 13.5-14.5 12.5-13.5 12-14 Average ees ttn irene (15.1) (14.1) (12.9) (13) Ratio to body length (%).... 21.6 18.9 19.1 18.0 Head‘lengths3...cc ee 17-19 17.5-19 16.5-18 17-19 IAVETASOS et Pataca we eye aarke (17.9) (17.8) (17.3) (18.4) Ratio to body length (%).... 25.7 23.8 PASS | 25.5 Head height 2:2 3.24.23 sate 9-11 8.5-10 6.5-9 7-8 IAVerage aa cn. aati hays (10.4) (9.1) (7.7) (7.6) Ratio to body. length (%).... 14.9 VAR 11.4 10.5 Trunk lengthicenat ct. wae 30-36 33-39 29-39 33-39 AVOETAGC yoo ee on neo acie (83.0) (87.4) (82.7) (85.6) Reto inhaly length (%).... 47.4 50.1 48.5 49.3 Length of foreleg............ 20 .5-22 19-21 17.5-20 19-21 AVOTAGO™ cone ns ela (21.1) (20.1) (19.1) (20) Ratio to body length (%).... 30.3 26.9 28.3 A feat f Length of hind leg........... 20 .5-23 22-25 21-24 22-25 A Veragen se oe Seah ee (21.9) (23.2) (22.7) (24.1) Ratio to body length (%).... 31.4 31.0 33.7 33.4 Maiidengthysie.e arate vse 66-75 59-71 59-68 70-81 AVETAGO A eae ce a ieee (70.1) (67.2) (64) (74.8) Ratio to body length (%).... 100.7 90 95.0 103.6 Paiiwidthe ner ee 9-10 7.5-8.5 6-7 7-8 Averaged) osc oe one (9.4) (7.9) (6.05) (7.7) Ratio = body length (%).... 13.5 10.6 9 10.7 DPailiheight ce. verge ccs ee 9-10.5 9-11 8-9.5 7-8 AVETAGO sa ota eh at (9.7) (9.9) (8.4) (7.6) Ratio iD body length (%).... 13.9 13.2 12.5 10.5 Halitat and habits.—Batrachuperus pinchonii is an aquatic salamander adapted to cold mountain streams from 5,000 to 13,000 feet in altitude. Speci- mens are found in the water or under stones near the margins of streams. This species may occur together with others of the same genus, especially with karl- schmidti, and is occasionally associated with Aelurophryne glandulata in larger mountain streams. B. pinchonii is most abundant in smaller mountain streams. Larvae and the recently metamorphosed young are commonly found near the upper reaches of the streams, especially where springs emerge from beneath stones. Breeding behavior.—The breeding habits of this species are not well known. Eggs.—A number of egg-cases were collected in July, 1948, in association with many adults, on Nine Peaks Mountain, Szechwan. These egg-cases are very similar to those of karlschmidti, described below (p. 92). Larvae.—Batrachuperus larvae of different stages, which were collected from small mountain streams along Mengtunkou, especially around 9,000 feet altitude, LIU: AMPHIBIANS OF WESTERN CHINA 87 Batrachuperus pinchonti: Measurements! of Larvae Ten fully Four specimens Measurements developed specimens with gill stumps Body length seca wre siterscrjasth § hee ieee 26.0-30.0 (27.26) 27.4-30.0 (28.2) Head widthts .cosecna wily oss tea cree 5.7- 7.0 (6.26) 6.2- 7.0 (6.7) Ratio: to, body length (%)s...23..5204.24.05. 22.9 rer f Head length ore 1, vas vars won earners ee 6.8- 8.8 (7.78) 6.8- 8.5 (7.4) Ratio-to body length:(%)s <2 sade fe 28.3 26.2 Head NON: jos neeaw ess Pugh eat. ce ROS t 3.6- 4.2 (8.94) 3.4- 4.0 (8.7) Ratiosco pody, length: (%))sec3 ese 14.4 14.1 Drunks lengh ans oe erise ahaa ced vesene yas 10.6-18.2 (12.08) 11.3-18.3 (12.2) Ratio.to body wength: (0G)... te te 44.2 43.2 Length of foreleg a3 ee ae ee 6.8 8.0 (7.29) Ratio:tospody lengthr(%) 2 eee eee 26.7 ueek Length OF hind Wee soso ce, o8 sta da ahead 6.0- 9.3 (7.72) 6.6- 8.0 (7.7) Ratio:to;bod yalengthe( Sayin es ea 28.2 VA fos Daal engin 55s eas oe 5 Siege actys ober Be Sts 19.5-23.5 (21.3) 20.4-24.7 (21.9) Ratio to: body dength(%)s.sesaesee acest 78.1 77.6 ali wid thes Aeste nec Seer eae ene rig 1 8="252 (220) TS 8=02259 (222) Ratio to. body:length: (%) 2. s3..05.eseceses {Ee 7.8 Pea NOG eG tia veinhet aati. tae elena 3 3.8- 5.2 (4.63) 3.8 4.4 (4.2) Ratio:to' body. lensth:(%) 2.26.52 6: bs bees 16.9 14.8 1 Average in parentheses. differ from those of B. karlschmidti in smaller size and the condition of the gills. Ten fully developed larvae of pinchonii have been carefully studied and measured. In most dimensions they are much smaller than karlschmidti. The most conspicu- ous difference between pinchonii and karlschmidti larvae lies in the gills: pinchonit has fewer filaments, and these are arranged in one row only, except on the third gill (number of gills and gill filaments on both sides I:8-8, II:11-11, III:12-10), while, for the fully developed larvae of karlschmidti, the gill filaments are in two rows on each gill; for Paohsinghsien specimens the gill formula is I:9-9, II:14—-12, III:18—16; for Lifanhsien material the gill formula is I:16—14, II :22—22, III:17-21; and for Luho specimens the gill formula is I:21-20, II:34-36, III:37-36. The fleshy bases of the gills are well developed in karlschmidti and only poorly in pinchonii. The length of the filaments is also different in these two species. In pinchonii, the longest filaments reach not more than half way to the forelimb in the neck region, whereas in karlschmidti they reach the middle upper arm region. The larvae of pinchonii, with only stumps of the gills left, are much smaller than karlschmidti larvae of the same stage; the body length of pinchonir ranges from 27.4 to 30.0 mm., mean 28.2 mm.; in karlschmidti the range is from 39.0 to 50.3 mm., and the average 43.6 mm. Batrachuperus karlschmidti sp. nov. Type.—No. 49879 Chicago Natural History Museum, from Chiala (11,000 feet altitude), Luhohsien, Sikang. Adult female, collected August 10, 1943, by Ch’eng-chao Liu. 88 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 Diagnosis.—A distinct species of Batrachuperus, differing from cochranae by having labial folds, and from all other species of the genus in the total lack of spots and marblings (fig. 12), and in the wide separation of the vomerine tooth patches, the space varying from two-thirds of the length to the full length of one of the tooth patches. Description of type.—Body cylindrical and rather stout, with thirteen costal grooves. Snout squarish rather than rounded, head length 23.8 per cent of the body length, its width 16.6 per cent and its height 10.7 per cent of the same Fic. 12. Batrachuperus karlschmidti; adult (xX 1). measurement; labial folds well developed; length of eye greater than the distance from the anterior corner of the eye to the nostril; interorbital space flat; angle of the mouth reaching a point below the posterior corner of the eye; distance from the angle of the mouth to the postrictal vertical groove about equal to the length of the eye; gular fold extending to the dorso-lateral sides of the neck, visible from above; tongue small and elongate, with sides free; vomerine teeth 4-5, in two small patches between the choanae and separated by a space about equal to the length of one vomerine tooth patch. Limbs short; when appressed, the tips of the longest digits separated by more than two costal spaces; fingers 2-8-4-1 and toes 3-2—4-1 in order of length; a large pad between the bases of the first and second fingers and another between the bases of the first and second toes; tips of digits covered with dark brown horny epidermis; no horny epidermis on the palms and soles. Tail strong, little shorter than the body, cylindrical at the base, flattened gradually toward the tip; only the posterior part of the dorsal side of the tail fin thin, as is the ex- treme tip of the tail; the ventral side of the tail thick and rounded. Vent quad- rangular with a groove behind. Chaetura black on the back and much lighter below, all stippled with dark color. Measurements of type (percentage of body length in parentheses).—Body length 84 mm.; head width 14 mm. (16.6); head length 20 mm. (23.8); head height 9 mm. (10.7); length of eye 5 mm. (5.9); trunk length 45 mm. (53.5); length of foreleg 18.5 mm. (22.0); length of hind leg 24 mm. (28.5); tail length 82 mm. (97.6); tail width 9 mm. (10.7); tail height 8.5 mm. (10.1). LIU: AMPHIBIANS OF WESTERN CHINA 89 History of species.—In 1945 I published the information gained on various collecting trips under the title of Life History of Batrachuperus pinchonii. In this paper, I mentioned two more possible subspecies besides the large common form that was identified as B. pinchonii. These were a smaller spotted form from Lifanhsien and a long weak-tailed and marbled species from Yenyuanhsien, Sikang. Study of the literature and conferences with Schmidt, Pope, Dunn, and Netting have convinced me that the large form is a distinct new species, that the smaller spotted one is B. pinchonii, and that the form collected from Yenyuanhsien is another new species. The large, uniformly colored salamander is named after my friend and colleague Mr. Karl P. Schmidt, Chief Curator of the Department of Zoology at Chicago Natural History Museum. Distribution and collection data.—B. karlschmidti is a common salamander in small mountain streams ranging from 6,000 to 13,000 feet altitude in the moun- tains of the Szechwan-Sikang boundary and on the plateau of Kangshu, Sikang. In 1938, 1940, and 1945, adults as well as young ones were collected from the White Dragon Pool on the top of Mount Omei. From August 11 to 15, 1939, egg-cases, young, and adults were obtained in small mountain streams at Liang- hokou (12,500 feet altitude) of Paohsinghsien, Sikang. During the summer of 1941, large numbers of young and adults were collected from Tsakunao, Meng- tunkou, and Tsingshan of Lifanhsien, Szechwan. From July 8 to September 5, 1943, many egg-cases, young of different stages, and adults were collected from Kangting, Taining, Taofu, Luho and Kantze of Kangshu, Sikang Province. No. 92318 in the Museum of Zoology, University of Michigan, collected by D. C. Graham from Jedo, Sikang, is a specimen of this new Batrachuperus. B. karlschmidti is generally found together with B. pinchonii on Mount Omei and in the mountain streams between northern Szechwan and north- eastern Sikang. Inastream at Lianghokou, pinchonti, karlschmidti and cochranae were collected on the same day. Only karlschmidti was obtained on the high plateau of Kangshu. Variation.—The natural coloration of karlschmidti is nearly uniform, vary- ing slightly in intensity, and without spots or marbling. Most individuals are Chaetura Black or dark neutral gray above and lighter below; many have dark olive or olive green on the back, and in a few cases the back is yellowish olive. After preservation in formalin, they are usually Chaetura Black or dark gray. The dimensions of the different parts of the body in proportion to its length have no positive or negative correlation with size. Measurements of ten speci- mens of each sex from three localities are given (see table). The tail is relatively longer and the body shorter in the male than in the female. The longer body of the female may be related to the requirements of reproduction, by providing space for developing eggs. The number of costal grooves varies from eleven to thirteen, the usual number being twelve. The shape and the number of the vomerine teeth vary greatly. Nine specimens from Luho, chosen at random, have been studied. One has two round patches between the choanae, with four 90 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 Batrachuperus karlschmidti: Measurements Paohsing (Muping) Luho Lifan Measurements Ten males Ten females Ten males Ten females Ten males Ten females Body length.... 88.3-96 82.5-96 74.0-88 78.0-90 80.0-98 75.0-89.8 Average: i.50. . 91.0 91.0 80.8 84.8 86.1 83.2 Head width.... 11.5-18.9 14.6-16.5 18.3-15.5 14.0-16.5 15.0-19 13.4-16.0 Average........ 16.7 15.5 14.4 15.0 17.0 14.8 Ratiots. 2. 2222. 18.4 pl fane L7<8 17.5 19.8 17.8 Head length. 19.5-23.6 17.4-21.5 16.0-20 17.8-21.5 20.0-24.5 17.2-21.5 Average........ 2153 19.7 1Se7 19.3 22.0 19.3 Rao hou 23.8 21.6 Zove 23.6 25.6 23.2 Head height. 7.5-9.6 7.0- 8.7 7.0-9.0 7.49.38 8.0-18 7.5-10.5 Average........ 8.7 8.1 7.8 8.2 9.8 8.8 Ratios. 2ec6c5: 9.6 8.9 9.6 10.0 11.5 10.5 Length of eye 4.8-5.5 4.85.5 4.2- 5.0 4.2-5.3 4.0-5.2 4.0- 4.8 Average........ 5.2 5.0 4.6 4.8 4.4 4.4 Ratiows> =. 4." 5.7 5.5 5.8 Ban Bal 5.38 Trunk length 41.4-42.2 43.0-50.5 36.448.8 37.6-48.7 36.0-49.0 38.0-47.5 Average? 2.5. 44.9 47.3 41.3 44.4 42.2 43.4 IRaWOn 2 et 49.3 52.0 51.1 52.3 49.0 52.2 Length of foreleg....... 24.3-27.3 21.0-26 18.0-24.0 18.5-21.8 20.0-26 18.0-22.2 Average........ 25.6 23.8 20.6 20.9 23.8 20.2 Ratove-h os 28.1 25.6 25.5 24.0 27.0 24.3 Length of hind leg...... 27.0-380.6 25.0-28.6 20.7-27.5 23.0-27 25.0-29 23 .7-27 Average........ 28.6 26.5 23.9 25.7 24.5 26.1 Ratiostcstc-sc- 31.4 29.2 29.6 30.0 28.4 31.4 Tail length..... 93.0-108 77.5-95 77.0-102 79.0-99 88.0-122 70.5-93.4 Average........ 99.7 19.3 84.7 84.6 98.7 82.4 Ratios =o 109.5 87.1 104.8 98.5 114.5 98.9 Tail width... ;:.. 8.6-11 6.4- 9.8 7.3-10 8.0-10.5 9.0-13 6.6- 9.6 Average........ 10.0 8.0 8.6 8.8 10.3 8.4 Ratios asses 11.0 8.8 10.5 10.7 12.0 10.0 Tail height..... 9.2-11.5 8.0-10.3 9.411 8.0-11.0 10.0-15 8.0-11.4 VeTAge 655s 10.3 9.1 9.9 9.6 i by yf 9.9 Ratlou- Clee Lies 10.0 12° 2 1% 13.6 ab key 1 Percentage of body length. teeth in the left patch and five in the right one. Five specimens have two oblique rows anterior to the choanae with the posterior edge near the choanae, teeth 3-3, 3-4, 4-3, 4-4, and 6-6; one specimen has two curved patches, teeth 5-5, median to the choanae; two have a rounded patch on one side and an oblique patch on the other, teeth 2-3 and 3-3, all anterior to the choanae. The shape and position of the patches and the number of the teeth are not reliable as taxonomic characters. In the male, the vent is a transverse crescentic opening with a large light- colored papilla in the middle, with a longitudinal groove posterior to it. The vent of the female varies greatly with preservation; in this sex there is never a transverse crescentic opening with a large papilla. With experience, one can tell the sex in spite of differences of preservation, the vent being a longitudinal LIU: AMPHIBIANS OF WESTERN CHINA GL slit (as is normal in salamanders) sometimes contracted so as to appear quad- rangular or puckered, with a short slit anterior to the opening and a long one behind. The vent is more prominently swollen in the female, the opposite of the normal salamander sex difference. Habitat and habits.—Batrachuperus karlschmidti is a salamander adapted to cold water and found under stones of small mountain streams from 6,000 feet to 13,000 feet above sea level. Specimens are more abundant under somewhat flat stones than under pebbles or small boulders, and they are most often found under stones near the margins of streams. I occasionally found specimens hiding under decayed logs or under roots of vegetation on the banks. The larvae and small metamorphosed individuals are generally found under stones in the upper reaches of small streams, especially in springs, which are the principal breeding sites. The cold-adaptation of this salamander is well known to the monks of Mount Omei, but they interpret it in a different way. The monks told me that one cannot bring such a salamander down the mountain for if one tries to do so, the animal either mysteriously disappears on the way or dies. My experience indicates that these salamanders die from an abrupt rise in temperature. Analysis by number of items of stomach contents of six specimens collected September 17, follow: 61.2 per cent gammarids; 24.3 per cent stone fly larvae; 11.3 per cent caddis fly larvae; 1.6 per cent fly larvae; and 1.6 per cent muscoid fly. A piece of moss was found in one of the stomachs. It is evident that this salamander eats aquatic animals. Superstitions and economic value.—Near the White Dragon Pool on the top of Mount Omei there is a small temple consecrated to the worship of this sala- mander. The pool is so named because the monks and the people of that moun- tain call the salamander the “white dragon” and believe that the White Dragon King is in the pool. They say that if one captures and kills one of these sala- manders a storm will follow. The people of the high mountains of Sikang and western Szechwan believe in the use of this salamander for the cure of stomach trouble. The herb col- lectors usually collect the various species of Batrachuperus and dry them. They sell the dry salamanders to drug stores, or may keep them. I was told that when one has a stomach ache one should grind the dry animal into powder and drink it in boiling water. Breeding.—The breeding season probably lasts from May to the first part of August. Young with external gills were collected on Mount Omei on July 25, 1938; young animals of different stages and three egg-cases with fully developed embryos were collected at Lianghokou on July 21, 1939; many young individuals at different stages and some empty egg-cases were obtained from Tsingshan, Lifanhsien, on August 12, 1941. On August 10, 1943, many larvae just hatched and others at different stages of metamorphosis, as well as many egg-cases con- taining embryos of various stages, were secured in a very small mountain stream near Chiala and Chuwo of Luhohsien, Sikang. 92 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 Egg-cases (fig. 18, A) are attached to the under surfaces of large stones, or to the stream-bed under or by the sides of large stones, where the water flows through. They are found mostly in very small brooks, especially near their sources where spring water seeps out of the ground or from under stones. The end of the egg-case attached to the stone is flattened and sticky; the body is a cylindrical tube, largest in the middle and becoming smaller toward the free end, which is smooth and transparent. Longitudinal striations are found on the transparent wall of the body, which has some traces of a creamy yellow color in the fresh condition. The smooth, cup-like cap at the free end is even more delicate than the rest of the egg-case. Young animals free themselves through a hole at the tip, where the cap is forced off by the movement of the fully developed embryos and the weight of the liquefied jelly inside. In five egg-cases collected from Luho, the average length was 91.6 mm., the range from 75 to 96 mm. The average greatest diameter was 16.6 mm., with a range from 14 to 19mm. The number of the embryos in each egg-case varied from seven to twelve. The embryos lie at right angles to the long axis of the egg- case. Eggs dissected out from the female are yellowish in color, and the average diameter is 3.7 mm. Eggs laid under natural conditions must be larger than this. How many eggs and egg-cases are laid by a single female is unknown; more than forty-five eggs at the same stage of development have been dissected out from a single specimen. Larvae.—Young specimens collected at the time of hatching (fig. 13, B) are different from adults. The color is light gray on the back, yellowish on the belly. Labial folds are discernible, but there is no indication of a movable eye- lid. The top of the head is smooth and the parotoid glands are indistinct. A light mid-dorsal line is present on the head. Pores of the lateral line organ are clearly to be seen on the head. External gills are well developed. There are four gill slits on each side of the neck, the first the largest, the last the smallest. Three gill arches are developed, with short rod-like gill filaments on the internal lateral surface of each arch, the outer lateral side of which bears a fleshy base on which two rows of gill filaments are developed; the filaments of the external row are regularly arranged and with pigmented cords, whereas those of the inner row are fewer in number, irregularly arranged and colorless. The filaments of the outer row are antero-posteriorly flattened, those of the inner row laterally flattened, so that they occupy less space when they fold like a fan. A thin colorless membrane is developed on the ventro-median side of each arch. A similar membrane, continuous with the side of the throat, covers the first gill slit, and the last gill slit has such a membrane at its posterior margin. The most anterior gill, near the throat, has an outer regular row of eleven or more long filaments, and from three to five short filaments median to the outer row. The second gill is dorso-posterior to the first, with thirteen or fourteen filaments in the outer row, and eight or nine in the inner. The third gill is dorso-posterior to the second, the outer row of filaments containing from eight to twelve, the inner row from nine to eleven, all about equal in length with those of the outer LIU: AMPHIBIANS OF WESTERN CHINA 93 row. The vertebral groove is distinct but limited to the anterior region of the trunk. The anterior limbs have three fingers well developed, with an indication of the fourth. On the hind limb only the third toe is indicated. The tail is much deeper than that of an adult. The dorsal fin, more pigmented than the ventral, reaches nearly to the shoulder region. The measurements of ten hatchlings are tabulated. The body length (from the tip of the snout to the vent) ranges from 15 to 16 mm. (average 15.3). | if ll if Fic. 18. Batrachuperus karlschmidti. A. Egg-case (X 11%). B. Larva just after hatching (Xx 2). C. Fully developed larva (x 13/5). The head length of the larva is proportionately larger than that of the adult, 35.8 per cent of the body length as compared with 23.4 per cent for the adult. The height of the head is 18 per cent of the body length in the larva and only 9.8 per cent in the adult; its width is 23.7 per cent in the larva and about 18 per cent in the adult. The hind limb is shorter than the fore limb in the larva, and the reverse is true in the adult. The tail of the larva, with a much higher tail fin, is much shorter than that of the adult. Its length is 80 per cent of the 94 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 Batrachuperus karlschmidti: Measurements of Larvae and Young Ten specimens Ten fully Ten specimens at time of developed Ten specimens just after Measurements hatching larvae with gill stumps metamorphosis Bodysiength,. asco 3 15-16 36-43 39-50.3 34.8-47.5 IAT AICS arsene hertinre cee 15.33 39.59 43.57 42.6 Headiwidth:2 3.2 22s). 3.5- 4.0 7.8 8.5 8.0- 9.4 7.3- 9.9 AVETAGO ns Renta 3.64 8.13 8.65 8.31 Ratiolicc een 23.7 20.5 19.9 19.5 Headilength. 2 Sssse..: 4.7-6.0 10.0-11.4 10.3-13.6 9.3-12.2 AVETAVO Ler accres hae 5.49 10.63 11.61 10.88 RatiOncn sane mot 35.8 26.8 26.7 25.5 Headtheight:<:2e. 72600: 2.7- 3.0 4.3- 5.0 3.5- 4.5 3.4- 4.8 AVOTAGC Food Heancts crave Zo 4.71 4.21 4.32 Ration eee ee. 18.0 11.8 9.7 10.1 ength'of eye 2:2 Sas Sica 2.3- 2.8 2.5- 3.4 2.5- 3.7 IAVOCTAGE Ean kPa wear! ptihcest 2.54 2.719 2.9 Ratio oo ae ian ee 6.4 6.4 6.8 Trunk length carn osse see rae a 16.0-23.5 19.8-23.0 17.0-23.4 AVOrage rice Steers ihe texan 19.47 21:97 20.67 RatOs a. 2 ee eG cee eee 49.1 50.4 48.5 Length of foreleg........ 2.7- 3.7 10.5-138.5 11.3-13.8 10.6-14.9 Average....... 3.15 12.18 12.55 12.54 Ration scar re nc es 20.5 30.6 28.8 29.4 Length of hind leg...... 2-3 11.0-13.6 13.2-16.7 10.0-16.5 Average: 05.600 c 2.55 12.84 14.43 13.27 Ratiovt eaten corte 16.7 32.1 33.1 31.1 Tatlilength <<. st.acws cs. 12.0-12.8 31.0-44.5 33. 7-48 .7 28 .2-42.0 AVeragenn sitet aes LAE 36.41 40.83 34.4 Ratios yas eee es 80.0 91.9 93.7 98.7 Pailtwidthyws*-\ atte each 3.3-.4.4 3.5- 5.0 2.6— 4.3 AVGEAGOUE Sse een nes ae, 8 Tyne 3.65 4.07 3.36 IRatiO Rk Gem tetas: Moe Fat hae 9.2 9.3 7.8 Mailsheighties: + anes 3.5- 4.0 5.8- 7.0 4.3- 6.5 3.3- 5.0 AVETAVE hee eats 3.75 6.4 5.37 4.21 Ration ead ee: Oa WS 16.4 iP aeR: 9.8 1 Percentage of body length. body length in the larva and its height 24.5 per cent; in the adult, its length is 104 per cent and its height only 12 per cent. In ten fully developed larvae from the same locality (see table), the average body length from tip of snout to vent is 39.6 mm., ranging from 36 to 43 mm. At this stage the animal (fig. 13, C) is nearly black above, with a lighter bluish-black belly. Pores of the lateral line organ are even more conspicuous than in earlier stages, as the ground color is much darker, especially on the head. The median dorsal interparotoidal groove, parotoids, and other prominences are still inconspicuous. The eyelids are well developed, but the membrane covering the eyeball is still connected with the margins of the upper and lower eyelids. Gill filaments and their arrangement are as before, except that the color is lighter than the surrounding tissue. LIU: AMPHIBIANS OF WESTERN CHINA 95 The number of gill filaments varies greatly, but the first gill always has fewest filaments and the third gill the greatest number; for example, in one specimen the first gill has two rows of filaments, the exposed row having twelve filaments, the inner row only one; the second gill has seventeen filaments, fourteen in the outer row and three in the inner, whereas, in the third gill, the outer row consists of thirteen filaments and the inner of ten. In another specimen, the num- ber of gill filaments is much greater, twenty-one on the first gill, with fourteen in the outer row and seven in the inner; thirty-four on the second gill, with twenty- one in the outer row and thirteen in the inner; on the third gill there are thirty- six filaments, the separation into two rows being indistinct. Furthermore, the number of filaments is different on the two sides of the same animal. The filaments are flattened obliquely, in correlation with the axis of the fleshy base of the gill, and the inner gill filaments are flattened at right angles to the outer row. They may also overlap each other, so that the filaments of the outer row and the inner accessory filaments fold down like a fan on the sides of the neck region. When they fold down, the tips of some filaments of the first gill reach to just anterior to the base of the arm, those of the second gill to the middle region of the upper arm, and those of the third gill to the dorsal side of the elbow, or in some specimens even to the third costal groove. The fingers as well as the toes are fully developed in the metamorphosing individual, but the arms and legs are longer in proportion to the body than those of the adults. It is difficult to correlate this difference with adaptation. Costal grooves are well formed. The vertebral groove is evident only anteriorly, as the dorsal tail fin reaches the middle of the body. The head is still larger in proportion to the body than in the adult but much smaller than that in the young at the time of hatching. The tail differs from that of the adult, its length being 91.1 per cent of the body length and its height 16.4 per cent, whereas in the young at the time of hatching, the tail is shorter and higher, and in the adult it is longer and lower. During metamorphosis, the gill filaments become black and shrivelled. Ten specimens at this stage were measured (see table). The body length ranges from 39 to 50.3 mm., averaging 43.6 mm. In young specimens nearing the completion of metamorphosis, body proportions approach those of the adult. The coloration at this stage varies greatly; usually there are indistinct marblings or spots on a dark bluish black or dark brownish black ground color. The belly is much lighter. The pores of the lateral line organ have disappeared. The median dorsal interparotoidal grooves, parotoids, and other prominences are clearly evident. The vertebral groove reaches the level of the posterior aspect of the hind limbs, and in some specimens a lighter line is indicated along the groove, this being much deeper and wider on the back, above the hind limbs. The length of the head in proportion to that of the body is less than in the early stages but still larger than the head of the adults. The limbs of the young at this stage are proportionately the same as those of adults. The most conspicuous change occurs in the gills. Not only the filaments, but also the fleshy bases of 96 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 the gills are blackened and shrunk in the advanced larvae, and the fleshy bases are covered by a fold growing posteriorly from the sides of the neck. The gill filaments are not only shrivelled but have also lost their regular arrangement and are curved and more or less entangled. The tail is still shorter and weaker than that of the adult. The animals of this stage are generally found with the adults in the larger mountain streams. Near or just after the completion of metamorphosis, the adult form has been attained except that the head is still larger and the tail still shorter. Gills have either completely disappeared or there may be an indication of a small, dark, depressed spot on each side of the neck. The animals of this stage are found together with the adults in the larger mountain streams. Batrachuperus tibetanus Schmidt Batrachuperus tibetanus Schmidt, 1925, Amer. Mus. Nov., No. 175: 5—Tibetan border of Kansu, at about Lat. 33 N., southwest of Titao, 9,000 feet altitude, type in Chicago Natural History Museum; idem, 1927, Bull. Amer. Mus. Nat. Hist., 54: 554. History of species.—Two female paratypes collected by Robert B. Ekvall, with the same data, agree with the type (an adult female) in all essential characters. Eleven specimens (five adult females, one adult male, two young, and three metamorphosing individuals) of B. tibetanus are in the United States National Museum, also collected by Mr. Ekvall at the type locality in 1924. One of this series was exchanged to the Museum of Zoology of the University of Michigan. These eleven specimens agree in all essential characters with the type. Schmidt (1927) suggested that a male, M.C.Z. No. 2848, from ‘‘Lianghokou (12,000 feet altitude), western Szechwan,” collected by Zappey and recorded by Barbour (1912) and described in detail by Dunn (1923), belongs to the species tibetanus. This locality is undoubtedly the Lianghokou of Paohsinghsien (Muping) in Sikang. Examination of this specimen convinces me that this is its correct allocation. Chang (1936) put tebetanus (“‘thibetina’’) in the synonymy of pinchonti, as he found some young specimens in the Paris Museum collected from the type locality of pinchonii and also without horny palms and soles. Boring (1936) examined the specimens in the Hoangho Paiho Museum in Tientsin from Hweihsien, Kansu, and could not find any characters to distinguish them from pinchonti, thus extending the range of that species to include the region of the type locality of tibetanus. Pope and Boring (1940) remarked that none of the characters listed by Schmidt is easy to use for diagnosis, and they concluded that tibetanus should be placed in the synonymy of pinchoniit. From my field studies in the mountains and plateaus of western China, it is clear that the amphibian fauna of that region is extremely complex and rich in distinct species. Study of the types and topotypes of tibetanus proves that, though it is difficult to distinguish from pinchonii, it is in fact a valid species. LIU: AMPHIBIANS OF WESTERN CHINA 97 Distribution and collection data.—Batrachuperus tibetanus is known only from the type locality, which lies well north of the known range of pinchonit. Comparison with related species.—Batrachuperus tibetanus (pl. 1, figs. 1, 2, 3) can be distinguished from all other species of the genus by its thirteen costal grooves and more posteriorly located vomerine teeth. The absence of palmar and solar tubercles separates tibetanus from pinchonii and yenywanensis; the presence of a labial fold distinguishes it from cochranae; the spotted back and tail differ from those of the more or less uniformly colored karlschmidti. Original description.—‘‘Costal grooves, 14; the adpressed toes overlap; head width 6 times, and head length 41% times in length from snout to vent; eye as long as its distance from tip of snout; a prominent labial fold on the upper jaw; a slight fold on the lower jaw; a shallow groove back of the eye; a well-marked groove from eye to gular fold; no groove to angle of jaw; gular fold extends on sides of neck to dorsal surface; limbs well developed, overlapping when adpressed; fingers 2-3-4—1 and toes 3—-2—4—1 in order of length; tips of digits covered with a thick horny epidermis, which is absent from the palms and soles; tail cylindrical at the base, flattened gradually to the tip, only the last one-fourth very flat; vent formed by the confluence of five grooves; much swollen; vomerine teeth 5-5, in slightly arched series beginning well within and slightly behind the internal nares and extending diagonally forward to the level of their anterior border; distance between the vomerine series about equal to the length of one of them; olive-gray above with indistinct black marbling. “‘Measurements.—Body 85 mm.; tail 80 mm.; length of head 18 mm.; breadth of head 14 mm.; arm 23 mm.; leg 27 mm.” Diagnosis.—Closely allied to Batrachuperus pinchonii, from which it is distinguished by the absence of tubercles and horny covering of the palms and soles, the weak and low tail, smaller head, longer trunk, and more posteriorly situated vomerine teeth. Description of type.—Body cylindrical and moderately stout, with thirteen costal grooves and a narrow vertebral groove from the postero-dorsal side of the head to the base of the tail. Snout round from above and below, head length 22.0 per cent, head width 16.2 per cent of the body length; labial fold well de- veloped; angle of the jaw on level with the posterior corner of the eye; distance of the postrictal vertical groove from the angle of the jaw slightly less than the length of the eye; gular fold extending on the sides of the neck; tongue large, oval, extensively free on the sides; vomerine teeth 5-5, in slightly arched series beginning well within and slightly behind the choanae and extending diagonally forward to the level of their anterior border; distance between the vomerine tooth patches less than half of the length of one series. Limbs slender, the tips of the longest digits not meeting when limbs are appressed; fingers 2-3-4-1 and toes 3-2-4-1 in order of length; tips of digits covered with thick horny epidermis, which is lacking on palms and soles; no tubercles on palms and soles. Tail shorter than body, with weaker tail muscle, the width near its base only 98 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 9.8 per cent of the body length, whereas in pinchonii it is 13.5 per cent in the male and 10.6 per cent in the female; the height of the tail only 9.8 per cent of the body length, while for pinchonii the tail height is 13.9 per cent for the males and 13.2 per cent for the females. Light olive grayish brown above with black marbling, ventral side light in color with indistinct black marbling. Measurements of type (percentage of body length in parentheses).—Body length 86 mm.; head width 14 mm. (16.2); head length 19 mm. (22.0); head height 7 mm. (8.1); length of eye 4 mm. (4.6); trunk length 44 mm. (51.1); length of foreleg 21 mm. (24.4); length of hind leg 25 mm. (29.0); tail length 79 mm. (91.8); tail width 8.5 mm. (9.8); tail height 8.5 mm. (9.8). Variation.—Five adult females and one adult male collected by Robert B. Ekvall from the type locality, Titao, Kansu, in 1924, closely resemble the type. All have thirteen costal grooves. The condition of the vomerine tooth patches varies from specimen to specimen but in most cases the patches are widely separated and posterior to the choanae. The vomerine tooth patches in No. 69093 are curved, separated by a space about equal to the length of one patch, nearly posterior to the choanae and with teeth 5-6; in No. 69094 the vomerine teeth are similar, 4-4; in No. 69095 the patches are between the choanae, teeth 3-3; in No. 69096 the patches are separated by a space about three-fourths of the length of one patch, and the teeth are 4-5; in No. 69097 the tooth patches are as in the last, with teeth 6-7. Abnormality of the digits is uncommon in any species of Batrachuperus except yenywanensis and tibetanus. There are only three fingers on the right hand of No. 69094; in No. 69095 the first and second fingers of the right hand are rudimentary, and on the left hand the second finger is very long, the first very small, and the third rudimentary and fused with the fourth; in No. 69096 the fingers and toes are normal but rather long; and in No. 69097 the third finger of the right hand is short, and the first and second toes of the left foot are abnormal and very short. Batrachuperus tibetanus: Measurements! No. 69094 No. 69096 No. 69093 No. 69095 No. 69097 Female Female Female Female Male Ratio Ratio Ratio Ratio Ratio to body to body to body to body to body length length length length length Measurements MM. (%) MM. (%) MM. (%) MM. (%) MM. (%) Body: length? = 3.2<...25.. 93 Arce 87 peer 84 ene 84 EReeate 82 Reve: Head‘ width: 2:52 2c 142150) ebro 4 Oe 2 14 ed Oe Oe 146 Headslengthiine ee CON ele NO. 21 R92 22-6 Gee ols Be 20 Headvheight:37 4... 6 6.4 6 6.8 6 (fel 6 6.8 is 8.5 ‘rank lengths) 3:4: 52: 85079) =46. “b228: 47 bbe9. 844: 552° 8 d2, cb I1RZ Length of foreleg...... Dok DO On eA. IZA eas SZOn iol 2k.) oan I2OsS Length-of hind) leg. 226: = 227.9" 2i. S10 ar {S820b —26) S3l 0 “26r ols railplength a= tee Sa = 9053" <805- 91.97 rT4d= = 88.0 2749 9y = 7b <9 4 ailawidthe uc ce 7 T.5 8 9.1 9. 1037 8 9.1 92 21059 Taiheight: -3.).2.:2,.08 tf U0 7 8.0 8 9.5 9 8.0 8 eth ' All specimens in United States National Museum. LIU: AMPHIBIANS OF WESTERN CHINA 99 A juvenile specimen with shrivelled gills is 65 mm. long, with body length 35 mm. and tail 30 mm. It has all the adult characters and has a very distinct light-colored vertebral groove running from the postero-dorsal side of the head to the base of the tail. Batrachuperus yenyuanensis sp. nov. Type.—No. 49370 Chicago Naturai History Museum from Peilinshan, Yenyuanhsien, Sikang, China, 14,500 feet altitude. Adult male, collected July 4, 1942, by H. W. Chang and Y. W. Kao. Diagnosis.—Batrachuperus yenywanensis (fig. 14) is a distinct species, dis- tinguished from all others by its long and weak tail, and especially by the dorsal Fic. 14. Batrachuperus yenyuanensis. A. Adult female (x 1). B. Oral cavity (X 21%). C. Ventral view of hand (x 24%). D. Ventral view of foot (x 214). - tail fin, which reaches the base of the tail. The posterior two-thirds of the tail is thin, and the last quarter is a knife-like fin. The two tubercles on the palms and soles distinguish this new form from all other species of the genus except pinchonit. The depressed head distinguishes it from all except tibetanus. Description of type-—Body slender, dorso-ventrally depressed, with twelve costal grooves. Head greatly depressed; snout rounded; labial folds strongly developed; skin on the throat very loose, with many longitudinal folds; tongue 100 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 small, elongated, with narrow free edges; two short, slightly curved patches of vomerine teeth, four teeth in the right patch and five in the left; head length four times in body length; head height about ten times, and head width about six times in body length; angle of the mouth reaching beyond the posterior corner of the eye, postorbital horizontal groove indistinct, postrictal vertical groove not visible, gular fold extending to the sides of the neck; eye as long as its distance from the nostril. Limbs well developed; longest digits meet when appressed; fingers 3-2-4—1, toes 83-2-4—1 in order of length; tips of fingers and toes covered with dark thickened horny epidermis; two darker cornified tubercles on the palm, one between the first and second fingers and one at the base of the fourth finger; two darker cornified tubercles on the sole, one between the first and second toes and the other at the base of the fourth toe. Tail longer than body, 112.6 per cent of the body length, cylindrical at the base, flattened toward the tip, last two-thirds very flat, tail fin thin and high and with last one-fourth knife-like, the edge of the dorsal tail fin especially thin. Vent crescentic, trans- verse, with a papilla-like swelling at the middle of the anterior edge, and with a shallow posterior longitudinal groove. Dark brown above, indistinctly stippled with black, light gray below, dis- tinetly stippled with black. Measurements of type (percentage of body length in parentheses).—Body length 79 mm.; head width 14.4 mm. (18.2); head length 19 mm. (24.5); head height 7.3 mm. (9.2); length of eye 4.5 mm. (5.6); trunk length 39.5 mm. (50.0); length of foreleg 23 mm. (29.1); length of hind leg 24 mm. (80.3); tail length 89 mm. (112.6); tail width 7 mm. (8.8); tail height 12.4 mm. (15.6). Discussion of paratypes.—Fifty-nine specimens of yenywanensis were col- lected from a large pond on Peilinshan at an altitude of 15,000 feet between Yenyuanhsien and Yenpienhsien, Sikang, on July 6, 1942, by H. W. Chang and Y. W. Kao. The series is uniform in coloration, body form, and shape of tail. The coloration exhibits indistinct marbling on the back, belly, and tail. The head is depressed, as is the body. The tail is weak, with a thin, high fin reaching the base of the tail dorsally, and with a knife-like tip. There are eleven or twelve costal grooves. The vomerine teeth are in two oblique patches on a line between the choanae, the number in each patch varying in the ten specimens studied as follows: 3-4 in one, 44 in two, 4-5 in three, 5-5 in two, 5-6 in one, and 6-6 in one. The digits are much more flattened and pointed than in any other species of Batrachuperus. They exhibit abnormalities in num- ber, shape, and size, fifty-one out of fifty-nine specimens showing some combina- tion of abnormalities. The tubercles on palms and soles are very distinct. The average body length of ten males is 82.9 mm., and of ten females 74.8 mm. The trunk of the female is proportionately longer than that of the male. The male has a transverse crescentic vent, with a lighter-colored papilla in the middle of the anterior lip; in the female the vent is a longitudinal slit with swollen lips. LIU: AMPHIBIANS OF WESTERN CHINA 101 Batrachuperus yenyuanensis: Measurements of Ten Male and Ten Female Paratypes Ratio to Measurements Sex Range Average __ body length (%) Bodyslength fesse 2 ae ee roe 75.6-91.5 82.8 a , Q 68 .2-82.2 74.8 aes Head twid thee sence ace ets 24 pte: of 14.0-16.7 15.0 18.0 Q 11.6-18.5 12.9 fer? Heads lengthens este ee fof 17.8-23.0 2053 24.4 Q 16.2-19.0 uy e 23.6 HeadsheightwsSsc5 oi ricttar hea ates 8 roe 6.6— 8.3 7.4 8.9 Q 6.2— 7.8 7.0 9.3 eNO Uh Ol evens arnt atera ee oh a tre J 4.2- 5.2 4.6 5.7 Q 3.9% 5.0 4.3 5.6 ERunieleng Ghetto eer ee ene ref 84.7-45.6 40.4 48.8 Q 34.5-48.0 40.0 52.0 Lengthvof: foreleg: 725 se asin ane ot 23 .0-26.9 25.0 30.1 Q 20.5-24.0 22.0 29.5 enpth-ofshindwWegsss shines os anaes oe 24 .0-28.4 26.4 31.7 Q 21.8-27.4 24.0 32.0 Pail length ee. Acc tee, race cats so ou 89-120 98.6 118.9 Q 67-95 79.9 106.6 anliwid Gites cate ner ern koe ref 5.7- 8.4 6.6 (032 Q 4.6- 6.3 5.4 U2 Wailheights: teers csi, ete roe 10.4-138.5 1 Ss 13.6 Q 8.3-10.8 9.4 a ee Batrachuperus cochranae sp. nov. Type.—No. 49378 Chicago Natural History Museum, from Lianghokou, Paohsinghsien (Muping), Sikang, China, 12,500 feet altitude. Adult male, collected August 9, 1939, by Ch’eng-chao Liu. Diagnosis.—A smaller Batrachuperus, distinguished from all other species by the lack of labial folds and the presence of pitted and thickened skin on the throat; mouth large, with the rictus as far back of the posterior corner of the eye as the distance from the nostril to the anterior corner of the eye; maxilla much better developed than that of other species of Batrachuperus; without cornified epidermis on the palm or sole; the tips of the digits slightly cornified, palmar and plantar tubercles lacking; vomerine teeth seven to nine; the vomerine tooth patches curved and nearly meeting each other. Description of type.-—Body cylindrical and moderately stout, with eleven costal grooves. Head long, snout oval as seen from above and below; head length 25 per cent, head width 17.1 per cent, and head height 10 per cent of the body length; eye longer than the distance from the nostril to its anterior corner; no labial folds; maxillary and mandible well developed, with rictus as far behind the posterior corner of the eye as the distance from the nostril to the anterior corner of the eye; top of the head flat; postorbital horizontal grooves, and postrictal grooves very strongly developed, with a distinct depression where they meet; gular fold extending on to the sides of the neck; skin on the throat thick, without the folds of other species of the genus; mouth large, tongue large, 102 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 rounded, with free edge; vomerine teeth well developed, 9-7, the tooth patches curved and nearly meeting each other; maxillary teeth well developed; when appressed the tips of the longest digits do not meet by a distance less than a costal space; fingers 2-3-4-1 and toes 2-3-4-1 in order of length; no dark, thickened horny epidermis on palms and soles or at the tips of digits; tubercles on palms and soles wanting. Tail as long as body, cylindrical at the base, flattened gradually toward the tip; tail fin very thick, only the dorso-posterior portion thin. Vent T-shaped externally; the opening proper crescentic, with a small papilla at the middle of the anterior lip, and posteriorly with a shallow longitudinal groove. Skin thick and smooth but with conspicuous pits, especially on the head; skin of throat and belly also greatly pitted. Back dark brown, with three ill-defined, black, marbled lines; throat, belly and the ventral sides of the tail and limbs lighter in color and uniform. Measurements of type (percentage of body length in parentheses).—Body length 70 mm.; head width 12 mm. (17.1); head length 17.5 mm. (25.0); head height 7 mm. (10.0); length of eye 4 mm. (5.7); trunk length 32 mm. (45.7); length of foreleg 18 mm. (25.7); length of hind leg 20.5 mm. (29.2); tail length 70 mm. (100.0); tail width 9 mm. (12.8); tail height 9 mm. (12.8). Discussion of paratype.—The single paratype, a smaller specimen, was collected with the type. It has a very poorly developed labial fold, and fewer vomerine teeth (6-7). In all other characters it is similar to the type. Genus TYLOTOTRITON Anderson KEY FOR IDENTIFICATION OF SPECIES OF TYLOTOTRITON I. Costal tubercles indistinct and squarish. A. Three or more wide clay-colored bands on the body and a clay-colored tail. kweichowensis. AA. No clay-colored bands on the body, which is uniformly black; tail black with a SCarlet-eaged! Ventral tallitin 5.4.) 1ht on tara te ecu eetemen Cains feet: taliangensis. 11.2 Costaltubercles-very;.distinctiand rounded (55-6 esas ee cc ins ee ae eee verrucosus. Tylototriton kweichowensis Fang and Chang Tylototriton kweichowensis Fang and Chang, 1932, Sinensia, 2, (9): 112—Kungchishan, Tatinghsien, western Kweichow. History of species.—The type and seven paratypes were collected on Sep- tember 8, 1930, from Kungchishan (about 2,000 meters altitude) of Tatinghsien by Mr. Y. S. Ling and described (1932) as Tylototriton kweichowensis. Distribution and collection data.—Besides the type series, ten specimens were collected from Shihfangin, Kweichow, July, 1934, by D. C. Graham, and in the same year he collected four specimens from Shihmenkan, in the same province. All these specimens are in the United States National Museum. T'ylototriton kweichowensis is endemic in Kweichow, from about 6,000 feet altitude. Comparison with related species.—Tylototriton kweichowensis can easily be distinguished from andersonii, verrucosus, and asperrimus by its costal tubercles, LIU: AMPHIBIANS OF WESTERN CHINA 103 which are indistinct and squarish, whereas in the three other species the costal tubercles are rounded and distinct. The coloration sharply distinguishes kwei- chowensis from taliangensis. Preserved specimens of kweichowensis (pl. 2) have three wide clay-colored bands on the back, and a clay-colored tail, and in some cases two clay-colored bands on the sides of the body. Original description (type No. 4664 Academia Sinica, adult male).—‘‘Habit rather stout; head flat, depressed broader than long; snout short, obtusely rounded; lateral cranial porous ridge subtriangular, prominent, very slightly depressed at anterior connection on the tip of the snout, posterior end curved inward, like a scroll, in front of the parotoid; a faintly marked median ridge beginning from a little behind the apex inside the triangle, running posteriorly to end at a point 3 mm. anterior to vertebral porous ridge in a prominent knob with a depression before it; parotoids also prominent, auricular, slightly concave above, as long as the distance between two scrolls of lateral cranial ridges; eye lateral and moderate, orbit diameter about equal to its distance from the nostril or to the internasal space; upper eyelid granular and rather broad, about one- third of interorbital and visible from above; lower eyelid smooth, semitransparent and narrower than the upper one; nostril oval, rather small, with a small valvular flap in it, close to the anterior extremity of the lateral cranial ridge and half way between the top of the snout and the labial edge, separated from its fellow of the opposite side by a wide internasal space which is about half the width of the interorbital; vomero-palatine teeth in two oblique longitudinal series, meeting anteriorly and commencing in front of (or on a line with), and between the anterior borders of choanae, nearly parallel to each other at first, then diverging latero-posteriorly, and sometimes sharply diverging behind and along the inner elevation of the eyeball, ending at a point slightly before a line between the mouth corners; tongue nearly circular, moderate, its diameter slightly less than half the width of the mouth, slightly free at lateral sides; a gular fold from parotoid to parotoid transversely across the under side of the nape; a series along each side of the body of somewhat squarish knob-like porous glands separated from each other by narrow grooves, about eighteen in number, com- paratively prominent and distinct anteriorly, evanescent and confluent to each other posteriorly, the first starting a little behind and quite above the axilla on a level with the parotoid, the last terminating at the root of the tail just on the level of the posterior border of the vent; a flattened vertebral porous ridge nearly as wide as the lateral series beginning on a line with the scrolls of the lateral cranial ridges and terminating at the same level as that of the lateral series; glandular pores on the anterior cranial ridges readily seen externally; distance from the tip of the snout to the gular fold more than one-third of the distance from the gular fold to the anterior border of the vent; tail long, slightly shorter than head and body together, laterally compressed, with a sharp dorsal edge commencing just behind the vent, tapering from the base and ending in a blunt point, but curved up or more vertically expanded at its mid portion, slightly higher than the base; limbs rather stout, moderately long, the tips of 104 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 the fingers reaching much beyond the nostril to the tip of the snout, those of the toes much beyond the middle of the distance between the axilla and the groin when both fore and hind limbs are stretched forward, and interpalmating when both are appressed against the flank; phalanges rather short, with blunt tips and without webs; fingers four, the third one longest, the fourth subequal to the first; toes five, the third one the longest, the fourth very slightly shorter than the third, the first and fifth the shortest; no metacarpal or metatarsal tubercles; vent a short longitudinal slit with transverse folds or papillae along the edges, which are slightly swollen up. Top of head, dorsal and lateral sides of body and tail covered with glandular porous tubercles of various sizes, irregu- larly distributed; ventral surface of body transversely wrinkled and also with glandular tubercles which are much smoother than those of other parts of the body; tail with its dorsal and ventral margins vertically wrinkled, the dorsal margin more or less with discriminable minute tubercles, the ventral margin rather smooth, without tubercles; snout, lips, digits, palms and soles as well as lower eyelids free from tubercles. ‘Ribs sixteen in number, one stout sacral, fifteen dorsal, the first two under the scapula, the third to the sixth each bearing a small latero-posterior process, each process ending in one of a knob-like lateral series of glands that begin on the end of the process of the third rib; the extremities of the remaining ribs not associated with, but beneath these glands and terminating in the pleuromuscles; first four (or more or less) caudal vertebrae with their transverse processes projected, also terminating in the knob-like glands.” Measurements of type (after Chang; percentage of body length in parenthe- ses).—Total length 155.8 mm.; body length 82.0 mm.; head length 18.6 mm. (22.6); head width 18.3 mm. (22.3); length of foreleg 27.5 mm. (33.5); length of hind leg 31.5 mm. (87.8); tail length 73.8 mm. (90.0). Variation.—Variation in Tylototriton kweichowensis is great. In preserved specimens there are two groups: one is blackish brown on the head, the body, the limbs, and a small part of the sides of the base of the tail; the other has three or five wide clay-colored bands on the body, this color covering nearly the whole tail, and present on the lower eyelids, the corners of the mouth, the posterior ends of the cranial ridges, most of the parotoids, the sides of the neck, the an- terior aspects of the bases of the arms and legs, all the fingers and toes, and the region around the vent. The three dorsal bands, and the color of the tail and the parotoids are constant, the coloration being otherwise variable. Of eleven specimens, five had three clay-colored bands on the back, with scattered spots of the same color near the groin region, and the basal region of the vent blackish brown; four individuals had five clay-colored bands, three on the back and two wide ones on the sides of the body, with the vent region entirely clay- colored; two had three bands on the back and two narrow interrupted ones on the sides. The vomerine teeth of five specimens do not meet at their anterior ends between the choanae, and their anterior portions are parallel; they are angularly LIU: AMPHIBIANS OF WESTERN CHINA 105 bent laterad opposite the middle or posterior parts of the ocular prominences. The anterior ends of the vomerine tooth rows meet anteriorly between the choanae in three specimens and diverge posteriorly along the curves of the ocular prominences. In one specimen the anterior ends of the two vomerine tooth rows are fused into a single row for a short distance posterior to the choanae, after which the two rows are divided, diverging along the curves of the ocular prominences. I took X-ray pictures of five specimens, and four were dissected and carefully studied with the binocular microscope. The X-ray pictures show very definitely that all the specimens have fifteen ribs instead of sixteen, as stated by Chang, and this is substantiated by the dissections. CONDITION OF RIBS Rib U.S.N.M. U.S.N.M. U.S.N.M. U.S.N.M. No. No. 95523 No. 95525 No. 95560 No. 95562 1 No spine and under Nospine and under Nospineand under No spine and under 2 (vv) scapula Small spine near apical end of rib, partly under scapula Small spine more apical Large spine near apical scapula No spine and partly under scapula Large spine at middle Large spine near distal scapula No spine and partly under scapula Small spine more apical Large spine near apical scapula Spine near apical end of rib, under scapula Small spine more apical Large spine near apical 5 Small spine more Small spine more Normal Small spine more apical distal distal 6 With apical dorsal Normal Normal Normal ridge 7 With apical dorsal Normal Normal Normal ridge 8 With apical dorsal Normal Normal With apical dorsal ridge ridge 9 With apical dorsal Normal Normal With apical dorsal ridge ridge 10 Normal Normal Slightly broad and With apical dorsal flattened ridge 11 Normal Normal Slightly flattened Forked at free end and forked at free end 12 Free end curved up- Freeend curved up- Broad, flattened Forked at free end ward ward and forked at free end 13 Free end curved up- Freeend curved up- Broad, flattened Forked at free end ward ward and forked at free end 14 Free end curved up- Freeend curved up-_ Very broad, flat- Greatly flattened, ward ward tened and forked not forked at free end 15 Sacral rib Sacral rib Sacral rib Sacral rib 106 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 Although all have fifteen ribs, no two specimens have the same modifications in all of the ribs (table, p. 105). Nos. 95560 and 95562, collected from Shimenkan, have most of the posterior ribs forked. This condition is not found in Nos. 95523 and 95525, collected from Shihfangin. Tylototriton kweichowensis: Measurements of Ten Specimens Ratio to Measurements Range Average _ body length (%) Total lengthitn toe ee nr eee 158-174 163.4 sere Body length sisi ae coed ee ee 82-94 86.9 ee HMeadlengther tn. kate eee isi oe iec mes 19-22 20.2 23.2 reads width tres tits es a es eres 18-20 18.8 21.6 Head height 15.5 S-8 es et ee ee eee ea ects 9-14 11.5 13.2 Palclength cen fa ee eee oe eee 65-88 76.5 88.1 ail wid Gots a eee ee te tyres eae 7-10 8.6 9.8 Pailsheighty - selec aor ca ee tak chee et eee 10-16 1351 15.0 The proportions and dimensions vary greatly. The length of the tail of No. 95522 is only 84.0 per cent of its body length, whereas it is 103.5 per cent in No. 95560. The tail height of No. 95560 is 8.2 per cent of its body length and 10.7 per cent in No. 95522. The preservation has something to do with the height of the head and the diameter of the tail muscle but does not greatly affect the length of the body and the tail. Four specimens out of nine have abnormal digits: in No. 95521 the two outer fingers of the right hand are fused together, and the outer finger of the left hand is very small; No. 95523 has six toes on the left foot; No. 95560 has four very short flattened fingers on each hand; and No. 95562 has four toes on the right foot, and the fourth toe on the left foot as long as the third, both flattened. Tylototriton taliangensis sp. nov. Type.—No. 49388 Chicago Natural History Museum, from Pusakang, Fulinhsien, Sikang, 8,700 feet altitude. Adult male, collected July 25, 1942, by H. W. Chang. Diagnosis.—A Tylototriton differing from verrucosus and asperrimus in having the costal glands very indistinct and squarish. 7’. verrucosus has fourteen ribs, the last one the sacral, and has fifteen or sixteen large, distinct, rounded and light-colored costal glands. TJ. asperrimus has fifteen ribs, the last two being sacrals, and has fifteen or sixteen small, rounded costal glands, of the same coloration as the body. T’. taliangensis (pl. 3, fig. 3) can easily be distinguished from kweichowensis by its uniformly black body and long black tail, whereas kweichowensis has light-colored longitudinal bands on the back, and a relatively short, light-colored tail. Description of type-—Body moderately stout. Head depressed, longer than broad, its height about half of its width; snout squarish and projecting beyond the mouth; cranial ridges prominent, with a broad, somewhat convex space on top of the head; nostrils at the corners of the squarish snout; length of eye equal LIU: AMPHIBIANS OF WESTERN CHINA 107 to the distance from the nostril to the anterior corner of the eye; upper eyelid with small warts, the lower free from warts; an elongated oval elevation on the top of the head with a free posterior edge in a line with the posterior ends of the cranial ridges; light-colored parotoid glands well developed, their caudal ends curved mesad in the neck region; mouth large, rictus in line with the posterior ends of the cranial ridges; tongue small and rounded; vomero-palatine teeth in two long oblique longitudinal series, near each other between the choanae, diverging posteriorly, and ending in a line at the middle of the quadratojugal bone; gular fold distinct and limited to the ventral side. Body cylindrical with very indistinct squarish costal grooves; ribs sixteen, the first two under the scapula and the last one modified as a sacral. Arm long, the tip of the longest finger reaching beyond the snout; fingers flattened, 2-3-4-1 in order of length; no subarticular or palmar tubercles; when the arm and leg are appressed, the fingers and toes overlap. Legs well developed, with five well-developed toes; toes flat and long, 3-4—2-5-1 in order of length, the third only slightly longer than the fourth; no subarticular or metatarsal tubercles. Tail weak, thin and long, 117.2 per cent of the body length, low, at the highest place only 11.7 per cent of the body length; the thin, sharp-edged dorsal tail fin reaching farther forward than the anterior margin of the vent; the ventral tail fin rounded and thick, reaching the posterior edge of the vent; the tail tip rounded, but very thin and narrow. The skin very rough, with numerous small and large warts scattered over the body, except on the very margin of the jaws, around the nostrils, the lower eyelids, the ventral and dorsal surfaces of the digits, and the ventral edge of the tail; besides warts, the muscular portion of the tail has many vertical folds, and the belly many transverse folds. Vent a long slit, its length 9.8 per cent of the body length, surrounded by numerous folds. Color in life-—Uniformly black, the dorsal side darker, in ground color; scarlet on the parotoid and the posterior ends of the cranial ridges, around the vent, on the ventral margin of the ventral tail fin, from the vent to the ventral side of the tip of the tail, and on the dorsal and ventral sides of the distal parts of all the digits, except that the tips of the digits are black. After preservation in formalin there is no change in the dark color, but the scarlet parts all become old ivory. Measurements of type (percentage of body length in parentheses).—Total length 178 mm.; body length 81 mm.; head length 17 mm. (20.9); head width 15.5 mm. (19.1); head height 8 mm. (9.8); length of foreleg 27 mm. (33.3); length of hind leg 29 mm. (35.8); tail length 95 mm. (117.2); tail width 6 mm. (7.4); tail height 9.5 mm. (11.7); length of vent 8 mm. (9.8). Remarks.—Only three specimens of this exceedingly rare species are known, all of these collected from two localities during the summer of 1942. One speci- men was obtained near Lolokou, Chaochiaohsien, and two were collected in the vicinity of Pusakang, Fulinhsien, Sikang Province. All these specimens were found on land. 108 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 Tylototriton verrucosus Anderson Tylototriton verrucosus Anderson, 1871, Proc. Zool. Soc. Lond., 1871: 423, pl. 76, fig. 6, and pl. 77—West Yunnan; Boulenger, 1882, Cat. Batr. Grad. Brit. Mus., p. 29; Brown, 1910, Rec. Ind. Mus., 5: 193; Annandale, 1911, op. cit., 6: 215; Boulen- ger, 1920, Bull. Soc. Zool. France, 45: 98; Schmidt, 1927, Bull. Amer. Mus. Nat. Hist., 54: 555; Pope, 1931, op. cit., 61: 430. Tylototriton verrocosa Fang and Chang, 1932, Contr. Metro. Mus. Nat. Hist. Acad. Sinica, 2: 121. ‘ Tylototriton verrucosa Pope and Boring, 1940, Peking Nat. Hist. Bull., 15, pt. 1: 20. History of species—Anderson (1871) described the new genus and species from the Nantin, Momien (Tengyueh), and Hotha valleys of western Yunnan. In 1882, Boulenger reported another specimen from Yunnan; Brown (1910) stated that verrucosus is common around Tengyueh; Annandale (1911) reported this species from Yangpi about ten miles west of Talifu at an altitude of 5,200 feet, and from Tengyueh; Schmidt (1927) reported two specimens from Teng- yueh and one from Chenkang, a town in Yunnan southwest of Shunning; and Pope (1931) reported another specimen from Yunnan. Distribution and collection data.—This species is limited to the western part of Yunnan in China and ranges to the Kakhien Hills of Upper Burma, to Darjiling, Sikkim, and to Chieng Dao, northern Siam. The vertical distribution is from 4,500 to 6,000 feet altitude. I have collected no material. Comparison with related species.—T ylototriton verrucosus can be distinguished from kweichowensis and taliangensis by its distinct rounded costal tubercles, these tubercles being indistinct and squarish in kweichowensis and taliangensis. Original description.—‘‘The lateral cranial ridge subtriangular; the median ridge running backwards from the inside of the apex of the triangle, but not reaching so far posteriorly as the lateral ridge, the extremities of which curve inwards like a scroll in front of the parotoids. The parotoids slightly concave above, and somewhat resembling the outline of an upturned human ear. The nostrils close to the extremity of the rounded snout, but with a considerable interval between them, semicircular, closed by a small valvular flap of skin. Eye of moderate size; upper eyelid large, granular. Fifteen knob-like glands along the side of the body; the first a short distance behind and about the axilla and on a level with the parotoids; the last three behind the leg when it is extended at right angles to the body. The vertebral glandular ridges begin on a line with the scroll-like extremities of the cranial crest, and terminate at the root of the tail. An obscure line of pores, larger than those of the rest of the body, from below the arm to the groin, rather toward the under surface of the side. A series of pores behind the angle of the mouth along the lateral cranial ridge to the top of the snout, on the loreal region behind the eye, and along the mandible and internal to it. The chin and throat thickly covered with small, smooth, porous glandular tubercles of nearly uniform size. The sides and upper parts of the body and of the tail are densely covered with glandular tubercles (porous) of various sizes and irregularly distributed. The ventral surface transversely LIU: AMPHIBIANS OF WESTERN CHINA 109 wrinkled and covered with very minute porous glands, which scarcely project above the level of the skin. The upper margin of the tail sharp, and commencing with the last lateral knob. Under surface rather rounded. Numerous folds on the inner margin of the vent. Forearms extend the length of the fingers beyond the snout; the legs reach halfway to the axilla. “Uniform blackish brown, paler on lips, snout, chin, throat, and under- surface of the limbs, all of which are of a brownish-olive tinge. Under-surface of the tail dull orange-yellow, fading to lightish brown on the sides. “Length from tip of snout to vent 3 to 4 inches; vent to tip of tail 3 to 334 inches. “Hab. Nantin, Momien, and Hotha valleys, Western Yunnan, China. “T first met with this remarkable newt in the flooded rice fields about the little Chinese town of Nantin, where, however, it was not very common. In the more elevated and subtemperate valley of Momien, at about a height of nearly 5,000 feet above the sea, and in the high sequestered valley of Hotha (5,000 feet) it was far from uncommon.” Triturus wolterstorffi Boulenger' Molge wolterstorfi Boulenger, 1905, Proc. Zool. Soc. Lond., 1905, pt. 1: 277, pl. 17— Yunnanfu. Triturus wolterstorfii Dunn, 1918, Bull. Mus. Comp. Zool., 62: 451. Hypselotriton wolterstorfi Herre, 1934, Verh. Deutsch. Zool. Ges., 1934: 158; Wolter- storff, 1934, Zool. Anz., 108: 257; Pope and Boring, 1940, Peking Nat. Hist. Bull., 15, pt 123: Cynops wolterstorffi Chang, 1935, Bull. Soc. Zool. France, 60: 426; idem, 1936, Contr. Etude Morph. Biol. Syst. Amph. Urodéles Chine (Paris), p. 110. History of species—The above synonymy accounts for the history of the species, which has not been adequately studied. I got four living specimens from Professor Tieng, who studied the embryology of this species in Peip’ei, Szechwan, by means of living specimens transported from Yunnanfu. Distribution and collection data.—Triturus wolterstorffi (pl. 3, fig. 2) is an endemic form in Yunnan, inhabiting the lake outside of Yunnanfu. I have not collected specimens. Comparison with related species.—The species wolterstorffi has been referred to the distinct genus Hypselotriton because its quadrate is directed antero- laterally instead of postero-laterally, as in Triturus sensu str. Triturus wolter- storffi is larger and has a somewhat more laterally compressed body than is usual in related forms. I have followed Dunn and others in the use of T'riturus in a broad sense. Original description.—‘‘Fronto-squamosal arch bony, thick. A chevron- shaped series of palatine teeth, the apex on a line with the choanae. Tongue 1 For detailed synonymy, see Chang, 1936, p. 110. 110 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 small, subelliptical, the sides slightly free. Head without grooves, once and one- fourth to once and one-third as long as broad, its length to base of tail; the greatest width of the head behind the eyes, which are rather small and feebly prominent; snout broadly rounded; labial lobes much developed. Body rounded in both sexes; no dorsal crest, no vertebral ridge. Limbs moderate; fingers and toes depressed, free; outer carpel and tarsal tubercles small but distinct, the latter sometimes very prominent. ‘Tail strongly compressed, blade-like, with upper and lower crest, obtusely pointed, its length about that of head and body. Cloacal lips as in M. vulgaris. Skin perfectly smooth and shiny; a strong gular fold. Blackish olive above, with an orange or orange-vermilion vertebral stripe, with or without round or roundish spots or dots of the same color; orange- vermilion beneath, with black spots or marblings, which may be confluent into longitudinal bands; lower edge of tail vermilion-orange. Male Female Male Female mm. mm. mm. mm. Totallerigth? 2. escent lO 140 Bore Jim Ds oon sis estes 19 22 From snout to cloaca..... 55 70 Hind’hmbse se. vad 24 Head length] eae ere ee 16 19 PGi ran es eta teas Sh 55 70 Width of hea eer chm ile 15 “Of the six specimens sent by Mr. Graham, all except the male, of which measurements are here given, have preserved the external gills—another instance of the neoteny already observed in tailed Batrachians living at great altitudes. The altitude of Yunnanfu, where the specimen was obtained, is about 6,000 feet. The skull in these branchiferous specimens is fully ossified and has all the features of the mature state. The female is full of ripe spawn. “In the structure of the skull and the absence of crest or digital web in the male this new species approaches the Spanish-Portuguese M. boscae Lat., and the Chinese-Japanese M. pyrrhogaster Boie, the affinity of which I pointed out many years ago.” SALIENTIA : Bombina maxima Boulenger Bombinator maximus Boulenger, 1905, Ann. Mag. Nat. Hist., (7), 15: 188, pl. 13, text fig. 2—Tungchuan, Yunnan. Bombina maxima Schmidt, 1927, Bull. Amer. Mus. Nat. Hist., 54: 556; Liu, 1945, Jour. W. China Border Res. Soc., 15, (B): 56, figs. 1-5. Distribution.—Bombina is Eurasian, with two species in Asia and two in Europe. This genus has a curious discontinuous distribution. Bombina maxima is the species common in western and southwestern China, characteristic of the mountain regions of Yunnan, of southwestern Szechwan, southeastern Sikang, the western part of Kweichow and Kwangsi and northern Tonking. Bombina orien- talis is a northern species of fire-bellied toad and is a low mountain form. It is found in very small mountain streams along the seacoast of northeastern Shan- tung, and in Manchuria, Korea, and adjacent areas. LIU: AMPHIBIANS OF WESTERN CHINA 111 Comparison with related forms.—The various species of Bombina can easily be distinguished from all other Salientia of China by the red or orange yellow and black patches on the belly. The common name “‘fire-bellied toad” refers to these colored patches. Bombina maxima (fig. 15) can be easily separated from Bombina orientalis by the body length, the size of the warts, and the colored patches on the belly. Bombina maxima is much larger in size than orientalis, with an average body length from snout to vent of about 60 mm., while in orientalis it is only about Fic. 15. Bombina maxima; male (x 1). 45 mm. The body of maxima is rather stout and heavily built; in the north- eastern form, the body is rather slender and more or less dorso-ventrally de- pressed. This difference may be related to their different breeding habitats, since maxima lays its egg-masses in open water, while orientalis attaches its eggs to the under surface of stones in small mountain streams. The warts of maxima are much larger and less uniform in size than those of orientalis. In maxima the colored patches of the belly are very large, while in orientalis the patches are smaller and much more numerous, so that its belly seems much more marbled. It is especially noteworthy that the warning behavior of the fire-bellied toad is a generic character as well fixed as are skeletal or other structural features. Collection data.—For the study of the life history of this toad I made a collecting trip to Sichang, Chaochiao, and Yenyuan in Sikang from March to August of 1942. The altitude of the city region of Sichang is 5,700 feet, and the maximum temperature is about 92° F. from the end of April to the end of July. The city of Chaochiao is about 150 miles northeast of Sichang. The highest mountain between these two cities reaches an altitude of 11,000 feet, but the altitude of the city of Chaochiao is not more than 7,200 feet, and the tempera- ture range was about 60°-78° F. from May 7 to 15. The best place for collect- ing is on the high mountain between Hsuanshenpa (8,700 feet) and Lolokou 112 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 (10,000 feet). Bombina maxima is found midway between these two villages, and also on the way to the city of Chaochiao. Its vertical distribution in this region is from 7,800 to 10,000 feet. The city of Yenyuan is about 120 miles southwest of Sichang, and has an altitude of 9,600 feet; from June 22 to July 9, 1942, a collecting trip was made to study the Amphibia of that locality. Bombina maxima is very abundant from 9,000 feet upward in Yenyuanhsien. Original description.—‘‘Vomerine teeth in two small groups or short trans- verse series close together behind the level of the choanae. Tongue large, circular, scarcely free at the sides and behind. Head broader than long; snout rounded, not quite as long as the diameter of the orbit; no canthus rostralis; nostril equally distant from the eye and the tip of the snout; interorbital space narrower than the upper eyelid, nearly equal to the distance between the nostrils. Fingers short, obtusely pointed, first shortest, third longest, fourth a little longer than second; no subarticular tubercles; two round palmar tubercles, inner larger and more prominent. Tibio-tarsal articulation reaching the shoulder, tarso-meta- tarsal articulation reaching the eye; tibia as long as the femur, the heels meeting when the legs are folded at right angles to the rhachis; foot as long as the tibia; toes short, obtuse, flattened, only half-webbed; no subarticular tubercles; a small, rounded inner metatarsal tubercle. Upper parts covered with small warts intermixed with very large glands studded with pores, similar to the parotoids of toads; the largest are situated behind the eyes (true parotoids), on the tibia, on the tarsus, and on the back, where they form a pair of curved or angular chains behind the head, with the convexity turned inwards. Lower parts nearly smooth; a more or less distinct gular fold. No horny spines on any part of the body. Blackish olive above, with rather ill-defined black markings forming vertical bars on the upper lip and cross-bars on the limbs; only the inner finger and toe with a light tip; a more or less distinct light transverse spot on the back, just behind the head. Lower parts marbled bright orange and black, in about equal proportions, or the black predominating; greater part of palm and sole orange, this colour involving the inner digit; the orange of the lower surface of the arm either extending across the breast, or widely separated from a pair of pectoral spots; plantar, tarsal, and tibial orange spots continuous or interrupted; the orange not extending on the back of the thighs. MM. MM. SHOE GOMVeN Gs sas te eke esac eerie 68 Ore IMD hoes ecser ete ee 32 Pengthror heads enc sors. terteeeraacion 19 Hind timbers: soe age oe te eee eee 73 Widthroft head vi. i5 see Seow ie oe 23 Tibia isc caeeectre ce ae = ope ok Diameter’ of eyes: 5). sigs oan ste 6 MOOG Areas any he et ae ee 24 Interorbitalwidth: 3225.54: scead cee 3 “The three specimens here described are females.”’ Coloration.—The variation in coloration has no connection with sex. The adult toad is stippled with silver gray or silver tinged with golden on the cinnamon brown ground color of the head, back of the body, and dorsal sides of the limbs, and mostly has a green area on the dorso-median part of the shoulder region. LIU: AMPHIBIANS OF WESTERN CHINA 113 Each wart on the back bears numerous fine black spines, and these spines are more numerous in males than in females. This coloration varies slightly accord- ing to the nature of the surroundings. Under bright light it is lighter, but in the dark the silver appearance may be lost, and the animal may become a golden cinnamon brown. Dark olive or black bars are developed on the limbs, but those on the upper arms and legs are not constantly present and are rather incon- spicuous, while those on the lower arms, fingers, tarsus, foot and toes are much better developed, especially those on the digits. The pupil is triangular in shape, with a single angle pointed ventrad, amber in color. A bright golden ring en- closes the pupil and the rest of the iris is golden, stippled with black. The throat, the belly and the ventral sides of the limbs are black or dark gray variegated with scarlet, orange, or orange yellow. There is always a patch of scarlet or orange color on the palms and soles, and a transverse patch of the same color extends on the ventral side of the thighs. All this makes the several species of fire-bellied toads unlike any other salientians. The brilliant colors are unquestionably effective in warning off predators, as is known to be the case with the European species of Bombina. Bombina maxima: Measurements of Twenty Male and Nine Female Adults Ratio to Measurements Sex Range Average body length (%) Body: length ttc te eee roe 53-69 62.2 2 (snout to vent) Q 54-73 61.8 2 Head) length serine reise keane roe 17-22 19.9 32.0 (angle of jaws to tip of snout) Q 18-22 19, Sieg Head widthe erase cst erate eee of 17-25 20.7 33.3 (at angle of jaw) Q 20-23 PA Ve 34.2 Rowerarmivlengthtse. ese oe of 20-29 23.8 38.2 (elbow to tip of third finger) Q 23-29 24.4 39.5 Diameter of lower arm.............. ret 7-12 9.6 155 (greatest diameter) Q 6.0-8.5 ffl 11.4 Length of hand) .2205 ct ee sa oe 11-15 13.1 21.0 (base of palm to tip of third finger) 92 12-15 12.8 20 27 length. of hinds limbs ana S2: ee roe 76-107 82.6 132.8 (vent to tip of fourth toe) Q 70-90 (Ase 122.5 Mengthiotiblad cee cae 22-33 26.1 42.0 Q 21-27 22.9 otal Length of foot and tarsus........... roe 32-49 37.1 59.6 Q 32-44 SB hyate: 57.9 Hengthvottootes pant sper reese fof 19-31 23.3 BY (er. (base of inner metatarsal tubercle to 2 17-25 20.6 Sono tip of toe) Secondary sex characters.—The average body length of females is 62 mm., while in males it is the same. In males, a large diffuse nuptial pad covers the in- ner side of the arm, extending from the distal portion of the upper arm, covering the inner face of the forearm, and two-thirds of the inner metacarpal tubercle (fig. 16, B). The inner dorsal aspects of the first, second, and third fingers are provided 114 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 with similar horny nuptial pads. There are also two patches of nuptial pads on the thorax near the base of each arm (fig. 16, A). The arm of the male is dis- tinctly larger than that of the female, with the hand strongly bent inward. The fingers are much shorter in the males, especially the first, which is angularly bent. The fingers of the male are fringed with a skin fold, rather wide at the base, diminishing towards the tip. The metacarpal tubercle is larger in the males. The skin of the male is more rugose, with minute spines on the back, the sides, and the limbs. The number of warts is much greater in the male. Thus Fic. 16. Bombina maxima; male. A. Ventral view of head and thoracic region (xX 1). B. Ventral view of hand (xX 2). C. Ventral view of foot (xX 2). the females have a generally smoother appearance, though covered with large warts. The hind limb is distinctly longer in the males, and the glands on the tarsus and the toes are much better developed. The toes are nearly fully webbed in the male (fig. 16, C), and slightly more than half in the female. Habitat and habits.—The adults of this fire-bellied toad are found in small pools and ponds, especially those with dead vegetation, and also less commonly in small mountain streams of running water. Both adults and young are found under stones at the edges of very small mountain streams, and of pools or ponds formed in connection with the streams. Just by turning a stone, very often one or more individuals are obtained. Some specimens were even found in a well by Mr. Kao at Lolokou. Fire-bellied toads are very inactive, and it is easy to catch them by hand. This animal has a distinctively warning coloration, for the black and red, orange-red, or orange-yellow patches on the lower side and on the palms and soles are very conspicuous. When disturbed it does not attempt to escape, but remains in place and turns its limbs over its back, with the head and posterior extremity of the body bent upward, so as to show as much as possible of the bright colors on the lower surface. The palms and soles of the hands and feet are turned outward and upward, so that four red spots are shown conspicuously. LIU: AMPHIBIANS OF WESTERN CHINA 115 Breeding behavior.—The breeding season is rather short. I found many paired individuals in pools near the top of the mountain of Yenwot’ang on May 4, 1942, on the way to the city of Chaochiao, but when I came back, on May 12, only eggs and embryos at different stages of development were collected from those pools. Eggs and tadpoles at different stages were collected near the city of Yenyuan on June 29 by H. W. Chang and Y. W. Kao. Although there is no vocal sac in the male, yet a weak croak like the sound of a very young chicken or the cry of a newly born puppy was produced at twilight, near the breeding ground, or when they were carried around in the collecting bags. The male grasps the female just in front of the base of the thigh, as she is floating on the water. Eggs.—Eggs are laid in masses, which mostly sink to the bottom of the pools, where they are covered by a thin layer of mud from the red soil. If there is any vegetation floating on the surface of the water, the egg-mass is attached to it and suspended from it. The eggs are coated with three layers of jelly, the middle layer being the thickest and the inner layer the thinnest. In the living condition the jelly is usually in masses, closely connected with the corresponding layer of the other eggs, but sometimes there are isolated eggs. The animal pole of the egg is brownish gray, with a creamy yellow color on the vegetal pole. The eggs are rather large. Ten eggs were measured, the average diameter being 3.18 mm., with a range from 3.0 to 3.4 mm. Tadpoles and development.—A newly hatched individual is rather large and greatly flattened, as it is 9.8 mm. long from the snout to the tip of the tail and 3.7 mm. high in the middle region of the body; but it is only 1.5 mm. in diameter in the thickest region of the body. That is why the animal at this stage always lies on its side. The color in this stage is uniformly grayish brown, with evenly distributed, light-colored granules. Nasal pits and proctodeum are clearly shown, with two dark-colored ventral suckers on the ventral side of the head region. Six short gill-filaments are developed on a single large fleshy base. The tail crest is strongly developed, reaching the nuchal region. When the tadpole reaches 12 mm. in length, the color of the body is darker and the tail crest is much lighter. The eyes are seen externally as black spots, the mouth is well developed, and the ventral suckers are clearly shown. The number of external gill-filaments is generally twelve. They are without fleshy bases, and are located near the ventral side of the neck region. The abdomen, which is larger than before, becomes much more rounded and the center of gravity is established in the mid-ventral region. Thus at this stage the body assumes a normal position. The tail crest is tremendously developed, reaching the dorso-posterior region of the head. The external gills are covered up by a skin fold when the tadpole reaches 14 mm. in length. Vestiges of the ventral suckers are still visible. Black lines that form a network by crossing each other are developed all over the body and at the tail crest, but the black granules are limited to the body and the muscular 116 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 axis of the tail. The tail crest is still very high and reaches to the middle part of the trunk. A median ventral spiracle is formed posterior to the thoracic region. At the morphologically defined stage, when the tadpole has developed a hind limb-bud (pl. 10, fig. 8), it is a pale creamy color on the head, the gill region, and the tail. The length at this stage varies according to environmental condi- tions. The body is a dark brownish gray, stippled with yellow. Most charac- teristic of the tadpole of this species are the sharp black lines crossing each other in a definite fashion on the body, especially conspicuous on the tail crest, where there are no other pigment granules. The whole animal is semi-transparent, so that the different parts of the brain, the circulation of the internal gills, and the viscera can be seen through the body wall. For this reason, the tadpole seems very delicate and very much like a mass of jelly. The pupil is round and sur- rounded by a wide black iris stippled with gold. The fully developed tadpoles are darker than the young ones. Ten tadpoles with hind legs fully developed were measured. The average body length from snout to vent was 16.9 mm., ranging from 16 to 18 mm. As this tadpole is a quiet-water form, the body is stout and the tail is short, with a delicate tail crest. The width of the head is very nearly equal to the height of the body, and nearly equal to the height of the tail. The eyes are dorso- laterally located and quite close together, the space between them being only 25 per cent of the body length. The tail is very weak, delicate, and short, re- sembling that of Nanorana pleskei. It is interesting that this body feature is common to these two unrelated forms, both living in similar bodies of water in high mountains. Obviously this is a body form adapted to quiet water. The length of the tail, with its rounded tip, is 122.6 per cent of the body length, though the diameter of its muscular part is only 16.1 per cent of the body length. The weakness of the tail muscle is regarded as an adjustment to quiet water. The spiraculum is median ventral in position, with a transverse slit-like opening a little bit anterior to the central coil of the intestine. The anal opening is rather small, median ventral in position, at the base of the ventral tail crest. Bombina maxima: Measurements of Ten Tadpoles Ratio to Measurements Range Average __ body length (%) Body length (snout to vent)................. 16-18 16.9 Deter Body height (at highest place)............... 8.2-9.2 8.5 50.5 Body width (greatest diameter).............. 9.6-10.8 10.4 61.2 Head height (at orbital region)............... 5.8-7.2 6.3 37.1 Head width (at orbital region)............... 7.2-8.8 8.1 47.9 Mouth width (including lips)................ 3.9-4.2 4.0 yA ar | Space DELWOEN CYOR. ii 55 chm otras inl 4.0-4.5 4.3 25.0 SHOwt LO SpIrACUUUIN hoje eens eee eee ete 9.0-9.8 9.4 55.1 Tail length (vent to tip).. SF aftes SOT an LOS ee 20.8 122.6 Tail height (at highest place)... BRD Ay Pats ie 7.8-9.0 Sel 47.5 Thickness of tail muscle (near tail base)....... 2.5-3.0 Zi 16.1 The mouth is small and ventral in position. Its average width is 4.04 mm., ranging from 3.9 to 4.2 mm. A single row of small papillae are found around LIU: AMPHIBIANS OF WESTERN CHINA 117 the mouth. The labial teeth of the tadpole are the type characteristic of Bombina. Each row generally consists of two sub-rows of short pointed teeth. In some specimens extra teeth may be present between the two sub-rows, making the arrangement of the teeth more or less irregular. Ten tadpoles with well-developed hind limbs were selected at random, and were carefully studied as to their tooth characters. Six individuals had two continuous rows on the upper lip and three rows on the lower lip but the innermost row was very narrowly interrupted; and four tadpoles had two continuous rows on the upper lip and three continuous rows on the lower lip. Under normal conditions the inner row of the upper lip and the middle row of the lower lip are the longest for the upper and the lower lips, respectively. The mandibles are rather weak, with fine teeth on their margins. Metamorphosis.—As soon as the hind limbs become fully developed, the fore limbs come out through the skin, and dark marks on the limbs and the warts on the body and limbs are conspicuously developed. In the centers of the palms and soles a patch of light orange-yellow color is developed. As soon as half of the tail has been resorbed, the light orange-yellow patches become much more conspicuous. The ventral side of the limbs and the belly also develop dark and yellowish-orange patterns. The body is 14 mm. long from snout to vent, im- mediately after the completion of metamorphosis. Genus AELUROPHRYNE Boulenger The first species of Aeluwrophryne was described by Giinther in 1896 as Bufo mammatus, from specimens collected by the Russian explorer Potanin in the mountains and on the Kham Plateau (Tungsolo) on May 16 and July 3, 1894. Giinther’s specimens were in a poor state of preservation, and he could not recognize the shape of the pupil. Boulenger (1919) received a single young specimen, still with a vestige of the tail, collected by his son Captain C. L. Boulenger in Ladakh Valley, Kashmir, at an altitude of 12,000 feet, in August, 1919. The combination of a vertical pupil with the absence of teeth in this 25 mm. specimen suggested to Boulenger the examination of the vertebral column and the pectoral girdle of one of the type specimens of Bufo mammatus in the British Museum. He found that the sacral vertebra, with unusually widely dilated transverse processes, articulated with the coccyx by a single condyle, as in Pelobates and Megophrys, with which it agreed also in the structure of the pectoral girdle. Boulenger accordingly proposed the generic name Aelurophryne for Bufo mammatus, in allusion to the cat-like pupil. Aelurophryne is closely allied to Scutiger, differing from it in the absence of a notch in the posterior border of the tongue, which is oval, and in having the tympanum present, though hidden under the skin, and the eustachian tube moderately large. In 1917 Annandale described a tadpole from Lake Kreshen, Kashmir, as that of Rana pleskei, and in 1919 Boulenger suggested that this must be the tadpole of Aelurophryne mammata. Zarevsky (1925) 118 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 recorded Bufo mammatus from western China, describing his specimens as similar to those of Giinther’s original series. Aelurophryne gigas was described by Zarevsky in 1925, based on a breeding male and female collected in September, 1900, from the Chamchu River, Kham Plateau, Tibet. His new frog is markedly distinguished from Aelwrophryne mammata by its much greater size and by absence of glands on the chest. Six specimens identified by Stejneger as Aelurophryne mammata included three collected from Sungpan by D. C. Graham, July, 1924, and three from Gieolonghsien (=Chiulung), Sikang, also collected by Graham, August 1-5, 1930. Careful examination of my own collection, of the collections of Chicago Natural History Museum, and of the six specimens in the United States National Museum, discloses a considerable amount of taxonomic confusion. The tadpole described by Annandale as Rana pleskei (1917) and regarded as a specimen of Aelurophryne mammata by Boulenger (1919), must be quite another species of Aelurophryne, for the labial tooth formula described is entirely different from that of tadpoles from Sikang, which I regard as certainly true mammata. The Kashmir tadpole has the labial tooth formula I:4-4/1:3-3 and most of the mammata tadpoles have the formula I:5-5/I:5-5. I have now carefully studied three species of Aelurophryne tadpoles, and none have three interrupted rows of teeth on the lower lip, as described by Giinther. Therefore, I consider Giinther’s tadpole to represent still another species of Aelwrophryne. Boulenger’s young specimen from Kashmir may represent the adult of the Kashmir tadpole, and in any case is not likely to belong to the species mammata. A young adult specimen in the United States National Museum is not mammata but belongs to my species tainingensis, as shown by the rounded glands on the posterior median region of the belly, the longer slender arms, and the elongated depressed body. Another adult male in the same museum is not a mammata and is clearly a distinct new species characterized by the shape and location of the chest glands and the number and arrangement of their spines. Two further new forms, glandulata and brevipes, without spines on the chest glands, are represented in my own collection. According to the nature of the chest-gland spines the three large species of Aelwrophryne of western China can be assorted into two groups; mammata has strong spines, and this form has a geographical distribution limited to the western and southern parts of the Kham Plateau, whereas glandulata and brevipes, without spines on the glands, are limited to the northeastern corner of Sikang and the northwestern part of Szechwan. It has been suspected that the spines on the chest glands of the males may be present only during the breeding season. The present study does not support this hypothesis, as males of Aelurophryne brevipes, collected July 29, have on the thorax very finely granulated chest glands without spines or the scars of spines; whereas true Aelurophryne mammata, collected July 21, have very strong black spines on the chest glands. Furthermore, the eggs of brevipes were col- LIU: AMPHIBIANS OF WESTERN CHINA 119 lected with the adult in the same locality and at the same time; July must accordingly be the breeding season of this species of Aelurophryne. All the males of brevipes have spines on their first and second fingers, like those of mammata, but have neither spines nor the traces of spines on the chest glands. The shape of the chest glands and the space between the chest glands in the males of brevipes are quite different from those of mammata; it is evident that they are distinct species with well-defined specific characters. It now appears that the presence or absence of spines on the chest is not the result of seasonal change. Aelurophryne glandulata from Hopachai, Lifan, Szechwan, is closely related to brevipes. Aelurophryne tainingensis is an independent and distinct species not closely related to the four species discussed above. Its smaller size, slender depressed body, absence of spines on the chest glands and on the fingers, and rounded glands on the belly distinguish it fully. The types of Aelurophryne mammata were not sexed. Giinther described only the gland near the axilla, as “an oval white or pink gland on each side of the chest, immediately behind the shoulder-joint.”’ This is not the chest gland of the male, but the axillary gland of the female. The axillary gland is distinct in the female specimens collected by Herbert Stevens and indistinct in his male specimens. The chest glands, either with or without spines, are always present and prominent in the males of the various species of Aelurophryne, and it would be impossible to overlook such a conspicuous structure. Giinther’s specimens thus appear to be females. Zarevsky’s Aelurophryne gigas was distinguished from mammata by its much greater size and by absence of glands on the chest. Size alone is quite unreliable as a character to distinguish closely allied species when only one specimen of each species has been measured, especially when the difference amounts only to 10 mm., Giinther’s type measuring 60 mm., and Zarevsky’s specimen 70 mm. Zarevsky reports the “‘breeding male with callosities forming brushes of black prickles on the upper parts of first and second finger and with two asperities on each side of the chest.”” The axillary glands are indistinct in male specimens of Aelurophryne mammata in the large series collected by Stevens, and the absence or presence of axillary glands, unless the sex of the specimen is specified, does not distinguish allied species. It appears to me that Giinther described the female of mammata and Zarevsky gave a description of a male of the same species. The difference in relative leg-length stated in the descriptions of Giinther and Zarevsky appears to be explainable by differences of preserva- tion of the material and a different terminology (see description of Aelwrophryne mammata below). Within the relatively primitive family Pelobatidae the species of Aeluro- phryne form a specialized group in which the maxillary teeth are entirely lost and the jaws are feeble. The ecological valence of Aeluwrophryne is much limited, as the several species live only on the high plateau or high mountains and are 120 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 evidently adjusted to low temperatures. The wealth of species of Aelurophryne in western China appears to represent a rejuvenation of the genus after its estab- lishment as a relict in the unfavorable plateau environment. KEY FOR IDENTIFICATION OF ADULTS OF AELUROPHRYNE I. Size large, with warts on the soles and with nuptial spines on the first and second fingers of the male. A. Strong spines on chest glands, no pre-humeral glandular areas........ mammata. AA. Chest glands without strong spines, with pre-humeral glandular area. B. Fourth toe long, two inner toes two-thirds webbed, inter-glandular space on chest small, about 1.2 per cent of body length.................. glandulata. BB. Fourth toe short, two inner toes one-third webbed, inter-glandular space on chest wider, about 3 per cent of body length..................... brevipes. II. Size small, without warts on the sole and with nuptial pads on the first and second fingers of the male. A. Warts on belly, two inner toes one-third webbed....... ......... tainingensis. AA. Belly without warts, two inner toes one-half webbed................ maculata. KEY FOR IDENTIFICATION OF TADPOLES OF AELUROPHRYNE I. Upper labial teeth six rows. ‘A... Labial: tooth formula:mostly [:5-5/1 55-5. 354s a cies ees Ls mammata. II. Upper labial teeth less than six rows. A. Labial tooth formula mostly I:3-3/1:4-4.............0...0...0.... glandulata. AA’. --Labial:tooth: formula. mostly 124-4/0- 526i s 5 2 asc 3 cen erect dec brevipes. Aelurophryne mammata Giinther Bufo mammatus Giinther, 1896, Ann. Mus. Zool. Acad. Sci. St. Petersbourg, 1: 208— Tungsolo [Tungolo], Kham Plateau, Sikang, China. Aelurophryne gigas Zarevsky, 1925, Ann. Mus. Zool. Acad. Sci. USSR, 26: 77—Cham- chu River, Kham Plateau. History of species in China.—The type locality of Aelurophryne mammata formerly was in western Szechwan, near Tibet. It is now in central Sikang. Specimens collected from the Kham Plateau have sometimes been recorded as from Szechwan and sometimes as from Tibet. Chicago Museum has 52 speci- mens of mammata, including adult males and adult females, collected by Herbert Stevens from Hlagong, near Yakiang, west of Kangting, July 21, 1929. Hlagong is on the Kham Plateau and it seems clear that this series may be regarded as essentially topotypic. These specimens afford the opportunity for a more detailed description of the species than has hitherto been possible. Comparison with allied species.—Aelurophryne mammata (fig. 17) differs from Aelurophryne glandulata, brevipes, and tainingensis in having strong spines on the chest glands, which are only granulated or may have some fine black pigment in the three latter species; axillary glands are indistinct in the males of mammata and well developed in the males of glandulata, brevipes, and taining- ensis. The warts on the soles of the feet of mammata distinguish this frog from maculata and tainingensis. LIU: AMPHIBIANS OF WESTERN CHINA 121 Distribution.—The range of Aelurophryne mammata is the Kham Plateau, Kangshu, Sikang. Tadpoles of various stages were collected in a small stream near Hsintuchiao, which is close to the type locality, Tungsolo (Tungolo). Original description.—‘‘Crown without bony ridges. Head broad, snout obtuse, with the bones of the jaws remarkably feeble. Inter-orbital space as broad as the eye-lid. Tympanum small, entirely hidden under the skin. Fingers rather slender, the first and the second subequal in length; legs thin; the hind- ii Fic. 17. Aelurophryne mammata; male (x 1). A. Dorsal view. B. Ventral view. limb being carried forwards, the tibiotarsal articulation reaches the eye; meta- tarsus with an elliptical, flat tubercle; subarticular tubercles very indistinct. The three outer toes not half webbed, but the web between the three inner ones is as broad as that between the three outer ones. Upper surfaces with very flat, glandular patches, that representing the parotoids not more prominent than the others. Lower parts smooth, but there is an oval white or pink gland on each side of the chest, immediately behind the shoulder joint. Legs smooth, with small simple glands disseminated over the hinder part of the thighs. Uni- form black, or dark brown with the glandular patches on the back of a darker shade. Length of body 60 mm., length of third finger 12 mm., length of hind- limb 84 mm., length of fourth toe with metatarsal 27 mm.” Description.—Adult male, C.N.H.M. No. 15199, Hlagong, near Yakiang, west of Kangting, Sikang, collected July 21, 1929, by Herbert Stevens. Body stout (fig. 17), head depressed, broader than long; snout round, depressed, and slightly projecting beyond the mouth, as long as the length of the eye; canthus rostralis very obtuse; loreal region oblique and slightly concave; nostrils about halfway between the eyes and the tip of the snout; interorbital space 122 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 flat, about equal to the width of the upper eyelid; tympanum hidden; parotoid glands flat, with indistinct mesial and posterior boundaries, with a latero-ventral fold running obliquely to the arm; jaws weak, without teeth; tongue thick, oval, with very slight indentation behind; no vomerine teeth. Arm very strong, fingers slender, tips rounded, the first and second fingers equal in length, much stronger and broader than the third and fourth fingers and with strong black sharp spines on the dorsal aspect of their basal segments; third and fourth fingers slender, the third one longest and the fourth much longer than the first two fingers; no subarticular tubercles; two large palmar tubercles of which the inner is larger than the outer. Legs short, tibio-tarsal articulation reaching the posterior part of the parotoid gland, heels not meeting when hind limbs are placed at right angles to the body; toes fringed, with very small rounded tips except the first; first and second toes about one-half webbed, and the third and the fourth toes about one-third webbed; subarticular tubercles wanting; inner metatarsal tubercle oval, flat and without free edge; no outer metatarsal tubercle. The skin extremely loose with numerous wrinkles or folds, and a large quantity of white spongy connective tissue between the skin and the muscle, especially prominent on the back of the body; warts extremely flat so that the skin is quite smooth; warts dark in color on account of their dark-colored pits; small rounded warts at postero-ventral sides of the thighs just below the anus; small black spines on the sides of the head, especially near the tympanic region; black pigmented areas on the ventral margin of the jaws; some small rounded warts developed behind the angles of the jaws. Throat skin smooth with fine wrinkles; the skin of the belly and the ventral sides of the thighs smooth, with loose, wrinkled, and folded skin; chest glands with strong black spines forming two elongated patches on the chest (fig. 17, B) similar to the strong black spines on the inner dorsal sides of the first and second fingers. Color (in alcohol) on the upper side of the body nearly uniform deep mouse gray; when the specimen is submerged in water, a darker mark with a lighter margin may be seen extending from the middle part of the interorbital space backward to the shoulder region, where it becomes inconspicuous; flattened warts covered with black pits, the top of the snout much lighter in color than the region below the canthus rostralis; pale mouse gray on the throat, the belly and the ventral sides of the limbs; black pigmented areas on the margin of the jaws; spines of chest glands and of the first and second fingers coal black; tips of digits lighter in color. The skin, the warts, and the coloration vary very slightly among the male specimens. The most noticeable variation is in the length of the hind limb; the tibio-tarsal articulation may extend to the angle of the mouth or only to the shoulder region. The fringing and webbing of the toes, though usually as described above, is subject to some variation; in one, the toes are more extensively fringed, and the two inner toes are about two-thirds webbed instead of one-half. LIU: AMPHIBIANS OF WESTERN CHINA 123 The tips of the fingers and toes may be covered by black cornification but usually are lighter in color. The secondary sex characters of Aelwrophryne mammata. are remarkably developed. Sex dimorphism in size is slight. In ten of the largest specimens of each sex, the average body length of the males is 72.0 mm., while for the females it is 67.1mm. The skin of the males is extremely loose, with numerous wrinkles and folds, whereas in the females the skin is not loose and is more rugose on the dorsum, with various-sized warts, very numerous small warts on the sides, and only the throat and belly smooth. A large quantity of white spongy connec- tive tissue is present between the skin and the muscles in the males; in the females this is not conspicuous. There is no conspicuous color dimorphism in alcoholic specimens except that the throat of the female is greatly marbled by deep mouse gray on a much lighter background. In males, there are two well-developed chest glands covered by strong spines. The axillary glands present in the arm pits of the females are not developed in the males. In the males, very strong black spines are present on the inner dorsal sides of the first and second fingers, which are much stronger and broader than the corresponding fingers of the female. Aelurophryne mammata: Measurements of Ten Male and Ten Female Adults Ratio to Measurements Sex Extremes Average __ body length (%) IBodyslengthes-- - h eee of 63-75 72.0 ee Q 62-75 67.1 eee Headtlensth a soon ere ee of 19.5-24.0 220 Sire Q 20.0-23.0 Dike SL. Head widthiciten cee ieeias a eens rot 22-27 24.5 34.0 Q 21.0-23.5 21.8 32.4 Imterorpital swith... ee oe roe 5.0-6.5 Hol. 7.9 Q 4.5-5.5 ys Catt Lengthiof:chest-gland sacs 2 ose roe 7.5-12.0 9.4 13.0 Ore eaters o Neanh bie Width of: chest: gland:2: (2-2 -<4-- = roe 2.5-6.5 4.8 6.6 OM hi aoe ae Rare Meee Length:of lower arm)s,,.505 4.04550 fou 28-37 33.4 46.3 Q 27.5-35.0 3022 44.9 Diameter of lower arm.............. oe 10.5-17.0 13:3 18.4 Q 6-8 6.7 9.9 engthcoishandes cee cs ees, roe 16.0-20.5 17.9 24.8 Q 15-19 159 23.6 ength of legs ca Nees Mad Tomas of 82-104 91.4 126.9 Q 72-89 81.8 121.9 Itength’ of tiblavnss 2 ee ere 25-31 27.8 38.5 Q 22 .0-23.3 22e0 BR Length of foot and tarsus........... roe 41-49 44.6 61.9 Q 35-45 38.8 57.9 Isength OL loots: 2eert ae ay atme roe 29-37 Slo 43.6 Q 25-32 26.9 40.0 Length of inner metatarsal tubercle... o& 4.0-5.5 4.7 6.5 Q 3.5-5.0 4.0 5.9 124 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 The arm is distinctly stronger and bent inward in the males. The ratio of the diameter of the lower arm to the body length is 18.4 per cent in ten males; this ratio is only 9.9 per cent in ten females. The hind limb of the male is longer and stronger than that of the female, especially in the tibial region. The toes are much more webbed and strongly fringed in the males than in the females. Lineae masculinae and vocal sacs are wanting. Measurements (adult male; percentage of body length in parentheses).— Body length 76.0 mm.; head length 25.0 mm. (82.8); head width 27.0 mm. (35.5); interorbital space 6.0 mm. (7.8); length of chest gland 10.0 mm. (13.1); width of chest gland 5.0 mm. (6.5); space between two chest glands 3.5 mm. (4.6); lower arm length 37.0 mm. (48.6); diameter of lower arm 16.0 mm. (21.0); length of hand 20.5 mm. (26.9); length of leg 101.0 mm. (182.8); length of tibia 32.5 mm. (42.7); length of foot and tarsus 50.0 mm. (65.7); length of fourth toe and sole 34.0 mm. (44.7); length of inner metatarsal tubercle 4.2 mm. (5.5). Tadpoles.—Five tadpoles of Aeluwrophryne mammata were collected by Stevens at Hlagong. One of these five tadpoles is fully developed, with hind limbs measuring 10 mm. Ten tadpoles from Hsintuchiao were measured and studied. The coloration of the alcohol-preserved tadpole is uniformly dark gray on the back and sides and the muscular portion of the tail; the belly and the crests of the tail are cream. This coloration is similar to that of preserved tadpoles of glandulata and brevipes. The tadpole of A. mammata is a typical running-water form, the body being elongated and dorso-ventrally depressed, with a long tail, strong tail muscle, and low thick tail crest with a blunted tip; the head is depressed, with a rounded snout. The mouth is ventral, with strong mandibles. There is a single row of papillae on the margin of the lips, with additional papillae, some with teeth, at the corners of the mouth. The number of rows of labial teeth varies, but all are supported by thick fleshy ridges. Three of the tadpoles from Hlagong have Aelurophryne mammata: Measurements of Ten Tadpoles from Hsintuchiao Ratio to Measurements Extremes Average body length (%) BOGY NEN OCR ss crc ft se neeat rds Socteie rniahig is OS 25-28 26.5 ne BOOVsNCIGN Ge nee min cs mc sm ce ae ead tee 11-12 11-2 42.2 BOG Y) Width ccs, Sin see ae ee tae RE 11-16 14.9 56.3 Headiheightt nie tee, ne ee rs ws Se 7.5-9.0 7.9 29.7 Heddswidthtrnscei cis cerca etc aeeeres 11-12 LST 44.0 Mouthiwidth er narandiwis.. tee tecenat on neice 7-8 at 28.9 Spacenrom’eye'to 6ye: aes). cacecie. ce eeashas 7-8 Tez 27.2 Space from:nostril:to nostril 714103. 56. eee 4.5-5.0 4.9 18.7 Tap otisnout to.spiracwlum oo <.o.2 ca oe oes 15-18 16.9 64.0 Malllengthicrsee sewer rapa es Ere 41-52 46.1 174.3 PalltReIGh Gs erred edt ade eee on eatery 11-14 12.0 45.6 Diameteroktailumuscle:. te peo ier sce on ee 6-7 6.7 25.3 icengt hots hind wim biel pare eee oe on ees 7-12 8.6 32.4 LIU: AMPHIBIANS OF WESTERN CHINA 125 the labial tooth formula I:5-5/I:5-5; one has teeth I:5-6/1:5-6; and one I:4-4/I:5-5. On the upper lip, the outermost row is very short and continuous, partly filling the gap on the mid-dorsal part of the lip where no papillae are developed. The remaining rows on the upper lip are all interrupted, the outer- most row the longest, and the innermost the shortest. On the lower lip there are usually six rows of teeth, the outermost the shortest and continuous, and the inner five all interrupted, the innermost the shortest. In the Hsintuchiao tadpoles, five have the formula I:5-5/1:5-5; two have the formula I:4-4/1:5-5; one has [:4-4/1:5-4; one has I1:6-6/1:5-5; and one has I:6-6/1:6-6. Aelurophryne brevipes sp. nov. Type.—No. 49393 Chicago Natural History Museum, from Taining, 11,500 feet altitude, Sikang, China. Adult male, collected in July, 1943, by Ch’eng-chao Liu. Diagnosis.—Closely related to Aelurophryne glandulata, but with shorter feet, about 41 per cent of the body length; chest glands large and oval, the Fic. 18. Aelurophryne brevipes; male. A. Ventral view of head and thoracic region (xX 1). B. Ventral view of hand (x 2). C. Ventral view of foot (x 2). anterior half of their inner margins parallel, and posteriorly diverging; axillary glands with two-thirds of their anterior inner margins in contact with the chest glands; the inner two toes about one-third webbed, and the third, fourth and fifth toes with webs only at the base; prehumeral glandular area developed. Description of type-—Body stout, somewhat depressed; head depressed, slightly broader than long, snout rounded, depressed, slightly projecting beyond the mouth, and much longer than the length of the eye (fig. 18, A); canthus rostralis obtuse; loreal region very oblique and slightly concave; nostrils about mid-way between the tip of the snout and the anterior corner of the eye; inter- 126 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 orbital space about equal to the width of internasal space or upper eyelid; tympanum hidden; jaws feeble, without maxillary teeth; no vomerine teeth; tongue large, oval, and slightly notched behind; no vocal sac. Arm strong; fingers moderately long, the first and second fingers equal in length but thicker and shorter than the fourth finger, and the third finger the longest; tips of the fingers lighter in color; nuptial spines developed on the inner dorsal -sides of the first and second fingers; subarticular tubercles indistinct, Fic. 19. Aelurophryne brevipes; male (xX 1). palmar tubercles prominent, a larger rounded inner one and an outer slightly smaller one; palm not rugose, with a few inconspicuous small warts (fig. 18, B). Hind limb short and weak, the length of the hind limb only 127.7 per cent of the body length, the tibio-tarsal articulation reaching the shoulder region, the tibia only 31 per cent of the body length, the foot shorter than those of other species of Aelurophryne; toes fringed, with tips dilated and light in color; the two inner toes about one-third webbed, and the other toes webbed only at the base; the subarticular tubercles indistinct and easily confused with warts on the toes; the inner metatarsal tubercle well developed, elongated oval, with a free edge (fig. 18, C). Skin rough, with large flattened warts more or less in rows on the dorso- lateral sides of the back (fig. 19); many small warts on the sides of the body and on the dorsal sides of the limbs; larger warts pitted, especially on the back; parotoids large, distinct, flat, and with a strong ridge running from the posterior corner of the upper eyelid toward the base of the arm; many rounded and light- colored warts on the median postero-ventral aspects of the thighs. Skin of the throat, chest, belly, and the ventral sides of limbs smooth, except for the develop- ment of glandular areas on the margins of the jaws; two pairs of flattened glands developed on the thoracic region—chest glands—large, oval and flat, with the anterior half of their inner margins parallel and the posterior half diverging; the axillary glands small, with two-thirds of their anterior inner margins in LIU: AMPHIBIANS OF WESTERN CHINA 127 contact with the postero-lateral edges of the chest glands; a definite glandular area with scattered small rounded warts antero-laterad from each chest gland (fig. 18, A); skin of belly wrinkled in preserved specimens. Measurements of type (percentage of body length in parentheses).—Body length 72.0 mm.; head length 23.0 mm. (81.9); head width 24.0 mm. (83.8); interorbital space 5.0 mm. (6.9); length of chest gland 13.0 mm. (18.0); width of chest gland 7.0 mm. (9.7); space between two chest glands 2.0 mm. (2.7); length of axillary gland 7.0 mm. (9.7); width of axillary gland 2.5 mm. (8.4); length of lower arm 32.0 mm. (44.4); diameter of lower arm 12.0 mm. (16.6); hand length 17.5 mm. (24.3); leg length 92.0 mm. (127.7); length of tibia 22.5 mm. (31.2); length of foot and tarsus 42.0 mm. (57.5); length of foot 29.5 mm. (40.8); length of inner metatarsal tubercle 4.6 mm. (6.3). Coloration in life-—Yellowish olive stippled with gold on the back and the upper sides of the limbs; much lighter on the anterior region of the top and the sides of the head and on the large, pitted, flattened glands; sides of the body greenish yellow, with lighter-colored small warts; yellowish gray on the throat, chest, and anterior region of the belly; more yellowish on the hip region, the posterior part of the belly, and the under sides of the limbs; dorsal sides of the inner fingers and toes greenish yellow; chest gland pinkish gray; axillary glands yellowish, stippled with gold; the color of the eye similar to that of Aelurophryne glandulata. After preservation, the back and dorsal sides of the limbs are dark, the snout being lighter in color. The coloration of living tadpoles of Aelurophryne brevipes is usually Chaetura Black on the body, with the muscular portion of the tail stippled with gold. The snout and the sides of the body are Benzo Brown, stippled with gold. The tail crest is much lighter in color, near Ecru Drab, with scattered black stippling. The belly is light pinkish gray. The pupil is round and black enclosed by a golden ring, and the iris is golden stippled with black. The preserved tadpole of brevipes is black, with the belly light gray and the tail crest colorless. Collection data.—Kangting, also known as Tatsienlu, the capital of Sikang Province, has an altitude of about 8,000 feet. The temperature was 52° to 60° F. on July 9, 1948. Tadpoles of a species of Aelwrophryne were collected from a river in the city. In one of these tadpoles nearly through metamorphosis, the coloration of the back was dark gray stippled with gold and four ill-defined rows of elongated warts with lighter centers were developed on the back. This character is very much like that of adult Aelurophryne brevipes collected from Taining on July 29 of the same year. Tadpoles of different stages collected from small mountain streams of Tangnipa (12,600 feet altitude), northwest of Kangting, on the way to Taining, July 25, probably belong to this species. On July 29 many fully developed tadpoles of A. brevipes were collected from mountain streams east of the city of Taining at 11,500 feet altitude. Many adults were found under stones along the same mountain streams. In such mountain streams Batrachuperus pinchonii is also common, but no other salien- 128 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 tians appear to be present. Aelurophryne brevipes is an inactive frog. Individuals of this species hide under stones at the sides of mountain streams or in the water. Turning such a stone discloses the frog, and one can scarcely miss catch- ing it. Egg-masses were obtained in the upper reaches of the same stream where adults and tadpoles were found. On the way from Taining to the city of Luho (11,100 feet) tadpoles of this new Aelurophryne at different stages were collected but no adults were seen. On August 8, 1943, adult males and females were collected from a mountain stream west of the city of Luho together with tadpoles of various stages and some specimens of Batrachuperus pinchonui. From August 12 to 14 more specimens of adults and tadpoles of Aelurophryne brevipes were obtained from a small mountain stream at Chuwo, the largest village in the west border of Luhohsien, at the same altitude as the city of Luho. In the stream at Chuwo we collected, besides Aelurophryne brevipes, Batrachuperus pinchonii of various stages and some cascade frogs (Staurois). This was one of the few places where I found true cascade forms inhabiting the same mountain stream as running-water types. The dark brown mask between the eyes extending to the back of the shoulder region is generally very distinct in young specimens and indistinct or entirely lost in large adults. The background coloration on the back varies from yellowish olive to golden. Two large males collected by the author are uniform muddy golden in color without any markings or pattern on their backs. Sex dimorphism.—Sex dimorphism in size is very marked. Ten large males and nine large females were measured. The average body length of the males is 71.4 mm., ranging from 68 to 80 mm., while in females it is 64.6 mm., with a range from 58 to68 mm. The skin of living males is looser than that of females, but after preservation this condition is much less evident than in A. mammata. The chest glands of the thoracic region of the male are without spines or pigmented granules, in contrast with Aelurophryne mammata and glandulata. Axillary glands are found in both sexes but are larger in males. A well-defined area, covered with thinner skin and with scattered small warts antero-ventral to the base of the arm and antero-lateral to the chest gland, is found only in males. Such an area is present in males of all species of Scutiger and in the males of Aelurophryne glandulata and tainingensis, but not in the male of mammata. A conspicuous male character is the development of strong black spines on the inner dorsal sides of the inner two fingers. These two fingers, especially the first, much stronger than the third and fourth fingers, are flattened and curved toward the palm. The arm of the male is distinctly stronger than that of the female, and bent inward. The average diameter of the lower arm of the male is 13.6 mm. with a ratio to the body length of 19 per cent, while in the female this ratio is only 10 per cent. The hind limb of the male is longer and stronger than that of the female, and the toes are much more webbed and more strongly fringed. LIU: AMPHIBIANS OF WESTERN CHINA 129 Aelurophryne brevipes: Measurements of Ten Male and Nine Female Paratypes Ratio to Measurements Sex Extremes Average _ body length (%) Body lengths. oor 2 eee teens ef 68-80 71.4 ei Q 58-68 64.6 Sar Headilengthe.ecxs ns. nace cate eee rou 21-27 23.3 32.7 Q 19-23 21.4 33.1 Head widthe.2 ate34 rt eee of 23-27 DAT, 34.6 Q 20-23 2253 34.5 Interorbital spaces. os. 3. seen oe roe 5-6 5.6 7.8 Q 5-6 5.4 8.4 ength of lowerarm=..e ce hee 32-36 S3i50 46.8 Q 28-32 30.6 47.4 Diameter of lower arm.............. of 10-16 136 18.9 Q 6-7 6.5 10.1 Lengthiofshand? 3:2 = ean see of 16-20 18.6 26.1 Q 15.5-17.0 16.6 25.7 ength of lege cee ae ee ee roe 90-105 95.6 132.9 Q 77-88 82.2 12723 eng th vot tibia jiecceri.yiant-rns nes chee roe 27-32 Pate | 40.8 Q 23-25 24.0 Stet Length of foot and tarsus........... roe 43-50 46.7 65.4 Q 37-41 39.1 60.5 henpth of footss mwas een) anaaw, tee 31-37 33.2 46.5 Q 26-30 28.5 44.1 Length of inner metatarsal tubercle.. 5-7 5.9 8.3 Q 4-5 4.6 {teak Habitat and habit.—Aelurophryne brevipes is a high mountain form. Frogs of this species are found under stones at the edge of the water in small or medium- sized mountain streams from 9,000 to 13,000 feet altitude. Tadpoles were very abundant in mountain streams from Kangting to Kantze, but not many adults were found in or near the same streams. This fact indicates that Aelwrophryne brevipes is not essentially an aquatic running-water form. Before and after the breeding season, these frogs appear to live in the crevices of stones on hillsides and only come out at twilight to feed. In a dry season the individuals near a stream may enter the water and hide themselves under stones. The short and weak hind limbs and the poorly developed webs, the well-developed palmar tubercles, much like those of the common toad, and the rareness of the adults in the mountain streams where their tadpoles are so abundant, all indicate that this species must be a terrestrial form. Breeding behavior.—From my data, it appears that the breeding season of this species is rather long. Fully developed tadpoles were found in mountain streams in the middle of July. On July 29, 1943, two egg-masses and newly hatched tadpoles were collected from the upper reaches of a mountain stream. I conclude that the breeding season of this new frog may extend from the middle of May to the middle of July. The breeding season may of course vary from year to year according to temperature and the rainy season. The upper reaches of mountain streams, where large and small stones are numerous, with little vegeta- tion, and only a small amount of water, are the site for egg-laying. 130 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 Eggs are laid in masses and attached to the under surface of the larger stones. The eggs are large (3.5 mm.) and white, with their outer jelly layers connected with those of other eggs. In my limited material, the eggs at the center of the egg-masses were mostly spoiled and only the outer eggs developed into tadpoles and hatched. Tadpoles.—Newly hatched tadpoles stay with the jelly mass from which they hatched. At this stage they attach themselves to the jelly or swim in the Fic. 20. Aelurophryne brevipes; tadpole. A. Lateral view (Xx 2). B. Mouth (x 10%). shallow water. The body of the newly hatched tadpole is gray, the tail crest is much lighter, and the belly is nearly colorless. Large tadpoles are not found in the places where the eggs are laid. They inhabit the larger mountain streams and the quieter waters at the sides of mountain rivers. They generally hide themselves under stones and one rarely sees them swimming in open water. They are good swimmers and very sensitive to any disturbance. Ten tadpoles from Taining and ten from Pamei, selected at random, were measured and carefully studied. The general body form, the size of the different parts of the body and the mouth parts are relatively constant (see table). The detailed description is based on tadpoles from Taining, where most of the adults were obtained, tadpoles from other localities being mentioned only for comparison. The tadpole of Aelurophryne brevipes (fig. 20, A) is rather large in size and is a typical running-water form. The average body length of ten tadpoles of LIU: AMPHIBIANS OF WESTERN CHINA 131 Taining is 24.8 mm., ranging from 23 to26mm. The body is somewhat depressed dorso-ventrally, the average body height being 10.6 mm. and the width 13.7 mm. The length of the tail is 168 per cent of the body length, with rather low tail crest and strong tail muscle. The head is depressed, its average height about 28 per cent of the body length, and its width 48 per cent. The snout is rounded; the nostril, enclosed by a light-colored ring, is about mid-way between the tip of the snout and the anterior corner of the eye. The eyes are dorso-lateral in position, and the space between them is greater than that between the nostrils. The spiraculum is sinistral, ending in an attached tube, directed upward and backward, scarcely visible from above and below. It is nearer to the vent than to the tip of the snout. The vent is dextral, with a large oblique opening under a skin fold that overlies it and is continuous with the ventral tail crest. The tail is long and strong, with its muscular part strongly developed. The tip of the tail is blunt, with the tail crest about equally developed dorsally and ventrally. The dorsal tail crest is thickened at its anterior end and only indicated by a very low ridge near the base of the tail. The ventrally placed mouth is large and strong (fig. 20, B). A single row of papillae, variable in size, is developed on the margin of the lips, except for a Aelurophryne brevipes: Measurements of Tadpoles Ratio to Measurements Localities Range Average _ body length (%) Body length (snout to vent)...... Pamei 24-26 25.4 eas Taining 23-26 24.3 ee Body height (highest region)..... Pamei 10-12 10.7 42.3 Taining 10-12 10.6 43.6 Body width (greatest diameter)... Pamei 13-16 14.4 56.5 Taining 12.5-15.0 13.7 56.4 Head height (at eye region)...... Pamei 6.0-8.1 7.3 2520 Taining 6.0-7.0 6.9 28.4 Head-widthets arcs cere ances Pamei 10.5-138.0 14 44.9 Taining 9.0-11.5 10.4 42.8 Mouth width (including lips)..... Pamei 6-7 6.8 26.8 Taining 6-7 6.2 25.5 Space between eyes.............. Pamei 6-7 6.4 2520 Taining 6-7 6.6 27.0 Space between nostrils........... Pamei 4-5 4.3 16.7 Taining 4-5 4.3 17.4 Tip of snout to spiraculum....... Pamei 14.5-17.0 15.8 6252 Taining 14.0-15.5 14.6 60.0 Length of tail (vent to tip)....... Pamei 40-46 43.8 1 PAR Taining 37-46 40.8 168.0 Height of tail (highest part)...... Pamei 10-12 10.9 43.1 Taining 9-11 10.1 41.6 Diameter of tail muscle.......... Pamei 6.0-7.0 6.4 VASEr- (near tail base) Taining 5.5-7.0 6.0 24.7 Length of-hindvlimb:.5253 49.5 Pamei 6-14 aa Taining 5-9 6.6 Number of specimens measured: 10 from Pamei, 10 from Taining 132 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 median gap above. Papillae on the mid-dorsal margin of the upper lip usually have pigmented centers and the last two papillae near the mid-dorsal notch are usually largest. Additional papillae, mostly with teeth, are found at the corners of the mouth. The number of rows of labial teeth varies greatly, but there are usually five rows on the upper lip and six on the lower. All the labial teeth are supported by thick fleshy ridges. The uppermost row is very short and continuous, and partly fills up the gap where no papillae are developed. The remaining four rows on the upper lip are all interrupted, the innermost row the shortest. On the lower lip, with six rows of labial teeth, the outermost row is continuous and longer than the outermost row of the upper lip. The other five rows on the lower lip are all interrupted, the innermost row being the shortest. The mandibles are thick and strong, with large sharp denticulation of their edges. The number of rows of labial teeth varies greatly in the tadpoles of this species, especially in those collected from different localities. Ten tadpoles selected at random from each of three different localities were examined. In the Taining group, the tooth formula is as follows: eight have I:4-4/1:5-5; one has I:5-5/1I:5-5; one has [:5-5/1:6-6. In the Pamei group, four have 1:4-4/1:5-5; four have I:5-5/1:5-5; two have I:6-6/1:6-6. Aelurophryne tainingensis sp. nov. Type.—No. 49395 Chicago Natural History Museum, from Taining, 11,500 feet altitude, Sikang, China. Adult male, collected July 29, 1943, by Ch’eng- chao Liu. Diagnosis.—A very distinct Aelurophryne (pl. 4, fig. 4) with many rounded light-colored warts on the back, marbled with many smaller ones scattered among them; crowded rounded warts on the belly of the male, confined to the posterior belly region in the female; size small, body slender and depressed; a pair of squarish flattened chest glands narrowly separated from each other, and a pair of oval axillary glands with two-thirds of their inner margins in contact with the lateral margins of the chest glands in the male; the inner two fingers of the male with nuptial pads, without strong black asperities; tongue notched behind, no vomerine or maxillary teeth; no prehumeral glandular area; no warts on the soles of the feet. Description of type-—Body slender and dorso-ventrally depressed; head scarcely longer than broad, depressed; snout rounded, slightly projecting beyond the mouth, and longer than the length of the eye; canthus rostralis obtuse; loreal region oblique, concave; nostril nearer to the tip of the snout than to the anterior border of the eye; interorbital space as broad as the space between the nostrils and greater than the width of the upper eyelid; tympanum hidden; jaws rather weak; no vomerine teeth; no vocal sacs in the male. Fingers long and rather slender (fig. 21, B), the first as long as the second, with light-colored nuptial pads on the inner dorsal sides; the fourth finger longer than the first and second but much more slender; the third the longest and also LIU: AMPHIBIANS OF WESTERN CHINA 133 slender; subarticular tubercles indistinct; palmar tubercles developed but very flat; tips of fingers lighter in color. Hind limb short and weak, its length 125 per cent of the body length, the tibio-tarsal articulation reaching the base of the arm, the heels not meeting when placed at right angles to the body; tibia 38 per cent of the body length; toes (fig. 21, C) slightly fringed, their tips rounded and slightly dilated, the two inner toes about one-third webbed at base; subarticular tubercles present but Fic. 21. Aelurophryne tainingensis; male. A. Ventral view of head and thoracic region (xX 1). B. Ventral view of hand (Xx 2). C. Ventral view of foot (xX 2). indistinct; inner metatarsal tubercle elongated, with a free edge, outer metatarsal tubercle wanting. Skin rough, with many large and small pitted warts, more or less evenly distributed on the back; smaller warts on the exposed parts of the limbs; a strong glandular fold from the posterior corner of each upper eyelid, descending obliquely to the forelimb; large warts near the anal region; numerous warts varying in size on the posterior aspect of the thighs, extending distad to a large light-colored wart at about the middle of the thigh. Belly with numerous smooth warts, with smaller ones scattered on the inner and ventral sides of the arms. Two pairs of large glands on the thoracic region (fig. 21, A); the chest glands, the inner pair, slightly longer than broad, rounded anteriorly, the posterior margin squarish, with indentations; the outer pair, the axillary glands, oval, the anterior two-thirds of the inner margin in contact with the posterior lateral margin of the chest gland; both pairs smooth and without black asperities or spines; light-colored nuptial pads developed on the inner dorsal sides of the first two fingers. Measurements of type (percentage of body length in parentheses).—Body length 51.0 mm.; head length 16.0 mm. (81.3); head width 16.0 mm. (81.3); interorbital space 4.4 mm. (8.6); length of chest gland 5.0 mm. (9.8); width of chest gland 5.5 mm. (10.7); space between chest glands 0.33 mm.; length of axillary gland 4.5 mm. (8.8); width of axillary gland 2.5 mm. (4.9); length of lower arm 25.0 mm. (49.0); diameter of lower arm 6.0 mm. (11.7); length of 134 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 hand 13.5 mm. (26.4); length of leg 63.5 mm. (124.5); length of tibia 19.5 mm. (38.2); length of foot and tarsus 33.0 mm. (64.7); length of foot 23.0 mm. (45.0); length of inner metatarsal tubercle 4.0 mm. (7.8). Aelurophryne tainingensis: Measurements of Five Male and Two Female Adult Paratypes Ratio to Measurements Sex Range Average body length (%) Bodytlengths 5 Ass en oe rou 47-52 48.8 ae 9 56-57 56.5 Bin Headwiength::.% mtg sce ale eee: ou 15-17 16.0 32.9 9 16.5-17.0 16.7 29.6 Head! widths. 000242075 whee es of 15.6-17.0 16.2 33.2 ce} 17.5-18.0 b by bey 31.6 Interorbital space 2s. <5ciceeawen as of 3.8-4.0 4.0 8.1 ie) 4.5 4.5 7.9 hength ofiowerarm :.. ance. J 22 ..5-24.5 23.9 48.9 9 26 26 46.0 Diameter of lower arm.............. roe 5.4-6.0 5.8 11.8 Q 4.5 4.5 7.9 Bengthvon handes: 9 ananassae roe 12.0-13.5 12.9 26.4 Q 14.5-15.0 14.7 26.1 TAngtniOrieg vino. ce ee cee roe 63.0-64.5 63.7 130.5 Q 62-67 64.5 114.3 Bength: Or tibiae: cece ck se eas roe 17.0-19.5 18.6 38.1 Q 18.5-19.0 18.7 33.1 Length of tarsus and foot........... roe 28-32 30.1 61.7 Q 32 32 56.6 Length: of foot sin ccicascts ihe sears roe 21.0-22.5 22 45.1 Q 22 oe 38.9 Length of inner metatarsal tubercle... & 3.2-4.0 3.6 7.5 Q 3.5-4.0 3.7 6.6 Coloration in life-—Light brownish golden anteriorly and on the sides of the head, yellowish olive green on the back and dorsal sides of the limbs, with a dark brown mark from between the eyes extending posteriorly and becoming indistinct in the shoulder region; a dark brown band from the tip of the snout through the nostrils to the anterior corner of the eye, and descending from the posterior corner of the eye below the light-colored glandular fold to the shoulder. Below this dark brown band, on the sides of the jaws, light brownish golden color; some brownish marblings on the limbs. Warts lighter in color than their sur- roundings and mostly greenish yellow. Yellowish creamy on the throat and chest and yellowish on the belly, with many yellowish warts; ventral sides of limbs also yellowish. The axillary glands yellowish creamy, and the chest glands dark gray. The pupil black, vertically oval, enclosed by a golden ring; the dorsal part of the iris lighter in color, the ventral part darker, stippled with gold. After preservation, dark gray on the back with a distinct dark brown pattern between the eyes extending backward and becoming indistinct in the shoulder region; warts light gray; light dirty gray on the throat and chest, and gray on the belly, with many lighter-colored warts. LIU: AMPHIBIANS OF WESTERN CHINA 135 This new frog is not acommon form. In July, 1988, M. C. Chang of West China Union University got one specimen from Taining. Description was re- served for additional information and material. From the end of June to the middle of September, 1943, I had the opportunity to go to Sikang for field studies, and on July 29, 1948, I collected three adult males, two females, and eleven young specimens from a small mountain stream east of the city of Taining. On August 24, 1943, I collected two adult males and one female from the upper reaches of a small mountain stream east of “Eight Beauties Village,’ Pamei. The altitude of this locality is 11,000 feet, with a temperature of 42° to 60° F. on that date. Pamei is about fifteen miles southwest of the city of Taining. Description of paratypes.—Variation in structure and coloration is very slight in five adult males and two females. Sex dimorphism in size is marked, the average body length of the males being 48.8 mm. (47-52 mm.) and that of the females 56.5 mm. (56-57 mm.). The female is darker than the male. The male is much more rugose than the female. The warts on the back of the female are fewer in number, much larger, and mostly more elongated than in the male. In the female there are two lateral rows of large rounded warts extending from the shoulder region above the base of each arm posteriorly to the groin, and lateral to this there are many light-colored and rounded warts. At the mid- posterior aspect of each thigh of the male there is a very light-colored gland, not found in the female. Enclosing the latero-ventral sides of the anus in both sexes there are elevated glands not found in other species of Aelurophryne. Chest glands are found only in males, axillary glands in both males and females. The whole belly is covered by large rounded warts in the male, whereas in the female such warts are confined to the posterior and middle of the belly. In the male, the margins of the jaws bear glandular areas. The arm is distinctly stronger, longer and inwardly bent in the male. The ratio of the diameter of the lower arm to the body length is 12 per cent in the male and only 8 per cent in the female. The hind limb of the male is distinctly longer than that of the female. The webbing between the toes is similar in both sexes. No vocal sac or linea masculina is developed in the male. Habitat and habits—Aelurophryne tainingensis is a high mountain form. According to our present knowledge it is an endemic species of the high plateau of Sikang, found around Taining (11,500 feet). The habitat of this species is in the upper reaches of very small mountain streams where the water trickles through beneath the stones. There is no vegetation where I found my specimens, otherwise the habitat would be very much like that of Scutiger schmidti. Aeluro- phryne tainingensis is not a good swimmer, as its legs are short and weak and its webs poorly developed. It may be a more terrestrial frog, hiding itself under stones for moisture. The depressed body, short legs and rather well-developed inner metatarsal tubercle are fitted for terrestrial life under stones or in crevices of stones on hillsides. The time and site for breeding of this new frog are still unknown. The breeding season of tainingensis may be earlier than that of Aelwrophryne brevipes, 136 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 as egg-masses of brevipes were collected on July 29, 1943, when no eggs that could be those of tainingensis had been seen. Aelurophryne maculata sp. nov. Type.—No. 55869 Chicago Natural History Museum, from Hasa, 11,000 feet altitude, Kantze, Sikang, China. Juvenile male, collected August 19, 1943, by Ch’eng-chao Liu. Diagnosis.—A small species of Aelurophryne (fig. 22) closely related to A. tainingensis, differing in being without warts on the belly; in having the warts Fic. 22. Aelurophryne maculata. A. Dorsal view (xX 1). B. Ventral view of hand (xX 4). C. Ventral view of foot (x 4). on the back very small and dark; toes about two-thirds webbed; no solid dark brown triangular mark between the eyes and on the anterior part of the back; no large light-colored flat wart at the mid-posterior aspect of each thigh; sub- articular tubercles on fingers and toes well developed; fringes on toes very strongly developed; no warts on soles of feet. Description of type-—Tongue slightly notched; no vomerine teeth; jaws weak and without maxillary teeth. Body moderately stout; head scarcely broader than long, depressed; snout rounded, slightly projecting beyond the mouth, about as long as the eye; canthus rostralis obtuse, with loreal region oblique and concave; nostril nearer to the tip of the snout than to the anterior corner of the eye; internasal space narrower than the interorbital space, which is wider than the upper eyelid; tympanum hidden. Arm moderately long, fingers (fig. 22, B) slender and long, the first slightly shorter than the second, the fourth longer than the second and the third the LIU: AMPHIBIANS OF WESTERN CHINA 137 longest; palmar tubercles prominent, the inner larger and flat, the outer small but with elevated center; subarticular tubercles well developed; finger tips much lighter in color. Hind limb short and weak, the length of the hind limb 128 per cent of the body length; tibio-tarsal articulation reaching the base of the arm, heels not meeting when placed at right angles to the body; tibia 39 per cent of the body length; toes (fig. 22, C) greatly flattened, with wide fringes, tips with very small dilations, about two-thirds webbed; subarticular tubercles well developed; inner metatarsal tubercle flattened, elongated oval, with a free edge; no outer meta- tarsal tubercle. Skin moderately rugose, with very small warts on the anterior part of the back, with much larger ones on the sides and posteriorly and with some small warts on the upper sides of the limbs; medium-sized warts developed around the anal region; skin smooth on the throat, the belly, and the ventral sides of the limbs. A thick glandular fold from the posterior angle of the upper eyelid extend- ing posteriorly and obliquely to the shoulder. Measurements of type (percentage of body length in parentheses).—Body length 49 mm.; head length 16 mm. (82.6); head width 17 mm. (84.6); inter- orbital space 4 mm. (8.3); length of lower arm 24 mm. (48.9); diameter of lower arm 4 mm. (8.3); length of hand 13 mm. (26.5); length of leg 63 mm. (128.5); length of tibia 19 mm. (388.6); length of foot and tarsus 32 mm. (65.3); length of foot 22 mm. (44.8); length of inner metatarsal tubercle 3.7 mm. (7.5). Coloration in life-—Olive green on the back and greatly marbled with dark brown, which usually encloses the olive green warts; limbs similar but the con- cealed parts of the hind limbs lighter and more pinkish; the throat, the belly, the ventral sides of the limbs and the inner dorsal sides of the hands and the feet pinkish cream. After preservation, gray is the ground color on the back and on the upper sides of the limbs, marbled with darker; colorless on the throat, the belly and the ventral sides of the limbs and the inner dorsal sides of the hands and feet. Distribution.—This new frog was found under a stone at the edge of a small mountain stream close to Hasa, about 11,000 feet altitude. Many tadpoles and a few adults of Aelurophryne brevipes and adults of Rana temporaria chen- sinensis were collected along the same stream. After I discovered the new frog, we worked long and hard, but we failed to find another specimen. The new Aelurophryne is notably distinct in the field in its remarkable coloration. Aelurophryne glandulata sp. nov. Type.—No. 49392 Chicago Natural History Museum, from Hopachai, Lifanhsien, Szechwan, China, altitude 8,500 feet. Adult male, collected August 9, 1941, by H. C. Chang. Diagnosis.—An Aelurophryne (pl. 4, fig. 6) with a long foot, nearly 51 per cent of the body length; foot rugose beneath; tibio-tarsal articulation reach- 138 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 ing the shoulder region; the inner two toes more than two-thirds webbed, the third and fifth toes about one-half webbed; chest glands without strong spines, large, elongate oval, diverging posteriorly; axillary glands long and prominent, the anterior third of their inner borders in contact with the chest glands; pre- humeral glandular area distinctly developed. Description of type-—Body stout; head depressed, slightly broader than long; snout rounded, slightly projecting beyond the mouth and much longer Fic. 23. Aelurophryne glandulata; male. A. Ventral view of hand (xX 2). B. Ventral view of foot (x 2). than the length of the eye; canthus rostralis obtuse; loreal region very oblique, only slightly concave; nostrils about mid-way between the tip of the snout and the anterior corner of the eye; interorbital space wider than the distance between the nostrils, and broader than the width of the upper eyelid; tympanum hidden; jaws weak, without teeth; tongue oval, slightly emarginate behind; no vomerine teeth; no vocal sac. Arm strong; fingers (fig. 23, A) slender and long, first and second equal, thicker, and shorter than third and fourth, the third finger the longest; nuptial spines developed on inner dorsal sides of the first and second fingers; tips of fingers lighter in color; subarticular tubercles present but not differentiated from other tubercles on the fingers, much as in the Chinese common toad; palmar tubercles large and prominent, the inner larger and flat, the outer much smaller; palm rugose with small warts. LIU: AMPHIBIANS OF WESTERN CHINA 139 Hind limb short and weak, its length about 142 per cent of the body length; tibio-tarsal articulation reaching the shoulder region; tibia 42 per cent of the body length; foot (fig. 23, B) longer than those of other species of Aelurophryne, about 51 per cent of the body length; toes prominently fringed; the two inner toes about two-thirds webbed and the third and fifth toes about half webbed; tips of toes rounded, slightly dilated, lighter in color; subarticular tubercles present, but not distinct, like those of the fingers; inner metatarsal tubercle well developed, elongated oval, with a free edge. Skin rough, with flat, irregularly shaped and pitted warts on the back and rounded pitted ones on the sides of the body; numerous small warts without pits Fic. 24. Aelurophryne glandulata; male. Ventral view of head and thoracic region (x 1). scattered among the rounded warts, fewest on the back; the pitted warts dark brown in color; small warts on upper sides of limbs; a large, ill-defined, flattened parotoid gland behind the eye; a strong glandular fold at latero-ventral margin of each parotoid, extending obliquely to the base of the arm; many rounded light- colored warts behind and at the base of the thighs; skin of the throat, chest, belly and the ventral sides of the limbs smooth, except for glandular areas on the margin of the jaw (fig. 24); two pairs of flattened glands on the chest, the inner pair oval, large, and flat, with fine black spines on some of the granules of the gland, 16 mm. long, about 20 per cent of the body length; the lateral pair (the axillary glands) smaller, elongate oval and much more elevated than the chest glands; latero-dorsal to the chest gland and ventro-anterior to the base of the arm, a definite area covered by thinner skin with scattered small warts; belly much wrinkled in the preserved specimen. Coloration in life-—The ground color of the head, the back and the sides of the body and the dorsal sides of the limbs is olive. A dark brown bar crosses the posterior interorbital space. The top of the snout is much lighter than the other parts of the body. The anterior region of the upper eyelid and its margin are golden stippled, more so than other parts of the body. The snout has dark brown marblings and a brown band extends from the snout through the nostril 140 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 and the edge of the upper eyelid to the latero-ventral side of the glandular folds, ending at the base of the arm. Irregular dark brown spots are scattered on the back and sides of the body and the dorsal sides of the limbs and digits. The exposed parts of the lower arm and leg have more golden stipples and the digits are lighter in color, especially the two inner fingers and toes. Webs are colored like the corresponding digits, stippled with dark brown. Pupils vary in shape with the intensity of light, but are always vertically elongated; the lower part of the iris is darker than the upper, depending on the amount of golden stippling. A bluish membrane is sometimes visible on the ventral inner margin of the upper eyelid. The throat is gray, with some indication of purple, and lightly marbled with ochraceous buff. The large anterior median glands of the chest are purple- gray, stippled with bluish black, and the lateral pair is cream-buff stippled with gold. Belly is brownish ashy. There are round greenish-yellow warts on the ventral side of the base of the thigh and the vent. The coloration of the ventral sides of the arms and legs is like that of the belly, but the palms and soles corre- spond in color with the upper sides of the hands and feet. The coloration varies slightly among the five specimens available. The sides of the body, especially the groin region and the latero-ventral sides of the limbs may be greenish yellow or light olive green. After preservation, the back is dark gray, with scattered dark brown patches, and the dorsal sides of the limbs have small whitish warts. Throat, belly, and under sides of limbs are light grayish cream. Distribution.—This new frog was discovered at Hopachai, about 45 miles from the city of Lifan. The altitude of Hopachai is 8,500 feet, and the tempera- ture was only 35° F. at 6:00 A.m., August 9, 1941. On this morning Mr. H. C. Chang found the first specimen under a stone in a small mountain stream by the side of a larger stream. We turned over nearly all the stones of that small stream, and an additional adult male, an adult female, two young specimens, and tadpoles, were obtained. On August 10, while we were on the way to the top of Tsingshan, we collected another female from a side branch of a mountain stream at an altitude of about 9,500 feet. No specimens were found during four days spent in a lumber company house near the top of Tsingshan, at 11,000 feet. On August 15, when we came back to Hopachai, we collected an egg-mass and tadpoles at various stages from another very small mountain stream. Comparison with related species.—Aelurophryne glandulata can easily be distinguished from mammata and other species by the chest glands, as in glandulata these glands are very large and without the spines of the other species. It differs from A. tainingensis and A. maculata in size, the length of the body being about 80 mm., while in tainingensis and maculata it is never over 60 mm. In tainingensis there are rounded warts on nearly the whole belly in the male, and on the posterior central region of the belly in the female, whereas in glandulata the belly in both sexes is smooth. The foot is rugose beneath in glandulata and smooth in tainingensis and maculata. A. glandulata is more closely related to A. brevipes than to other species of the same genus, but in glandulata the two chest glands are separated by a space 1.2 per cent of the body LIU: AMPHIBIANS OF WESTERN CHINA 141 length, and in brevipes this space amounts to 3 per cent; the axillary glands are longer and wider in glandulata than in brevipes; and the anterior third of the ‘axillary gland is in contact with the postero-lateral side of the chest gland in glandulata, this contact being about two-thirds in brevipes. The hands and feet are much longer in glandulata than in brevipes, the length of the hand being about 30 per cent of the body length in the former while it is only about 25 per cent in the latter. The length of the foot is about 51 per cent of the length of the body in glandulata, while in brevipes it is 41 per cent. There are more spines on the first and second fingers (27-37) in glandulata and fewer (17-24) in brevipes. Aelurophryne glandulata: Measurements of Adults Male (type) Male Female Female Ratio Ratio Ratio Ratio to body to body to body to body length length length length Measurements MM. (%) MM (%) MM. (%) MM. (%) Body. length® 0235.52 2c SION ac OO See OL Om ets EMU On eae: Headblengthi sere. terse 2hsObinos- 0 LAU e216 ua 2Oc On moa le et ote Oem ollg Head! width: 2.22320 he5 2580" SO 5 Zouk s Sosa 205 Ole tease ete O) o520 Interorbital space......... 7.0 8.6 5.0 7.4 7.0 8.6 525 fea Length of lower arm...... AZ 2b: 4b2s4 320. 52625 74100" 5076 74050) 5129 Diameter of lower arm.... 14.0 17.2 835.7 1226 8.4 10.3 S508 eles ILength.ot:leg. 2.3324 sees 11520 141295 49-8820" 13s e100" 1352 8:-10220) 13254 Mength. of tiblasisc.J 42455 DAD y SAD De atom AOS doleO- wioS42s Sec Oe ooo Length of foot and tarsus.. 60.0 74.0 46.0 68.6 55.0 67.9 52.0 .67.5 Pengthvorfoote: A320 05 a Alor sol. 2. Noac08 Alek ste" 45.02 236.54 -41e4 Length of inner metatarsal GUDERCIE? Sao Renee. 6.3 of Cesi 4.4 6.5 C20 8.6 4.5 5.8 Sex dimorphism.—Sexes do not differ in size; body length of males is from 67 to 81 mm., while that of females is from 77 to 81 mm. The arm of the male is much stronger than that of the female. On the ventral region of the lower jaw of the male specimens there are many glandular areas (fig. 24), the median ones well developed and tending to be fused. Black pigment is irregularly scattered on the purplish glandular areas. In the males there are two pairs of flattened glands, one pair on the chest (the chest glands), and the other near the axilla (the axillary glands). In the female, only axillary glands are present. The first and second fingers are provided with spines in the male (fig. 23, A) but not in the female, and these two fingers are obviously stronger in the male than in the female. Habitat and habits.—The adults of Aelurophryne glandulata are found under stones of the side branches of mountain streams from 7,500 to about 9,000 feet altitude. Their behavior—their slow awkward movements and their way of walking and jumping—is like that of the common toad. When it is found under a stone, the frog does not jump or move away. We turned over hundreds of stones along the river from 7,500 to 11,500 feet altitude, and also stones on the bank of the river, but only five animals were found. The real home of this frog may be on the hillsides in holes under stones near streams. 142 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 Breeding.—The breeding season may be from July to August, as we collected tadpoles of different stages, including newly hatched individuals and tadpoles with four limbs well developed. Spoiled egg-masses were also obtained. On August 10, 1941, one large and three small egg-masses were found under boulders in a very small slow-running stream near Hopachai. The eggs were not measured, B Fic. 25. Aelurophryne glandulata; tadpole. A. Ventral view (xX 214). B. Mouth (Xx 16). as they were in a late stage of development and most of them were already spoiled; they are large and creamy white in color, enclosed in jelly that connects them strongly to each other and attaches them to the under surface of the boulder, The development is similar to that of Rana boulengeri, the embryo being rather small, with the tail bent to one side and applied to the surface of the large yolk of the egg. After hatching, the tadpoles escape by making a hole in the jelly capsule at the side facing the water. The just-hatched tadpoles may stay with the jelly capsules. One case perplexed me very much; one egg-mass was not attached to the stone, but was found on the bottom of a pool with many tadpoles accumu- lated on it. This mass of jelly may have become detached from a stone, as in some cases in Staurois chunganensis. The jelly may supply food for the young tadpoles, as many of them are found on the capsules. LIU: AMPHIBIANS OF WESTERN CHINA 143 Tadpoles.—The young tadpoles are found in small pools in brooks and also in side pools or in branches of the main stream. In the small brooks they aggre- gate together on the jelly capsules under stones or on the bottom of the water; those in larger bodies of water, as in the main stream, are scattered and hide under stones. The larger tadpoles scatter and hide under stones, mostly near the bank or in temporary pools of the main stream. The tadpoles are very sensi- tive; when disturbed they swim very quickly through the crevices between the stones. At the time of metamorphosis, the tadpoles are found under stones at the margin of the water. It seems likely that the tadpoles found in the water of the main river had been washed down from small brooks where the eggs are laid, as no egg-masses were found in the main river. The young tadpole is dark brownish gray stippled with gold; the snout region is lighter; and the tail fin is pearl gray stippled with brownish gray, the dorsal fin darker than the ventral fin. The tadpole with hind limb-buds developed is similar in coloration, but with much more golden stippling on the body. Con- spicuous patches of golden stippling are present on the back of the head and the dorsal part of the muscular portion of the tail. The pupil is round and black, with a black-marked golden iris. The tail fin is darker than that of the young tadpole. The belly is colorless. . ; Twenty tadpoles (pl. 10, fig. 7; text fig. 25) with hind limb-buds ranging from 2 to 7 mm. long were measured; the average body length from the snout to the base of the tail is 23.8 mm., ranging from 21.5 to 25 mm. The form of the body varies with the habitat. The tadpoles collected from running water or slow- running water by the side of the river have a somewhat elongated and cylindrical body, and those found in stationary water with very rich vegetation have a very clumsy body with the abdominal region greatly enlarged by food, the head being greater in width than in height. The ratio of head width to body length is 48 per cent and that of the height of head is only 33 per cent. The length of the tail is 178 per cent of the body length, with a moderately developed tail fin and tail muscle. The snout is rounded, with the mouth more ventral than anterior and about one-fourth of the body length, like that of Megophrys oshanensis. The lips are long. The nostrils are nearer to the snout than to the eye, and are surrounded by lighter-colored rings. The eyes are latero-dorsal in position, the space between them being 23.2 per cent of the body length. The spiraculum is sinistral, ending with a very short tube directed upward and backward and visible from above and below. It is nearer to the base of the tail than to the tip of the snout. The vent is dextral, with a broad oblique opening overlapped by a skin fold continuous with the ventral tail fin. The tail is moderately long, with a blunted tip, and the dorsal tail fin is slightly deeper and darker than the ventral fin. The dorsal fin is very low and thick near the base of the tail and the ventral fin extends to the vent. The thickness of the muscle near the base of the tail is one-fourth of the body length. 144 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 Aelurophryne glandulata: Measurements of Tadpoles Ten Specimens from Running Water and Ten from Quiet Water Ratio to Measurements Range Average __ body length (%) Body length since a7 scents Running water 21.5-24.5 23.3 ie: Quiet water 22 .5-25.0 23.4 eee Bodyheighes .:ccixesecc. = 23 Running water 9-11 10.1 41.2 Quiet water 9-13 11.0 47.0 IBOOYeWIGth's 303 thse tes | Running water 10.2-13.0 11.6 49.7 Quiet water 11-17 13.5 57.6 Head height.c0c he Sk Running water 7-9 7.8 33.4 Quiet water 7-9 7.9 33.3 Head width: acc. cance es: Running water 9.5-11.5 10.5 45.0 Quiet water 11-13 11.8 50.4 Mouth width: 225.0325. .5-.: Running water 6.0-6.8 6.2 25.7 Quiet water 5.5-7.0 6.1 26.0 Space from eye to eye...... Running water 4.8-5.8 5.4 23.1 Quiet water 5.2-6.0 5.5 23.5 Space from nostril to nostril. Running water 4.04.5 4.1 17.5 Quiet water 4.0-4.5 4.1 17.5 Tip of snout to spiraculum.. Running water 13.0-15.5 14.3 61.3 Quiet water 13.0-15.5 13.7 58.5 Taillength eA ssc.ce ocvo ah Running water 38-46 41.1 176.3 Quiet water . 39-45 41.9 179.0 sPallphelgntz.20 ive, sats nese Running water 9.5-10.8 9.9 42.9 Quiet water 9-12 10.7 45.7 Diameter of tail muscle.... . Running water 5.5-6.2 5.8 24.8 Quiet water 5-7 6.3 26.9 Length of hind limb........ Running water 3.8-8.0 5.3 Quiet water 2-7 4.2 The variation of the labial teeth and papillae (fig. 25, B) in twenty tadpoles was studied. There is a row of papillae on the margin of the upper and lower lips, without or with a very small space on the mid-dorsal side of the upper lip; the papillae at the sides of the lip are larger than those of the lower lip; in some specimens, the smaller ones bend inward and give the appearance of two rows of papillae. A few additional papillae are found at the corners of the mouth, and in many cases there are two or more papillae located lateral to the second and third rows of labial teeth of the lower lip. Normally there are four rows of labial teeth on the upper lip, the first the shortest, and complete, the second the longest and narrowly interrupted, and the other two widely interrupted. On the lower lip, there are four or five rows of labial teeth, the outermost row short and complete and the other four rows all interrupted. In the twenty specimens studied, the frequency of tooth formula is as follows: ten specimens have tooth formula I:3-3/1:4—4; six have I:4-4/1:4-4; two have I:3-3/1:5-5; one has I:3-4/1:5-4; and one has I:4-4/1:5-5. The last four groups have the outer- most row of the lower lip interrupted. The mandible has a heavily serrated edge. Genus SCUTIGER Theobald Usage of the names Scutiger, Aelurophryne, and Cophophryne has become confused by the assumption of Boulenger (1887, p. 405) that Scutiger Theobald, LIU: AMPHIBIANS OF WESTERN CHINA 145 1868, in the Amphibia, is preoccupied by the earlier Scutigera Latreille of 1802, acentipede. According to the subsequent ruling of the International Commission on Zoological Nomenclature, this is not the case, but Procter (1922, p. 583) continued the use of the Boulengerian substitute name Cophophryne, which is a pure synonym of Scutiger. Procter, furthermore, refers Aelurophryne to Copho- phryne (and thus to Scutiger). Noble (1931, p. 493) correctly distinguished the toothless Aelurophryne from the toothed Scutiger, and correctly uses Scutiger for the Himalayan genus, but without adequate explanation. The discovery that Scutiger includes several distinct species re-enforces the desirability of maintaining it as a distinct genus. KEY FOR IDENTIFICATION OF ADULTS OF SCUTIGER I. Tibio-tarsal articulation reaching beyond the eye. A. Rounded black warts on back; very distinct black bars on limbs; webs hardly STOWE era cea Re eo Te Hc ert Ie, Piven PPE Tee ee pope. AA. Warts irregular, color dark as background; bars on limbs rather indistinct; toes one-fourth webbed ie Aas o see eee hia han ined tine at wetet tates & rugosa. II. Tibio-tarsal articulation not reaching beyond the eye. A. Skin smooth, color light, toes distinctly webbed........................ pingit. AA Skin rough, color: dark. toesinot wepbed hee sacs coats octactchawiete becca: schmidti. KEY FOR IDENTIFICATION OF TADPOLES OF SCUTIGER I. Body and tail uniform black, tooth formula I1:4-4/1:4-4..................... rugosa. II. Tail much lighter in color than the body. A. Color not uniform, with dark spots or light markings; tail muscle strong, diameter about one-third of body length. B. Black spots on the body, diameter of tail muscle about 34 per cent of the bodyslengthciaasska ah een Re or ete hace eer an ens Lungtung sp. BB. No black spots on the body but with a light mark on the dorsum near the base of the tail (very distinct in living specimens). C. Diameter of tail muscle about 35 per cent of body length. Mount Omei sp. CC. Diameter of tail muscle about 28 per cent of body length. Panlungshan sp. AA. Color uniform, no spots or marks; tail less than twice as long as the body. B. Tooth formula I:4-4/1:4-4; tail short and weak, its length about 155 per cent of the body length, tail muscle 19 per cent.................... pingit. BB. ‘Tooth formula more than I[:4-4/1:4-4. C. Size small (19-21 mm.); tail short and weak, its length about 168 per cent of the body length, diameter of tail muscle about 23 per cent. schmidti. CC. Size large (23-29 mm.); tail long, about 196 per cent of the body length, diameter of tail muscle about 27 per cent..................245. popet. Frogs of the genus Scutiger are rare Himalayan pelobatids first discovered by Blyth in 1854. The first species was described as Bombinator sikkimensis, from two specimens preserved in the Calcutta Museum. In 1887 Boulenger redescribed this rare frog as Cophophryne sikkimensis from three specimens (a male, a female and a juvenile) collected by W. T. Byntan at the foot of Yakla 146 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 Pass, Sikkim (altitude 13,000 feet). These three specimens in the British Mu- seum are apparently immature, as the measurement of the largest one was given by Boulenger as 50 mm. and Pope measured it as 48 mm. Pope’s manu- script notes record two more British Museum specimens, mature males collected from the Burma-Sikang border. Scutiger alticola was described by Procter (1922) as Cophophryne alticola from a single female specimen, collected from the Kharta Valley, Tibet (altitude 16,500 feet). The fully webbed toes and granulated abdomen of this species are not found in any of the Chinese species of Scutiger since discovered. The apparently rare frogs of the genus Scutiger now prove to be well repre- sented in China, as I have discovered seven additional species, four species with adults and tadpoles, and three species thus far known only from tadpoles and young. Scutiger sikkimensis and S. alticola have not been found in Szechwan or Sikang. Scutiger pingii Liu Scutiger pingiit Liu, Jour. West China Border Res. Soc., 14, (B): 35, fig. 5, 1948— Yenwot’ang, Chaochiao, Sikang; idem, op. cit., 14, (B): 71, 1948. History of species.—Scutiger pingii was discovered in 1942 and is still known only from the type locality. Comparison with related species.—Scutiger pingii (pl. 4, fig. 1) agrees with schmidti in having the tibio-tarsal articulation extending only to the angle of the mouth, thus differing from the longer-legged pope and rugosa; pingii differs from schmidti in having the toes distinctly webbed, the color lighter, and the skin smoother. Of all the species of this genus in western China only pingit has spines on the outer toes in the male. The body is stouter in pingii than in schmidti, popei or rugosa. Scutiger pingii is much lighter in color and less distinctly spotted on the back than the other three species, which have darker dorsal patterns or spots. The tadpole of pingii has a short, stout body and a short and weak tail, in association with its habitat of quiet or very slow-running water. Collection data.—On May 4, 1942, fourteen specimens were collected at two places between Hsuanshenpa and Lolokou, in Chaochiao, Sikang. Four more specimens were collected from the same locality on May 17 and eight more on June 14 and 15 of the same year. The altitude of these localities was from 9,000 to 11,000 feet, with the temperature from 44° to 68° F. on May 6 and May 12. Tadpoles in different stages and young just after metamorphosis were obtained from the same localities and on the same days that the adults were collected. Original description.—‘“‘Tongue elliptical, free behind and slightly notched, vomerine teeth lacking [fig. 26, B]. Head moderately large, slightly broader than long; snout rounded; canthus rostralis feebly indicated; loreal region oblique and flat; nostrils rather near to the eye and the internasal space about equal to LIU: AMPHIBIANS OF WESTERN CHINA 147 the interorbital space, which is slightly broader than the upper eyelid; length of eye about equal to the internasal space; tympanum lacking. “Fingers [fig. 26, C] long and slender, the first barely longer than the second, and both with nuptial spines on the inner dorsal sides. Subarticular tubercles developed; an inner large and an outer smaller palmar tubercle well developed. Arm rather long, length of the lower arm to the tip of third finger nearly half of the body length and its diameter about one-tenth of the body length; many small, rounded, light-colored warts on the antero-ventral sides of the upper and lower arms; small warts with black spines on the dorsal sides of the upper and lower arms, especially at the elbow region. “Hind limb rather short, tibio-tarsal articulation reaching the angle of the mouth; heels not meeting when placed at right angles to the body; tibia about Fic. 26. Scutiger pingii; male. A. Ventral view of head and thoracic region (xX 1). B. Oral cavity (x 2). C. Ventral view of hand (x 2). D. Ventral view of foot (x 2). 42 per cent of the body length. Webs between all the toes [fig. 26, D], about one- third to one-fourth webbed; all the toes fringed, with a row of black spines on the outer and inner fringes of the fifth toe and only the outer fringe of the fourth toe; subarticular tubercles developed but not very distinct; inner metatarsal -tubercle elongated, rod-like and greatly elevated, its length about 6.3 per cent of the body length; outer metatarsal tubercle lacking. “Skin of the back with small flattened warts; warts on the shoulder region and on the dorsal sides of the limbs with black spines. Length from snout to vent 43 mm. Fine black spines developed on the chest glands. “The dorsum is rufous brown with many indistinct darker spots that usually coincide with warts. The throat and the belly and the under sides of limbs are fleshy-purple as in Kaloula. The color of the upper sides of the limbs is like that of the back, with indistinct bars only on the femur. There is a golden- creamy axillary gland on the chest at the base of each arm. On the middle posterior aspect of each femur there is a more or less round, light-colored, flattened 148 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 gland. In living specimens, the pupil is changeable in shape, but always more or less vertical, with golden iris stippled with black, especially at the lower side. “There is no sexual dimorphism in size. The average body length of the two mature males is 43.5 mm., ranging from 43 to 44 mm.; and the average body length of the nine females is 40.4 mm., ranging from 37 to 46 mm. The arm of the male is moderately enlarged and much thicker than that of the female. In the male [fig. 26], there are many black nuptial spines on the inner dorsal side of the first finger, extending to the medial side of the wrist, and then less numerous on the inner dorsal side of the second finger. There are many black spines on the ventral margin of the lower jaw and many spines aggregated together on the antero-median ventral side of the floor of the mouth of the male. Two elliptical areas of fine black spines are developed on the chest glands of the male; many strongly developed spines on light-colored bases are scattered on the shoulder and tympanic region and on the dorsal sides of the limbs. It is very interesting to notice that in the male of this species characteristic spines are found on the lateral margins of the fifth toe and on the outer margin of the fourth toe. The three inner toes are about one-fourth webbed, and webs between fourth and fifth are rudimentary in the female and all the toes of the male are about one-fourth to one-third webbed.” Scutiger pingii: Measurements of Two Males and Nine Females Ratio to Measurements Sex Range Average __ body length (%) Body. lengths <2: ecxtracacrsaanene haere fof 43-44 43.5 eae Q 37-46 40.4 aes Headslength* nen tos reece rot 13.5-14.5 14.0 32.2 Q 13-16 14.4 35.6 Headewidth). .saas ern tek oe rot 15.0-15.5 15.3 35.1 Q 13-16 14.4 35.6 interorpital space... s2+4: cc ee cece rot 4 4.0 9.2 9 3.6-4.8 4.0 9.8 Length of lower arm................ roe 20-22 21.0 48.5 Q 18-24 19.9 48.9 Width Onlower arm cea sac eee ee rot 5.0-5.5 5.3 12.1 Q 3.5-4.0 3.8 9.3 Length; of hand s< scas023c6e08.555408 fou 12 12.0 27.6 Q 9.5-13.0 10.9 27.0 ihength of leg acces sae rot 60-64 62.0 142.5 f°) 51-67 55.8 137.9 Lengtivotibigst..00 ee te. a 18-19 18.5 42.5 9 15-20 16.8 41.5 Length of foot and tarsus........... roe 29-31 30.0 69.0 Q 20-32 26:7 66.0 Mength’ of 100t 3.545 cle use es fof 20-21 20.5 47.1 9 15-22 18.2 45.1 Length of inner metatarsal tubercle.. 2.5-3.0 2.8 6.3 ) 2.0-2.6 2.1 5.2 Measurements of type (percentage of body length in parentheses).—Body length 43 mm.; head length 15 mm. (34.9); head width 15 mm. (34.9); interorbital LIU: AMPHIBIANS OF WESTERN CHINA 149 space 4 mm. (9.3); length of lower arm 21 mm. (48.8); length of hand 12 mm. (27.9); length of hind limb 54.5 mm. (126.7); length of tibia 18 mm. (42.0); length of tarsus and foot 29 mm. (67.5); length of foot 20 mm. (46.5); length of inner metatarsal tubercle 2.7 mm. (6.3). Habitat and habits—Scutiger pingii is a high mountain form found under stones in slow-running water of small mountain streams or under stones on the Fic. 27. Scutiger pingii; tadpole. A. Lateral view (xX 3). B. Mouth (x 12). margins of small pools at about 10,000 feet altitude. The adults are very inactive and behave very much like those of Kaloula. They were found to be more abundant the first part of May in quiet or running water and in the first part of June very few specimens were found near water. This may indicate that the breeding season is early in May, although no eggs were found. Tadpole.—Tadpoles (fig. 27, A) of this new Scutiger are rather sensitive. Under undisturbed conditions, they swim singly or in small groups in slow-running water, but when disturbed they hide in the cracks between stones or under stones. They are bottom feeders with strong mouth-parts (fig. 27, B). Their coloration varies slightly in different stages. The background color of the back and the sides of the body is dark gray, densely stippled with gold and brown. The tail is gray and much lighter toward its tip, where it is also stippled with gold and brown. Toward the time of metamorphosis, the color becomes lighter. The tadpoles are rather large. Ten tadpoles with hind limbs ranging from 2 to 12 mm. in length were measured. The average body length from the tip of 150 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 the snout to the base of the tail is 25 mm., ranging from 23 to 26.5 mm.; for the tail, the average length is 39 mm., with a range from 36 to 43 mm. The width of the head is 43 per cent of the body length and its height is only 31 per cent. The body is somewhat flattened dorso-ventrally; its height is 42 per cent of the body length and its width is 50 per cent. The length of the tail is about one and a half times the body length and its height two-fifths. Scutiger pingit: Measurements of Ten Tadpoles Ratio to Measurements Range Average _ body length (%) IBOdylength er tee a aes 23 .0-26.5 25.0 Fei Body heights: fice eee ay re Oe 9.5-11.0 10. 42.0 Bod yswidth errno asec ec ee 11.5-13.0 12.6 50.4 Head heignt.-:0..-0ek Ag SAT onal MOT ee 7.5-8.0 7.8 31.2 HMeadawidth misters rc ae 10-11 10.8 43.2 Mouth widths2.. rine ee eee 5.2-6.0 5.7 22.8 Space: between:.eves %.3.. 55.5 Se et ae es 5.5-7.0 6.4 25.6 SNOULLO SpIFACUIUM saceetios Coie ae ee ee 14-16 15.3 61.2 Tailtiength Wee soe ent ert ce oy ec 36-43 38.8 155.2 Taitheiphts ver. arene ne ee aah noe 9-11 9.9 39.6 Diameter of tail muscle (near base of tail)... 4-5 4.7 18.8 Wengthiot hind Hm ch eee oh or 2-12 ere ae The snout is rounded, and the nostril, enclosed by an elevated ring, is about mid-way between the tip of the snout and the eye. The eyes are dorso-lateral in position but more dorsal than lateral, with an average space between the two eyes of 26 per cent of the body length. The spiraculum is sinistral, ending in a semi-tube directed upward and backward and visible from above and below. It is 15.3 mm. from the tip of the snout to the spiraculum, which is nearer to the base of the tail than to the tip of the snout, as the distance from the snout to the spiraculum is 61 per cent of the body length. The vent is dextral, with a large oblique opening, under a broad skin fold that overlies it and is continuous with the ventral tail crest. The tail is weak. The tip of the tail is blunt, with the tail crest about equally developed dorsally and ventrally. The dorsal tail crest is thickened at its base, but is not continued onto the body. The mouth is ventral, with an average width of 5.7 mm., which is 23 per cent of the body length. A single row of papillae is developed on the margin of the lip. There are five rows of teeth on the upper lip. The outermost row is short and continuous, mostly filling up the gap where there are no papillae. The remaining four rows are all interrupted, the first row next to the short continuous row being the longest, and the innermost row the shortest. On the lower lip, there are also five rows of teeth. The outermost row is short and continuous, but it is longer than the outermost row of the upper lip. The other four rows are all interrupted, the fourth row the longest, and the innermost row the shortest. Teeth are also found on some of the additional papillae at the corners of the mouth. The mandibles are thick and heavy, with strong sharp teeth on the edges of both upper and lower mandibles. The number of rows of labial teeth is constant; the formula is I:4-4/I:4—4. LIU: AMPHIBIANS OF WESTERN CHINA 151 The young just after metamorphosis has all the characters of an adult except for the secondary sex characters. From snout to vent, it measures about 25 mm. Scutiger rugosa Liu Scutiger rugosa Liu, 1948, Jour. West China Border Res. Soc., 14, (B): 37, figs. 1-4— Yenwot’ang, Chaochiao, Sikang; idem, op. cit., 14, (B): 78, 1948. History of species.—Scutiger rugosa was discovered in 1942 in a small moun- tain stream near Lolokou, Chaochiao, Sikang. The known distribution of this new Scutiger is still limited to the type locality. Distribution and collection data.—The type series of this species was found under a stone near the margin of a small mountain stream at Chilipa, between Yenwot’ang and Lolokou, in Chaochiao, Sikang, on May 4, 1942. The altitude of the type locality is 11,000 feet. Scutiger rugosa is still known only from the type locality. Comparison with related species.—Scutiger rugosa (fig. 28, A) agrees with S. popet in having the tibio-tarsal articulation reaching beyond the eye, but differs in having the toes distinctly webbed. The dark bars on the limbs, very distinct in popei, are very indistinct in rugosa. The longer leg of rugosa dis- tinguishes it from both schmidti and pingii. In color rugosa is much darker than pingit. The body of rugosa is dorso-ventrally flattened, whereas it is stout in pingti. The tadpole of rugosa is black with black tail crest marbled with dark brownish gold. The tadpoles of other species of Scutiger are much lighter in color and always have a very light-colored tail crest. Original description.—‘‘Body somewhat flattened, 835 mm. long from snout to vent; vomerine teeth wanting, tongue free behind and distinctly notched [fig. 28, B]. Head large, broader than long, and dorso-ventrally flattened; snout rounded; canthus rostralis obtuse; loreal region concave; nostrils nearer to the tip of the snout than to the eye; internasal space smaller than interorbital space, which is equal to the width of upper eyelid; tympanum hidden. “Arm slender and long; lower arm about half of the body length; fingers [fig. 28, C] rounded, slender and long with light-colored rounded tips; first finger shorter than the second but the fourth finger longer than the second, the third finger the longest; subarticular tubercles indistinct; an inner large rounded palmar tubercle and an outer smaller elliptical palmar tubercle. “Legs moderately long and rather slender: the tibio-tarsal articulation reaching the middle of the eye, and the heels slightly overlapping when placed at right angles to the body; tibia 47 per cent of the body length from tip of snout to vent; toes [fig. 28, D] about one-fourth webbed and slightly fringed laterally; subarticular tubercles indistinct; tips rounded; inner metatarsal tubercle oval and highly elevated, about 5.7 per cent of the body length; outer tubercle lacking. “Skin rough on dorsal sides of head, body and limbs; many small warts with spines scattered on the head and limbs; larger rounded, oval or elongated warts 152 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 provided with spines on the back and sides of the body, a strong glandular fold from the posterior corner of the eye to shoulder; anal region with small rounded light-colored glands; a large oval gland enclosed by dark color on middle posterior aspect of each femur; throat, belly and ventral sides of limbs smooth. The color is dark brown-gray above; lower region of limbs with the same color but the upper arms and legs lighter, with indistinct dark-colored bars; digital Fic. 28. Scutiger rugosa (X 2). A. Young adult. B. Oral cavity. C. Ventral view of hand. D. Ventral view of foot. tips much lighter; jaws marked by dark bars; throat, belly and ventral sides of limbs much stippled with gray.” Habitat and habits—At first glance the living specimen of rugosa seemed very much like Scutiger schmidti in behavior and in appearance. Sexual behavior and secondary sex characters are still unknown. Tadpoles.—Tadpoles (pl. 10, fig. 3) were collected from the same locality and on the same date. Tadpoles of various stages, including a specimen nearly completely metamorphosed, were obtained from the same place on June 14, 1942. They were found under the stones of a small mountain stream. They are very active and because of their strongly built tails they can swim against the current. When disturbed they rush into the cracks between or under the stones in the water or hide themselves among vegetation. Under natural conditions they are black in color except for the dorso- posterior part of the body and the dorsal side of the base of the tail, where the color is dark brownish golden and forms a Y-shaped mark. This brownish golden color runs posteriorly along the dorsal ridge of the tail crest about half of the tail length, and then becomes discontinuous. The black tail crest is marbled with dark brownish gold. The belly is colorless and transparent, with four white spots, two postero-lateral to the heart and two near the anal region. The pupil is rounded and black, enclosed by a silver-golden iris stippled with black. LIU: AMPHIBIANS OF WESTERN CHINA 153 The tadpole of this species is rather large. Six tadpoles, with hind legs averaging 4 mm. and varying from 3 to 5 mm., have an average body length of 24.1 mm., ranging from 24 to 25 mm. The width of the head is 49 per cent of the body length while its height is only 34 per cent. The body is somewhat Fic. 29. Scutiger rugosa; mouth of tadpole (x 16). flattened, its height 49 per cent and its width 58 per cent of the body length. The length of the tail is 173 per cent of the body length, while its height is 53 per cent. The snout is rounded, and the nostril, enclosed by a light-colored ring, is slightly nearer to the tip of the snout than to the eye. The eyes are dorso- laterally located but are more dorsal than lateral. The space between the eyes is 30 per cent of the body length. The spiraculum is sinistral, ending in a semi- tube, directed upward and backward, visible from above and below. It is nearer to the base of the tail than to the tip of the snout. The vent is dextral, with an oblique opening, under a broad skin fold that overlies it and is continuous with the base of the ventral crest. The tail is very thick, with the muscular portion strongly developed. The lines formed by myocomma are very characteristic for this species, as they bend posteriorly in V-shape and bend again anteriorly at the base of the greatly thickened dorsal and ventral tail crests, especially at the anterior region of the tail. The tip of the tail is blunt, with the tail about equally developed dorsally and ventrally. The anterior region of the tail crests, especially the dorsal one, is greatly thickened. The mouth (fig. 29) is ventral, with an average width of 6 mm., one-fourth of the body length. A single row of papillae surrounds the mouth, except for a 154 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 small space in the middle of the upper lip. Papillae are larger toward the dorsal side of the upper lip, with a number scattered at the corners of the mouth, espe- cially near the upper sides of the lower lip. Some of these papillae are provided with teeth. Labial teeth and mandibles are remarkably developed and invariable. Five rows of labial teeth are found in the upper lip, the outermost the shortest and continuous and the other four rows all interrupted, the first interrupted row the longest. There are also five rows of teeth on the lower lip, the outermost row complete and much longer than the outermost row of the upper lip, and the rest all interrupted, the second interrupted row being the longest. Mandibles are enormously developed, with very strong large teeth on their margins. A young specimen with tail stump measured 23 mm. long from tip of snout to vent. Measurements of type (percentage of body length in parentheses).—Body length 35.0 mm.; head length 13.0 mm. (87.1); head width 13.5 mm. (38.5); interorbital space 3.8 (10.8); length of lower arm 18.5 mm. (52.8); diameter of lower arm 3.0 mm. (8.5); length of hand 10.5 mm. (30.0); length of hind limb 53.0 mm. (151.4); length of tibia 16.5 mm. (47.1); length of foot and tarsus 23.0 mm. (65.7); length of foot 15.5 mm. (44.2); length of inner metatarsal tubercle 2.0 mm. (5.7). Scutiger rugosa: Measurements of Six Tadpoles Ratio to Measurements Range Average __ body length (%) BOdyclengthss ak sacs sees iat ae etka cies 24-25 24.1 eat Bodyshelgnt 2337 rhc es Sunde 11.0-12.5 11.8 48.9 ‘Bod yawidtihrtwaak sone yc ries creme 14.0-14.5 14.1 58.5 Headshelgntscc ca co ski fece et sree ween 8-9 8.3 34.4 Head width ine lets Sh ee 11.5-12.0 11.9 49.3 Monuthswidthttg ce ree oe hy 6.0-6.5 6.1 25.3 Space ‘bet weeni eyes 3652202 :.. essa eee 7-8 TZ 29.8 Snoutitoispiraculum > sr. ears Seer ahs 15.5-16.0 15.6 64.7 Taillength: 34. se eo a es bees 40-45 41.8 173.4 EAI REIG NG es ie see an ee aor eee ee 12-14 12.8 5321 Diameterroftail’muséle 3252s eee oe 6 6.0 24.8 Length‘ot- hind tlimD:2oc4oe. cassis. as 3-5 4.0 ae Scutiger popei Liu Scutiger sikkimensis Liu, 1940, Jour. West China Border Res. Soc., 12, (B): 12; idem, 1948, op. cit., 14, (B): 51, pl. 1, fig. 1, pl. 2, figs. 6, 10 (not of Blyth). Scutiger popei Liu, 1947, Copeia, 1947: 125, fig. 2—Paohsing, Sikang. History of species.—Specimens collected from Mount Omei, Paohsing, and Panlungshan and described as Scutiger sikkimensis by myself in 1940 and 1943 and by Pope and Boring in 1940, must be referred to Scutiger pope (pl. 5, fig. 5) as still another endemic species of pelobatid frog from western China. Distribution and collection data.—Scutiger popei is widely distributed in western China. It is found in the mountain ranges between Szechwan and Sikang at altitudes ranging from 3,500 to 7,000 feet. During the summers of LIU: AMPHIBIANS OF WESTERN CHINA 155 1938 and 1940, tadpoles of this species at different stages were collected from pools behind large stones, under cascades or in side pools of small mountain streams under Changshouchiao, between Chiulaotung and Yuhsiensze at an altitude of 5,900 feet, on Mount Omei. On August 18, 1939, a large number of tadpoles at various stages were collected from a mountain stream near Lung- tung, Paohsinghsien, Sikang (3,400 ft.), and three adults were obtained from the same stream. From April 13 to May 8, 1941, only large tadpoles with hind limb- buds were found in small mountain streams of Shuimokou, Kwanhsien; and no i Fic. 30. Scutiger popet (xX 2). A. Oral cavity. B. Ventral view of hand. C. Ventral view of foot. eggs or young tadpoles were obtained. From June 28 to July 15, 1941, very young tadpoles and tadpoles at the time of metamorphosis were collected from the same stream of Shuimokou. On July 5, 1945, a dead matured male, a living young adult and various tadpoles were obtained from Changshoukou on Mount Omei. An egg-mass was collected in the same stream in its upper reaches but it could not be proved that the eggs were those of popet. Comparison with related species.—Scutiger pope can be distinguished from all other species of Scutiger, except rugosa, by its longer legs, as the tibio-tarsal articulation extends beyond the eye in popei and rugosa and only reaches the eye in alticola and the angle of the mouth in sikkimensis, schmidti and pingit. Scutiger popei differs from rugosa in the absence of spines on the sides of the shoulder and on the warts of the back, spines being well developed on the shoulder region and on the warts, especially those on the back of rugosa. The coloration of popei is very characteristic; there are distinct large black or dark brown spots on the back and conspicuous dark bars on the limbs, the two outer fingers, and the three outer toes. Scutiger popei has rounded warts without pores, and the tibio-tarsal articula- tion reaches beyond the eye, whereas in S. sikkimensis the warts are porous and 156 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 the tibio-tarsal articulation only reaches the shoulder or the angle of the mouth. The light triangular spot on the forehead of sikkimensis is not found in popei. The choanae are very small in sikkimensis and very large in popei. The tongue of popei (fig. 830, A) is deeply notched behind, whereas that of sikkimensis is rounded or only slightly notched posteriorly. S. popei is found at altitudes rang- ing from 3,500 to 7,000 feet; the specimens of sikkimensis in the British Museum were collected from Bhutan, Sikkim, at 13,000 feet. Scutiger popei (fig. 30, C) can be distinguished from rugosa by the absence of webs and the distinct black bars on limbs. S. alticola differs from popei in having the toes fully webbed. The shorter leg and the indistinct bars on the limbs of pingit and schmidti distinguish them from popei. Original description.—‘‘Body moderately slender; head somewhat depressed, as long as broad; tongue broad, free behind and deeply nicked; no vomerine teeth; choanae very large; snout rounded, slightly projecting beyond the mouth, and much longer than the length of the eye; canthus rostralis obtuse; loreal region very oblique, concave; nostrils nearer to the tip of the snout than to the anterior corner of the eye; interorbital space greater than internasal, and broader than the upper eyelid; tympanum hidden. “‘Arm strong, fingers slender and long, first and second about equal in length, but shorter and thicker than the third and fourth, the third finger the longest; nuptial spines number eleven on the first and eight on the second finger of the left hand, and respectively ten and five on the right hand; tips of fingers rounded and lighter in color, subarticular tubercles present; thickened skin between the subarticular tubercles, especially the two outer fingers, palmar tubercles large and prominent, a large inner one at the base of the first finger and a much smaller one opposite the base of the third. | “Hind limb slender and long, the tibio-tarsal articulation reaching beyond the eye; length of tibia 48.8 per cent of the body length; foot 49.2 per cent of the body length; toe slightly fringed with rudimentary webs, subarticular tubercles better developed than on the fingers, thickened ridges found between the sub- articular tubercles; inner metatarsal tubercle oval, with a free edge; no outer metatarsal tubercle. “Skin rough; small warts on the top and sides of the head, especially posterior to the angle of the mouth, and on the dorsal sides of the limbs; larger round warts more or less regularly scattered on the back and the sides of the body; fine warts on the inner anterior side of the arm, with a definite warty area on the antero-ventral region at the base of the arm; two well-developed chest glands, covered with fine spines, located on the thorax; the length of this gland is about 8 mm. and the breadth 4 mm.; rounded axillary glands make contact with the postero-lateral corners of the chest glands at the arm pit on each side; a large flattened gland, lighter at center and dark-colored at periphery, on the mid- posterior aspect of each thigh; skin smooth on the throat, the belly and the ventral sides of the legs. LIU: AMPHIBIANS OF WESTERN CHINA 157 “Coloration in life—Ground color of the top and sides of the head, the back and sides of the body and the dorsal sides of the limbs is brownish yellow; rounded black or dark brown spots with light centers and brown processes radiating out from each spot are correlated with the dorsal warts. The markings extend from spot to spot and form a network on the body; smaller black spots on the top of the head; black bars on the jaw, dorsal sides of the limbs, the two outer fingers, and the three outer toes; throat, belly, and ventral sides of limbs flesh-purple in color; throat, sides of belly and ventral sides of limb strongly marbled with brownish gray. The eye is very large and has a vertical pupil which can be changed from oval to slit-like form as in the cat, the size of which is regulated by the intensity of the light. The color and the shade of the iris is striking. The upper half of the eyeball is bright and the lower half is dark. After preservation the ground color of the back is light cinnamon drab, the black spots cover the warts, and the light brown network disappears.” Variation.—The specimens collected from Lungtung, and those collected from Panlungshan differ slightly in coloration and texture of the skin. Warts, especially those on the top of the head, are smaller and much more numerous in the specimens from Panlungshan than in those collected from Lungtung, so the skin of the former seems much more rugose than that of the latter. Dark brown or black stripes on the femur are different in number and in shape. The Panlungshan specimen has four or five complete narrow stripes alternating with incomplete ones on the thigh, while in the specimens from Lungtung there are only four wide stripes, with no incomplete ones. The coloration is apparently the same, except that the Lungtung specimens are lighter in color with a rather light brown network, formed by bands radiating out from the dark brown center that covers each wart. The background color is brownish yellow. There is no distinct network-like pattern on the specimens collected from Panlungshan, but the light-colored tips on the warts are much clearer than in Lungtung individuals. Scutiger popei: Measurements of Type and Paratypes Paohsing (Muping) Panlungshan, Kwanhsien Male (type) Female Female Female Ratio Ratio Ratio Ratio to body to body to body to body length length length length Measurements MM. (%) MM. %) MM. (%) MM. (%) Body lengthi3.33245 52%, BSC0NE eee, Dae. Ret BOLERO) een MOCO crete Head length? <2. 25-2. 230% “36.52 20S7-- 37.9: © 23.2) $3628 21.0% 236.2 Head:width + oe 230) HOG bee scl Ol, 3984s 6 Okt SS) marae Ree Interorbital space......... 6:6° — 10:5 6:4. 117 6.4 10.0 6.5) 11:2 Length of lower arm....... 30:0) 74726" 229.5) = 540° © 18253" 75035) 2724) 347-2 Width of lower arm....... 820) 1276. 5.2 O2bax 85D e2 Sati 525 9.4 Length of hand........... 162526: 1 Sib sOne2o6. 16 OL weet els we eual: Length of hind limb....... 99.0) 157-1 -91.0) 166.7 104.0 162°5.- 91-0: 151.7 Length: of tibiae: 232.0532 3028 AS-S. “2723-500 32205 -5020** 292.0" 5020 Length of foot and tarsus.. 46.0 73.0 40.0 738.2 47.4 74.1 41.0 70.6 ength ol foots see en: 3120) 4922 :28-5) 262.2) 23120) 24824)- 227.0" 4655 Length of inner metatarsal tubercle:o.2 es Se 4.0 6.3 2.6 4.8 3.0 4.8 3.0 5.2 158 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 Secondary sex characters.—The adult of Scutiger popei is a medium-sized frog with no sexual dimorphism in size, the body length of the male being 63 mm., while the body length of the females ranges from 54 to 64mm. The arm of the male is moderately enlarged and is much thicker than that of the female. In the male, from five to eight spines are developed on the dorsal side of the second finger and from ten to eleven on the inner dorsal side of the first finger; there are two well-defined patches of weakly developed nuptial spines on the breast. The length of the hind limb is apparently the same in both sexes. Webs are absent in the female and rudimentary in the male, in which the toes are flattened and laterally fringed. There are no vocal sacs and no lineae masculinae. Habitat and habits.—Scutiger popei is a species found on high mountains. In four years’ collecting only five adults were obtained, though from the num- ber of tadpoles found in so many places on Mount Omei, on Panlungshan, Chungkingchow, Szechwan, and at Lungtung in Paohsing, Sikang, it seems that adults should be abundant. All the specimens I secured were found in small mountain streams, either sitting on stones near the water or concealed under stones in the water. As webs are wanting, this frog seems not to be an aquatic form, especially in mountain streams. Occasionally one or two individuals may pay a visit to the water nearby, so by chance we got these few specimens after much hard work. I suspect that it is a terrestrial form living among vegetation, under stones or in holes on hillsides. The adults may come out in numbers during the breeding season, as does Stawrois chunganensis. We searched diligently among the vegetation and turned over thousands of stones along small and large mountain streams where tadpoles of Scutiger pope: were abundant, but not one specimen was found in this way. Therefore, the real home of this species is still unknown. It may be a good traveller, coming from a habitat distant from the streams where the eggs are laid. Tadpoles.—The presence of young tadpoles at the end of June indicates that eggs may be laid from the middle part of May to the early part of July. The breeding habits of this new frog are still unknown. The tadpoles are bottom feeders with strong mouth-parts (fig. 31). When disturbed, they get into cracks between stones to hide themselves, and also under stones, so it is hard to collect them in stony places. They can swim in the current of the stream as their tails are strongly built, especially the muscular portion. They are omnivorous, but rather predacious than herbivorous. They eat larvae of stone flies, caddis flies and such other animals as they can get in small mountain streams, and they also take plant food, especially algae growing on stones in the water. Tadpoles brought to Chengtu ate a lot of tadpoles of Bufo b. gargarizans every day. Tad- poles of Scutiger popei are strongly built, especially the mouth parts and the tail. If they touch a Bufo tadpole, they suck it and carry it from place to place with their strong mandibles and broad lips. Even though it struggles to escape it can rarely get away from the strong hold of the Scutiger tadpole. The coloration of the tadpoles varies greatly in different stages. Young tadpoles are black on the back and sides of the body. The pupil is black and LIU: AMPHIBIANS OF WESTERN CHINA 159 round, enclosed by a thin, whitish golden ring, which in turn is enveloped by a wide iris; this is black below and marked with golden dots on the antero-dorsal side. The muscular portion of the tail is light gray and the tail crests are very light yellowish gray. Above the muscular portion, but below the dorsal crest, there are two lines, the upper light yellowish brown, and the lower dark gray or black. These two lines start from the base of the tail and run beyond the Fic. 31. Scutiger poper; tadpole. A. Lateral view (xX 214). B. Mouth (x 10). middle portion of the tail, where they become indistinct. The belly is light gray in color. When the limb-buds are well developed the body becomes umber above, and on the sides this color darkens to bluish black lower down and then fades to dark gray on the belly. The hind limb-buds are grayish blue above and yellowish gray below. The tail is yellowish gray, the ventral tail fin lighter. The yellowish brown and dark gray lines between the dorsal tail fin and the muscular part of the tail have disappeared at this stage but their area is still darker in color than the other parts of the body. When the fore limbs come out, the body is greenish gray above with many round or oval warts enclosed by black rings. The dorsal sides of the limbs have the same coloration as the body, but with a number of discontinuous black stripes. The tadpole of this species is very large; individuals with hind limb-buds reach 75 mm. in length. In twenty tadpoles measured, in which the hind legs 160 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 averaged 5.2 mm., the average body length from snout to base of tail is 25 mm., ranging from 22.5 to 28.7mm. The width of the head is 54 per cent of the body length, while its height is only 36. The body is nearly cylindrical, its height 51 per cent and its width 56 per cent of its length. The length of the tail is about twice that of the body. Scutiger popei: Measurements of Twenty Tadpoles Ratio to Measurements Range Average _ body length (%) BOdY IGNgun ek rec nan, coe si ee 22 .5-28.7 25.2 orate Body heigntie eee tod eran ee ak ee 11.4-14.5 12.8 51.0 Body: width =. 22sec eka te aed ee 10.8-16.0 14.2 56.4 ead: heights ss osee ete ee ce es 8.4-10.2 9.1 36.2 Head: widths = 404.03 00s ee ee 12.0-15.5 13.6 54.2 Mouth’ widthei2c5-0 eee Cn ae 6.4-8.2 tek 28.1 Spaceibet ween \6vesnnsc.n caren ate 7.6-10.7 9.1 36.2 Snout; to spiracullimy.s.456 32 43s ss cose 14.3-18.0 15.8 62.8 aillengthin. 03 cas eee Ce aa eae mater 41.4-56.4 49.3 196.0 Pailtheight tence erect en tan eke ens 12.5-18.4 15.1 60.1 Diameterof tail'muscle.2. 3s os 5.5-8.5 6.7 26.6 Pengthsotshindsimbrerse see a eee ee 4.0-6.7 5.2 eae The snout is rounded; the nostril, enclosed by a light-colored ring, is nearer to the snout than to the eye. The eyes are latero-dorsal in position, more dorsal than lateral, with an average space between the eyes of 9 mm., which is 36 per cent of the body length from the snout to the base of the tail. The spiraculum is sinistral, ending in a semi-tube, directed upward and backward, visible from above and below. It is 16 mm. from snout to spiraculum, which is nearer to the base of the tail than to the tip of the snout, the distance from snout to spiraculum being 63 per cent of the body length. The vent is dextral, with a large oblique opening. The broad skin fold that overlies it is continuous with the ventral tail crest. The tail is thick, with the muscular part strongly developed. The tip of the tail is blunt, with the tail fin better developed dorsally. The dorsal fin is thickened at its base but is not continued on to the body. The mouth is ventral, with an average width of 7 mm., which is 28 per cent of the body length. There is a single row of papillae of varying size all around the mouth, except for a small free space on the mid-dorsal part of the upper lip. Additional papillae, with or without teeth, are found at the corners of the mouth, especially at the latero-dorsal corners of the lower lip. There are seven rows of teeth on the upper lip. The outermost row is short and continuous, partly filling up the gap where there are no papillae. The remaining six rows are all interrupted, and the inner row is the shortest. On the lower lip there are seven rows, with the outermost row continuous and longer than the outer row on the upper lip. The other six rows on the lower lip are all interrupted, the innermost row being the shortest (fig. 31). Teeth are also found on some of the additional papillae at the corner of the mouth. The mandibles are thick and heavy, with sharp strong teeth both above and below. The number of rows of labial teeth varies greatly in the tadpoles of this species. Seventeen tadpoles collected from LIU: AMPHIBIANS OF WESTERN CHINA 161 Lungtung, Paohsing, in Sikang, and twenty-two from Mount Omei were studied. For the Paohsing group, the tooth formulaeare as follows: ten have I:5-5/1:5-5; five have I:6-6/1:5-5; one has I:5-5/I:6—6; and one has I:6-6/1I:6-6. In the Mount Omei group, one has [:4-4/I:4—4; two have I:4-5/1:4-5; four have I:5-5/1:5-5; ten have I:6-6/I:6-6; one has I:6-7/1:6—7; one has I:6-6/1:7-7; one has I:7-7/1:6-6; and two have I:7-7/I:7-7. Ten tadpoles collected from Panlungshan were studied and the tooth formulae are as follows: nine have I:5-5/1:5-5 and one is abnormal. The majority of the Paohsing popei tadpoles have a tooth formula [:5-5/1:5-5; on Mount Omei the common type is I:6-6/ I:6-6. The process of metamorphosis is rather slow, but the species characters are clearly shown, even at the time when the fore limbs come out. At this stage small round warts are much more conspicuous and the color on and near the warts becomes much darker than before; dark stripes on limbs are clearly shown. When the absorption of the tail fin begins, warts with creamy centers and black margins are developed, and a row of small creamy warts appears on each dark stripe on the limbs. These stripes are darker and wider on the anterior and posterior sides of the femur and tibia; on the dorsal sides they are weakly developed. Between these strong ones, there are very weak and incomplete stripes. As soon as the tail is half gone, the ground color on the back becomes olive brown. Warts and color patterns become more distinct. At the ear region and the posterior angle of the jaw the skin is smooth and has a purplish coloration. The young just after metamorphosis has nearly all of the characters of an adult. The web between the toes of all the young individuals of Scutiger popei from Mount Omei, Panlungshan, and Lungtung, is not present in the adult. The coloration of the young is rather uniform. The top of the head is lighter and there is a creamy color at the angle of the jaw and the anterior base of the arm. The sides of the body are also lighter and are washed with scarlet. Many warts, with brownish ashy color enclosed by fuscous or black, are scattered on the back. The limbs are brownish ashy above but with more olive color crossed by black stripes. Margins of the jaw are cream-colored, marbled with black. The throat and the belly are gray. A round, flattened cream-colored gland, enclosed by fuscous, is located on the middle posterior region of the thigh; this is shown in the tadpole even before the fore limbs come out. The size of the young varies greatly; young adults from Mount Omei range from 33 to 33.5 mm. in length, those from Panlungshan from 29 to 32 mm., and those from Lungtung from 28 to 29 mm. Our knowledge of the life history of this rare frog is still fragmentary, the breeding habits, eggs, and process of hatching being unknown. Very young tadpoles and tadpoles at different stages of metamorphosis were found on Mount Omei during the first part of August, 1938, and again during the summer of 1940. In the spring of 1941, from April 13 to May 3, I went to Panlungshan with the hope of finding Scutiger popei breeding and laying eggs, but only advanced tadpoles were found. The local people told us that they saw tadpoles during 162 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 winter. Again from June 30 to July 15 of the same year I made another trip to the same place and in the same stream I found very young tadpoles and ad- vanced tadpoles with two legs fully developed, four-legged tadpoles, and young near the completion of metamorphosis. The tadpoles in the process of meta- morphosis are evidently those found in the latter part of April, but the younger ones must be hatched from eggs laid from early May to the middle part of June. The breeding season of this frog is most likely from May to June. The fully developed tadpoles might be those of the previous year, as indicated by the study in April, by the slow process of growth and metamorphosis, and by the information from the farmer who reported them from the same stream during winter. Further investigation, during the winter, is essential to the solution of this problem. Scutiger schmidti Liu Scutiger schmidti Liu, 1947, Copeia, 1947: 123, fig. 1—Mount Omei, Szechwan. History of species—A small Scutiger tadpole was obtained from Maliuwan, Chungkingchow, Szechwan, during the summer of 1941, and remarked upon as an unknown species of Scutiger in 1948. During the summer of 1945, adults of this form, with additional tadpoles, were collected from Mount Omei, and the new species was named Scutiger schmidti in 1947. Distribution and collection data.—Scutiger schmidti is an endemic form of western Szechwan, found on Mount Omei, at Maliuwan, west of Kwanhsien, and near Shuimokou, Kwanhsien. On July 10, 1941, a number of tadpoles of different stages were obtained from side pools or pools beneath cascades on the main mountain stream of Maliuwan, 5,700-7,000 feet altitude. A tadpole of this species with hind limb-buds and one with four limbs fully developed were found in a pool beneath a cascade near Heifengt’ou near Shuimokou, at an altitude of about 5,500 feet. I found tadpoles of Scutiger schmidti together with some tadpoles of S. popei on June 6, 1945, at an altitude of 6,100 feet in a pool of a small mountain stream known as Pientankou between Chiulaotung and Yuhsiensze. No adult was found. On June 28, 1945, my collector found two egg-masses in this same small mountain stream and two tadpoles passing through the process of metamorphosis. The new Scutiger tadpole was found in a famous well, known as Lungwangching or Dragon King Well, at an altitude of 7,800 feet and temperature of about 60° F. on June 7, 1945, very near to Tachengsze. The discovery of this tadpole led me to adults of the new frog, as I heard a peculiar croaking coming from a shallow depression connected with the well. This depression was densely covered by shrubs and bamboo and with only a very limited amount of water running through among small flat stones, dead leaves and roots. Guided by the croaking, I secured my first specimen by turning over a stone. My assistant and I secured fourteen adults, all males, four masses of eggs and many tadpoles in different stages. During the afternoon of the next day (June 8), three females ready for egg-laying, each one watched by three males, were obtained by turning over the bamboo roots; some other males, three LIU: AMPHIBIANS OF WESTERN CHINA 163 egg-masses and a few tadpoles were also collected. Twenty-six males, seven masses of eggs with embryos developed, and a few tadpoles were collected from the same locality on July 22 of the same year. Comparison with related species.—Scutiger schmidti (pl. 5, figs. 1 and 2; text fig. 32) can easily be distinguished from all other species of Scutiger in China by its coloration. It differs from popei especially in the brown marking on the back, Fig. 32. Scutiger schmidti (x 2). A. Ventral view of head and thoracic region of male. B. Oral cavity. C. Ventral view of hand of male. D. Ventral view of foot. the unmarbled belly and the indistinct irregular brown bars on the limbs. S. ping differs from schmidti in having a rufous brown back without any large markings or pattern, and with only a few scattered warts, and in having the toes distinctly webbed. SS. schmidti differs from alticola and rugosa by the absence of webs between the toes, alticola having the toes fully webbed and rugosa having them about one-fourth webbed. The tadpoles of schmidti are the smallest of the various species of Scutiger, with a characteristic coloration, especially the dark color near the base of the body and the tail. Original description.—‘‘Tongue elliptical, free behind and distinctly notched; head moderately large, longer than broad; snout rounded and slightly projecting beyond the mouth; canthus rostralis feebly indicated; loreal region oblique and flat; nostril about mid-way between the tip of the snout and the anterior border of the eye; internasal space about equal to the interorbital space, which is broader than the upper eyelid; length of the eye greater than the orbital space; tympanum hidden; spines on the margins of the upper and lower jaws and a more or less circular spiny area on the median ventral side of the lower jaw; black spines strongly developed on the tympanic region. “Arm rather long; fingers slender and long, first finger equal in length to the second, both shorter than the fourth finger and the third finger about twice as long as the first two; nuptial spines developed on the inner dorsal sides of 164 FIELDIANA: ZOOLOGY MEMOIRS, VOLUME 2 the first and the second fingers; subarticular tubercles not evident; two large prominent palmar tubercles, the outer one smaller than the inner; warts with spines on the outer side of the arm and smooth warts on the inner sides; in front of the base of the arm a pre-humeral warty area. “Hind limb moderately long and relatively weak, tibio-tarsal articulation reaching the angle of the mouth; heels just meet when placed at right angles to the body; tibia 43.1 per cent of the body length, webs barely indicated and toes fringed; spines on the outer fringes of the fifth and fourth toes; no subarticular tubercles; inner metatarsal tubercle elongated, rod-like and flat, no outer meta- tarsal tubercle, many large and small spines scattered on the hind limbs especially on its dorsal side, each femur with a light-colored flattened gland on its postero- middle region. “Skin very rough with many small warts on the back of the body and the sides of the head and the body and all over the limbs; warts provided with many small spines especially on the back of the body, very much like those of Bufo bufo gargarizans; larger spines found along the margin of the jaws, tympanic region and the postero-ventral aspect of the thigh; some fine and a few large spines on the belly. “The dorsum is grayish brown in ground color, with dark brown or nearly black marks on the jaws and the antero-dorsal side of the snout; a similar colored mark starting from the anterior region of the interorbital space, including the eyelids and extending posteriorly covering the whole back to the vent; the posterior part of the back nearly black, from the presence of many small and a few large black spines; the exposed parts of the limbs also grayish brown with irregular dark brown bars; sides of the body lighter and with scattered still lighter colored warts; throat, belly and the ventral sides of the limbs purplish fleshy in color and semitransparent, as in Kaloula; digital tips very light in color; pupil black, vertically oval, sometimes even squarish, with black connections dorsally and ventrally; iris golden, stippled with black, the ventral part slightly darker than the dorsal portion.” Coloration.—The male is always much darker than the female and in the latter the grayish brown mark on the back is very conspicuous on a lighter ground color. The coloration of the males varies only slightly. The much lighter brownish ashy color in females has indications of green, and there are distinct grayish brown marks on the snout, head and body. The female has very few warts and no spines on any part of the body. The coloration of the limbs is similar to that of the body. The throat, belly and ventral sides of the limbs are still lighter than in the male and are spotted with pearly gray. A pair of flattened glands is present on the sides of the chest near the bases of the arms. The vent of the female is heavily spotted with creamy buff to pearly gray. After preservation in formalin, the color of the male specimens is darker and the mark of the back becomes more indistinct; in females, the much lighter color preserves the distinctness of the dorsal pattern. LIU: AMPHIBIANS OF WESTERN CHINA 165 Secondary sex characters.—In thirty males and three females measured, sexual dimorphism in size is evident, the average body length of the males being 438 mm., ranging from 40 to 47 mm., while for the females it is 51 mm., with a range from 48 to 54mm. Other secondary sex characters in males (fig. 32, A and C) consist of two elongated patches of closely packed fine black spines on the chest, fine black spines on the margin of the jaws and tympanic region, and a more or less circular spiny area on the median ventral side of the lower jaw; black nuptial spines are developed on the inner dorsal sides of the first and second fingers of the male. The arm of the male is stronger than that of the female, the diameter of the lower arm of the male being 10.5 per cent of the body length in the male and only 6.8 per cent in the female. The leg of the female is shorter than that of the male, 183 per cent of the body length as compared with 148 per cent in the male. Scutiger schmidti: Measurements of Thirty Male and Three Female Adults Ratio to Measurements Sex Range Average __ body length (%) Bodyslength® 02. ascsccicnck eee rot 40-47 43.1 pees Q 48-54 51.0 Se Héadvliengthigy-