U. S. NATIONAL MUSEUM BULLETIN 160 PLATE 1
BUFO ALVARIUS GiRARDIllustrating parotoifi, ulnar, and tibial glands, skin texture, and other identification points. Upperphotograph by Dr. Walter P. Taylor, Bureau of Biological Survey.
CJ\ re uuItvILlvi qSMITHSONIAN INSTITUTION ^f VUNITED STATES NATIONAL MUSEUM \^, \\\^ . \I\\iBulletin 160
MEXICAN TAILLESS AMPHIBIANSIN THE UNITED STATESNATIONAL MUSEUM
BYREMINGTON KELLOGGAssistant Curator, Division of MammalsUnited States National Museum
UNITED STATESGOVERNMENT PRINTING OFFICEWASHINGTON : 1932
For sale by the Superintendent of Documents, Washington, D. C.
AD^'ERTISEMEXTThe scientific publications of the National Museum include twoseries, known, respectively, as Proceedings and Bulletin.The Proceedings series, begun in 1S7S. is intended primarily as amedium for the publication of original papers, based on the collectionsof the National Museum, that set forth newly acquired facts in biol-ogy, anthropology, and geology, with descriptions of new forms andrevisions of limited groups. Copies of each paper, in pamphlet form,are distributed as published to hbraries and scientific organizationsand to specialists and otibers interested in the different subjects.The dates at which these separate papers are published are recordedin tie table of contents of each of the volumes.The series of Bulletins, the first of which was issued in 1875, con-tains separate pubUcations comprising monographs of large zoologi-cal groups and other general systematic treatises (occasionally inseveral volumes', fajimai works, reports of expeditions, catalogues oftype specimens ^nd special collections, and other material of similarnature. The majority of the volumes are octavo in size, but a quartosize has been adopted in a few instances in which large plates wereregarded as indispensable. In the Bulletin series appear volumesunder the heading Contributions jron\ the United States NationalHerbarium, in octavo form, published by the National Maseum since1902, which contain papers relating to the botanical collections ofthe Museum.The present work forms No. 160 of the Bulletin series.Alexaxdee Wetmore,Assistant Secretary, Smithsonian Institution.Washtn-gton, D. C, March 31, 1932.
CONTEXTS
Pa?eHistorical introduction 1Collecting work of Nelson and Goldman 1Other sources of National Museum material 3Material in other museums 9Acknowledgments 15Abbreviations 15Class Amphibia Linnaeus: Order Salientia Laurenti 16Key to families of Mexican Salientia 17Family Pelobatidae Lataste 18Genus Scaphiopus Holbrook 18Key to Mexican species of Scaphiopus 19Family Rhinophrynidae Gunther 24Genus Rhinophrynus Dumeril and Bibron 24Family Bufonidae Hogg 27Genus Bnfo Laurenti 27Key to Mexican species of Bufo 31Family LeptodactyUdae Berg 74Key to genera of Mexican Leptodactylidae 76Genus Engystomops Espada 77Genus Leptodaclylus Fitzinger 81Key to Mexican species of LeptodactyJus 82Genus Eleutherodaciyhis Dumeril and Bibron 90Key to Mexican species of Eleuiherodadylus 92Genus Cauphias Brocchi USGenus Tomodactylus Giinther 120Genus Syrrhophus Coj>e 123Key to Mexican species of Syrrhophus 124Family Hylidae Gunther 129Key to genera of Mexican Hylidae 131Genus Gastrotheca Fitzinger 133Genus Pternohyla Boulenger 135Genus Diaglena Cop>e 137Genus Triprion Cof>e ' 138Genus Phyllomedusa Wagler 139Key to Mexican species of PhyUomedusa 142Genus Hyla Laurenti 147Key to Mexican species of Hyla 150Genus Hyldla Reinhardt and Lutken 179Key to Mexican species of HyleUa 180Family Bre%icipitidae Cope 182Key to Mexican Brevicipitidae 182Genus Hypopachus Keferstein 183Genus Gastrophryne Fitzinger 1S6Family Ranidae Bonaparte 189Genus Rana Linnaeus 189Key to Mexican species of Rana 190Index -17m
ILLUSTRATIONS
Figures PageFiQUKE 1. Diagrams illustrating structural conditions found in the sacralvertebra and coccyx of Mexican tailless amphibians 182. Terminal phalanges of the toe in the foot of Hyla 183. Diagrammatic ventral view of shoulder girdle of Bufo ameri-canus .- 274. Lateral and dorsal views of Bufo alvarius from GuadalupeCanyon, Sonora, Mexico ._ 365. Lateral and dorsal views of type specimen of Bufo canaliferus,from Tehuantepec, Oaxaca, Mexico 406. Lateral and dorsal views of type specimen of Bufo coccifer, fromArriba, Costa Rica 417. Lateral and dorsal views of Bufo cognatus from Nebraska 428. Lateral and dorsal views of cotype specimen of Bufo compadilisfrom Pesquieria Grande, Nuevo Leon, Mexico 459. Lateral and dorsal views of Bufo cristatus from Jalapa, VeraCruz, Mexico 4910. Lateral and dorsal views of Bufo debilis from Delaware Creek,Culberson County, Tex .. 5011. Lateral and dorsal views of Bw/o mannws from Gatun, Panama. 5412. Lateral and dorsal views of Bufo marmoreus from mountains 12miles north of Santo Domingo, Oaxaca, Mexico 5813. Lateral and dorsal views of Bufo punctatus from La Paz, LowerCalifornia 6114. Lateral and dorsal views of Bufo simus from Valley of Mexicoand from Dolores, Nayarit, Mexico 6415. Lateral and dorsal views of Bufo valliceps from Victoria, Tex.- 6916. Lateral and dorsal views of type specimen of Bufo woodhousii,from San Francisco Mountain, Ariz 7317. The T-shaped terminal phalanx of Me?^//ieroc/adt/iws 7618. Lateral and dorsal views of Engystomops pustulosus from SantoDomingo, Oaxaca, 'Mexico 7919. Fore feet of Central American liylids, illustrating the extent ofthe web between the fingers 14120. Hind feet of Mexican hylids, illustrating the extent of the webbetween the toes 14121. Diagrammatic ventral view of shoulder girdle of Rana cates-beiana 19522. Diagrammatic dorsal view of skull of Rana catesbeiana 19623. Diagrammatic ventral view of skull of Rana catesbeiana 19624. Opened mouth of Rana pipiens, showing position of vomerineteeth 204Pl/ATESPlate 1. Bufo alvarius Girard FrontispieceI?
MEXICAN TAILLESS AMPHIBIANS IN THE UNITED STATESNATIONAL MUSEUM
By Remington KelloggAssistant Curator, Division of Mammals, United States National Museum
HISTORICAL INTRODUCTIONIn 1926 the writer was instructed by Dr. E. W. Nelson, at thattime chief of the Bureau of Biological Survey of the United StatesDepartment of Agriculture, to examine and prepare a report on theherpetological collections made by Doctor Nelson and Edward A.Goldman in Mexico. This work has been completed since the writerbecame a member of the staff of the United States National Museum.In the course of this study the scope of the report was enlarged toinclude all the Mexican material that was available in the NationalMuseum and other American institutions. Inasmuch as it was orig-inally intended to deal with the Mexican collection of Nelson andGoldman, a few facts in regard to the inauguration of their field workand the extent of their explorations may be of interest.COLLECTING WORK OF NELSON AND GOLDMANUpon completion of field work by the Death Valley expedition andthe consequent disbanding of the party at Visalia, Tulare County,Calif., the various members scattered to take up other assignments.Doctor Nelson traveled southward through the San Joaquin Valley tothe Goldman ranch at Alila (nowEarlimart), Calif., where he mentionedto E. A. Goldman's father his intention of obtaining an assistant.The father spoke of his son's interest in natural history and remarkedthat he was then in Fresno. As a result of this conversation, thesenior Goldman went to Fresno and brought Edward back with himto the ranch for Nelson's appraisal. On October 10, 1891, Goldmanleft his father's ranch as Nelson's personal assistant, the understandingbeing that he was to be given $30 a month and his board. This waspaid by Nelson out of his own salary of $150 a month. They traveledin a buckboard outfit, which was the object of no little curiositybecause of two 42-pound bear traps that hung one from each side.For some reason Nelson had chosen the name "Scalops" for onemember of their team, while the other went by the name of "Johnny."1
2 BULLETIN 160, UNITED STATES NATIONAL MUSEUMNelson and Goldman drove through the southern end of the SanJoaquin Valley to a site 8 miles west of Tejon Pass and thence tothe head of San Emigdio Canyon. Leaving this camping site theycrossed the southern end of the valley, collecting at various localitieson their northward trip to San Luis Obispo, Calif. From this placethey worked northward to San Simeon and the mountains near by,and in a few days drove southward along the coast, collecting seabirds particularly, to Santa Paula, where, about January 4, 1892,they abandoned the buckboard and field outfit.Orders had been received from Dr. C. Hart Merriam, chief of theBureau of Biological Survey, for Nelson to start on what it wasanticipated would be a three months' trip to western Mexico. Nelsonand Goldman traveled by rail from Santa Paula to San Franciscoand sailed on the Pacific mail steamer Acapulco for Manzanillo,Colima, where they landed on January 24, 1892. From the time ofsailing until March 1, 1892, Goldman, who had previously been apersonal assistant of Nelson, worked for the experience without pay,and then he received his first official appointment from the BiologicalSurvey, which authorized his employment at $75 a month, out ofwhich he was to pay his board and traveling expenses. Instead oflasting three months, the field trip was extended, until Goldman hadpassed four years in Mexico, after which he received permission to visithis home.In September, 1893, Nelson left Chihuahua City for Washington,D. C, but returned to the City of Mexico in December, 1893, wherehe was met by Goldman, and they continued the field work together.Again on August 15, 1896, Nelson left Carneros, Coahuila, for Wash-ington. The same year, Goldman received permission to visit hishome in California, and he accordingly left Santa Rosa, Guanajuato,on November 18, 1896, for Alila, Calif., but resumed field work atRio Verde, San Luis Potosi, on January 7, 1897. Nelson rejoinedGoldman at Ameca, Jalisco, on February 6, 1897. Field work forthat season was completed on September 17, 1897, at Bolaiios,Jalisco, where Goldman left to revisit his home in California andNelson to return to Washington. Goldman, on his return to Mexico,went to the Valparaiso Mountains in Zacatecas, where he begancollecting on November 26, 1897. Doctor Nelson rejoined him atAltamira, Tamaulipas, on April 24, 1898.About September 20, 1898, Nelson left Goldman at Parral in Chi-huahua and returned to Washington, while the latter proceededacross the Sierra Madre. Again, on June 3, 1899, Nelson met Gold-man at Casas Grandes, Chihuahua, and the two continued to collectuntil October 7, 1899, when they discontinued field work for the seasonat Sierra en Media, Chihuahua. Goldman again left for Californiaand while there did some collecting. About December 30, 1900, hearrived in Arizona and began field work near Winslow, the main
MEXICAN TAILLESS AMPHIBIANS 6
object of the trip being to obtain a series of wild turkeys. About themiddle of January, 1900, he went to Washington for the first tune andremained there for about six weeks before returning to Mexico. Heresumed field work at Frontera, Tabasco, on March 3, 1900, andafter a trip into the interior he returned there to meet Nelson on April20, 1900. This season's field work ended at Carrizal, Vera Cruz,about May 15, 1901, and then both Nelson and Goldman returnedto Washington. Early in the winter of that year Goldman receivedinstructions to return to Mexico, and accordingly he resumed fieldwork on December 19, 1901, at Nuevo Laredo, Tamaulipas. Thisrail journey was somewhat unusual in that it marked a departurefrom the previous practice of the Biological Survey in regard to thepayment of traveling expenses, for Goldman's expenses were paidto the Mexican boundary from Washington. Nelson met Goldmanabout February 6, 1902, at Matamoros, Tamaulipas, and after aboutfour months in the field both of them left Eagle Pass, Tex., on June8, 1902, for Washington.After spending a brief period at headquarters, Goldman was senton a field trip to California, where he began collecting on July 17,1902, at Paraiso Springs in Monterey County. The last entry in thefield catalogue was on September 10, 1902. During part of this tripDr. C. Hart Merriam was with liim. Returning to Washington,Goldman remained a few weeks before being ordered again to Zaca-tecas in Zacatecas, where he resumed field work on December 19, 1902.Nelson joined Goldman on December 23, 1902, at Ocotlan in Jalisco.On June 28, 1903, they terminated their collecting at La Barca,Jalisco, and returned to Washington. This was Nelson's last tripon the mainland of Mexico.The following year Goldman visited Mexico City on his way toJaltipan in Vera Cruz, where field work was begun on February 3,1904. This season's field work ended in Jidy, 1904, at Lerma in theState of Mexico, and Goldman returned to Washington. In thecourse of these 12 years, since the beginning of their partnership,Nelson and Goldman had visited every State and Territory in themainland of Mexico, and in some instances had crossed and recrossedthem several times.OTHER SOURCES OF NATIONAL MUSEUM MATERIALThe United States National Museum has received Mexican speci-mens of tailless amphibians from a number of other sources. Mostof the early acquisitions came from individuals wdth whom Prof.Spencer F. Baird had corresponded. Numerous letters on file in thearchives of the Smithsonian Institution reveal Baird's active interestin building up a representative collection of Mexican animals. Hewas instrumental also in effecting the appointment of qualified
4 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
naturalists on the various expeditions and surveys of the United StatesGovernment, and made an effort to get in contact either personally orby letter with everyone who showed any interest in natural history.From Dr. Thomas H. Webb, secretary and surgeon of CommissionerJohn Russell Bartlett's party of the United States and MexicanBoundary Commission, the Smithsonian Institution received its firstcollection from Mexico. Upon the completion of the Gila Riversurvey, Bartlett's party left Tucson, Ariz., on July 17, 1852, for ElPaso, Tex. Their route traversed northern Sonora and continuedthrough the villages of Tubac, Santa Cruz, Agua Prieta, Janos, andCorralitos, the party arriving at El Paso, Tex., on August 18, 1852.Owing to the lack of an adequate military escort, Bartlett acceptedthe offer of one from the Mexican Boundary Commissioner, and thenproceeded to Ringgold Barracks, Tex., through the Mexican States ofChihuahua, Durango, Coahuila, and Nuevo Leon. The party leftEl Paso, Tex., on October 7, 1852, en route to the city of Chihuahua,and arrived there on October 22, 1852. The road taken passedthrough Guadalupe, Carrizal, Encinillas, and Sauzillos. The typespecimen of Bvjo insidior Girard, collected by Dr. Thomas H. Webb,came from Chihuahua, possibly from the city itself, but this isuncertain. Bartlett states that when Doctor Webb's wagon brokedown south of the Laguna de los Patos, the members of the partyamused themselves by collecting insects, reptiles, and other objectsof natural history while it was being repaired. Bartlett's partyremained 10 days in the city of Chihuahua, and it is to be presumedthat Doctor Webb utilized his leisure to make collections, as there isample evidence in the commissioner's narrative that the surgeonavailed himself of every opportunity to collect minerals and natural-history specimens. Leaving Chihuahua on November 1, 1852,Bartlett's party passed through Saucillo, La Cruz, Las Garcas, SantaRosalia, Jimenez, Cerro Gordo, San Pedro del Gallo, Quincam^, andLa Pena, arriving at Parras on November 27, 1852. They reachedSaltillo, the capital of Coahuila, on December 7, 1852, Santa Catar-ina on December 11, and Monterey, the capital of Nuevo Leon, onDecember 12. They left Monterey the next day by a route thatpassed through Marin, Carrizitos, and Cerralvo, and on December19, 1852, they arrived at Mier. The following day the party passedthrough Camargo and thence across the Rio Grande to RinggoldBarracks in Texas.John H. Clark, a collector employed by the United States and Mexi-can Boundary Survey under the direction of Commissioner Bartlett,accompanied Col. J. D. Graham, chief astronomer, from Fort Davis,Tex., to Copper Mines, N. Mex., where they arrived on August 2,1851. On August 28, Bartlett and his personal party set out forSanta Cruz, Sonora, to replenish their supplies. On September 18,
MEXICAN TAILLESS AMPHIBIANS OColonel Graham caught up with Bartlett at the Calabassa Haciendaon Santa Rita Mountain, Sonora. Both parties then proceeded ontheir way to Santa Cruz. Although Bartlett does not specificallystate that Clark accompanied Colonel Graham on this trip, he musthave done so, for the type specimen of Hesperomys sonoriensis Le Contewas collected at Santa Cruz in Sonora, about 6 miles south of theMexican boundary line. In that event the cotypes of Hyla afinisBaird were collected somewhere along the route from Copper Minesin New Mexico to Santa Cruz in Sonora. Baird published the descrip-tion of this tree frog in 1854, and since the Boundary Commission, ofwhich Clark was the principal assistant and surveyor, did not resurveythe Sonora boundary line until 1855, these cotypes must have beencollected in September or October, 1851. Additional details in regardto Clark's field work will be found in the remarks under Bufopundatus (p. 62).Arthur C. V. Schott was appointed a collector and assigned tothe party of Lieut. A. W. Whipple by Commissioner Bartlett. Sub-sequently Schott was transferred to the party working under thedirection of Maj. William H. Emory. During the period betweenSeptember, 1851, and October, 1854, Schott evidently was assigned towork along the Rio Grande. One of the cotypes of Bufo debilis wascollected during this period. In October, 1854, when the BoundaryCommission was again reorganized, Schott was assigned as an assist-ant to Lieut. N. Michler, and this division proceeded through NewMexico to California. After establishing the initial point of theboundary line on the Colorado River, Michler's party journeyed toTucson, Ariz., along the wagon road that followed the Gila River.Pending the arrival of Major Emory, the party remained in Tucson,and it was not until June 26, 1855, that the survey of the obliqueline from Los Nogales to the Colorado River was comm.enced. Twoof the cotypes of Bufo alvarius were collected by Schott during thesurvey of this line.In July, 1853, the Smithsonian Institution received another collec-tion of amphibians from Mexico. The records show that ProfessorBaird corresponded with Maj. William Rich at Mexico City as earlyas September, 1852, and in the following year a small miscellaneouscollection from him was received. Among these specimens were twotree frogs that became the cotypes of Flyla eximia and a small seriesof frogs that formed the basis for Rana montezumae.To Lieut. Darius Nash Couch the Museum is indebted for one ofthe largest Mexican collections received in the early days of theSmithsonian Institution. He conducted the exploring expedition thatsurvej^ed northern Mexico for a Pacific railroad and that set out onMarch 1, 1853, from Matamoros in Tamaulipas and passed throughNuevo Leon and Coahuila to at least as far as the Rio Nasas inDurango. The type specimens of Bufo speciosus Girard, Scaphiopus
6 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
couchii Baird, Scaphiojms rectifrenis Cope, and Bufo debilis Girardwere collected on this trip. In addition to the specimens collected byhimself, Lieutenant Couch purchased the entire collection of Dr. LuisBerlandier and presented it to the Smithsonian Institution. The RioNasas was reached in June, 1853, and on August 1, 1853, he was backagain in San Diego, Nuevo Leon. The exact itinerary of this expe-dition is not known to the writer, for the report although written wasnever published. A more extended account of this expedition ispublished in the remarks under Bufo debilis (p. 52).On April 7, 1854, a small collection made in Chihuahua was receivedfrom John Potts. Subsequent collections received from Potts in 1854and 1855 contained a number of additional species, including the typespecimens of Eleutherodactylus longipes (Baird) and Scaphiopus multi-plicatus Cope.An exchange with the Museum d'Histoire Naturelle in Paris sometime in 1855 resulted in the transfer of a few of the Mexican amphib-ians collected by Adolphe Boucard. In 1860, duplicates of Mexicanamphibians and reptiles were received in exchange from the Academyof Natural Sciences of Philadelphia. Included among them wereseveral amphibians collected by R. Montes d'Oca at Jalapa in VeraCruz. One of these was a tree frog with a small tympanum, whichlater became a cotype of Hyla miotympanum Cope.In 1859, Professor Baird began to correspond with Dr. CarlosSartorius, who with his son Florentin collected a number of specimensin the vicinity of Mirador and Orizaba, including the types of Hylamiotympanum Cope, H. muricolor Cope, and H. gracilipes Cope.From Charles Laszlo, of Tabasco, the Smithsonian Institution in 1860received a small collection of alcoholic specimens.From the Smithsonian annual report for 1861,' one learns that acollection of reptiles from "Guadalaxara" had been received fromJ. J. Major. This shipment presumably included the type of Gas-trophryne usta (Cope), a small narrow-mouthed toad described byCope in 1866.Following a brief expedition to the interior of Mexico, Louis JohnXantus de Vesey, a Hungarian, and one or more of his companionssettled on a homestead in Iowa, but as might have been anticipated,Xantus soon tired of this. Fortunately for all concerned he had inthe meantime become acquainted with Dr. John F, Hammond,U. S. A., who, with the assistance of Professor Baird, obtained forXantus the appointment as hospital steward in the United StatesArmy, For a part of his enlistment, Xantus w^as stationed at FortRiley, Kans., and later was transferred to Fort Tejon, Calif. Speci-mens collected at these localities were transmitted to Baird, whobecame more and more interested in his natural ability along these
> Ann. Rep. Board Reg. Smithsonian Inst, for 1861, p. 66, 1862.
MEXICAN TAILLESS AMPHIBIANS 7lines. On the expiration of his enHstment in the Army, in April,1859, Xantus was placed in charge of the tidal station of the UnitedStates Coast Survey at Cape San Lucas, Lower California, at theearnest solicitation of Professor Baird, and he remained there until1861.Through the influence of Professor Baird, Xantus was appointed in1863 as United States consul at CoUma, Mexico, and in return madelarge collections in that region, including the type specimens ofPhyllomedusa dacnicolor Cope and Bufo argillaceus Cope. Thisappointment was revoked some time previous to August 11, 1863,because of certain indiscreet understandings that Xantus had withlocal "politicos," but he remained at Manzanillo for some monthsafterwards as receiver for wrecked vessels. Part V of his Mexicocatalogue was mailed to Professor Baird on February 29, 1864.Xantus was intensely interested in all branches of natural history, asis shown by the quantity and variety of material he collected for theSmithsonian Institution.In 1863, Francis Sumichrast, of Orizaba, Vera Cruz, began sendingspecimens to the Smithsonian Institution, and additional collectionswere received from this region until 1867. Sumichrast, who wasborn in Yvorne, Switzerland, on October 15, 1828, accompanied M.de Saussure of Geneva, the grandson of the celebrated geologist, onhis voyage to Mexico in 1855-56. The Mexican revolution madetraveling difficult, and Saussure after a year's stay returned to Genevawith their collections. Sumichrast elected to remain in Mexico anddevote his attention to the natural history of that country. Duringhis residence in the State of Vera Cruz, he collected the type specimensof Hyla spilomma Cope, H. staufferi Cope, H. bistincta Cope, Syrrho-phus cystignathoides (Cope), and Eleutherodactylus rhodopis (Cope).In the Smithsonian annual report for 1868,^ there appears the fol-lowing statement: "Under the joint auspices of the SmithsonianInstitution and the Kentucky University, at Lexington, ProfessorSumichrast, a well-known naturalist, long resident in Mexico, hasgone to the Istlmius of Tehuantepec to make observations andcollections in natural history * * *." His first collecting seemsto have been done at Juchitan, Tehuantepec, in July, 1868. Whilethe full details of his movements are not known, it is certain that hewas at Iztaltepee, Oaxaca, on April 30, 1870; at Rancho de Cacoprietoin Chiapas in April, 1872; and in December, 1872, evidently he hadsettled down at Santa Efigenia, Oaxaca, where he made his homeduring the remaining years of his life. In his explorations Sumichrastvisited the States of Vera Cruz, Puebla, Mexico, Oaxaca, and Chiapasand sent collections of natural-history specimens to the SmithsonianInstitution, the Academy of Natural Sciences of Pliiladelphia, and
J Ann. Rep. Board Reg. Smithsonian Inst, for 1868, p. 25, 1869.
8 BULLETIN 160, UNITED STATES NATIONAL MUSEUMthe Museum of Comparative Zoology at Cambridge, as well as tomuseums in Switzerland, Germany, and France. A number of am-phibians were sent to the Jardin des Plantes in Paris, where they werestudied by Paul Brocchi while he was preparing his "fitude desBatraciens de I'Amerique Centrale," pubHshed in 1882 as a part ofthe contribution made by the "Mission Scientifique au Mexique etdans I'Amerique Centrale." Among the numerous specimens col-lected by Sumichrast during his residence on the Istlmius of Tehuan-tepec in the State of Oaxaca are the t3^pes of Hylella sumichrastiBrocchi, H. platycephala Cope, Cystignathus perlaevis Cope, C.lahialis Cope, Eleutherodactylus rugulosus (Cope), Syrrhophus leprusCope, and Bujo canalijerus Cope. Sumichrast died of cholera onSeptember 26, 1882, at Tonala in Chiapas.A few toads and frogs collected inl863 and 1864 by Dr. H. Berendtin Tabasco, chiefly at San Juan Bautista, were presented by him tothe Smithsonian Institution.At the solicitation of Governor Jose Salazar y Larregui, ArthurSchott, who had previously been employed as a naturalist on theUnited States and Mexican Boundary Commission under the directionof Maj. W. H. Emory, was appointed to the Comision Cientifica deYucatan. By March 20, 1865, he had collected 300 specimens, accord-ing to a letter written from Merida, Yucatan. The type specimenof the peculiar Triprion petasatus (Cope) was collected by Schott nearthis locality. A letter received by Baird on June 6, 1865, states thatSchott had made a trip that included Sisal, Campeche, and Celestun.On November 21, 1866, Schott returned to Washington, D. C, butlater went back to Yucatan and collected there until 1868. Schottcollected also in Sonora in 1871.From Mazatlan in Sinaloa, specimens collected by FerdinandBischoff were received by the Smithsonian Institution in 1868. Dr.Gideon C. Lincecum, an entomologist, presented amphibians col-lected by himself in 1869 at Tuxpam, Vera Cruz. There is a specimenof Rhinophrynus dorsalis in the collection that was taken by Dr. T. H.Richardson at Cordoba in Vera Cruz on October 19, 1874.From time to time, beginning in 1877 and continuing until 1887,Alfred Duges sent to the National Museum specimens labeled ascoming from Guanajuato, though Dr. E. W. Nelson is of the opinionthat some of them at least were given him by students and that thespecimens actually may not have come from that locality. This natu-ralist also supplied the Museum d'Histoire Naturelle in Paris withsimilar collections. Professor Duges collected the type specimensof Scaphiopus dugesii Brocchi, Cystignathus microtis Cope, Eleuther-odactylus aljredi (Boulenger), E. augusti (Duges), Syrrhophus guttilatus(Cope), BuJo monksiae Cope, and Rana montezumae concolor Cope.From C. T. Hoege, a collector previously employed by F. D.Godman, the National Museum received amphibians collected in
MEXICAN TAILLESS AMPHIBIANS 9the Valleys of Mexico and Toluca in 1884 and 1885. AlphonsoForrer, another one of Godman's collectors, sent specimens collectedin 1885 at Ciiidad in Durango and at the Presidio de Mazatlan inSinaloa.During March and April, 1891, William Lloyd, a field naturahstemployed by the Bureau of Biological Survey, obtained a few amphib-ians near the Mexican boundary in the States of Nuevo Leon andTamaulipas. Pierre Louis Jouy, an ornithological collector employedto obtain specimens for the National Museum exhibit at the ChicagoWorld's Columbian Exposition, brought back a few specimens takenin February, 1892, at Lake Chapala in Jalisco.Maj. E. A. Mearns was detailed by the Vv^ar Department to actas the medical officer of the International Boundary Commission, andin the course of his work he traversed the entire boundary line andcollected a number of amphibians at localities along it. He reportedfor duty at El Paso, Tex., on February 1, 1892, and completed thework of the biological section on July 20, 1894, at San Diego, Calif.A few amphibians taken in Sonora in 1893 were received from anarmy hospital steward, Ernest C. Merton.Dr. Edward Palmer, a well-known professional botanical collector,in 1896 forwarded specimens collected in Durango. Dr. CharlesHaskins Townsend collected at Frontera in Tabasco during April,1897. General herpetological collections were made in Chihuahuaduring 1895 by H. H. and C. S. Brimley. The late J. N. Rose, abotanist employed by the National Museum, and his assistant J. H.Painter collected in Jalisco in 1903.From 1908 to 1923, a few amphibians were received from entomolo-gists, as follows: In 1908, Frederick Knab collected at Cordoba, VeraCruz; in 1918, J. A. Kusche at Venodio, Sinaloa; in 1920, W. S.Blatchley at Orizaba, Vera Cruz; and, in 1923, Dr. Wilham M. Mannat Tepic, Nayarit. A Phyllomedusa helenae taken at Berta, VeraCruz, in 1910 was purchased from Charles R. Orcutt. Surg. J. C.Thompson collected a Hyla baudinii at San Bias, Nayarit, in 1913. Afew specimens taken at Nogales, Sonora, in 1919, were received fromAmerican Consul Francis J. Dyer.MATERIAL IN OTHER MUSEUMSAmerican.?The writer was enabled, while a member of the staffof the Bureau of Biological Survey, to visit several American insti-tutions for the purpose of studjdng their collections of Mexicanamphibians. A large number of important distributional recordswere obtained from these collections.The Mexican herpetological collection of the American Museum ofNatural History in New York ranks next to that of the United StatesNational Museum in size and includes specimens received from
10 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
several sources. The bulk of this collection, however, was gatheredby two men: Dr. Alexander G. Ruthven, who made a herpetologicalcollection for this museum in Vera Cruz, and Paul D. R. Ruthling,who during 1919 and 1920 assembled a large and varied collection.Ruthling's itinerary covered Mexican States as follows : In 1919 collec-tions were made in Colima in April; in the Federal District in Mayand July ; in Vera Cruz in June ; field work was carried on in Guana-juato during the first week in August; the remainder of August, Sep-tember, and October were spent in Jalisco; in November and Decem-ber field work was carried on in Nayarit and Sinaloa; during May,June, and the first part of July, 1920, amphibians were collected inOaxaca, while Puebla was visited during the latter part ofJuly. Tliismaterial for the most part is well preserved, so that a critical deter-mination of some doubtful points of external form of some of theampliibians, which hitherto had been known only from specimenscollected many years ago, was possible.The Museum of Comparative Zoology, at Harvard College, Cam-bridge, Mass., possesses a number of Mexican amphibians. From1878 to 1880, Dr. Edward Palmer, a botanical collecter, seems to havebeen somewhat interested in the natural history of Tamaiilipas, SanLuis Potosi, and Coahuila, as specimens from these localities weresent to Cambridge. In 1911-12, members of a party engaged inMayan archeological work in Yucatan, particularly L. J. Cole,Edward H. Thompson, and O. Ricketson, sent specimens to thismuseum. A few specimens were collected by J. L. Peters in theState of Quintana Roo. From the State of Hidalgo, this museumhas specimens collected by Dr. William M. Mann. An ornithologi-cal collector, W. W. Brown, preserved a few amphibians in the courseof his field work in Tamaulipas, San Luis Potosi, and Sonora. Dr.E. R. Dunn collected in the State of Vera Cruz and in the FederalDistrict in 1921. On this trip Doctor Dunn collected the typespecimens of Eleutherodactylus dunnii Barbour and Syrrhopus mys-taceus Barbour near Jalapa in Vera Cruz. Several other individuals,including Dr. G. O. Rogers, P. Townsend, D. B. van Brunt, G.Gliickert, and T. J. Potts, have presented Mexican amphibians to theMuseum of Comparative Zoology.Along with his collections of fishes, S. E. Meek included a smallnumber of amphibians found in the States of Tamaulipas, Guana-juato, Federal District, and elsewhere in Mexico in Ms shipments tothe Field Museum of Natural History, Chicago, 111. Two mammalcollectors, Edmund Heller and C. M. Barber, likewise collected afew amphibians in the State of Vera Cruz. A few specimens werepresented by Dr. C. H. T. Townsend, and a collection made bySenor F. Ferrari-Perez, director of the museum at Tacubaya, Mexico,
MEXICAN TAILLESS AMPHIBIANS 11in 1889 was purchased. From these sources were obtained practi-cally all the Mexican amphibians belonging to this museum.In 1910, about 15 species of Mexican amphibians were collected forthe University Museums, University of Michigan, by Dr. A. G.Ruthven on a low plain at the foot of the San Andreas TuxtlaRange in southern Vera Cruz. From H. B. Baker this museum in1926 received a small number of amphibians collected in Puebla andVera Cruz.During July, 1925, Joseph R. Slevin made general herpetologicalcollections for the California Academy of Sciences, San Francisco, inVera Cruz, Federal District, and Oaxaca.European.?Herpetologists of Europe have described 54 species ofMexican tailless amphibians, and the majority of the types andCO types of these forms are still extant in European museums. Thewriter therefore made an effort during April and May, 1930, to locateand examine as many of these original specimens as possible.The nucleus of the Mexican herpetological collection in the Museumfiir Naturkunde in Berlin seems to have been obtained from threebotanical collectors. Included in the herpetological collections madeby Ferdinand Deppe, Christian Julius Wilhelm Schiede (who died inMexico in 1836), and Graf von Sack were the types of the toads ofthe genus Bujo described by Dr. Arend Frederick August Wiegmannin Oken's Isis for 1833. These toads, Bwfo compaciilis, B. cristatus,B. horribUis, B. marmoreus, and B. valliceps, formed a part of whatappears to be the first Mexican herpetological collection received byany European museum, and presumably led Wiegmann to publishin 1834 his "Herpetologia Mexicana." In 1870, Dr. Wilhelm C. H.Peters published a report on a collection made by Berkenbusch nearMatamoros and at other localities in Puebla. In reporting upon thiscollection, Peters described the following new species: Hyla microtis,Hylodes berkenbuschii, Liuperus nitidus, and Engystoma mexicanum.A new leptodactylid collected at Haunusco [ = ? Huatusco in VeraCruz] by Doctor Hille was named Phyllobates verruculatus by Peters.Fifteen supposedly new species of Mexican amphibians w^ere basedupon specimens belonging to the Museum National d'Histoire Natu-relle, in Paris. In their "Erpetologie Generale" published in 1841,Dum^ril and Bibron recognized two new amphibians from Mexico,one a peculiar burrowing narrow-mouthed toad (Rhinophrynus dor-salis), which had been collected by Auguste Sall6, and the other awidely distributed tree frog (Hyla baudinii). In 1853, Dumeril dis-covered the broad-headed cave frog (Eleutherodactylus laticeps) amongspecimens collected by Arthur Morelet in Yucatan. The Frenchnaturalist Pierre Marie Arthur Morelet made an extended field tripthrough southern Mexico and Central America during the years 1847and 1848, and presented his collection to the Paris Museum.
12 BULLETIN 160, UNITED STATES NATIONAL MUSEUMFollowing the example of Napoleon I after the conquest of Egypt,Napoleon III promulgated a decree, dated February 27, 1864, thatformally established the Mission Scientifique au Mexique et dansI'Amerique Centrale. Firmin Bocourt received an appointment asnaturalist to this commission. Adolphe Boucard seems to have beenattached to the commission in some capacity, as his name appearsamong those appointed as voyageurs. The exploratory work of thiscommission extended from 1865 to 1867. Boucard had visited Mex-ico at least once prior to this, for in 1855 he and Auguste Salle weremaking collections in Vera Cruz. After the death of Auguste Dume-ril in 1871, during the siege of Paris, Firmin Bocourt was given thetask of completing the report upon the herpetological collections.The aid of Paul Brocchi having been obtained, he was commissionedto prepare the report on the amphibians, and this was published in1882 under the title of "fitude desBatraciens de I'Amerique Centrale.
"
Preliminary descriptions of the new species were published in 1877and 1879. Two of those described as new, Leiuperus mexicanus andCauphias crassus, were collected by Adolphe Boucard. The types ofHijla plicata and Leptodactylusfragilis were found by Firmin Bocourt.In addition to the material assembled by the personnel of the mission,the Museum National d'Histoire Naturelle received herpetologicalcollections from Auguste Sall^, Consul Ghiesbreght, Baron Fredericde Miiller, Francis Sumichrast, and Alfred Duges. Rhinophrynusrostratus and Exerodonta sumichrasti were based upon specimensreceived from Sumichrast. Sall^ collected the type of Bufo mexicanusand Duges sent Brocchi the type of Scaphiopus dugesii. During theyears 1896 and 1897, L6on Diguet made collections in Mexico for theParis Museum, particularly in the State of Oaxaca, in the vicinity ofGuadalajara, Jalisco, and in the Territory of Tepic (Nayarit). Inreporting upon these collections in 1899, Mocquard recognized threenew species, which were named Rana trilobafa, Hyla rudis, andHyliola digueti.The British Museum of Natural History has a large representativecollection of Mexican amphibians, including the type specimens ofthe species described by Giinther and Boulenger, In tabulating thedescribed species, the writer found that in this collection alone speci-mens from Mexico have formed the basis for 27 species of taillessamphibians.Previous to 1858, the British Museum had acquired by purchase,gift, or exchange a small number of Mexican amphibians, amongwhich were several amphibians taken by Auguste Sall6 in Vera Cruzand Oaxaca. On April 16, 1855, Sall^ and his companion AdolpheBoucard were located at Tospam near Cordoba, Vera Cruz, wherethey were visited by Francis Sumichrast. At Cordoba in Vera Cruz,
MEXICAN TAILLESS AMPHIBIANS 13SalM collected cotypes of Hyla euphorbiacea Giinthev, H. lichenosaGiin-ther, and Bufo sternosignatus Giinther, and the type of Gastrophryneelegans (Boulenger). One of the cotypes of Eleutherodactylus sallaei(Giinther) was also received from Salle, and while Mexico is the onlylocality given, it too was probably collected somewhere in the Stateof Vera Cruz. The follo^\dng specimens, purchased from Emile Par-zudaki, a dealer at Paris, lack definite localities: Cotypes of Hylaeuphorbiacea Giinther, H. lichenosa Giinther, and Bufo sternosignatusGiinther, and the type of B. anomalus Giinther. The specimensacquired from Hugh Cuming, including cotypes of Hyla euphorbiaceaand H. lichenosa, were collected in the State of Vera Cruz, probablyby Auguste Salle, as the former secured a number of birds from thiscollector.In 1876, Frederick du Cane Godman and Osbert Salvin conceivedthe idea of promoting a monumental work entitled "Biologia Centrali-Americana," which was begun as a private undertaking, though later(beginning in 1881) the collections were presented to the BritishMuseum. To Dr. Albert C. L. G. Giinther were assigned the sec-tions on reptiles and amphibians, and collectors were employed tosecure specimens that would supplement available material. Amongthese collectors was Herbert H. Smith, who collected at Atoyac inVera Cruz, Teapa in Tabasco, Omilteme and Amula in Guerrero, andCuernavaca in Morelos. From Omilteme came the cotypes of Ranaomiltemana Giinther, Syrrhaphus omiltemanus Giinther, and Eleu-therodactylus calcitrans (Giinther); from Amula the type of Tomo-dactylus amulae Giinther; and from Cuernavaca the cotypes of Hylanana Giinther {H. smithii Boulenger). Dr. A. C. Buller collectedon the plateau of central Mexico, and at the Hacienda el Florencioin Zacatecas he obtained the cotj^pes of Borborocoetes mexicanusBoulenger. Godman collected in Mexico during 1887 and 1888, andhe is listed in the records of the British Museum as the collector of thetypes or cotypes of five of the species described by Giinther. C. T.Hoege and an Indian assistant of Godman, Mateo Trujillo, collectedat Jalapa and Misantla in Vera Cruz. At Jalapa, Hoege collected acotype of Hyla nigropunctata Boulenger, while Trujillo secured thetype of Hylella picta Giinther and the cot3^pes of Hyla iaeniopusGiinther. Godman obtained the types of Eleutherodactylus venustus(Giinther) and E. plicatus (Giinther) at Jalapa, while the cotypes ofHyla godmani Giinther came from Misantla and Jalapa.Alphonso Forrer collected on the Tres Marias Islands and at severallocalities in western Mexico, particularly at Presidio near Mazatlanin Sinaloa, from where Forrer forwarded specimens that became thetypes of Ranaforreri Boulenger and Pternohyla fodiens Boulenger andthe cotypes of Diaglena spatulata (Giinther) and Hypopachus oxyr-66785?32 2
14 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
rhinus Boulenger. Forrer collected also the type of Rana pustulosaBoulenger at Ventanas in Durango. In 1888 George F. Gaumerworked in Yucatan and visited Cozumel, Ruatan, and the BonaccaIslands.The cotypes of Eleutherodactylus alfredi (Boulenger), which werepresented by Alfred Duges, came from Atoyac in Vera Cruz. Dr.Hans Gadow and his wife on one of their visits to Mexico collectedthe cotypes of Eupemphix gadovii Boulenger at San Mateo del Marnear Tehuantepec, and those of Eleutherodactylus beatae (Boulenger)at La Perla near Orizaba. The cotypes of Rana tarahumarae fromthe Sierra Tarahumari formed part of a collection acquired fromDoctor Gadow.So far as known only one of the Mexican types
?
Bufo lateralis~islocated in the Naturhistorische Museum at Vienna, and that wasdescribed by Dr. Franz Werner from a toad collected in Tehuantepecin 1872 by T. K. Salmon. A number of Mexican amphibians collectedby Dr. Hans Gadow are in the Vienna museum.The type of the Mexican toad Bufo occipitalis, which was de-scribed by Prof. Lorenzo Camerano in 1879, should be in the museumat Turin, Italy, but it could not be found at the time of the writer'svisit there in 1930. The type of Hylodes augusti remained in thepossession of Dr. Alfred Duges at Guanajuato, Mexico, and its presentwhereabouts are unknown.The tailless amphibians of Mexico are not known to be less abun-dant than those found within the borders of the United States, andyet some of these species are very meagerly represented in Americancollections. For some of the species only a single individual wasavailable, and in one or two instances the species is known to thewriter solely from the diagnosis published by the original describer.Practically all the types of species desciibed by Baird, Barbour, Bo-court, Boulenger, Brocchi, Cope, Giinther, Mooquard, Peters, Werner,and Wiegmann have been studied by the writer. The majority of thespecimens examined have been preserved in alcohol from 30 to 100years, and the vicissitudes of time, including evaporation and actionof the preservatives, as well as the bleaching effect of light, havehindered and in some instances prevented critical comparisons.Among the many published reports that deal with or refer to Mexicanamphibians, there are a number of such a general nature that noreference is made to them in the text. Shorter articles and particu-larly those containing either descriptions of new species or supplemen-tal data on those already described are cited in the synonymy.
MEXICAN TAILLESS AMPHIBIANS 15ACKNOWLEDGMENTSDr. Leonhard Stejneger and Miss Doris M. Cochran, of the NationalMuseum, have facilitated the preparation of this report in many ways,and to them the writer is greatly indebted for criticisms and sug-gestions. Grateful acknowledgment for the privilege of includingspecimens under their care in this report is hereby extended to Dr.G. K. Noble, of the American Museum of Natural History in NewYork; Dr. Thomas Barbour, director of the Museum of ComparativeZoology in Cambridge, Mass.; Dr. Alexander G. Ruthven, directorof the University Museums of the University of Michigan in AnnArbor; Alfred C. Weed and K. P. Schmidt, of the Field Museum ofNatural History in Chicago; Joseph R. Slevin, of the CaliforniaAcademy of Sciences in San Francisco; Dr. Ernst Ahl, of the Museumfiir Naturkunde in Berlin; Dr. Otto Wettstein, of the Naturhistori-sche Museum in Vienna; Dr. Laura Gambetta, of the IstitutoZoologico R. Universita in Turin; Dr. Louis Roule, director, andF. Angel, of the Museum National d'Histoire Naturelle in Paris;and W. H. Parker, of the British Museum of Natural History inLondon. To Dr. E. R. Dunn, of Haverford College, Haverford, Pa.,the writer is indebted for comments and notes on specimens that hehad examined in American and European museums. The drawingsfor the figures of toads of the genus Bufo, prepared under the author'ssupervision, were loaned by the Bureau of Biological Survey.ABBREVIATIONSThe following abbreviations are used in the lists of specimensexamined during the preparation of this report and in the text wherecatalogue numbers are given:A.M.N.H American Museum of Natural Historj^ New York,N. Y.A.N.S.P Academy of Natural Sciences of Philadelphia,Philadelphia, Pa.B.M British Museum of Natural Histor}-, London,England.C.A.S California Academy of Sciences, San Francisco,Calif.C.G.E.M Coraision Geografica Exploradora de Mexico,Mexico City, Mexico.F.M.N.H Field Museum of Natural Iliitory, Chicago, 111.M.C.Z Museum of Comparative Zoology, Cambridge,Mass.M.H.N.P Museum National d'Histoire Naturelle, Paris,France.M.N.B Museum fiir Naturkunde, Berlin, Germany.N.M.W Naturhistorische Museum, Vienna, Austria.U.M.U.M University Museums, University of Michigan, AnnArbor, Mich.U.S.N.M United States National Museum, Washington, D. C.
16 BULLETIN 160, UNITED STATES NATIONAL MUSEUMClass AMPHIBIA Linnaeus1758. Amphibia Linnaeus, Systema naturae, ed. 10, vol. 1, p. 194.The amphibians have a skull that is characterized by severalmorphological features. The chrondrocranium is platybasic; thequadrate is immovably united with the cranium by an otic process,which joins the otic capsule, and by an ascending process, whichreaches to the pterygoid; the stapes occupies the fenestra vestibuliof the otic capsule; and there are two occipital condyles.Of the three recent orders included in this class, the Salientia(frogs and toads) are far more numerous as regards genera and speciesthan either the Caudata (salamanders) or the Apoda (caecilians).The Salientia alone are treated in this report.Order SALIENTIA Laurenti1768. Salientia Laurenti, Synopsin reptilium, p. 24.The members of this order have the hind limbs developed forleaping, and they are much longer and stronger then the forelegs.The most striking skeletal feature found in the Salientia is the fusionof the caudal vertebrae into a single rod, the coccyx or urostyle. Theskull is characterized in part by the absence of basioccipital, supra-occipital, and supratemporal bones, by the ankylosis of the frontaland parietal bones, and by the lack of any connection between thestapes and the quadrate. Vomerine and palatine bones are presentbut are not united with each other, and the latter bounds the choanaposteriorly and joins the pterygoid externally. The astragalus andcalcaneum are elongated and form a distinct section of the hind limb,the tarsus.The classification of the Salientia adopted is based on slight modi-fications of internal structure, and some of the forms can only bedistinguished from closely related species by close scrutiny. Theminute details of internal structure, such as the relations of theseveral elements that compose the ventral portion of the shouldergirdle, the shape of the sacral diapophyses, the articular relationsbetween the sacral vertebra and coccyx, the number of presacralvertebrae, the shape of the terminal phalanges, and the presence orabsence of intercalary disks, can only be determined by dissection.Other characters employed are the presence or absence of maxillary,vomerine, and parasphenoid teeth, the attachment of the tongue, thepresence or absence of parotoid glands, the presence or absence oflumbar glands, the shape of the pupil of the eye, the extent of theweb between the digits of the fore and hind feet, the characters fur-nished by the texture of the skin, and the color patterns.The present report is based on the critical study of nearly 2,200specimens from the mainland of Mexico. Seven families of Salientiaare represented in Mexico by a total of 19 genera and 65 species.
MEXICAN TAILLESS AMPHIBIANS 17KEY TO FAMILIES OF MEXICAN SALIENTIA
1. Sacral vertebra procoelous 2Sacral vertebra biconvex, with double condyle for coccyx (fig.1, a) ; eighth vertebra biconcave, preceded by seven procoelousvertebrae; without ribs; sacral diapophyses cylindrical ordilated; pectoral girdle firmisternal (fig. 21, p. 195) 62. Sacral vertebra ankylosed to coccyx (fig. 1,6); presacral vertebraeeight, uniformly procoelous (rarely opisthocoelous) ; withoutribs at any stage in development; sacral diapophyses dilated;pectoral girdle arciferal; pupil of eye vertically elliptical; soleof foot smooth, but provided with a large inner tubercle withblack cutting edge Pelobatidae (p. 18)Sacral vertebrae free (fig. 1, c), with double condyle for coccyx;presacral vertebrae eight to five, procoelous; without ribs; pec-toral girdle arciferal (fig. 3, p . 27) 33. Neither maxillary nor vomerine teeth present; parotoid glandsgenerally visible 4Maxillary teeth present (except in Engystomops) ; no visibleparotoid glands; vomerine teeth present or absent 54. Head small and unusually narrowed; no distinct neck; size ofbody disproportionate to size of head; mouth conspicuouslysmall and narrow; snout truncated in front, forming a flatcircular disk; eyes rather small; pupil vertical; parotoid glandsflattened and not raised above skin on shoulders; no tym-panum; fingers free; toes fully webbed, the tips not dilated;outer metatarsals united; inner toe tuberclelike; a very largeshovellike inner metatarsal tubercle; epicoracoid cartilagesnarrow, scarcelj'^ overlapping; no omosternum; sternum ves-tigial; sacral diapophyses noticeably dilated; terminal phalan-ges simple Rhinophrynidae (Rhinophrynus) (p. 24)Head not unusually narrowed; mouth large; parotoid glandsdistinct; terminal phalanges T-shaped or simple (very rarelyclaw-shaped); sacral diapophyses cylindrical or dilated;presacral vertebrae eight Bufonidae (p. 27)5. No intercalary cartilage or bone between ultimate and penulti-mate phalanges of each digit, supporting the claw-shaped orT-shaped terminal joint (fig. 17, p. 76); sacral diapophysescyUndrical or dilated Leptodactylidae (p. 74)An intercalary cartilage or bone supporting terminal phalanges(fig. 2), which are generally claw-shaped; sacral diapophysesdilated Hylidae (p. 129)6. Head distinctly narrowed and mouth small; generally a fold ofskin on top of head behind the eyes; a dermal ridge extendsacross the palate between the choanae, and another in frontof the esophagus; tympanum indistinct or not visible; vom-erine teeth absent; sacral diapophyses somewhat dilated; noomosternum Brevicipitidae (p. 182)Head not unusually narrowed; mouth large; tympanum distinct;maxillary and vomerine teeth present; tongue bicornuate ordeeply notched behind; sacral diapophyses cylindrical orslightly dilated Ranidae (p. 189)
18 BULLETIN 160, UNITED STATES NATIONAL MUSEUMFamily PELOBATIDAE Lataste1865. Scaphiopodidae Cope, Nat. Hist. Rev., London, p. 107, Jan.1879. Pelobatidae Lataste, Compte Rendu Assoc. Frang. Avanc. Sci., for 1878,pp. 761, 762.?BouLENGER, 1882, Catalogue of the Batrachia Salientia s.Ecaudata in the collection of the British Museum, 2d edit., p. 432.Sac.
Figure 1.?Diagrams illustrating structural conditions found in the sacral vertebra and coccyx ofMexican tailless amphibians: a, Rana. ptpi?ns (U.S.N.M. No. 541); sacral vertebra {Sac^ free andbiconvex with double condyle for the coccyx (Coc); 6, Scaphiopus ftawimondii (U.S.N.M. No.60359); sacral vertebra procoelous, that is, with an anterior concave articular surface for corre-sponding presacral vertebra, and ankylosed posteriorly with coccyx; c, Hyla baudinii (U.S.N.M.No. 16563); sacral vertebra free, procoelous, and with double condyle for coccyxThe genera comprising tlie true spade-foot series of toads have anearly continuous distribution over most of the Palaearctic region,and a second series
?
Megalophrys andAelurophryne?have a restricted range insouthern Asia and the neighboring islands.Doctor Noble ^ has recently studied Sooglos-sus and Nesomantis, which are indigenousto the Seychelles Islands in the IndianOcean, and concludes that his dissectionsshow that this family did penetrate intothe Southern Hemisphere. This is generallyconsidered to be one of the primitive families of living Salientia
Figure 2.?Terminal phalanges ofthe toe in the foot of Hyla, i indi-cating the position of the inter-calary phalanx
Genus SCAPHIOPUS Holbrook1836. Scaphiopus Holbrook, North American herpetology, Philadelphia, ed. 1,vol. 1, p. 85. [Genotype, Scaphiopus solitarius Holbrook, idem, vol. 1,p. 85, pi. 12 (Carolina, Georgia, and Tennessee) = fiano holbrookii Harlan,1835, Medical and physical researches, p. 105 (South Carolina).]
' Noble, G. K., An analysis of the remarkable cases of distribution among the Amphibia, with descrip-tions if new genera, Amer. Mus. Nov., no. 212, pp. 8-12, 1926.
MEXICAN TAILLESS AMPHIBIANS 191866. Spea Cope, Journ. Acad. Nat. Sci. Philadelphia, new ser., vol. 6, pt. 1,p. 81, July. [Genotype, S[caphiop7is] bombifrons Cope, 1863, Proc. Acad.Nat. Sci. Philadelphia, vol. 15, p. 53, Mar. (type locality not designated,but the cotypes were obtained at Fort Union on the Missouri River, latitude48? N.; on the Platte River, 200 miles west of Fort Kearney; and at LlanoEstecado, Tex.).]The genus Scaphiopus is the only representative of the familyPelobatidae in North America. The several species breed intemporary pools after rains, and the ropelike egg mass is attachedto water weeds. Wright has found that the eggs of Scaphiopushammondii are occasionally stalked. The egg has two capsules inaddition to the vitelline membrane. In one species, *S. holbrookii,the intracapsular development is very short, the larvae hatch in animmature condition, and the frontal gland initiates the hatchingprocess, the tadpoles emerging before the gills are formed. Thetadpoles of Scaphiopus have simple teeth.KEY TO MEXICAN SPECIES OF SCAPHIOPUS
1. Glandular enlargements or parotoid glands variable in theirpresence, generally limited to a thickening of the skin inscapular region; skin of upperparts generally conspicuouslywarty, covered with closely spaced large tubercles, especiallyon sides and shoulders; a conspicuous pair of light-coloredsupra-anal spots; vomerine teeth in two clusters either be-tween or at level of posterior margin of choanae; tympanumvisible or concealed by hypertrophy of parotoid; canthusrostralis scarcely indicated; head flat or slightly concavebetween the eyes; cephalic ossification incomplete; underpartssmooth, with a few rounded areolae; fingers in order ofdecreasing length, 3, 1, 4, 2; palm smooth except for con-spicuous elongated inner metacarpal tubercle and less dis-tinct smaU outer metacarpal tubercle; fingers essentially free,with a vestigial web on each side of second finger rarelypresent; first and second fingers of breeding males with hornynuptial excrescences; heels not meeting when legs are flexed;web complete or strongly scalloped, in which case the distalphalanx is largely free on all toes but fourth, on which the twodistal phalanges either are or are not bordered with dermalfringe; sole smooth except for large external metatarsaltubercle with black cutting edge; toes in order of decreasinglength, 4, 3, 5, 2, 1; color pattern of upperparts variable;darker coloration restricted chiefly to V-shaped interorbitalband and anastomosing blotches or wavy cross bands inthoracic and sacral regions, which may or may not interruptdistinct or indistinct longitudinal light stripes; generally apair of blotches of same color in scapular region; longitudinallight-colored stripes when present 2 to 4 in number, the innerpair of which are on a line with inner margins of orbits, and theouter with tympanum; vertical light-colored stripe on tip ofnose; fore and hind limbs with transverse dark crossbars; head-and-body length of adults, 55 mm hammondii multiplicatus (p. 22)
20 BULLETIN 160, UNITED STATES NATIONAL MUSEUMNo glandular enlargement on post-tympanic region; a supra-scapular dermal enlargement above and in front of tympanum;parotoid glands generally absent, or small and indistinct; skinof upperparts rough, with scattered low tubercles, more closelyspaced on sides and shoulders than elsewhere; light-coloredsupra-anal spots smaller and generally indistinct; vomerineteeth in two clusters behind or at level of posterior margin ofchoanae; tyinpanum large, more than half the diameter of theeye, but anterior margin often indistinct; canthus rostralisrounded; head rounded anteriorly, depressed or flattenedbetween the eyes; cephalic ossification complete; skin of under-parts slightly roughened, smoother anteriorly; fingers in orderof decreasing length, 3, 1, 2, 4; palm smooth except for twolarge metacarpal tubercles; fingers free; first and second fingersblack and horny internally in breeding males; heels not meet-ing when legs are flexed; foot large, web strongly scalloped, thefourth digit having the two distal phalanges bordered by dermalfringes; sole smooth except for large external metatarsal tuberclewith black cutting edge; color pattern of upperparts irregular,asymmetrical, but with intricate arrangement of spots, or net-work of brown bands; upperparts variable in color?green,olive, or brown?and streaked, spotted, or blotched more orless irregularly with some shade of yellow; a dark V-shapedinterorbital band; pustules black or yellow; underparts lightcolored; fore and hind limbs barred or spotted with somedarker color; head-and-body length of adults, 65 mm couchii (p. 20)SCAPHIOPUS COUCHII BairdCouch's Spade-foot Toad1854. Scaphiopus couchii Baird, Prcc. Acad. Nat. Sci. Philadelphia, vol. 7, p. 62,Apr.
?
Baird, 1859, Report on the United States and Mexican BoundarySurvey, vol. 2, Reptiles, p. 28, pi. 35, figs. 1-6.1863. S [caphiopus] rectifrenis Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 15,p. 53, Mar. (Tamaulipas; Rio Nazas, Coahuila).Type locality.?Rio Nazas, State of Coahuila, and Matamoros,State of Tamaulipas, Mexico.Range.?In Mexico from Matamoros southwestward through theStates of Tamaulipas, Coahuila, and San Luis Potosi to Acaponeta inNayarit.According to Strecker,^ Couch's spade-foot toad inhabits the entireRio Grande Valley from Brownsville to El Paso, Tex., and, althoughits range in Texas is still imperfectly known, it does not appear torange east of the 96th parallel. The northernmost record is 50 milessouth of Fort Worth.Remarks.?Baird gave Coahuila and Tamaulipas as the habitatof *S'. couchii, but neglected to cite the catalogue numbers of the co-types. Cope, however, in revising this genus in 1863 removed one(U.S.N.M. No. 3714) of Baird's original cotypes of S. couchii and
< Strecker, J. K., Notes on the life history of Scaphiopus couchii Baird. Proc. Biol. Soe. Washington,vol. 21, pp. 199-206, Nov. 24, 1908.
MEXICAN TAILLESS AMPHIBIANS 21transferred it to his new species redifrenis. This specimen wascollected on the Rio Nazas in Coahuila by Lieut. D. N. Couch andunfortunately can not now be located in the collection. In thissame paper, Cope cited a specimen (U.S.N.M. No. 3713) obtainedby Couch at Matamoros in Tamaulipas as the basis for the diagnosisof S. couchii. The other cotype of S. redifrenis (U.S.N.M, No. 3715),which was collected in Tamaulipas by Couch, is marked in the recordbook of the division of reptiles as having been destroyed. Thecollections that Cope had in his possession for study so many yearswere returned to the National Museum after his death, but so far asknown not one of these cotypes was ever found.This night-prowling spade-foot toad has a somewhat bloated appear-ance in life and frequents lowlands as well as plateaus. It is mostfrequently observed during the time of the spring rains, when itcomes forth to breed in temporary pools and at other times whenheavy rains drive it from its burrow. It is well adapted for a subterra-nean and nocturnal life, and at certain seasons may be found halfburied in moist earth under some old log. From 350 to 500 eggs arelaid in strings, the number in each string varying from 45 to 125. Theeggs are attached to grftss, weeds, or roots in shallow water, generallyless than 6 inches in depth, and they hatch in 8 to 10 days. Thegrowth of the tadpole is very rapid, and within three weeks the meta-morphosis to the land stage has taken place. Doctor Nelson observesin his field catalogue that hundreds of these spade-foot toads weremating at Acaponeta on June 26, 1897, in a pool of water near theriver bank.Specimens examined.?Thirty, as follows:Scaphiopus couchii
Museum
22 BULLETIN 160, UNITED STATES NATIONAL MUSEUMSCAPHIOPUS HAMMONDII MULTIPLICATUS (Cope)Mexican Spade-foot Toad1863. S[caphiopus] multiplicatus Cope, Proc. Acad. Nat. Sci. Philadelphia, vol.15, p. 52, Mar.1879. Scaphiopus dugesii Brocchi, Bull. See. Philom. Paris, ser. 7, vol. 3, no. 1,p. 23.
?
Brocchi, 1882, Mission scientifique au Mexique et dans I'AmeriqueCentrale, recherches zoologiques, pt. 3, sect. 2, p. 24, pi. 9, fig. 4 (Mexico;.Type locality.?"Valley of Mexico," State of Mexico, Mexico.Range.?From Sonoyta in northwestern Sonora south throughwestern Chihuahua and Durango to northern Jalisco; east to Mi-quihuana in western Tamaulipas, and south through San Luis Potosito Guanajuato, Mexico, and Federal District. North of the Mexicanboundary, the species S. hammondii and its geographic races aredistributed from the northern Okanagan Lake district (Vernon) ofBritish Columbia, south through western Washington, Oregon, Neva-da, and California, to northern Lower California (Ensenada) ; eastto the western margin of the great plains in northwestern NorthDakota (Fort Union); and south along the great plains to westernKansas, the panhandle district of Texas, and to San Antonio andEl Paso.Remarks.?The type (U.S.N.M. No. 3694) was sent by JohnPotts to the Smithsonian Institution. The actual preservation of thetype is fairly good, notwithstanding numerous incisions made byCope. There is a transverse incision across the right suprascapularregion, one across angle of jaw, a longitudinal and two crosswise cutsalong the sacrum, and an oblique cut on the upper surface of theright tibia. There are lengthwise and transverse cuts on the throat,and the viscera have been exposed by a long crescentic incision. Theskin has been freed from the upper surface of the head.The two cotypes of Scaphiopus dugesii (M.H.N. P. No. 281a,parchment labels 86-287, 86-288) are both young individuals, withhead-and-body lengths of 32 mm. and 33.5 mm., respectively. Blackspots are present on the upperparts and the black inner metatarsaltubercle is quite distinct. Both of these cotypes are typical youngindividuals of this Mexican form. Alfred Duges is listed in thecatalogue as the collector of the cotypes of S. dugesii.Specimens from northern Sonora have a relatively smoother skinon the dorsal surface of the body than those taken farther south.Some of those from northern Sonora differ in no appreciable mannerfrom California and Nevada specimens unquestionably referred to(S. hammondii. Along the southern border of the known range ofScaphiopus in Mexico, individuals with very warty skins and with anappreciable thickening of the skin in the parotoid region predominate.Between these two extremes are various intermediate individuals,and no satisfactory method for determining the status or relations of
MEXICAN TAILLESS AMPHIBIANS 23
such extreme types is apparent. Additional material may show thatthese spade-foot toads with skins grading from relatively smooth towarty are separable into at least two groups, and that a more satisfac-tory disposition of this puzzling question will result from the allocationof those taken along the northern border of Mexico to the subspecieshammondii and those with unusually warty skins from farther southto the subspecies multiplicaius
.
The spade-foot toad may be very abundant wherever it is found,though it may escape observation because of its nocturnal habits. Itis commonly assumed that it remains in its burrow most of the yearand that it comes out only to deposit its eggs in June and July inpools formed by heavy rains. The spade-foot makes circular holesin the ground near the bases of shrubs and clumps of weeds, and yetin very sandy soil it is very difficult to find the place where it hasburrowed, for in most instances it appears as if the hole has beenpulled in after the toad. After the breeding season is over, thesetoads seem to take more pains in constructing their burrows, as theyare well-rounded inside and resemble somewhat an earthen jar with anarrow top. The surface around the neck of the burrow is often quitesticky when the spade-foot is discovered inside. In Montana duringJune, 1916, I found that these toads made their appearance aftermidnight on dark nights, and when hunted with a flash light theyburrowed from sight in a very short time. The time from the layingof the eggs to the transformation of the tadpoles and their leavingthe water does not exceed 30 days. This rapid development is ne-cessitated by the tendency of the breeding pools of rain water to dryup very quickly.This spade-foot toad can be distinguished almost instantly by itstrim and neat body appearance, by its moist and usually wartyskin, if it has not been hopping about long enough to fill the poreswith fine sand and dust, and, like its relatives, by the catlike verticalpupils of the eyes. In life the upperparts are usually olive-coloredwith splotches of black or purplish black, and with orange-red tuberclesscattered over the skin.Specimens examined.?Seventy-one, as follows:Scaphiopus hammondii multiplicatus
Museum CatalogueNo. Num-ber ofspeci-mens Locality collected Date collected By whom collected orfrom whom received
U.S.N.MDo...Do...M.C.Z....Do...
62253218012659710038-398363-66
Sonora:NogalesSonoytaChihuahua: Meadow Valley.Tamaulipas: MiquihuanaDuRANGO: Durango (streamsnear city).
July 27, 1919Jan. 20, 1894July 18, 189819241921
F. J. Dyer.E. A. MearnsH. H. and C. S.ley.W. W. Brown.P. Townsend.
24 BULLETIN 160, UNITED STATES NATIONAL MUSEUMScaphiopus hammondii multiplicatus?Continued
Museum
MEXICAN TAILLESS AMPHIBIANS 25The absurd-appearing Rhinophrynus , locally known in Mexico as"poche" or "po-chi," is dark brown in life, with a yellow or orangevertebral stripe in addition to smaller spots or blotches of the samecolor. The head is narrow and unusually small for so large an animal,the skin on the oval-shaped body is loose and flabby, and the limbs arerather short. When disturbed the body is distended and the limbsare drawn up beneath. The small nostrils are located on the top ofthe snout, nearer the eyes than the end of the muzzle. Being noctur-nal, the small eyes of this toad have vertical pupils. No vestige of anexternal tympanum is discernible. Neither maxillary nor vomerineteeth are present. Although the tongue is attached posteriorly andcan not be flung forward like that of the common toad, it is said to beremarkably adept in licking up insects. It has a foot remarkablywell adapted for burrowing, with an inner horny metatarsal tubercleon the calcar fully 10 millimeters in length and a secondary shovelliketubercle about half as long on the end of the flrst toe.Correlated with its burrowing habits are certain anatomical modifi-cations. Inasmuch as the rather brief published diagnoses of thistoad do not adequately describe its structural peculiarities, the follow-ing diagnosis is given :Superior plate of sphenethmoid covered by the completely ossifiedfronto-parietals; sphenethmoid septal walls ossified to end of muzzle,and separating the nasals ; nasals large, entirely in contact with fronto-parietals, separated by median process of the latter, and by thesphenethmoid septum; pterygoid straight, thin, with a short maxil-lary suture; no wing of pterygoid extending inward to parasphenoid
;
exoccipital nearly completely covered by parasphenoid; single fissurefor anterior cranial nerves; neither maxillary nor vomerine teeth;no tympanum; tongue attached behind; coracoid short, expanded atextremities, and slightly curved; clavicle narrow, flattened, andstrongly curved upward; clavicles and coracoids attached mediallyto a single curved or epicoracoid cartilage; no omosternum; scapulashort and wide; suprascapula thin, constricted medially, and withdistal dilation; suprascapula normally in close proximity to and incertain positions capable of sliding over pro-otic; humerus with flathead and high Imifelike deltoid crest; radius and ulna fused; pre-sacral vertebrae eight; sacral vertebra free, procoelous, and withdouble condyle for coccyx; sacral diapophyses short, dilated; femurwith flat head, slender shaft, mner and outer crests on posterior surface,and greatly expanded distal end; tibia and fibula fused, shorter thanfemur; two very short, flattened metatarsals separated by medianoval aperture; large flattened shovellike calcar; first toe with verti-cally expanded terminal phalanx; remaining terminal phalangessimple.
26 BULLETIN 160, UNITED STATES NATIONAL MUSEUMRHINOPHRYNUS DORSAUS DnmerU and BibronBurrowing Toad1841. Rhinophrynus dorsalis Dumeril and Bibron, Erp^tologie g6n6rale, vol. 8,p. 758, pi. 91, figs. 2, 2a.
?
Gunther, 1858, Catalogue of the Batrachia Sali-entia in the collection of the British Museum, p. 127, pi. 12.1897. Rhynophrymis dorsalis Duces, La Naturaleza, Mexico, ser. 2, vol. 2, pp.98-100, pi. 5, figs. 1-5. [Colored plate with correct spelling.]1877. Rhinophrynus rostratus Brocchi, Bull. Soc. Philom. Paris, ser. 7, vol. 1,no. 4, p. 196.
?
Brocchi, 1882, Mission scientifique au Mexique et dansI'Amerique Centrale, recherches zoologiques, pt. 3, sect. 2, pp. 98, 99, pi. 9,fig. 1 (Tehuantepec, Mexico).Type locality.?Vera Cruz, State of Vera Cruz, Mexico.Range.?From Tuxpam in northeastern Vera Cruz, south throughTabasco (reported by Duges from Chiapas), Yucatan, and QuintanaRoo to Guatemala; on the west coast in the region of the Isthmus ofTehuantepec.Remarks.-?The type of Rhinophrynus dorsalis (M.H.N.P. No.693, parchment label No. 743) was collected by Auguste Sall^. Itis a young individual and is fairly typical of the species. The head-and-body length is 44.5 mm.Remarks.?One of these burrowing toads was found by Hugo Fuickon October 19, 1874, in the neighborhood of Cordoba in a lagoon atan altitude of 2,100 feet. Among the field notes of Dr. E. W. Nelsonare the following observations on this interesting toad
:
They occur in thickets and woods near Tuxtepec in the state of Oaxaca. Onecame into camp at night during a rainstorm and at the same time the muffledcall notes of others, which were more like the hoarse calls of some bird than ofa toad, were heard. These toads live in the ground and are called "Po-chi."When held in the closed hand they try to escape by moving backwards, and bya kind of a hitching motion of the hinder parts they exert a surprising amount offorce. The hard callosities on the inside border of the feet are used to secure ahold and no doubt serve in this way when the animal is burrowing underground.Specimens examined.?Fifty-eight, as follows
:
Rhinophrynus dorsalisMuseum CatalogueNo. Num-ber ofspeci-mens Locality collected Date collected B y whom collected orfrom whom receivedU.S.N.M.M. C.Z..U.S.N. M....C.Q.E. M..M.H. N. P.U.S.N.M...-Do..Do-DoDo-.-.M. C. Z-...U.S.N.M...M. C. ZDo
82628458-8506
69330271-737040446920-2147312473132421662227722S56
Vera Cruz:CordobaHacienda Cerro del Qallo,near San Cristobal delLlave.TuxpamVera Cruz
....doOaxaca:Tehuantepec CitydoTuxtepecTabasco:Montecristo
....doNo definite locality
..._doYucatan: Chichen Itza sa-cred cenote.
I Quintana Roo: Xcopen
Oct. 19, 18741921
Apr. 9, 1894May 8, 1900May 29, 19001895186019111912
Hugo Fuick.E. R. Dunn.Gideon C. Lincecum.Auguste Sallfi.Francis Sumichrast.Do.Nelson and Goldman.Do.Do.T. J. Potts.Ch. Laszlo.E. H. Thompson.J. L. Peters.Type.
MEXICAN TAILLESS AMPHIBIANS 27Family BUFONIDAE Hogg184L Bufonidae Hogg, Ann. and Mag. Nat. Hist., ser. 2, vol. 7, no. 45, p. 36LThe family Bufonidae is here retained for the genus Bwfo, whichcomprises numerous species in the Old and New Worlds. The major-ity of the species are readily recognized, and a few of them have anextensive range in their respective regions. Adults of the genus Bujolack maxillary teeth. There is some evidence, however, that Bujo isa derivative of a toothed ancestor. The presence of a tooth ridgeand a tooth furrow in the upper jaw of a Bv^o tadpole is regarded byOeder ^ as conclusive evidence that the ancestors of this genus hadteeth. Doctor Noble ^ has examined the morphological details of theseveral families of living Salientia andhas concluded that the bufonids areinseparable from the leptodactylids.The accumulating evidence seemsto indicate that a satisfactory schemeof classification can not be basedsolely on dentition characters. Copeattempted to outline a natural classi-fication based largely on the pectoral ^ ? ^. . , , . ,? "^ '^ FiGUEE 3.?Diagrammatic ventral view ofgirdle, Nicholls employed characters shoulder glrdle of Bufo amerkanus; arciferalfurnished by the vertebral column, 'I ''?''"';, '^^' T?T ^^^"''' '^ S'?^ ' shoulder girdle, or pectoral arch, are capablewhile more recently Noble has of movement with contraction and expan-stressed the importance of the thigh ?'?? ^^ '^^ *^?'-'^^; '^^ epicoracoid cartaages^ overlapmusculature. As might be expected ^l., clavicle; Co.. cora<;oid; Ep., epicora-there are exceptions to all these cold; Mes., sternum or mesostemum; Sc,1 PI -r* i- rni- j. ii. scapula; X., xiphisternumschemes 01 classification, i he tooth-less toads of the genus Bufo have long been recognized as a naturalgroup, and while the known morphological evidence may not warrantthe retention of the family Bufonidae solely for this genus, someallowance must be made for the fact that relatively few species havebeen critically studied by those interested in classification schemes.Genus BUFO Laurent!1768. Bufo Latjrenti, Synopsin reptilium, p. 25. [Genotj'pe, Bufo vulgarisLaurenti, idem, p. 28= [Rana] bufo Linnaeus, 1758, Systema naturae,ed. 10, vol. 1, p. 210 (Europe).]1824. Oxyrhynchus Spix, Animalia nova, sive species novae testudinum et ranar-um quas in itinere per Brasiliam, etc., p. 49. [Included species: Bufo naricus,p. 49, pi. 14, fig. 2; B. nasutus, p. 50, pi. 14, fig. 3; B. semilineatus, p. 51,pi. 21, fig. 1; B. granulosus, p. 51, pi. 21, fig. 2; B. aculirosiris, p. 52, pi. 21,fig. 3; B. proboscideus, p. 52, pi. 21, fig. 4 (all from Brazil).]
? Oeder, R., Die Zahnleiste der Krote. Zool. Anz., vol. 29, p. 538, 1905.
? Noble, Q. K., The phylogeny of the Salientia. Bull. Amer. Mas. Nat. Hist., vol. 46, art. 1, p. 20, 1922.
28 BULLETIN 160, UNITED STATES NATIONAL MUSEUM1828. Chaunus Wagler, Isis von Oken, vol. 21, p. 744. [Genotype, Chaunusmarmoratus Wagler, idem, p. 744 = Bu/o granulosus Spix, op. cil., p. 51,pi. 21, fig. 2 (habitat in Provincia Bahia, Brazil).]1838. Osilophus Tschudi, M6m. Soc. Sci. Nat. cie Neuchatel, vol. 2, pp. 52, 89.[Genotype, Osilophus typhonius CvviEn= [Rana] typhonia Linnaeus, 1758,Systema naturae, ed. 10, vol. 1, p. 211 (America).]1843. Phrynoidis Fitzinger, Systema reptillum, fasc. 1, p. 32. [Genotype,Bujo asper KvHh= B[ufo] asper Gravenhorst, 1829, Deliciae musei zoologiciVratislavieusis, fasc. 1, p. 58 (Java).]1843. Phrynomorphus Fitzinger, Systema reptilium, fasc. 1, p. 32. [Genotype,Bufo leschenaultii Dumeril and BiBRO]sr= Bu/o leschenaultii Tschudi, 1838,M6m. Soc. Sci. Nat. de Neuchatel, vol. 2, p. 89 (India?) = [i5?/o] guttatusSchneider, 1799, Historiae amphibiorum naturalis et literariae, fasc. 1,p. 218 (India Orientali) .] (Preoccupied by Phrynomorphus Curtis, 1833,Hemiptera.)1843. Leptophryne Fitzinger, Systema reptilium, fasc. 1, p. 32. [Genotype,Bufo cruentatus? Bufo cruentatus Tschudi, 1838, Mem. Soc. Sci. Nat. deNeuchatel, vol. 2, pp. 52, 88 (India Orientali).]1843. Docidophryne Fitzinger, Systema reptilium, fasc. 1. p. 32. [Genotype,Bufo agua Daudin, 1803, Histoire naturelle des rainettes, des grenouilles etdes crapauds, p. 64, pi. 37 (Brazil) ==[flana] marina Linnaeus, 1758, Systemanaturae, ed. 10, vol. 1, p. 211 (America).]1843. PeUophryne Fitzinger, Systema reptilium, fasc. 1, p. 32. [Genotype,Bufo peliocephalus Dumeril and Bibron= S?/o peUocephalus Tschudi, 1838,M6m. Soc. Sci. Nat. de Neuchatel, vol. 2, pp. 52, 89 (Cuba).]1843. Otolophus Fitzinger, Systema reptilium, fasc. 1, p. 32. [Genotype, Bufomargaritifer DAVDm= Rana margarilifera Laurenti, 1768, Synopsin rep-tilium, p. ZO= [Rana] typhonia Linnaeus, 1758, Systema naturae, ed. 10,vol. 1, p. 211 (America).]1843. Eurhina Fitzinger, Systema reptilium, fasc. 1, p. 32. [Genotype, Bufoproboscideus Wagler=jBw/o proboscideus Spix, 1824, Animalia nova, sivespecies novae testudinum et ranarum quas in itinere per Brasiliam, etc., p. 52,pi. 21, fig. 4 (habitat ad flumens Solimoens, Brazil) = Bufo typhonius (Lin-naeus).]1843. Chilophryne Fitzinger, Systema reptilium, fasc. 1, p. 32. [Genotype, Bufod'orbignyi Dumeril and Bibron, 1841, Erpetologie g6nlrale, vol. 8, p. 697.(Montevideo, Uruguay).]1843. Phryne Fitzinger, Systema reptilium, fasc. 1, p. 32. [Genotype, Bufovulgaris Laurenti, 1768, Synopsin reptilium, p. 28= Rana bufo Linnaeus,1758, Systema naturae, ed. 10, vol. 1, p. 210 (Europe) = 5m/o bufo (Lin-naeus).]1845. Anaxyrus Tschudi, Untersuchungen iiber die Fauna Peruana, Herpetologie,p. 78. [Genotype, Anaxyrus melanchoHcus Tschudi, idem, p. 78, pi. 11,fig. 5= Bufo compadilis Wiegmann, 1833, Isis Von Oken, vol. 26, pt. 7,p. 661 (Mexico).]1849. Schmismaderma Smith, Illustrations of the zoology of South Africa,appendix, p. 28. [Genotype, Schismaderma lateralis SMn:B.= Bufo carensSmith, 1848. idem, pi. 68, fig. 1.]1860. Adenomus Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 12, p. 371, Sept.[Genotype, Adenomus badiofiavus Cope (Ceylon) = jBu/o kelaartii Gunther,1858, Catalogue of the Batrachia Salientia in the collection of the BritishMuseum, p. 140, pi. 10, fig. A (Ceylon).]1862. Rhaebo Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 14, p. 358, Sept.[Substitute name for Phrynomorphus Fitzinger, 1843.]
MEXICAN TAILLESS AMPHIBIANS 291863. Incilius Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 15, p. 50, Mar.[Genotype, Chilophryne conifera Cope= Bw/o coniferus Cope, 1862, Proc.Acad. Nat. Sci. Philadelphia, vol. 14, p. 158, Mar.] (See also Cope, 1865,Nat. Hist. Rev., p. 102.)1865. Epidalea Cope, Nat. Hist. Rev., p. 102, Jan. [Genotype, Epidalea calamitaCoFE= Bufo calamita Laurenti, 1768, Synopsin reptilium, pp. 27, 119(Europe) .]1868. Otaspis Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 20, p. 312, Nov.[Genotype, Otaspis empusa Cove= Peltaphryne empusa Cope, 1862, Proc.Acad. Nat. Sci. Philadelphia, vol. 14, p. 344, Sept. (Cuba).]1870. Nannophryne GtJNTHER, Proc. Zool. Soc. London, p. 402, June 9. [Geno-type, Nannophryne variegata GtJNTHER, idem, p. 402, pi. 30, fig. 2 (PuertoBueno, Port Grappler, and Eden Harbour, in Territorio Magallanes, westcoast of South America).]1870. Ansonia Stoliczka, Journ. Asiatic Soc. Bengal, vol. 39, pt. 2 (Nat. Hist.),p. 152, June 7. [Genotype, Ansonia penangensis Stoliczka, idem, p. 152,pi. 9, fig. 4 (Penang Island, Straits Settlements).]1876. Cranopsis Cope, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8, pt. 2,p. 96. [Genotype, Cranopsis fastidiosus Cope, idem, p. 96, pi. 23, figs. 1,la (2,500 feet elevation on slope of Pico Blanco, district of Uren, Costa Rica) .]1876. Crepidius Cope, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8, pt. 2,p. 97. [Genotj^pe, Crepidius epioticus Cope, idem, p. 97, pi. 23, figs. 2, 2a(5,000 feet elevation on Pico Blanco, Costa Rica).]1876. Ollolis Cope, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8, pt. 2,p. 98. [Genotype, Ollotis coerulescens Cope, idem, p. 98, pi. 23, figs. 3, 3a(3,000 to 5,000 feet elevation on Pico Blanco, Costa Rica).]1879. Dromoplectrus Camerano, Atti R. Accad. Sci. Torino, vol. 14, p. 882.[Genotype, Dromoplectrus anomalus Camerano= Bu/o anomalus GtJNTHER,1858, Catalogue of the Batrachia Salientia in the collection of the BritishMuseum, p. 57 {Mexico) = Bufo compactilis Wiegmann, 1833, Isis von Oken,vol. 26, pt. 7, p. 661 (Mexico).]Tliirteen species of Bufo are recognized in Mexico. One, B.alvarius, is semiaqaatic and has a rather smooth leathery skin, butmost of the species are mainly terrestrial except during the breedingseason. Three others, B. compactilis, B. cognatus, and B. woodhousii,have a large shovellilie inner metatarsal tubercle, which facilitatesburrowing. The extent to which cranial crests are developed con-stitutes an important diagnostic character for many species. Cranialcrests are quite prominent in some, especially in B. valliceps, whichhas high sharp-edged bony ridges. Conversely, there are others,particularly B. punctatus, that have a nearly flat interorbital regionand normally are devoid of such crests. Abnormal development orpeculiarities in the conformation of the cranial crests are occasionallyobserved in a large series of any species of Bvfo. The Mexican toadB. cristatus may be an instance of this sort, for, with the exception ofthe peculiarly enlarged parietal crests, it conforms in most respectsto the valliceps type of toad. The nondevelopment of certain cranialcrests when associated with an exceptionally tubercular skin, suchas has been hereinafter described in the case of certain individuals66785?32 3
30 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
of B. compactilis and B. simus, often materially changes the generalappearance of the species. Nieden, in his key to the genus Bufo,often uses the occurrence of single or double subarticular tubercleson the toes as a diagnostic character. In many instances the numberof subarticular tubercles present at each joint varies from one to twoon the same digit, and 3^et the number of such tubercles will be con-stant in the majority of the specimens in any large series.Although most kinds of toads are squat, fat, and warty, there aresome smooth-skinned species, one of wliich {Bvfo alvarius) inhabitsthe arid sections of Southwestern United States and northwesternMexico. There is also a slender-bodied tropical toad {B. haematiticus)that reminds one of a frog. Of the many kinds of toads living to-dayin different parts of the world, both the largest and the smallestbelong to the American fauna. The largest, B. marinus, ranges fromnorthern Mexico southward to Patagonia, and within the UnitedStates and northwestern Mexico is a form nearly as large, which islocally known as the Colorado River toad {B. alvarius). Largeindividuals of B. marinus mfij have a body length of 9 inches and amouth large enough to swallow small birds, but insects constitute themajor portion of their food. Probably the smallest living toad is theoak toad, B. quercicus, of the Southeastern United States; this speciesfeeds largely on ants. The little green toad, B. debilis, is the smallestMexican member of the genus.Most toads have the skin covered with warts, which are moreclosely aggregated on the sides of the neck than elsewhere, and these,together with a pair of large glandular masses, the parotoid glands,situated behind the eyes, secrete a milky poisonous fluid wheneverthe animal is molested. This secretion is an acrid irritant, causingpain in cuts and producing a bitter astringent sensation in the mouth.Some investigators have advanced the idea that the female lacks thisfluid during the breeding season, while the male at the same timehas an overabundant supply of it.According to Madame Phisalix, toads possess two distinct kinds ofglands, which secrete venom of different properties, one type pro-ducing a mucous and the other a granular venom. While the mucousglands are distributed over the entire body, they are principallyfound on the abdomen. The mucous secretion produced is an alka-loid that acts as a narcotic, its effect being confined to the nervecenters. Glands situated on the back, together with the parotoidglands on the shoulders and connected with them, secrete a granularacid venom that acts as a convulsive.It is not unusual to find hundreds of toads belonging to the samespecies congregated in a small pond, temporary pool, or shallowstream during the breeding season. Some species are far less numer-ous, and a dozen pairs may comprise the breeding population for
MEXICAN TAILLESS AMPHIBIANS 31some small temporary pool. Under normal conditions, if mated inthe water, the female begins laying at once, and according to speciesmay lay from 600 to 16,000 eggs, the process usually being completedwithin one day unless there should be a sudden drop in the tempera-ture. The eggs are generally laid in long spiral strings of jelly, whichare attached to or which come in contact with some submerged object.One species, B. pundatus, does not deposit the eggs in strings, butsingly or in a scattered mass. The inner tube of the egg string isabsent in B. compactilis but present in B. valliceps. The length ofthe hatching period depends upon the temperature. Below 65? F. itrequires 8 to 12 days, and above that temperature 3 to 8 days. Warmweather speeds up the rate of development and shortens the totaltime in the egg and tadpole stages. Toads are thus born in the waterand in it spend their early life in a larval, fishlike state, breatliing bymeans of gills. The transformation of the tadpole to a young toadtakes place some time between 30 and 65 days after hatching, andimder abnormal conditions 200 days have been required.KET TO MEXICAN SPECIES OF BUFO
1. Elongated gland extending full length of ulna; oval gland onupper surface of thigh and an interrupted gland on superiorsurface of tibia; parotoid gland long, oval, and nearly aslarge as side of head, descending on shoulder; tympanumlarge, two-thirds the diameter of orbit; supraciliary cranialcrest forming regular curve with more elevated postorbitalcrest; a narrow preorbital crest; one or more prominentwarts posterior to rictus oris; first finger longer than second;smooth leathery skin, with scattered warts; grayish orbrownish green above; underparts light; size large, head-and-body length, 125 to 160 mm alvarius (p. 35)No elongated glands on ulna and tibia 22. Parotoid gland subtrlangular in outline and nearly as large as orlarger than side of the head 3Parotoid gland smaller than side of the head 43. Parotoid glands divergent, very large, each extending backwardto a point beyond shoulders, and covered with small tubercles;a small indistinctly margined tympanum almost contiguous toorbit; short-legged, small-bodied, with rough granular ortubercular skin on upperparts and top of head; supraciliaryand postorbital crests indistinct; greenish or bluish slateabove, with small blackish spots; eyelids with oblique cross-bar; tarsal fold with row of warts; metatarsal tuberclesinsignificant; head-and-body length, 40 to 75 mm debilis (p. 50)Parotoid glands divergent, unusually large, and studded withlarge pores; a distinctly margined tympanum, about one-halfwidth of eye; skin of upperparts warty, interorbital regiongenerally smooth; prominent cranial crests, includingcanthal, preorbital, supraciliary, parietal, postorbital, orbito-tympanic, and supratympanic; interorbital space wider thanupper eyelid; coloration of upperparts variable, ranging
32 BULLETIN 160, UNITED STATES NATIONAL MUSEUMthrough various shades of brown, including yellowish,Beddish, or even blackish, and sometimes greenish olive;upperparts with or without large black insuliform spots,which when present are usually edged with pale yeUow; a lightvertebral line occasionally visible; arms and legs of im-mature individuals usually banded with dark brown; under-parts dingy white or yellow; first finger longer than second;toes half or two-thirds webbed; double subarticular tubercleson distal phalanges of fourth toe; a distinct tarsal fold;head-and-body length, 115 to 185 mm marinus (p. 53)4. Inner metatarsal tubercle large and shovellike, with sharp,horny, and usually black edge 5Inner metatarsal tubercle neither large nor shovellike nor withsharp cutting edge 75. Coloration of upperparts leopardlike, with conspicuous blotches,which vary from green to brown, margined with white or paleyellow; a bony elevation or boss between and considerablyabove level of nostrils, from which thick, blunt-edged cranialcrests diverge backward; postorbital crest thick; interorbitalregion narrower than upper eyelid; reniform parotoid glandalmost in contact anteriorly with postorbital crest; firstfinger longer than second; toes one-half webbed; doublesubarticular tubercles on distal phalanges of fourth toe; athick tarsal fold or ridge; head-and-body length, 65 to95mm cognatus (p. 41)Coloration of upperparts not leopardlike 66. A light vertebral streak extending from transverse interorbitalband to vent; upperparts generally grayish or dull yellowishbrown, conspicuously blotched with a darker color or withscattered spots; limbs obscurely barred; underparts light yel-lowish, with or without black markings on breast; largestwarts red tipped and encircled at base with narrow black bor-der; cranial crests distinct, but not high; canthus rostralisgenerally distinct; interorbital space either concave or convex,depending upon approximation of supraciliary crests, whichmay either parallel one another or diverge slightly posteriorly;parietal crests generally lacking, but supraciliary crests maybe thickened at their posterior extremities; transverse post-orbital crest forms right angle with supraciliary crest; a well-developed preorbital, a less distinct orbito-tympanal, but nosupratympanic crest; parotoid glands in contact with post-orbital crests, about twice as long as broad, their maximumdiameter being longer than length of orbit; tympanum oval,higher than wide, and equal to half or more of length of orbit;toes one-third webbed; inner metatarsal tubercle large, withfree cutting edge; subarticular tubercles on distal phalangesof fourth toe single or double; tarsal fold indistinct orabsent; head-and-body length, 80 to 115 mm woodhousii (p. 72)No light vertebral streak; coloration of upperparts generallylight greenish gray or brown, irregularly spotted with darkeroolor; interorbital Ught band occasionally present; top ofxiead without or with poorl}' defined cranial crests; paro-toid glands reniform in outline, in contact anteriorly withpostorbital crest or upper eyelid; tympanum generally morethan half the diameter of orbit, rarely indistinct and par-
MEXICAN TAILLESS AMPHIBIANS 33
tially concealed by tubercles; first finger longer than second;subarticular tubercles on distal phalanges of fourth toe gen-erally single, occasionally double; a tarsal fold; head-and-body length, 60 to 87 mm compactilis (p. 44)7. Head broad, flat or shallowly concave, and thin through, with-out raised cranial crests, or with only vestiges of such; snoutshort, blunt; parotoid glands elevated, rounded or broaderthan long, and studded with minute tubercles; skin ofupperparts with closely set small tubercles; colorationranging from light reddish brown to gray or even malachitegreen; warts on upperparts tipped with red, orange, or yel-low and encircled or partially encircled at base by narrowblack border; obscure black markings occasionally present onlimbs; underparts uniform white or spotted with darker color;diameter of oval tympanum more than half that of orbit;first finger longer than second; subarticular tubercles on dis-tal phalanges of fourth toe single; a tarsal fold bearing rowof small tubercles; head-and-body length, 55 to 80 mm. punctatus (p. 60)Head narrow or broad, but with distinctly raised cranial crests 88. Parietal crests unusually swollen, merging with thickened post-orbital crests; supraciliary crests high, converging anteriorly;supratympanic crest short and connecting with large swollensubtriangular or ovoidal parotoid gland; interorbital regionconcave; snout pointed, prominent; tympanum small, lessthan one-half diameter of orbit and often rather indistinct;skin of upperparts and upper surfaces of fore and hindlimbs with scattered small conical warts; underparts stronglygranular; upperparts (in alcohol) sooty brown with a fewindistinct markings of some lighter color; throat and chestdusky or blackish; belly with irregular blotches of yellow;under surfaces of fore and hind limbs irregularly markedwith yellow and some dusky color; first and second fingerssubequal; toes about two-thirds webbed, the penultimate andlast phalanx of fourth toe free; metatarsal tubercles of mod-erate size; subarticular tubercles on distal phalanges of fourthtoe single or double; bicolored parotoid glands, black later-ally and grayish superiorly; head-and-body length, 48 to75 mm cristatus (p. 48)Parietal crests when present not unusually swollen or expandedlaterally [parietal crests of Bufo valliceps considerably swol-len in rare instances] 99. Parotoid glands elongate, in contact anteriorly either withpostorbital crest or with upper eyelid; no supratympaniccrest 10Parotoid glands not in contact either with upper eyelid or withpostorbital crest, but separated by an interval of varyingwidth; parietal crests generally present, occasionally ves-tigial; supratympanic crests present 1110. Parietal crests slender, short, and converging posteriorly;supraciliary crests convergent anteriorly; interorbital spacewider than upper eyelid; postorbital crests very short, ifpresent, and not extending more than halfway across to-ward tympanum; parotoid glands elongated, as large as orlarger than upper eyelid; tympanum cither distinctly out-lined or largely concealed by tubercular skin and closely set
34 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
conical warts; general coloration of upperparts grayish,greenish, or brownish, marbled with a darker shade of brown;an interrupted irregular light vertebral stripe and an irregu-lar lateral light band formed by anastomosing spots gener-ally present; a transverse ashy white or yellowish band ex-tending across upper eyelids and anterior interorbital region;underparts immaculate or spotted with some darker color;skin of upperparts with scattered small conical warts ordecidedly tubercular; first finger longer than second; toeshalf webbed; a tarsal fold with linear series of small conicalwarts; subarticular tubercles on distal phalanges of fourthtoe generally double; head-and-body length, 65 to 85 mm. simus (p. 63)No parietal crests present; low curving supraciliary crestscontinuous with short postorbital crests; parotoid glandselongate, larger than upper eyelids, and contiguous anteriorlywith postorbital crests; tympanum distinct, one-half thediameter of orbit; skin of upperparts rough, with numeroussmall warts; underparts coarsely granular; coloration ofupperparts light brown, with light vertebral stripe fromsnout to vent, on each side of which are large blackish brownangular spots; posterior interorbital region with inverteddark chevron, which ends on upper eyelids; a dark bandextending from orbit to thigh below light longitudinal band;underparts immaculate or grajdsh white; first finger longerthan second; toes half webbed; no tarsal fold; subarticulartubercles on distal phalanges of fourth toe single; metatarsaltubercles small; head-and-body length, 40 to 52 mm. canaliferus (p. 40)11. Cranial crests high, sharp edged, conspicuous; superior marginof upper jaw generally protruding 12Cranial crests less conspicuous; parietal crests often indistinctor absent; combined supraciliary and postorbital crests forma continuous curve from anterior margin of orbit to supra-tympanic crest; interorbital region shallowly concave, notwider than upper eyelid; preorbital crest indistinct or vestigial;supratympanic crest short and thick; snout short, bluntlytruncated, with angular canthus rostralis; tympanum dis-tinct, transverse diameter about half width of orbit; parotoidglands subtriangular, smaller or no larger than upper eyelid;skin of upperparts thickly set Avith small warts; a lateralrow of light-colored warts, but lateral fold is indistinct;coloration of upperparts brownish gray, marbled more orless distinctly with some darker color; whitish vertebralstripe of varying width, on each side of which are three largeand a greater number of small irregular light-margined darkblotches; whitish markings above and dark-brown markingsbelow lateral rov/ of warts form more or less confluent longi-tudinal bands of varying width; a continuous or mediallyinterrupted chevron band crosses upper eyelids and inter-orbital region; fore and hind limbs with dark crossbarsabove; underparts immaculate; first finger longer than sec-ond; a row of small closely spaced warts along tarsal fold;toes less than half webbed; subarticular tubercles on distalphalanges of fourth toe single; inner metatarsal tuberclesmall; head-and-body length, 55 to 75 mm marmoreus (p. 58)
MEXICAN TAILLESS AMPHIBIANS 35
12. Parotoid glands ovoidal in outline, descending on shoulders;no linear series of enlarged warts on side of body; first andsecond fingers subequal; subarticular tubercles on fourth toedouble; toes one-third webbed; no tarsal fold; two smallmetatarsal tubercles; skin of upperparts rough, tubercular;underparts coarsely granular; coloration of upperparts brownor brownish black with blotches of darker color on each side ofvertebral line; yellowish or whitish lateral stripe from paro-toid gland to thigh, below which is a broad dark-coloredlateral band; underparts usually yellowish, rarely spotted witha darker color; interorbital region and upper eyelids withanterior and posterior transverse medially interrupted darkbands; cranial crests high; canthal, preorbital, supraciliary,postorbital, and supratympanic crests distinct in adults;parietal crests vestigial or absent; tympanum smaU butdistinct, its diameter slightly more tlian half that of eye;head-and-body length, 63 to 71 mm coccifer (p. 41)Parotoid glands subtriangular in outline, bicolored, and as largeas or larger than upper eyelid; a linear series of large conicalwarts on side of body; first finger longer than second; subar-ticular tubercles on fourth toe generally single; toes one-third webbed; no tarsal fold; two small metatarsal tubercles;skin of upperparts rough, tubercular, but somewhat smootherin southern specimens; warts often spinose; underparts ?coarsely granular; coloration of upperparts quite variable,at one extreme yellowish or light grayish with distinctmarkings, and at the other extreme with markings subduedor obliterated by a dark slaty or blackish suffusion; prevailingcolor generally some shade of brown, with linear series ofconical warts extending along lateral line from parotoidglands to thighs, above which is a light-colored stripe andbelow which a dark-colored one; near level of middle ofparotoid glands an inverted black chevron bisects light ver-tebral line; back irregularly marbled or blotched with somedarker color; limbs cross banded; underparts unicolored orheavily marbled with black; a posterior interorbital trans-verse black band; cranial crests relatively high; canthalridges distinct; supraciliary crests high, diverging posteri-orly; parietal crests lower than the latter and convergingposteriorly; the thick transverse postorbital crest forms anobtuse angle with short supratympanic crest; distinct orbito-tympanal and preorbital crests; interorbital space deeplyconcave and much broader than upper eyelid; tympanumdistinctly outlined, ovoidal in outline, and equal in width toslightly more than half length of orbit; head-and-bodylength, 60 to 100 mm valliceps (p. 68)BUFO ALVARIUS GlrardPlate 1, frontispiece; Figure 41859. Bufo alvarius Girard, in Baird, Report on the United States and MexicanBoundary Survey, vol. 2, Reptiles, p. 26, pi. 41, figs. 1-6.
?
Cope, 1889, U. S.Nat. Mus. Bull. 34, p. 265, fig. 62.
?
Dickerson, 1906, The frog book, pp.106-108, figs. 107-111, col. pi. 5, fig. 4.?Nieden, 1923, Das Tierreich, Lief.46, Amphibia, Anura I, pp. 75, 129, fig. 182.
?
Storer, 1925, Univ. CaliforniaPubl. Zool., vol. 27, pp. 163-167, pi. 12, fig. 32.
36 BULLETIN 160, UNITED STATES NATIONAL MUSEUMType locality.?Valley of the Gila and Colorado Rivers.Range.?Vicinity of streams with permanent flow of water fromGuadalupe Canyon in northeastern Sonora north to Turkey Creeknear Prescott in central Arizona, and in the lower Colorado RiverValley from Fort Mohave to Fort Yuma; also occurs in ImperialCounty, Calif. Exact limits of range unknown. Reported fromGuaymas (Mocquard) and Cajon BonitoCreek (Meams), Sonora.Remarks.?This large toad frequentsthe hottest portion of the North Amer-ican Continent, and, although it hasnever been found in the strictly desertareas, it seems to be extending its rangewith the advent of irrigation. It hasbeen described as semiaquatic and, ex-cept in irrigated areas, is seldom ob-served during the dry season. The eggsare laid during the first heavy summershowers in temporary pools of water orin shallow streams.Exceptionally large individuals maybe rather uncommon, and the maximumsize attained by this species can notbe ascertained by the limited materialavailable. One specimen (U.S.N.M.No. 66734) measures 160 mm. in head-and-body length. The distinctive char-acters of this species are a smoothleathery skin, large size, and greenishcolor. Unlike other Mexican toads, it has a large elongate gland onthe forearm and several smaller ones on the hind limb.Major Mearns,^ in describing his experiences with this large toad,states
:
Of batrachians, a toad (Bufo) and a frog {Rana virescens hrachycephala Cope)were found at Warsaw Mills; and at Buenos Ayres, at the beginning of the sum-mer rains, Lieutenant Gaillard observed great numbers of a very large frog-liketoad, named Bufo alvarius by Girard. Nothing was seen or heard of them untilthe advent of the early summer rains, which formed a large shallow lake nearBuenos Ayres and about 10 kilometers (6 miles) north of the Boundary Line.These large toads then filled the air with their loud cries, which increased untila deafening roar was produced. Numbers of them were seen hopping about, buttheir rarity was not suspected by Lieutenant Gaillard, on which account nonewere collected. The range of Bufo alvarius Girard, extends from Monument
Figure 4.?Head of Bufo alp art us(U.S.N.M. No. 21063) from GuadalupeCanj'on, Sonora, Mexico: a, Lateralview; 6, dorsal view
' Meams, E. A., Mammals of the Mexican boundary of the United States. U. S. Nat. Mus. Bull. 56,pp. 113, 114, 1907.
MEXICAN TAILLESS AMPHIBIANS 37No. 73, in Guadalupe Canyon, to the Colorado River (Monument No. 205). Ifirst met with it in a dense growth of arrowwood {Pluchea sericea) on the edge ofthe Colorado River at Fort Mojave, Arizona, May 13, 1884. A specimen wascarried to Peach Springs and given into the charge of the hotel proprietor, whoallowed it to escape during my absence at the Grand Canj-on of the Colorado.No specimen of this toad was again seen until July 6, 1892, when LieutenantsGaillard and Irwin and the writer lay down to rest upon the damp grass besidethe San Bernadino Springs, near Monument No. 77. At dusk these huge greenbatrachians began to hop about us, occasionally lauding upon our faces. A fewwere caught and saved as specimens. No more were seen by me until October 3,1893, when Hospital Steward E. C. Merton brought me another that he had justcaught in a spring situated between Monument No. 73 and Cajon Bonito Creek,in Sonora, Mexico. Another was taken at Quitobaquita Springs, MonumentNo. 172, January 26, 1894.The potency of the toxins secreted by this large toad during thephysical stress of danger and pain has been recorded as follows byMusgrave : ^Just about sundown of that eveiiing [September 1, 1928] I was looking over aplanting in the front of our home when I discovered a large green toad in a stand-pipe used for irrigation purposes. I lifted the big fellow out and dropped himover the side. Our little wire haired fox terrier, standing near by made a dashat the toad, but I spoke to her and she stepped back. Immediately the toadswelled himself up, hissed at the dog and hopped a little way toward her. Thatwas too much, the dog immediately grabbed him and in one shake the toad wasdead. I was leaning over the standpipe and my face was perhaps 4 or 5 feetfrom the toad while she was shaking it.About this time a large police dog that is a visitor at our home ran up andtouched his nose to the toad; the little terrier snatched it away. I thought nomore of it and started back to the house, the big police dog following. He hadgone no more than a hundred feet when his front legs crumpled under him andhe pitched forward. However, he gathered himself and then tipped backwards,his legs and body being paralyzed.Immediately I realized that something was wrong and looking over to wherethe little terrier had been I saw her lying on the ground with her feet crumpledunder her and her face in the dirt. I ran over and picked her up and foundthat she had fallen on top of the toad as she was carrying it. I felt her heartand found the action slow, and although she gasped and did her best, she couldget no air into her lungs. Within two or three minutes from the time she firstbit the toad she died. Immediately after death, bloody foam oozed from hermouth and nose.About that time I became very sick myself, my head was swimming, and therewas a lifting feeling in my lung cavity. It affected me rather peculiarly, as Iwanted to walk and keep walking. I took a large dose of warm salt water andafter disgorging what I had in my stomach I felt better. However, the effectsdid not wear off for about 30 minutes. The old police dog also revived in aboutthree quarters of an hour.I do not know whether I got the effects of the poison while leaning over thestandpipe or while working with the little dog, as I pried her mouth open andtried to get salt water down her. I did not detect any odor whatsoever. I amsure that I did not get the poison from the toad before the dog attacked it,
? Musgrave, M. E., Bufo alvarius, a poisonous toad. Copeia, no. 173, pp. 96-98, Jan. 16, 1930.
38 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
although I did lift it with my hand, but this I have done on many occasionsbefore without experiencing any trouble. I also handled with no ill efiFect atoad of the same species that M'as later sent to the Bureau of Biological Survey,Washington, D. C.Recently I have had a communication from Miss May Noble, who lives inPhoenix and who within the past few days has had a similar experience. HerPekinese dog seized one of these green toads but Miss Noble got it away fromthe dog before he had hurt it to any great extent. However, the dog soon becameparalyzed and Miss Noble called two veterinarians, one of whom seemed toknow nothing of the effects of that sort of poisoning, but the other said it wasnot uncommon in this valley. Miss Noble informs me that she used ammoniawith good results and that the dog recovered within an hour after it had gottenthe poison. This dog, however, did not puncture the skin of the toad, onlypicking it up.Although it is not so stated in the original description, Girardseems to have based his description of Bufo aharius on specimensreceived from two collectors. The locality is cited as the "Valley ofGila and Colorado," and the collector is stated to be "A. Schott."Arthur Schott was an assistant of Lieut. N. Michler, who was incharge of the party surv^ejdng the United States and Mexican bound-ary line from Fort Yuma, Calif., to Los Nogales, Sonora. Michler 'sparty arrived at Fort Yuma from San Diego, Calif., on December 9,1854, shortly after which Schott made surveys of the Gila andColorado Rivers for short distances above their junction. After theinitial point of the boundary line at the Colorado River was establishedon April 21, 1855, preparations were made for the survey of the obliqueline. On May 5, 1855, the Mexican and American parties departedfrom Fort Yuma and began their journey along the wagon road upthe Gila Valley to Tucson, Ariz. A camp was maintained nearTucson during most of June. It was not until June 26, 1855, thatMichler 's party met the United States Commissioner, Maj. W. H,Emory, and resumed the boundary survey near Los Nogales. Thetopography of the boundary line between Los Nogales and theColorado River was completed on August 16, 1855. This itineraryshows that Schott actually traversed the area designated as the typelocality. The next point to be determined is whether specimens ofthis toad were collected in this area by Schott and transmitted tothe Museum.On the lower margin of the original set of drawings of Bufo alvariusreproduced on the plate in the report on the reptiles of the UnitedStates and Mexican Boundary Survey, Baird has written "Sierra dela Union." There is only one entry in the catalogue of the divisionof reptiles that could possibly apply to this specimen. The originalentry reads as follows: "Cat. No. 2571; Bu^o alvarius; Sierra de laUnion y Charcos de la Nariz; 2 specimens; received from MajorEmory; collected by A. Schott." This range of mountains is locatedon the divide and not in the drainage of the Gila River, but it is quite
MEXICAN TAILLESS AMPHIBIANS 39probable that Girard may have assumed that this range was sosituated. It would seem that specimens collected along the routebetween Monument No. 14 on Sierra de la Union and the water holesof the Nariz Mountains were placed in one container and so labeled.The Sierra de la Union is shown on modern maps as Sierra de Moreno,and this range of mountains crosses the boundary line betweenArizona and Sonora about 10 miles west and south of San Miguel insouthern Pima County. The Sierra de la Nariz crosses the boundaryline about 35 miles distant in a northwest direction. No other toadof this species collected by Schott is entered in the museum catalogue.An examination of the catalogue of the division of reptiles revealsthat another specimen (U.S.N.M. No. 2572) was available to Girard atthe time the description was written. This was collected at old FortYuma, Calif., by Maj. George Henry Thomas, and is designated inthe museum catalogue as the type of Bufo alvarius. Old Fort Yumais located near the junction of the Gila and Colorado Rivers and thuslies in the "Valley of Gila and Colorado." The fact that MajorThomas had no official connection with any of the parties workingon the survey of the boundary line does not suffice to explain theomission of this particular specimen, for a bat received in the sameshipment with this toad is listed by Baird. The collection made byThomas was received at the Museum in 1856.It is quite certain from the notation made by Baird that the speci-men figured on the plate accompanying the original description ofBufo alvarius came either from some spot between Sierra de la Union(Sierra de Moreno) and Charcos de la Nariz, or else from one or theother of these localities. These mountain ranges are located on thedivide between the drainage of the Gila River in Arizona and thatof the Rio Concepcion in Sonora. It is equally certain that the speci-men which was collected by Major Thomas at Fort Yuma, Calif.,had been received by the museum at least three years prior to thepublication of Girard 's description.It is unfortunate that the type or cotypes of Bufo alvarius are notdesignated by the original describer. The usual procedure is toselect as the lectotype or as the cotypes the specimen or specimensthat were seen by the original describer before his description waspublished. In this instance one has to assume that all the materialavailable in the Museum from the region covered by the boundarysurvey report was placed at Girard's disposal. If one takes Girard's
"Valley of Gila and Colorado" at its face value, then the specimenstaken by Schott and labeled as coming from "Sierra de la Union yCharcos de la Nariz" must be disregarded, inasmuch as they werenot obtained at the type locality. On the other hand, if one insiststhat the type or cotypes were collected by Schott as stated by Girard,then the specimen collected by Major Thomas at Fort Yuma in the
40 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
valley of the Gila and Colorado must be ignored, because it was nottaken by Scliott. The selection of either of the above as the basisfor Girard's description might lead to conflicting opinions, and sinceneither of these specimens has ever been formally designated as thetype, the two specimens (U.S.N.M. No. 2571) collected by Schottand the one (U.S.N.M. No. 2572) collected by Major Thomas arehereby designated as the cotypes of Bujo alvarius. The specimenscollected by Schott can not now be located. The one received fromThomas is quite well preserved and conforms in most details with theactual figures published by Girard.Specimens examined.?One (U.S.N.M. No. 21063), collected byE. C. Merton, October 3, 1893, in Guadalupe Canyon, Sonora, inbed of dry wash about 3 miles southwest of Monument No. 73.BUFO CANAUFERUS CopeFigure 51877. Bufo canaliferus Cope, Proc. Amer. Philos. Soc,vol. 17, uo. 100, p. 85, July 20.?Brocchi, 1882,Mission scientifique au Mexique et dans TAm^riqueCentrale, recherches zoologiques, pt. 3, sect. 2, p. 74,pi. 8, figs. 2, 2a-b.?NiEDEN, 1923, Das Tierreich,Lief. 46, Amphibia, Anura I, pp. 76, 131.Type locality.?Tehuantepec, State of Oaxaca,Mexico.Range.?Tliis species has been collected atlocalities in the States of Vera Cruz, Oaxaca,and Chiapas, the most northerly record beingOrizaba. It is reported from Tabasco (Dug6s)and from Cerro de Valtierra (Giinther) inChiapas. The exact limits of its range are unknown. The mostsoutherly record is Patulul in Guatemala.Remarks.?This is a small light-colored toad, with a whitish verte-bral stripe, on each side of which are dark-colored spots. It has anarrow head, low cranial crests, distinct tympanum, and elongateparotoid glands. The name chosen for this toad by Cope is in allu-sion to the once contemplated canal on the Isthmus of Tehuantepec.It seems to be a rare toad in American collections. There are nopublished observations on the habits and life history of this species.The cotypes of this species (U.S.N.M. Nos. 30315-24) were collectedby Francis Sumichrast. Notwithstanding their long immersion in apreserving fluid, most of them have a fairly well-preserved colorpattern and are otherwise in very good condition.Specimens examined.?Seventeen as follows:
Figure 5.?Head of Bufocanaliferus (U. S.N.M. No.30322) from Tehuantepec,Oaxaca, Mexico: a. Lateralview; b, dorsal view. Typespecimon
MEXICAN TAILLESS AMPHIBIANS 41Bufo canaliferus
Museum
42 BULLETIN 160, UNITED STATES NATIONAL MUSEUMType locality.?Arkansas River, Prowers County, Colo.Range.?In Mexico the range of tliis toad extends southward fromthe northern boundary to San Pedro in the State of Nayarit and tothe mountains of Alvarez in the State of San Luis Potosi. Reportedby Perez from Villa Lerdo, Durango. North of the Mexican bound-ary, the plains toad has been recorded from a wide extent of territory.It has been found in the Upper Missouri River drainage (ValleyCounty) and in southeastern Montana (Yellowstone, Rosebud, andCuster Counties); in north-central(McHenry County) and eastern(Jamestown, Stutsman County)North Dakota; the Missouri FloodPlain area of central South Dakota(Fort Pierre); south through west-ern South Dakota (Ardmore, FallRiver County); eastern Wyoming(Laramie County) ; central Nebraska(Fort Kearney) ; northeastern (Sedg-wick and Weld Counties) and south-central (Costilla County) Colorado;western Kansas (Trego, Gove, Wal-lace, and Morton Counties); north-eastern New Mexico (Koehler Junc-tion) ; central Oldahoma (Caddo andCleveland Counties); southwesternArkansas (Red River); the StakedPlains region from the northernboundary of Texas to Pecos River,the salt plains of El Paso County,and Jeff Davis County, Texas; central (Bernalillo County) and south-western (Grant County) New Mexico; southeastern Arizona (Cochise,Maricopa, and Pima Counties) and along the Colorado River at FortMohave; Salton Sea Basin (Mecca and Brawley) and along the Colo-rado River from Fort Yuma to Needles, Calif. ; and also in northernUtah (Utah and Emery Counties).Remarks.?In life this toad can be recognized instantly by theconspicuous greenish blotches on the upperparts. These irregularspots, as will be noticed on closer inspection, vary from green tobrown and are margined with white or pale yellow. The parotoidglands are short, broad, and widely separated; the bony crests on thetop of the head converge anteriorly and are united on the muzzle,forming a raised boss. The so-called leopard, or plains, toad frequents
Figure 7.?Head of Bufo cognatus (U.S.N.M.No. 4626) from Nebraska: a. Lateral view; 6,dorsal view
MEXICAN TAILLESS AMPHIBIANS 43the plains and the foothills as a general rule, though it may be foundoccasionally in the vicinity of streams and in bottomlands that aresubject to periodical overflow from some river. Although nocturnallike most toads, it leaves its place of hiding during heavy showers, andcomes forth in search of food. ^Say concludes his description of Bwfo cognatus with the followingstatement: "A specimen is placed in the Philadelphia Museum."This specimen is undoubtedly the type of this species and presumablyis the individual collected in July, 1820, in southeastern Colorado.The "Philadelphia Museum"^ was established by Charles WilsonPeale in 1784. The collections made by the expedition of MajorLong to the Rocky Mountains in 1819-1820 were deposited in thismuseum on March 23, 1821. Peale's museum was incorporated as astock company in 1820 and continued in existence until 1846, whenfinancial difficulties forced the sale of the collections at public auction.The natural-history collection, however, was kept intact and exhibitedin Masonic Hall, Philadelphia, until 1850, when Moses Kimballbought one-half for the "Boston Museum," and the other half wassold to P. T. Barnum's "American Museum" in New York City.Barnum's "American Museum" was destroyed by fire on July 13,1865. The bulk of the Boston Museum's share of the old natural-history collection of the Philadelphia Museum passed into the posses-sion of the Boston Society of Natural History in 1893 and the residuein 1899. After this collection reached the society's rooms, some of thespecimens were destroyed, but most of the birds were sold to C. J.Maynard in 1900. When the importance of this collection becamegenerally appreciated, the Boston Society of Natural History redeemedit from Maynard. Again, in 1914, the remaining relics of this collec-tion were transferred to the Museum of Comparative Zoology atHarvard College. The possibilities for the loss of the type of Bvfocognatus during the numerous transfers of the collections of the old
"Philadelphia Museum," or while it was in the possession of eitherBarnum's "American Museum" or Kimball's "Boston Museum," aretoo numerous to leave any serious hope for its future recovery insome unexpected depository.In 1876, Dr. F. V. Ha5^den, in charge of the United States Geologicaland Geographical Survey of the Territories, authorized an investiga-tion of the Judith River Lignite Formation. Prof. E. D. Cope wasplaced in charge of the expedition that fitted out at Fort Benton,Mont., for this exploratory survey. Among the published results ofthis expedition is the description by Cope in 1879 of a toad, which he
? Faxon, W., Bull. Mus. Comp. Zool., vol. 69, no. 3, pp. 119-148, July, 1915.
44 BULLETIN 160, UNITED STATES NATIONAL MUSEUMhad found "abundant on the plains north of the Missouri River eastof Fort Benton," as a new species, Bufo dipternus. The followingis quoted from the original description:There are two faint straight supraorbital ridges, and a postorbital but nosupratympanic ridge. The supraorbitals are united by the enlargement of theposterior part of the prefrontal bones, which forms quite a tuberosity in adults.In contradiction to this published description, Cope in 1889 statedthat a large young specimen of this species served as the type and that
"although an inch and a half long, this individual had not developeda trace of the cranial crests." Nothing is known in regard to thesubsequent history of the material upon which Cope based this name.Specimens examined.?Twenty, as follows
:
Bufo cognatus
Museum
MEXICAN TAILLESS AMPHIBIANS 45Type locality.?Mexico.Range.?From Mexican boundary south to Atemajac in Tabasco,to Tupataro in Michoacan, Xochimilco in Federal District, and toPotrero in Vera Cruz. Reported from Ciudad, Durango (Forrer).In Southern United States, disregarding a doubtful record for south-western Kansas, we may say that the range of this toad extends fromthe Red River Valley in southeastern Oldahoma (Frederick, TillmanCounty) south through the middle district of Texas (Upper WichitaRiver) west of the timber belt and east of the plains to McLennan,Burnet, Kendall, Austin, Colorado, Bexar, Goliad, Victoria, Refugio,Nueces, Starr, and Cameron Counties; northwest along the RioGrande drainage through Kinney, Val Verde, and Reeves Counties toEl Paso County, Tex.; northward in NewMexico along the Rio Grande to the Mim-bres Mountains and to Albuquerque; insoutheastern Arizona (Huachuca Moun-tains, Cochise County), as well as in Mari-copa, Yavapai, and Mohave Counties; andin extreme southwestern Utah (WashingtonCounty).Remarks.?The type of Wiegmann's Bufocompactilis (M. N. B. No. 3528) was col-lected in Mexico by two botanical collectors,Ferdinand Deppe and C. J. W. Schiede.The ground color of the upperparts of thetype is very light, although the mottling isdark; underparts white, with a few scattereddark spots on chest; vertical black bars on dupper lip quite distinct; interorbital band figurk s.?Read ot bu/o compactuisT 1 , T 1 ?!? J i u-i 1 (U.S.N.M. No. 2611) from Pes-hght colored; SUpraClliary and pOStorbltal quiena Grande, Nuevo Leon, Mex-CrestS distinct; a cluster of minute warts ico: a, Lateral view; 6, dorsal view.occupies the position of the parietal crest; otype specimenparotoid glands elongated and in contact with postorbital crests, not-withstanding the presence of a rather prominent knoblike supratym-panic crest; head relatively small; body wide and squat; skin ofupperparts warty, with numerous small tubercles; length of bodyapproximately 65 mm.Three specimens are designated in the National Museum catalogueas the COtypes of Girard's Bnfo speciosus. One of these (No. 2611)was collected by Lieut. D.N. Couch during April, 1853, at PesquieriaGrande, Nuevo Leon, Mexico, and was figured by Baird in his reporton the reptiles of the United States and Mexican Boundary Survey.The viscera of this specimen have been removed through an abdominalopening, and the lower jaws have been cut away from the head at the66785?32 i
46 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
angles of the mouth. It is an adult and is fairly well preserved. Thesecond cotype (U.S.N.M. No. 2608) was obtained in the vicinity ofRinggold Barracks [Rio Grande City, Starr County], Tex., by ArthurSchott, a surveyor attached to the United States and Mexican Bound-ary Commission under Maj. W. H. Emory. This is an adult femaleand may have been dissected by Cope. There is an incision on theabdominal wall, another across the right parotoid gland, and a thirdalong the under surface of the humerus. At some time the preserva-tive in which this cotype was kept may have been allowed to evapo-rate, for the skin is rather stiff. Capt. Stewart Van Vliet sent thethird cotype (U.S.N.M. No. 2610) to the Museum from Brownsville,Tex. It is quite young, and the head-and-body length is 27.5 mm.The black-margined inner metatarsal tubercle is well developed. Thebody is shrunken and somewhat distorted.The type of Giinther's Siz/o anomalus (B.M. No. 58. 9. 6. 12) is anadult male purchased from Emile Parzudaki, who was a dealer innatural-history specimens in Paris, and is labeled as coming fromMexico. The type does not have the cranial crests developed, theskin is rough and tubercular, and it is clearly an immature individual.The head-and-body length is 43.7 mm. The upperparts are lightgray with black spots, and the warts are dark at the base. It haslarge inner and small outer metatarsal tubercles.The type of Bufo levifrons could not be located in the Paris Museumat the time of my visit in April, 1930. Mexico is given as the typelocality for this species. A specimen from Mexico collected by Duges(M.H.N.P. No. 643b, parchment label No. 83-289) and labeled asthe type does not agree with the figure published by Brocchi. Thisspecimen has a head-and-body length of 53.5 mm. It is brieflydescribed as follows: The tibio-tarsal joint, when the hind limb iscarried forward along the body, reaches the posterior margin of theparotoid gland; skin of upperparts strongly tuberculate; cranial crestsnot developed ; tympanum covered with minute tubercles ; underpartslight colored with scattered dark spots; a large black inner and asmall light-colored outer metatarsal tubercle. The type of Bufomexicanus was received from Auguste Salle, according to Brocchi, butno entry of this specimen was found in the catalogue of the laboratoryof herpetology of the Paris Museum.This burrowing species has a rather compact body, and the skin isoften covered with closely set pustulose warts. As a rule it is irregu-larly spotted with dark, though there are occasional individuals thatexhibit little or no indication of darker spots. On many of the speci-mens there is a conspicuous white interorbital band that is almost ona line with the vertical white lip stripe below the eye. The underpartsare often white with small dark spots. On some of the specimens thecranial crests are lacking. From the variation observed in a series of
MEXICAN TAILLESS AMPHIBIANS 4742 individuals, it would seem that the development of the cranialcrests is too unreliable a character to have much of a diagnostic value.Fourteen of the specimens examined have no vestiges of cranial crestson top of the head. When present, the postorbital crests are generally-thick and blunt-edged. The supraciliary crests are often but faintly-indicated, though they may be quite prominent on some individuals.When supraciliary crests are developed, the interorbital region seemsmore strongly concave than those without such crests.In two instances the tympanum was covered with a tubercularskin, and on the others it was distinctly outlined and covered with asmooth skin. In the majority of the specimens examined, the paro-toid gland was separated from the upper eyelid by the blunt-edgedtransverse postorbital crest. On two individuals (U.S.N.M. Nos.47245 and 26596), however, the parotoid glands are almost in con-tact with the upper eyelids, and there are no vestiges of the postor-bital crests. One individual from McLennan County, Tex. (U.S.N.M.No. 57625) has subcircular parotoid glands in contrast to the normalelongate type. These glands are rarely very narrow, but they oftenhave a lateral enlargement that extends over upon the side of theshoulder.Specimens taken in Durango and Chihuahua occasionally have theunderparts spotted with dark brown or black. The skin of the upper-parts is strongly tubercular and is covered with low pustulose warts.Brocchi's Bufo levifrons represents a variation of this sort. Thefigure of Bufo mexicanus published by Brocchi resembles two toadscollected in Nuevo Leon (U.S.N.M. Nos. 46914-15) on which thewarts are low and flat, and the sldn feels smooth, as compared withthe variety named levifrons by Brocchi. Variations such as thesemay be expected in this species. This species has a conspicuouspalmar callosity and a large shovellike inner metatarsal tubercle.Specimens examined. ?Ninety-eight, as follows
:
Bufo compactilis
Museum CatalogueNo.
48 BULLETIN 160, UNITED STATES NATIONAL MUSEUMBujo compactilis?Continued
Museum
MEXICAN TAILLESS AMPHIBIANS 49directed black stripe from each parietal crest; underparts brownish,with light spots; head-and-body length, 63.5 mm.No. 3524: Parietal crests abnormally swollen, with internal pro-jections or buttresses, and separated posteriorly from parotoid glands;interval between supraciliary crests 7.8 mm.; parotoid glands pro-tuberant, somewhat elongate or ovoidal, and peculiarly marked withlight spots; parotoid glands measure 14 mm. in length and 9 mm. inwidth; subarticular tubercles distinct; numerous supernumerary tu-bercles on soles of fore and hind feet; a narrow blackinterorbital band; a light mark in middle of back;underparts brown, with numerous light spots; head-and-body length, 75 mm.A third specimen (N.M.W. No. 1869. I. 6) fromMexico was found in Vienna. This individual like-wise has very thick enlarged parietal crests, a heavysupratympanic crest, a thick supraciliary crest, andovoidal parotoid glands.The original specimen on which the descriptionof Bu^o occipitalis Camerano was based is thoughtto be in the Zoological Museum at Turin, Italy, andis said to be labeled as having come from "Mexico."The type specimen could not be located at the timeof my visit to this museum in April, 1930, In 1921,Dr. E. R. Dunn collected a fifth specimen of thisspecies (M.C.Z. No. 8362) at Jalapa in Vera Cruz.The characters used in the key for Mexican species of Biifo are basedon this specimen.The large swollen parietal branches of the supraciliary crests willat once distinguish this species from any other Mexican toad. Thesupraciliary crests are high and converge anteriorly. The postorbitalcrests are rather thick. The large protuberant ovoidal or subtriangularparotoid glaaids are often bicolored, black laterally and grayish above.Small conical warts are scattered over the skin of the upperparts andthe upper surfaces of the fore and hind limbs. The tympanum is small,less than one-half the diameter of the eye, and often is rather indistinctlyoutlined. This species seems to bemost closely related to Bufovalliceps.Specimens examined.?Four, as follows
:
Bufo cristalus
Figure 9.?Head of Bufocristatus (M.C.Z. No.8362) from Jalapa, VeraCruz, Mexico: a. Lat-eral view; 6, dorsal view
Museum
50 BULLETIN 160, UNITED STATES NATIONAL MUSEUMBUFO DEBILIS GirardFigure 101854. Bufo debilis Girard, Proc. Acad. Nat. Sci. Philadelphia, vol. 7, p. 87,May.1854. Btifo insidior Girard, Proc. Acad. Nat. Sci. Philadelphia, vol. 7, p. 88,May (Chihuahua, Mexico).
?
Baird, 1859, Report on the United States andMexican Boundary Survey, vol. 2, Reptiles, pp. 26, 35, pi. 41, figs. 13-18.1889. Bufo debilis Cope, U. S. Nat. Mus. Bull. 34, p. 264, fig. 61.
?
Dickerson,1906, The frog book, p. 112. Strecker, 1922, Sci. Soc. Sau Antonio Bull. 4,p. 10, fig.?NiEDEN, 1923, Das Tierreich, Lief. 46, Amphibia, Anura I, pp.73, 109.Type locality.?"In the lower part of the valley of the Rio Bravo(Rio Grande del Norte), and in the province of Tamaulipas, Mexico."Range.?From Mexican boundary southwest
^^jtsea^-gj^f^^M^''* through the States of Tamaulipas and Chihuahua^^^ l^^i^ij^^lP"'' to Mazatlan in Sinaloa, and south along Pacific^=^?s!^^>i' coast at least as far as Acaponeta in Nayarit.This small toad has a rather peculiar distributionin the United States. The northern limit seemsto be extreme southwestern Kansas (Morton andGrant Counties). It has been taken in the Wich-ita Mountains (Comanche County), Oklahoma^and in eastern New Mexico (Cuervo, GuadalupeCountj^). Cope reports that it occurs in thepanhandle district of Texas. It occurs in themiddle district of Texas from the Upper Wichita^ River south through McLennan, Burnet, Comal,Figure lo.-Head of Bufo Bcxar, Bee, Rcfugio, Duval, and Nueces Counties2624) from Delaware to thc Rio Grande, and along the Rio GrandeCreek, cuibersoncounty, fpom Brownsvillc (Camcron County) to the mouth
6, dorsal view ' of the Pccos RivcT (Val Vcrdc County). It hasbeen recorded from Delaware Creek (CulbersonCounty) in western Texas and also from Grant County in south-western New Mexico.Remarks.?Rough skin, small size, and greenish coloration usuallywill identify this toad in life, and doubtful specimens will be identifiedby transverse black bands across the upper eyelids. Although it is aburrowing species, the sole tubercles are insignificant; the body issmall, less than 2 inches in length; the parotoid glands are large, anddiverge posteriorly; and the legs are short.These little green toads seem to prefer the open country and aregenerally found on mesquite-covered flats. They emerge after darkfrom their burrows under the roots of the mesquite and other shrubs.They breed during April and May in shallow depressions or ditchesfilled by heavy rains.
MEXICAN TAILLESS AMPHIBIANS 51The cotypes of Girard's Bujo debilis are not designated in the cata-logue of the division of reptiles. On searching, it was found thatthere are only two entries that could possibly have been collectedprior to the publication of Girard's paper. One of these is a specimen(U.S.N.M. No. 2620) collected by Arthur Schott between the SaladaRiver (which empties into the Rio Grande near the town of Guer-rero in Tamaulipas) and Camargo in Tamaulipas. This locality i&in the lower part of the valley of the Rio Bravo (Rio Grande).Under the initial organization of the United States and MexicanBoundary Survey under Commissioner J. R. Bartlett, Arthur C. V.Schott was appointed a collector and assigned to the party of Lieut.A. W. Whipple. The commissioner's party left El Paso, Tex., inNovember, 1850, and arrived at Copper Mines, N. Mex., in May,1851. Col. J. D. Graham, the chief astronomer, did not arrive atCopper Mines until August 2, 1851, and this delayed the progress ofthe boundary survey considerably. The parties under the directionof Bartlett were then reorganized, and Schott must have receivedsome other assignment, for his name is not included among those wholeft Copper Mines on August 28, 1851. On September 13, 1851,Maj. William H. Emory was directed to proceed to El Paso, Tex.,to relieve Colonel Graham and take charge of the astronomical force.Observatories were established at Frontera (El Paso, Tex.), at SanElceario (Elizario, Tex.), and at Eagle Pass, Tex. It would seemthat Schott was subsequently assigned to Major Emory, as a birdskin was collected by him at Eagle Pass, in June, 1852. Bartlett'sexpedition did not return to Ringgold Barracks, Tex., until December20, 1852. When thework of the Boundary Commissionwas suspendedin December, 1852, Major Emory left for New Orleans. That Schotteither was left in Texas or was directed to report there in advanceof the main party of the new commissioner seems to be confirmed bythe birds collected by him at Sabanitas, Rio Grande, on September20, 1853, at Rio Grande, Tex., on October 13, 1853, and at Browns-ville, Tex., on October 25, 1853. The new commissioner. Gen.Robert B. Campbell, was appointed in March 1853, and work wasrecommenced in December, 1853. Arthur Schott was placed incharge of a surveying party under Major Emory and assigned thesurvey from Laredo to Ringgold Barracks, Tex. Although the dateof completion of this portion of the survey is uncertain, the boundarysurvey was again reorganized in October, 1854, under CommissionerWilliam Hemsley Emory. Schott was assigned as assistant to Lieut.N. Michler and this division proceeded through New Mexico to Cali-fornia, where camp was made on the initial point of the Rio Coloradoin April 21, 1855. Inasmuch as the entry in the catalogue for thisparticular toad indicates that Schott was working under the directionof Major Emory, the assumption is that it msij have been collected
52 BULLETIN 160, UNITED STATES NATIONAL MUSEUMsome time between October, 1851, and October, 1854, and in allprobability during the spring of 1854 while Schott was in charge ofthe party surveying the Rio Grande between Laredo and RinggoldBarracks, Tex. This specimen can not be located.One lot of eight newly metamorphosed young of Bujo debilis(U.S.N.M. No. 2621) was collected by Lieut. Darius Nash Couchat Matamoros in the State of Tamaulipas. In 1853, Couch, actingunder instructions received from the War Department, made a surveyfor a Pacific railroad in northern Mexico. Letters on file in thearchives of the Smithsonian Institution show that Couch made prep-arations for his forthcoming Mexican trip at Fort Miffhn, nearPhiladelphia, Pa., and that, although he received orders to proceedto Brownsville, Tex., in October, 1852, he was notified by ProfessorBaird on December 26, 1852, that the letter from the Mexican ministeraddressed to Couch in care of Professor Henry had been received.It is thus apparent that Couch did not reach Brownsville before 1853,and this assumption is confirmed in part by the fact that the firstspecimens collected at that locality are dated February, 1853.The exploring expedition conducted by Lieutenant Couch reachedMatamoros, Tamaulipas, on March 1, 1853. While there Couchpurchased the collection of Luis Berlandier and presented it tothe Smithsonian Institution. Baird acknowledged its safe arrivalin Wasliington on April 25, 1853. The party then proceeded west-ward through the States of Tamaulipas, Nuevo Leon, and Coahuila,at least as far as the Rio Nasas in Durango. The report of thisexploring expedition was never published, and it is difficult to deter-mine from the labels attached to specimens the route followed onthe return trip to Brownsville. A specimen labeled as having beentaken by Lieutenant Couch at Ringgold Barracks, Tex., is datedJuly 15, 1853, and this date would seem to indicate that the trip wasone of short duration. There are, however, other specimens thatare labeled as having been taken in August, 1853, in Nuevo Leon andTamaulipas. It is apparent, however, that all the Mexican speci-mens received from Couch were collected during 1853, since he arrivedin Washington some time prior to January 10, 1854. Couch re-mained in Washington for some months writing a report on theexplorations made by his party in Mexico, and was then orderedto Boston, Mass., and in October, 1854, to Fort Leavenworth, Kans.,where he remained until he resigned from the Army on April 30, 1855.These specimens are undoubtedly cotypes of Girard's species.Two specimens (U.S.N.M. No. 2622) from Chihuahua, collectedby Dr. Thomas H. Webb, are designated in the museum caWogue asthe cotypes of Girard's Bufo insidior. The preservation of thesespecimens is fair, but both are very much bleached. Doctor Webbwas the secretary and surgeon of Commissioner Jolm R. Bartlett's
MEXICAN TAILLESS AMPHIBIANS 53party of the United States and Mexican Boundary Commission.Returning from the Pacific coast, Bartlett's party followed the roadthrough the State of Cliihuahua, and, according to the commis-sioner's journal, was within the Hmits of that Mexican State fromOctober 8 until November 14, 1852. The entry in the museumcatalogue reads "Chihuahua, Mex.," which may or may not referto the city of Chihuahua.Specimens examined.?Twelve, as follows:Bufo debilis
Museum
54 BULLETIN 160, UNITED STATES NATIONAL MUSEUM(Duges); Michoacan (Duges); Morelos, Cuautla (Duges); Guerrero,Iguala, Tierra Colorado, and San Luis Allende (Gadow) ; Oaxaca, SanMateo del Mar, and Tehuantepec (Gadow); Vera Cruz, Tampico(Richardson), Coatzacoalcos (Street), Tetela (Gadow), Cuatotolapamand Lake Catemaco (Ruthven).Remarks.?Linnaeus gave the name Rana marina to the toaddescribed and figured by Seba,^? and merely states that its habitat isin America. In the caption for the Datch text, Seba refers to thisspecies as a "rare Virginia frog." In the Latin text, however, it islisted as "Rana, Marina, Americana, rara; Mas." The presence ofscrotumlike protuberances below the vent, which possibly were theresult of faulty preservation, led Sebato conclude that the specimen figuredwas a male. The type of Bufo mari-nus is therefore the specimen figuredand described by Seba. Informationfurnished by Dr. Leonhard Stejnegerindicates that the collection on whichthe ''Thesaurus" was based was pur-chased in 1716 for 15,000 Dutch guil-ders by "Peter the Great" of Russiaand taken to St. Petersburg. Thiscollection subsequently became theproperty of the St. Petersburg Acad-emy of Sciences, which, however, wasnot established by Catherine I untilDecember 21, 1725. A large part ofthe first collection ^^ assembled bySeba was destroyed by fire in St.Petersburg in 1747. Seba died in1736, and his second collection wassold at public auction in 1752 atAmsterdam. A portion of this collection was also purchased by the:St. Petersburg Academy of Sciences.Wallbaum, who published an extended description of this toad, con-cludes that it is a land type because the construction of the foot doesnot seem to be adapted for swimming in the sea. Hence, he arguesthat if this observation is corroborated, the Latin specific name Bufomarinus (Linnaeus) must be corrected, because it is misleading.Wallbaum proposes Rana gigas as a substitute name for the inappro-priate specific name marinus.
briGURE 11.?Head of Bufo marinus (U. S. N.M. No. 54119) from Gatun, Panama: a, Lat-eral view; 6, dorsal view
i? Seba, A., Locupletissimi rerum naturalium thesauri accurata descriptio, et iconihus artificiossimisexpressio, per universam physices historiam, Amsterdam, vol. 1, p. 120, pi. 70, fig. 1, 1734." Strauch, A., Das zoologische Museum der kaiserlichen Akademie der Wissenschaften zu St. Peters-burg ii} Seinem funfzig jahrigen Bestehen, St. Petersburg, pp. iv+372, 1889; Thomas, 0., Anu. and Mag.Kat. Hist., ser. 6, vol. 9, no. 63, p. 310, 1892.
MEXICAN TAILLESS AMPHIBIANS 55In describing Bujo agua, LatreUle seems to have confused the SouthAmerican Bufo marinus with the Cuban Bufo peltocephalus . It isstated that this enormous toad has been found principally in Braziland in Cuba. Reference is also made to the fact that the figure inSeba's "Thesaurus," which is the basis for Bufo marinus (Linnaeus),is hardly recognizable. Latreille does state, however, that his Bufougua is the same toad as the Bufo marinus of authors, or "L'epaule-armee." One might infer from these remarks that the figure anddescription of Bufo agua was based upon some specimen in the "Cabi-net d'Histoire Naturelle du Jardin du Roi " in Paris. It would appearfrom the synonymy listed under Bufo agua in Daudin's natural historyof frogs and toads that this author had failed to recognize the essen-tial characters of this species. In the eighth volume of his naturalhistory of reptiles, which was published during the preceding year,Daudin curiously enough does differentiate between "Le Crapaudfipaule-Armee " and "Le Crapaud Agua." The synonymy given for"Le Crapaud Epaule-Armee " is correct and includes Seba's "The-saurus," pi. 76, fig. 1. It should be noted that "Le Crapaud Agua" inthis last-mentioned publication of Daudin is a quite different batra-chian from Bufo agua Latreille.Wiegmann's Bufo horribilis is based on specimens collected byFerdinand Deppe and Graf von Sack in Mexico. The four cotypes(M.N.B. Nos. 3479-81) collected by Deppe are all toads ofmoderate size. The one collected at Misantla (No. 3479) has a head-and-body length of about 167 mm. ; an annotation on the label statesthat this species is No. 57 in the sale catalogue of Deppe's brother.Of the two (No. 3481) labeled as coming from Vera Cruz, the largesthas a body length of about 110 mm. The largest of the cotypes(No. 3480) is without definite locality and has a body length of about125 mm. The fifth cotype (No. 3493) is a yoimg individual collectedin Mexico by Graf von Sack and F. Deppe that lacks cranialcrests and has a head-and-body length of not more than 50 mm.Full-grown toads with a rather rough skin and with triangularparotoid glands as large as or larger than the side of the head andstudded with large pores may be referred without question to thisspecies. In general, the coloration of Bufo marinus is quite variable,ranging through the various shades of brown, including yellowish,reddish, or even blackish, and occasionally greenish olive; upperpartswith or without large black insuliform spots, which when present areusually edged with pale yellow; a light vertebral line occasionallyvisible; arms and legs of immature individuals usually banded withdark brown; underparts dingy white or yellow. The interorbitalspace is wider than the upper eyelid and the cranial crests are oftenquite prominent in old adults. Sexually mature males have numerous
56 BULLETIN 160, UNITED STATES NATIONAL MUSEUMlow spiny tubercles on the back. The range of individual variation inB'u;fo marinus has been discussed by Ruthven^^ and Noble. '^Adults of Bufo marinus are the largest of the true toads. Miranda-Ribeiro ^* states that specimens with a head-and-body length of 220mm. have been taken in Brazil and that this species ranges southwardto the Patagonian district. These large toads are nocturnal in theirhabits and hide under fallen tree trunks, matted leaves, and stones, orburrow into loose soil.Atmospheric conditions seemingly influence the time of breedingof this species. Egg laying may commence as early as February, ifthere has been plenty of rain, but may be delayed until July in regionsof continued drought. The breeding season also varies according tolocality. Tadpoles have been noticed in Barbados in the pools duringAugust and February, while in Bermuda the eggs are usually laidduring April. From the observations made by Ruthven,'^ it wouldappear that 45 days are required for development from egg to adult.The eggs that Mark ^? kept under observation hatched in a few hoursless than 4 days.The poisonous nature of the glandular secretions of some of thetoads has been investigated by Madame Phisalix,^'' and she has dis-cussed the kinds of toxins that are present. Few actual observationson the action of this secretion on other animals, however, have beenpublished. It is therefore of interest to review a short paper byDodds.^* Near the end of July, 1922, at Los Mochis in northernSinaloa, a small terrier dog was observed teasing a large toad, pre-sumably Bvfo marinus. At last the dog caught the toad by the backand "although the dog's mouth was not in contact with the toad formore than an instant, he immediately lost all interest in the animal.'^The dog refused water and "in about a minute's time showed signsof weakening and general paralysis. He sank to the ground with hislegs spread out, writhing and whining with pain, and unable to recog-nize his master. During this time he was able to push himself alongthe ground, gradually becoming weaker and very rigid, with eyesgreatly protruding and respiration and heart action exceedingly rapid.After twenty minutes he was somewhat quieter as if he was going todie." Castor oil was then administered to the terrier and an hourlater the dog apparently was quite normal.Specimens examined.?One hundred and four, as follows:
12 Ruthven, A. Q., The amphibians and reptiles collected by the University of Michigan-WalkerExpedition in southern Vera Cruz, Mexico. Zool. Jahrb. (Syst. Abt.), vol. 32, pi. 4, p. 309, 1912."Noble, G. K., Bull. Amer. Mus. Nat. Hist., vol. 38, art. 10, p. 333, 1918." Miranda-Ribeiro, A. N., Archiv. Mus. Nac. Rio Janeiro, vol. 27, pp. 134, 216, 217, 1926." Ruthven, A. Q., The breeding season of Bufo marinus (L.) in Demerara. Copeia, no. 31, pp. 43, 44>May 24, 1916.
'? Pope, P. H., The introduction of West Indian Anura into Bermuda. Bull. Mus. Comp. Zool., vol. 61,no. 8, pp. 123, 124, June, 1917." Phisalix-Picot, M., Animanx venimeux et veuins, vol. 2, pp. 1-174, 1922.
?* Dodds, C. T., A note on Bufo marinus. Copeia, no. 114, pp. 5, 6, Jan. 20, 1923.
MEXICAN TAILLESS AMPHIBIANSBufo marinus 57
Museum
U.S.X.M....DoDo
CatalogueNo.
DoF.M.N.H.U.S.N.M .M.N.B....C.O.E.MDoU.S.N.MDoF.M.N.H.A.M.N.H...M.N.B. .C.O.E.M. .A.M.N.H...U.S.N.M....Do.Do.Do.Do.Do.M.C.Z.
47243471694745030517136171054-5534791383035463521367123053481
IT.S.N.M . ..C.O.E.M ..F.M.N.HDoU.M.U.M...M.C.ZDoC.A.SW Do....U.S.N.DTDoA.M.N.H....C.A.SA.M.N.H....DoDoDoDoDoU.S.N.M .DoDoA.M.N.H. ..U.S.N.MA.M.N.H
13751-5218987-884790363855-671111363665-7157591395057592101379135814
1253105450405188-89
U.S.N.MDo.Do.Do_Do.
47466-6813920634562586261-626266-676269-70627462764736247465304346252-574; 08013906
46854-56
Do.Do.Do....Do....F.M.N.H.?!M.N.B.Do....
468474676347310466194757113907122881928054934803493
Num-ber ofspeci-mens
111
142U111141?212
Locality collected Date collected
east
1903
Sept. 10, 1918
1903June 3, 1919
Sonora: Camoa Nov. 8 1898DuRANGo: Guasamota... " Aug' 14*1897Tamaulipa.s: Hidalgo Mar. is! 1891Vera Cruz:CordobaEI Juile '_['_MiradorMisantla '_'_[Motzorongado_Orizabado .y/..PerezVera Cruz..dodoNayaeit: Puerta Azulof Santiago Ixcuintla.Jalisco: Barranca IbarraColima:ArmeriaColimado " """dododo ''.'.'.'.'."MoRELos;Puente de IxtlaTetecalaYantepecPuebla:AtlixcoVaso de Tenango, i,4'0(}-lV5"obmeters elevation.OUERRERO:Acapulcc. ...
---.do ."."I":Oaxaca:Cosolapa
..-.do -lllllllll'CuicatlanOuichicoviMixtequillo-doTehuantepeedo
By whom collected orfrom whom received
July ?,1918Dec. 23, 1919Apr. 21,1892Feb. ?, 1892July ?,186318631863July 27,19021916June ?, 1901Oct. 14,189019031903July 17, 1926
-do.
-do-
-do-
.do.
-do-
-do..
-do-.West of Tehuantepec RiverTuxtepecXadani and San Miguel delPuerto.Chiapas:La Trinidad, 800 feet alti-tude.TaoachulaTonala ITabasco:Monte Cristo-.
-.-.do ['/,Yucatan:Cozumel Island
18721872July 12, 1925July 20, 1925Oct. 8, 1894June 24, 1895May 14, 1920Aug. 24,1925July ?,1920dododo
---do
-.--doMay 28,1895May 1, 1895
E. A. Goldman.Nelson and Goldman.William Lloyd.(?) C. Sartorius.S. E. Meek.C. Sartorius.Ferdinand Deppe.
(?) C. Sartorius.Francis Sumichrast,S. E. Meek.Paul D. R. Ruthling.Ferdinand Deppe.S. Seniles.Paul D. R. Ruthling.P. L. Jouy.E. W. Nelson.John Xantus.Do.Do.Julius Hurter.G. Gluckert.Julius Hurter.S. E. Meek.Do.H. B. Baker.
MayApr./June 1. 19206, 1894
Hassler Expedition.Do.Joseph R. Slevin.Do.Nelson and Goldman.Do.Paul D. R. Ruthling.Joseph R. Slevin.Paul D. R. Ruthling.Do.Do.Do.Do.Do.Nelson and Goldman.Do:Francis Sumichrast.Paul D. R. Ruthling.Nelson and Goldman.8, 1920 It, , t^ t, -r. ^,.IJune 10, 1920 /P^*^ ^- K- Ruthhng.Nov. 10, 1895
-do-No definite locality
.....dodo(?) StateNo definite locality
-.-.do
Feb.Aug.MayMay 1, 18965, 18958,19009, 1900pan.IJan.
Nelson and Goldman.Nelson and Goldman.Do.E. W. Nelson.Nelson and CJoldman.Do.23,1885 1 Albatross, U. S. Fish29, 1885 / Commission.Arthur Schott.Do.
! George F. Gaumer.Ferdinand Deppe.Ferdinand Deppe andGraf von Sack.
Apr. 14, 1901
' Cotypes of Bufo horrtbUit,
58 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
-.-SS^i3"i
BUFO MARMOREUS WiegmannFigure 121833. Bujo marmoreus Wiegmann, Isis von Oken, vol. 26, pt. 7, p. 661.
?
GuNTHER, 1901, Biologia Centrali-Americana, Reptilia and Batrachia,p. 246, pi. 70, fig. A, Feb.?NiEDEN, 1923, Das Tierreich, Lief. 46,Amphibia, Anura I, pp. 76, 130.1868. Bufo argillaceus Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 20, p. 138,Mar. (Colima, Mexico).1894. Biifo lateralis Werner, Zool. Anz., Jahrg. 17, no. 446, p. 156, May 7(Tehuantepec, Mexico).Type locality.?Vera Cruz, Mexico.Range.'?From Mazatlan and Rosario in Sinaloa south through.Colima, Jalisco, and Guerrero to the Isthmus of Tehuantepec inOaxaca and to Tonala in Chiapas.Reported from: Guerrero, Cocoyul, TierraColorado, Cajones, Chilpancingo, Rio Balsas,and Iguala (Gadow); Oaxaca, Salina Cruz(Gadow).Remarks.?The cotypes of Bufo marmoreus(M.N.B. Nos. 3529-31) were taken at VeraCruz by Ferdinand Deppe, according to thelabel on the jar. These specimens have thelight-colored vertebral stripe with irregularlongitudinal light line. The supraciliary andpostorbital crests are very low. The darkspots on the upperparts have a narrow whiteedge; the parotoids are light colored aboveand dark laterally; the underparts are white;the metatarsal tubercles are small; and theskin of the upperparts is studded with minutewarts. The head-and-body lengths of thethree cotypes are 58.2, 56.5, and 56.5 mm.,respectively.John Xantus de Vesey was appointed United States consul atManzanillo, Mexico, in 1863. From March, 1863, to March, 1864, heremained in Colima, although his consular appointment was revokedprior to August 11, 1863. On February 29, 1864, his last shipmentfrom Manzanillo was assembled, and he states in Part V of his fieldcatalogue that this collection included 1,000 alcoholic specimenswithout labels. Evidently Xantus traveled a great deal while residingin Colima, as an entry in his journal dated June 25, 1863, states that hehad traveled 700 miles that month. He made numerous trips tothe city of Colima, and in all probability collected the cotypes ofBufo argillaceus (U.S.N.M. Nos. 27763-64) in the vicinity of thatcity. These cotypes are fairly well preserved, but are somewhatdistorted. It is possible that the preservative may have been allowed
Figure 12.?Head of Bufo mar-moreus (U.S.N.M. No. 47129)from mountains 12 miles northof Santo Domingo, Oaxaca,Mexico: a, Lateral view; 6,dorsal view
MEXICAN TAILLESS AMPHIBIANS 59to evaporate at some time when they were in the possession ofProfessor Cope.Professor Werner received the type specimen (N.M.W., WernerColl. No. 115, 1929) of Bujo lateralis from T. K. Salmon, who madecollections on the Isthmus of Tehuantepec in 1872. The followingnotes were made on the type: Head-and-body length, 59 mm.; thehind limb being carried forward along the body, the tibio-tarsaljoint does not reach to the level of the parotoid gland, and is at least7 mm. from the posterior margin of the tympanum; the parotoidglands are placed diagonally on the shoulders, the greatest diameterof the right one being 6.4 mm. and that of the left one 6.8; cranialcrests low; a short supratympanic crest; transverse diameter of tym-panum, 3.3 mm.; transverse diameter of eye, 7.5 mm.; transversediameter of head at level of angle of jaws, 21 mm.; subarticulartubercles on fourth toe double; soles of fore and hind feet with numer-ous supernumerary tubercles; inner metatarsal tubercle small, butlarger than outer; a row of minute warts along tarsal fold; throat,abdomen, and under surfaces of thighs granular; warts on upperpartslow and flattened ; a white dorso-lateral longitudinal stripe above linearseries of low warts; a lateral brown band extends from angle of paro-toid to thigh; coloration of upperparts much faded; a light vertebralstripe, on each side of which is an anterior and a posterior dark blotch;under surfaces of fore and hind feet black.The species Bnfo marmoreus seems to be somewhat closely relatedto B. canaliferus, a species that has a somewhat similar coloration,but is distinguished at once by its elongated parotoid glands.The habits and life history of Bnfo marmoreus are unknown.Collectors in the past have paid little attention to the ecologicalpreferences of Mexican toads. A large lot received from Xantuswould seem to indicate that they congregate in pools in considerablenumbers during the breeding season and that their breeding habitsare not unlike those of other toads. Their main center of abundanceseems to lie west of the Sierra Madre del Sur.Toads belonging to this species have the following characteristics:Ground color of upperparts brownish gray, marbled more or lessdistinctly with some darker color; a whitish vertebral line of varyingwidth, on each side of which are three large or a greater number of smallirregular light margined dark blotches; the whitish markings aboveand the brown markings below the lateral row of small warts formmore or less confluent longitudinal bands of varying width; a contin-uous or medially interrupted V-shaped band crosses the upper eyelidsand interorbital region ; fore and hind limbs above with dark cross-bars; underparts immaculate; cranial crests low, the most conspicu-ous of which are the combined supraciliary and postorbital crestswhich form a continuous curve from the anterior margin of orbit to
60 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
supratympanic crest ; orbito-tympanal crests short ; canthai and pari-etal crests vestigial or absent; supratympanic crest short and thick;parotoid glands nearly subtriangular, smaller or at most no largerthan upper eyelid; skin of upperparts rather thickly studded withsmall warts and tubercles ; and a row of small closely spaced tuber-cles in place of a tarsal fold.Specimens examined.?Two hundred, as follows
:
Bufo marmoreus
MEXICAN TAILLESS AMPHIBIANS 61Range.?From the Mexican boundary southward through Sonora,Chihuahua, Coahuila (Monclova and Castanuelas), and Tamaulipas(Sierra Nola) to San Luis Potosi and Guanajuato (Silao). In theUnites States the species is confined to the region west of the Mis-sissippi River, where it occurs locally as far east as the WichitaMountains (Mount Scott, Comanche County, Okla.). It has beentaken in southwestern Colorado (Basin Creek, San Miguel County)in a small alkaline stream whose waters reach the Colorado River byway of the San Miguel and Dolores Rivers. Along the San JuanRiver in southeastern Utah this toad has been found at a numberof localities (Blufi', Natural Bridges, and Caroline Bridge in San JuanCounty). In northern Arizona it has beenfound on the floor of the Grand Canyonbetween the Coconino and Kaibab Plateaus,and in southwestern Utah in the drainageof the Virgin River (Zion National Park,Washington County). It has been foundin eastern California in the PanamintMountains (Cottonwood Canyon, InyoCounty) and in Death Valley (FurnaceCreek). South of these localities it hasbeen reported in California from the TurtleMountains (San Bernardino CountjO^ FortYuma, the Imperial Valley, and in SanDiego County (Vallecito); and thencesouthward in Lower California to CapeSan Lucas. In west-central Arizona thered-spotted toad has been taken in thedrainage of Salt River and its tributaries(White River Canyon, Navajo County;and McMillenville, Gila County). It hasalso been recorded in Arizona from Mari-copa County (Cave Creek), Pima County (Catalina Mountains nearTucson), Santa Cruz County (Crittendon), and Cochise County(Ramsay Canyon, Huachuca Mountains). It is known to occur inextreme southwestern New Mexico (Dog Spring and Dog Mountains,Hidalgo County). With the exception of the panhandle region, thistoad is distributed locally over the western half of Texas. The recordsby counties are as follows: Dallas (Dallas), Somervell (Glen Rose),Tom Green (Fort Concho), McLennan (Waco), Burnet (SherrardRanch), Hays (San Marcos, Wimberley, and Devils Canyon), Kendall(Boerne), Bexar (San Antonio and Helotes), and Duval (San Diego).The occurrences along the Rio Grande River by counties are asfollows : Cameron (Brownsville), Starr (Ringgold Barracks), Val Verde66785?32 5
pRo o C(5(^cx
Figure 13.?Head of Bufo punctatus(U.S.N. M. No. 12661) from La Paz,Lower California: a, Lateral view;6, dorsal view
62 BULLETIN 160, UNITED STATES NATIONAL MUSEUM(Devils River and Cornstock), Brewster (Boquillas and Terlingua),and El Paso (El Paso). In the drainage of the Pecos River in Texasthis toad has been taken in Jeff Davis County (Fort Davis and DavisMountains) and in Reeves County (Cherry Creek). Farther norththere are records for Alamogordo in Otero County and Albuquerquein Bernalillo County, N. Mex.Remarks.?The three cotypes of Bujo pundatus (U.S.N.M. No.2618) were collected in Texas by John H. Clark, a member of Col.J. D. Graham's party of the United States and Mexican BoundarySurvey, on the Rio San Pedro (Devils River), a tributary of theRio Grande. According to the original entry in the catalogue of thedivision of reptiles, these cotypes were received from "J. D. G."(J. D. Graham). The name of the collector is not given in thecatalogue, but Baird in his report on the reptiles of the United Statesand Mexican Boundary Survey credits their discovery to J. H. Clark.Inasmuch as they must have been collected not only prior to thepublication of the original description, but also while Clark wasworking under the supervision of Colonel Graham, it will not bedifficult to fix the approximate date of collection. The party underthe direction of Graham followed well-known wagon roads used bythe United States Army in southern Texas. Leaving Indianola theypassed through San Antonio on their way to Fort Inge. From thatpost the old road turns westward toward the Rio Grande and crossesthe Rio San Pedro (Devils River) near its mouth, and then follows thevalley of the latter for some distance before going across to the PecosRiver. The road followed the western bank of the Pecos River tothe mouth of Delaware Creek and then taking a more westerlydirection it passed Salt Lagoons Spring, and ended at Frontera (ElPaso, Tex.). Graham's party arrived at Copper Mines, N. Mex.,on August 2, 1851. Work on the boundary survey was com-menced shortly after this date, and Clark accompanied Graham inSeptember, 1851, as far west as Santa Cruz in Sonora, Mexico. Onthe return trip, Clark spent some time collecting in the vicinity ofCopper Mines, N. Mex. Maj. W. H. Emory was not directed toproceed to El Paso, Tex., to relieve Colonel Graham until Septem-ber 13, 1851. Clark received a new assignment from Major Emory,and on February 12, 1852, we find him at the Presidio del Norte,which was near the mouth of the Rio Conchos in Chihuahua, and inMay, 1852, at Frontera. The cotypes thus were collected duringthe year 1851 and probably during July.Spotted toads are readily distinguished from other species by thebroadness and flatness of the top of the head, but vestigial cranialcrests may or may not be present. The general coloration of theupperparts ranges from light reddish brown to olive or even malachitegreen. The small sharply pointed warts on the upperparts are bright
MEXICAN TAILLESS AMPHIBIANS 63pink or vermilion at the base, and their bases are often encircled orpartially encircled by narrow black borders. The underparts arewhitish, irregularly speckled with black.The spotted toad seems to prefer desert areas and in many localitiesis an inhabitant of rocky gulches and canyons with seepages andsprings. During the hot midsummer it takes refuge under largeflat stones and bowlders in creek bottoms, under stones partlyimbedded in moist earth and sand, or in niches in rock bluffs.Although large numbers of them frequently congregate in pools ofwater during the breeding season, they are rarely found except afterdark. The breeding season varies according to locality and meteoro-logical conditions. In southern Texas, this toad breeds early inMarch but along the northern limits of its range the breeding seasonmay be delayed until April and May.Specimens examined/?Forty-five, as follows
:
Bufo punciatus
Museum
64 BULLETIN 160, UNITED STATES NATIONAL MUSEUMRange.?From Mazatlan in Sinaloa, Sierra Madre in Zacatecas, andOrizaba in Vera Cruz, south through Nayarit, JaUsco, Guanajuato,Hidalgo, Morelos, Puebla, and Guerrero to the Isthmus of Tehuan-tepec in Oaxaca; exact limits of range not known; occurs in theprovince of Bocas del Toro in northeastern Panama and in the Andesof Ecuador. Reported from: Vera Cruz, Tetela (Gadow); Sinaloa,Presidio (Giinther); Durango, Milpas (Forrer); Guanajuato, Guana-juato, Huazteca Potosina, and Silao (Duges); Puebla, Izucar deMatamoros (Perez); Guerrero, Omilteme and Chilpancingo (Gadow),Amula (H. H. Smith); Oaxaca, Totolapan (Gadow).Remarks.?The original description of Bufo simus is based solelyon young toads taken by J. von Warszewiez in shallow places in theChiriqui River near Bo-cas del Toro, Panama.It is assumed that theauthor refers to rivers orstreams descending fromthe Volcano Chiriquithrough the presentprovince of Bocas delToro and flowing intothe lagoon, which is con-nected with the Carib-bean Sea by the so-calledBocas del Toro. It isstated that the cotypesare in the ZoologicalMuseum of Krakow.According to the originaldescription, the bodylength of the largestindividuals (cotypes) is26 mm. There are no bony crests on the crown and on the hinderpart of the head. The tympanum is concealed. The parotoid glandsare medium sized and irregularly rounded. There is a slight web onthe hind foot, but the toes have dermal fringes. The skin on theback is studded with large and small warts, but the skin of theunderparts, with the exception of the thighs, is nearly smooth. Thecoloration of the upperparts is a lighter or darker gray, and that of theunderparts yellow-gray with irregular blackish spots.It has been generally believed that Bujo simus was closely relatedto Bufo intermedius, and that it differed from the latter only in havingthe tympanum hidden under a tubercular skin. Ten young toadsbelonging to the American Museum of Natural History and collectedat or near Orizaba in Vera Cruz were compared with the original
Figure H.?Heads of Bufo simus: a. Lateral view of specimen(U.S.N.M. No. 2607) from Valley of Mexico, Mexico; -b, dorsalview of same; c, lateral view of specimen (U.S.N.M. No. 47171)fromDolores, Nayarit, Mexico; d, dorsal view of same
MEXICAN TAILLESS AMPHIBIANS 65description of Bujo simus. These individuals have body lengthsranging from 22 mm. to 29 mm. A small indistinctly outlined tym-panum, which was covered with minute tubercles, was recognized onfive of these young toads. On the remaining five, the tympanum wasentirely concealed by numerous closely spaced tubercles. On all, theskin of the underparts is seen on close inspection to be distinctlygranular. Nevertheless the smaller individuals have a much smootherand a much more delicate skin on the underparts than have olderindividuals. A rather casual examination of such individuals mightpossibly have led Schmidt to infer that the skin on the underparts wasnearly smooth. Giinther,^^ however, states that the British Museumof Natural History received through Doctor Werner one of the cotypesof Schmidt's Bufo simus. This specimen (B. M. No. 98. 9.14. 6) hasa tarsal fold, and the tympanum is concealed by the tubercularskin. The head and body length is 15 mm. Although it is a veryyoung toad, the coloration of the upperparts, especially the light areas,the sides, and the upper lip, corresponds rather closely with the typeof color pattern most commonly observed in Mexican specimens. Theskin on the abdomen is distinctly granulated.The cotypes of Bui'o intermedius (B. M. Nos. 58. 9. 20. 3-6) werecollected in 1857 by Louis Fraser, a bird collector employed by Dr.P. L. Sclater, and are labeled as coming from Guayaquil and the Andesof Ecuador. Although the Andes locality is not stated, it is knownthat Fraser was collecting at Gualaquiza and at Zamora in the easternwooded region along the Rio S. lago and on the eastern slope of thewatershed at Cuenca, which is situated at an altitude of 8,200 feet onthe Rio Matadero. Two of the cotypes have the type of colorationthat is commonly present in Mexican specimens, one has prominentblack spots on the light ground color of the upperparts, and the otherhas the color pattern somewhat obliterated by a darker suffusion.In addition, the parotoid glands come in contact with the postorbitalcranial crests, the parietal crests are short, the subarticular tuberclesOjI distal phalanges of the fourth toe are double, and a linear seriesof small tubercles extends along the tarsal fold. The measurementsof these four cotypes are, respectively: Head-and-body length, 90.4,88, 76.4, and 76.7 mm.; transverse diameter of tympanum, 3.8, 3.1,3.7, and 3.2 mm.; transverse diameter of orbit, 8.5, 8.8, 8.1, and 6.9mm.; distance between anterior rim of eye and nostril, 6.3, 5.8, 5.0,and 4.3 mm. The specimen figured and described by Giinther bearsa very close resemblance to the toad that occurs in Mexico, and thereare no known characters that will separate the former from the latter.Camerano merely published De Filippi's manuscript name, Bvfo occi-dentalis, in the synonymy of Bufo intermedius, and it is strictly anomen nudum.
>? Gunther, A. C. L. G., Biologia Centrali-Amerieana, Reptilia and Batracliia, pp. 2.')4, 255, May, 1901.
66 BULLETIN 160, UNITED STATES NATIONAL MUSEUMThe type of Bufo monksiae (U.S.N.M. No. 9896) is a young toadthat measures 23.5 mm. from the tip of the snout to the vent. It wascollected at Guanajuato by Alfred Duges in 1877. The upperpartsare brownish, with a few small ashen spots and a vertebral stripe ofthe same color. A transverse light-colored band extends across theeyelids and the interorbital region. It is clearly a young individualof Bufo simus. It is possible that the coloration of this individualmay have been changed somewhat by the preservative or by the con-tainer.The general coloration of the upperparts of Bufo simus is light grayor olive, marbled with a darker shade of brown. A light-coloredinterorbital band is generally present. The spotted pattern is espe-cially noticeable on a specimen (A. M. N. H. No. 13896) from SantaCatalina in Puebla. Some of the specimens collected in Jalisco havenearly white underparts. A specimen (A. M. N. H. No. 13905) takenat Cafetal CapaUta in Oaxaca has an unusually warty skin. Youngindividuals of this species have a distinct light-colored vertebralstripe, and many have the belly and chin spotted with white. Twentyimmature individuals, ranging from 32 mm. to 61 mm. in length, andcollected in Vera Cruz, Jalisco, and Oaxaca, have the tympanumpractically hidden by closely approximated conical warts. On allthese the tympanum is covered with a tubercular skin, and on a num-ber of them the sole discernible vestige of the tympanum is locatedin a dermal fold bordered by numerous conical tubercles. The paro-toid glands are generally elongated, somewhat bean-shaped, and incontact with the upper eyelid. These glands are generally studdedwith minute pores and in rare instances are somewhat pustulose.The tarsal fold is more or less distinct and generally is furnished witha linear series of small closely spaced tubercles. Posteriorly divergingsupraciliary crests, short oblique parietal crests, and vestigial postor-bital crests are generally present, except on young or immature individ-uals. In a series of 50 individuals, all but two have double subartic-ular tubercles on the distal phalanges of the fourth toe.Considerable variation in the color markings on the upperpartswas observed in the series studied. The skin of the upperparts isgenerally covered with small warts, which are either conical or blunt.One individual (U.S.N.M. No. 47171), taken at Dolores in Nayariton August 6, 1897, by Nelson and Goldman, differs in a number ofrespects from the average toad of this species. The skin of theupperparts, top of the head, and surfaces of the fore and hind limbs isexceptionally tubercular and is covered with closely set small conicalwarts. The elongated parotoid glands are likewise covered withspinose warts. The posteriorly diverging supraciliary crests are dis-tinct, but there are no vestiges of the parietals and the postorbitals.The tympanum is indistinct and is covered with tubercular skin and
MEXICAN TAILLESS AMPHIBIANS 67
numerous small conical warts. There are single subarticular tuber-cles on the distal phalanges of the fourth toe. It would seem thatthe hind feet are deformed, for the fourth toe of the right foot islonger than the corresponding toe on the left foot, and both are consid-erably shorter than the normal. The tarsal fold has a linear series ofsmall conical warts.Specimens examined.?Eighty-four, as follows:Bufo simus
Museum CatalogueNo.
A.M.N.H.-U.S.N.M?DoDoDoA.M.N.H-U.S.N.M-.DoDoM.C.ZDoDoA.M.N.H-.DoU.S.N.M...DoA.M.N.H_.DoDoDoDoF.M.N.H..N.M.W...-U.S.N.M...DoA.M.N.H_DoDoU.S.N.M...DoA.M.N.H..DoDoDoU.S.N.M...A.M.N.H..DoU.S.N.M-A.M.N.H.C.G.E.M..F.M.N.H_U.S.N.M...A.M.N.H-U.S.N.M..Do
1325347449468264717047171132521135821160-61
39513953-559032-3313761-6213779-9530224-251926813764-6513776
13247629313896-9713591910-15469614682513251132501375464663468271376613248-491375313755479161390513890-954720013902-0423027470096259252582607
Num-ber ofspeci-mens Locality collected
Sinaloa:Ruined monastery north ofMazatlan.PlomosasZacatecas:Sierra MadredoNayarit:DoloresMarquesado..Guanajuato:GuanajuatododoHrDALGo:Guenero.
.._.do.VelascoVera Cruz:Orizaba
."-do.'.V."."--"-----!------'
....doNear OrizabaNorth of Rio Atoyae, nearPotrero.Federal District:3 miles east of IxtacalcoPuebla:PueblaSanta CatalinaMORELOS:Cuautla...CuernavacaJalisco:AtemajacColotlanEl Aquilar MineHostotipaquilloNorth of Hostotipaquillo...JacalaLa LagunaOn way to MagdalenaPlatanillo, 8 miles north ofMagdalena.Rio B lanco, north of Zapopan.West of TonalaGuerrero:TlalixtaquilloOaxaca:Cafetal CapalitaEjutlaJuquila, 5,000 feet altitude...Mountain descent into valleyof Miahuatlan.Oaxaca .-
Date collected
Nov. 25. 1919July 19,1897Aug. 17,1897Aug. 18, 1897Aug. 6,1897Nov. 14,1919Jan. 30, 18801877Sept. 24, 1913do.1913Apr. 25,1919June 25,1919Aug. 2, 1891Summer (?)1918.June 18,1918
Oct. 27,1918July 22,1920July 31,1920
.do.Mountains near Oaxaca,9,000 feet altitude.Tehuantepec.?State:No definite localityValley of Mexico
MayAug.Nov.Nov.Sept.Mar.Sept.Sept.Oct.
28, 189228, 18976, 19191, 191926, 19196, 189720, 189729, 191928, 1919Sept. 12, 1919Sept. 17, 1919Dec. 10,1894July 15,1920July 9, 1920Mar. 2. 1895July 7, 1920Aug. 10,1919
By whom collected orfrom whom received
June 20,18941920
Paul D. R. Ruthling.Nelson and Goldman.Do,Do.Do.Paul D. R. Ruthling.Alfred Dug6s.Do.Do.W. M. Mann.Do.Do.Paul D. R. Ruthling.Do.Francis Sumichrast.Willis S. Blatchley.Paul D. R. Ruthling.Do.Do.Do.Do.S. E. Meek.H. Gadow.Nelson and Goldman.Do.Paul D. R. Ruthling.Do.Do.Nelson and Goldman.Do.Paul D. R. Ruthling.Do.Do.Do.Nelson and Goldman.Paul D. R. Ruthling.Do.Nelson and Goldman.Paul D. R. Ruthling.J. Jarduno.C. H. T. Townsend.Nelson and Goldman.Paul D. R. Ruthling.John Potts.
* Type of Bufo monksiae. ? Young.
68 BULLETIN 160, UNITED STATES NATIONAL MUSEUMBUFO VALLICEPS WiegmannNebulous ToadFigure 151833. Bufo valUceps Wiegmann, Isis von Oken, vol. 26, pt. 7, pp. 657-659.
?
Peters, 1863, Monatsber. k. preuss. Akad. Wiss. Berlin, p. 81.
?
Cope,1889, U. S. Nat. Mus. Bull. 34, p. 292, fig. 73.?Dickerson, 1906, Thefrog book, pp. 108-110, figs 112-115, col. pi. 5, fig. 3.?Nieden, 1923, DasTierreich, Lief. 46, Amphibia, Anura I, pp. 76, 128.1833. Bufo trachypus MS. Wiegmann, Isis von Oken, vol. 26, pt. 7, pp. 657, 658,footnote. (Nomen nudum.)1852. Bufo granulosus Baird and Girard, Proc. Acad. Nat. Sci. Philadelphia,vol. 6, p. 173, Oct. [Nee Bufo granulosus Spix, 1824.)1854. Bufo nebulifer Girard, Proc. Acad. Nat. Sci. Philadelphia, vol. 7, p. 87,May.
?
Baird, 1859, Report on the United States and Mexican BoundarySurvey, vol. 2, Reptiles, pp. 25, 35, pi. 40, figs. 1-4.1858. Bufo sternosignatus [part] Gunther, Catalogue of the Batrachia Salientiain the collection of the British Museum, pp. 68, 69, pi. 5, fig. C (Cordovaand Mexico).1896. Bufo valliceps microtis Werner, Verhandl. k. k. zool.-bot. Gesell. Wien,vol. 46, no. 8, p. 350. (Honduras.)Type locality.?Mexico.Range.?From the Rio Grande, below the mouth of Pecos River,southwest through Tamaulipas, Nuevo Leon, Coahuila, and Durangoto southern Sinaloa (Mazatlan) and Nayarit (Acaponeta), and thencesouthward along both coasts as well as in the interior of Mexico tosouthern Costa Rica (Buenos Aires). Reported from Sinaloa, Pre-sidio de Mazatlan (Forrer); Vera Cruz, Tampico (Richardson), Ja-lapa (Hoege), Atoyac (H. H. Smith), Cuatotolapam (Ruthven), andPresidio, Tetela, and Agua Fria (Gadow).The range of this toad in the United States extends from the east-ern boundary of Louisiana (Como, Franklin Parish) west to Bosque,Llano, Kendall, and Val Verde Counties in Texas, and along the Gulfcoast from the mouth of Pearl River, La., to the mouth of the RioGrande, Tex.Remarks.?The so-called nebulous toad occurs in the vicinity ofcoastal marshes in southern Louisiana. In eastern Texas it has beenfound along the edges of pine forests. Along the western border ofits range in central Texas on the grand prairie region, this toad seemsto prefer small streams with rocky banks, and hides in fissures andniches in the rocks. In Guatemala, it occurs in considerable num-bers along the margins of forests, in the clearings around villages, andin lesser numbers in the savannas. No published information isavailable in regard to its breeding habits.It has very high cranial crests and a dark interorbital cross band.The coloration of the upperparts is quite variable; the lighter-coloredindividuals are generally yellowish or light gray, and the darker oneshave the color pattern subdued or obliterated by a dark slate or black
MEXICAN TAILLESS AMPHIBIANS 69
suffusion. The vertebral line is usually light colored, and the backis generally blotched with some dark color. The linear series of coni-cal warts extending along lateral line from parotoid gland to thighseparates the dorsal light-colored stripe from the lateral dark-coloredone. The parotoid glands are often bicolored.The normal coloration of this toad may be described as follows:Below lateral linear series of yellowish warts extending from paro-toid gland to thighs is a blackish stripe and above it is a light-colored,often yellowish, stripe. A black transverse bar extends across headbetween orbits and near origin of parietal crests; a black obliquelyplaced spot is placed in front of antero-internal angle of parotoidgland; a dark inverted V bisects light-colored vertebral stripe nearlevel of middle of parotoid glands ; behind this mark-ing and on each side of vertebral stripe are smallirregular dark-colored blotches; an irregular elon-gate black blotch in sacral region on each side ofvertebral stripe; warts conspicuous and numerous;cranial crests very distinct.One of the variants has the upperparts suffusedwith reddish brown. In another phase the upper-parts are very light colored, and are generally j^el-lowish or light gray. In this phase the dark blotchesor markings are often indistinct. figure 15.?Head of b?/oIn the dark phase the upperparts are slate vatucepsiv.s.NM no.^
,
x-r- r- 42351) from Victoria,colored or even blackish. All the markmgs are Tex.: a, Lateral view; 6,either somewhat subdued or less evident because ^"^^'^^ ^'?^of the general darkening of the entire ground color. The lateral dor-sal light-colored stripes are less noticeable because of an invasion ofa slaty suffusion. Some of the lateral linear series of warts may beslate colored.In some immature individuals, the black markings have a velvetyappearance. This condition is occasionally met with in subadultspecimens. The underparts are often unspotted, but in most instancesdark spots or blotches are present.It would appear from the footnote in Wiegmann's article that thebrother of Ferdinand Deppe had been selling specimens of this toadunder the name Bufo trachypus. It is a nomen nvdum and has nostanding in zoological nomenclature. The cotypes of Bujo valliceps(M. N. B. Nos. 3525-27, 3532) were collected in Mexico by Ferdi-nand Deppe, who was a member of a party of Germans engaged inmaking botanical collections in Mexico. The following notes weremade on these cotypes:No. 3525, two cotypes collected by Deppe in Mexico. Head-and-body length, 94 mm. and 100 mm., respectively; the former has whiteunderparts and the latter has white underparts with brown marbling.
70 BULLETIN 160, UNITED STATES NATIONAL MUSEUMThese specimens correspond to normal individuals of this species,and have heavy, high, and relatively thick cranial crests. The lateralrow of warts is well developed.No. 3526, one cotype collected by Deppe in Mexico. Head-and-body length, 84 mm. ; underparts white with brown marbling; a dark-colored individual with light blotches on upperparts; cranial crestssharp-edged; parotoid glands sub triangular in outline; a lateral rowof minute warts; inner metatarsal tubercle small, but larger than theouter.No. 3527, one cotype collected by Deppe in Mexico. Underpartswhite, Avith dusky spots; upper{)arts light colored, with dark inter-orbital band, dark spots and dark stripes; cranial crests sharp; innermetatarsal tubercle small, but larger than the outer.No. 3532, one cotype collected by Deppe at Vera Cruz, Mexico.Head-and-body length, 55 mm. Underparts white, with dark mar-bling; upperparts dark, marbled with light colors; cranial crestsblunt-edged and rather thick; parotoid glands subtriangular in out-line; a light-colored vertebral stripe and a light-colored interorbitalband.The type of Bujo granulosus (U.S.N.M. No. 2595) was collectedin 1851 by John H. Clark, a member of Col. J. D. Graham's party ofthe United States and Mexican Boundary Survey, somewhere alongthe old wagon road that extended from Indianola to San Antonio,Tex. Inasmuch as this name is preoccupied, Girard proposed Bu^onebulifer as a substitute name. The specimen (U.S.N.M. No. 2602)figured by Baird in his report on the reptiles of the United States andMexican boundary came from New Braunfels, Comal County, Tex.Giinther's Bufo sternosignatus was based on three adult toads fromVenezuela and two half-grown individuals from Mexico. One ofthese latter (B. M. No. 56. 3. 17. 25) was taken by Auguste Salle atCordoba in Vera Cruz, and the other, labeled as coming from Mexico,was purchased from Parzudaki (B. M. No. 58. 9. 6. 13). The last-mentioned specimen has a head-and-body length of 30 mm. Bou-lenger ^? has shown that the two half-grown Mexican toads are iden-tical with Bufo valliceps and has restricted the name Bufo sternosig-natus to the Venezuela form.More recently Werner has proposed to recognize the Honduran formas a distinct subspecies. The type of Bufo valliceps microtis (M. N.B. No. 13200) is an adult female with a small tympanum and wascollected by Schliiter. The following notes were made on this speci-men : Head-and-body length,73 mm. ; transverse diameter oftympanum,3.7 mm. ; transverse diameter of eye, 9 mm. ; cranial crests high, sharp-edged; thin, sharp-edged, and nearly vertical preorbital and postor-bital crests; parotoid glands protuberant; general coloration of upper-
? Boulenger, Q. A., Catalogue of the Batracbla Salientia s. Ecaudata in the collection of the BritishMuseum, 2d edit., p. 323, 1882.
MEXICAN TAILLESS AMPHIBIANS 71parts light; upper lip white; a wide white interorbital band, borderedanteriorly and posteriorly by a black stripe; white vertebral stripeinterrupted and irregular in width; a lateral white stripe; underpartsyellowish, with dusky marbling; a lateral row of small warts; skin ofupperparts tubercular; bony surface of top of head fairly smooth.In the southern part of the range of Bujo valliceps specimens will befound that have a much yellower ground color than typical individ-uals and these usually have rather conspicuous dark-brown spotsirregularly arranged on the upperparts. They often occur along withother specimens that exhibit the normal coloration of this species.Females generally have a smaller tympanum than males. In a largeseries of specimens considerable variation in the size of the tympanumwill be noted.Specimens examined.?One hundred and thirty-three, as follows:Bufo valliceps
Museum
M.C.Z.U.S.N.M..Do...Do.Do.Do.Do.DoF.M.N.H..M.C.ZA.M.N.H..B.MU.S.N.M..Do....Do.-..C.O.E.M..U.S.N.M..Do....A.M.N.H.Do....U.S.N.M..M.C.ZF.M.N.H...A.M.N.H...DoC.A.SDo....Do....Do....M.N.B--.-A.M.N.H.Do....Do....U.S.N.M..C.O.E.M.M.C.Z
Catalogue-No.
47168259947081-834691847053-5542501-04259114818447-51137565a 3. 17. 251217025094-96710564723030360-69137921376316548-5110298168713768-7213774-775083529954685563353212306-08122921234046948-492630
Num-ber ofspeci-mens Locality collectedCoahuila: MoDclovaNUEVO Leon:Cerro de la SillaSanta CatarinaTamaulipas:Alta Mira ?HidalgoJaumaveLa Barra, 8 kilo, east of Tam-pico.MatamorosVera Cruz:AchotalCerro del QalloCocos...Cordoba ?MiradordodoMotzoronga..Orizabadododo...Vicinity of OrizabaPanuco--PerezPotrero ?North of Rio Atoyac nearPotrero.TezonapaVera Cruzdododododo..Vera Cruz (2 miles south)...Natarit: AcaponetaPuebla: PueblaMorelos: Cuernavaca
Date collected
Mar. 27, 1902Apr. ?1853Apr. 23,1898Mar. 19,1891June 4,1898Mar. ?, 186319041921June 22,1919
Jan. 19, 1894June 25,1919do19231903June 19, 1919June 18,1919July 14,1925July 24,1925July 23,1925July 25,1926June 3, 1919June 2, 1919Mar. 1,1919/June 30,1897IJuly 2, 1897
By whom collected orfrom whom received
1 Young.
' Cotypes of Bujo sternosignatus.
? Cotypes of Bufo valliceps.
Edward Palmer.Nelson and Goldman.D. N. Couch.Nelson and Goldman.William Lloyd.Nelson and Goldman.Edward Palmer.D. N. Couch. andEdmund HellerC. M. Barber.E. R. Dunn.Paul D. R. Ruthling.Augusta Sallfe.C. Sartorius.Do.Do.Nelson and Goldman.Francis Sumichrast.Paul D. R. Ruthling.Do.Francis Sumichrast.W. W. Brown.S. E. Meek.Paul D. R. Ruthling.Do.Joseph R. Slevin.Do.Do.Do.Ferdinand Deppe.Paul D. R. Ruthling.Do.Do.[Nelson and Goldman.T. Barbour.
72 BULLETIN 160, UNITED STATES NATIONAL MUSEUMBufo valliceps?Continued
Museum
U.S.N.MC.A.SDo.DoDoDoDoDoU.S.N.M...-DoDoDoDo.Do.Do.Do.Do.Do.Do.Do.Do.Do.M.N.B 3525-27B.M I 58.9.6.13
CatalogueNo.
46967503850395091-975099-5104517051875190-9147117-184712347125-2647127-281001330180-8146766-69467614685147311377471228512292-93
Num-ber ofspeci-mens Locality collectedOaxaca:Choapam.Cosolapa..
....dodo
....do
....do
....do
.-..doGuichicovi.Santo Domingodo.-.Mountains near Santo Do-mingo.Tapana...TehuantepecChiapas:OcozocoautlaOcuilapaTumbalaValley of ComitanTabasco:TeapaFronteraYucatan:No definite locality
---.do? State:No definite locality
....do
Date collected
July 28,1894July 12,1925July 13,1925July 14,1925
-...doJuly 16, 1925July 19, 1925July 20, 1925/June 24, 1895-IJune 26, 1895June 11,1895
-...doJune 19, 1895Feb. ?1876
Aug. 20, 1895Aug. 27,1895Nov. 16, 1895Dec. 9, 1895
.\pr. 15,1900Mar. 12,1897
By whom collected orfrom whom received
Nelson and Goldman.Joseph R. Slevin.Do.Do.Do.Do.Do.Do.[Nelson and Goldman.Do.Do.Do.Francis Sumichrast.Do.Nelson and Goldman.Do.Do.Do.Do.C. H. T. Townsend.Arthur Schott.Do.Ferdinand Deppe.Emile Parzudaki.
' Cotypes of Bufo sternosignatus. > Cotjrpes of Bufo valliceps.BUFO WOODHOUSn GirardFigure 161852. Bufo dorsalis Hallowell, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, p181, Oct. (New Mexico). [Nee Bufo dorsalis Spix, 1824.]
?
Hallowell1853, Sitgreaves's report of an expedition down the Zuni and Colorado RiversU. S. Senate Doc. 59, 32d Cong., 2d Sess., p. 142, pi. 19.1854. Bufo woodhousii Girard, Proc. Acad. Nat. Sci. Philadelphia, vol. 7, p86, May.?Stoker, 1925, Univ. California Publ. Zool., vol. 27, pp. 199-203pi. 11, fig. 31b.1866. Bufo frontosus Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 18, p. 301Oct.?Cope, 1889, U. S. Nat. Mus. Bull. 34, p. 283.1875. Bufo lentiginosus frontosus Yarrow, Report upon geographical and geo-logical explorations and surveys west of the One Hundredth Meridian(Wheeler survey), vol. 5, chap. 4, p. 520.
?
Coues, 1875, ^de7n, chap. 5, pp. 627, 628.1889. Bufo lentiginosus woodhousei Cope, U. S. Nat. Mus. Bull. 34, p. 281, fig.69.?Dickerson, 1906, The frog book, pp. 91-93, figs. 68-74, col. pi. 4, fig.2.
?
Nieden, 1923, Das Tierreich, Lief. 46, Amphibia, Anura I, p. 126.Type locality.?"New Mexico" [= San Francisco Mountain, Co-conino County, Ariz.].Range.?From the Missouri River in Montana (north of Wilder)and in North Dakota (Goodall), west to near the junction of theUmatilla and Columbia Rivers (Umatilla); south through westernIdaho (Boise), northwestern Nevada (Humboldt County), south-western Utah (Washington County), to the Lower Colorado River
MEXICAN TAILLESS AMPHIBIANS 73(Yuma), the Salton Sea Basin (Agua Dulce), southern Arizona(Nogales), and northern Sonora; and east through the Rocky Moun-tain region to the Badlands of South Dakota, the Loup fork of thePlatte River in Nebraska, northeastern Kansas (Doniphan County),the Arkansas River in eastern Oklahoma (Okmulgee, Pushmataha,and McCurtain Counties), southern Texas (Haj^s and Bexar Counties)and along the Rio Grande west of the mouth of the Pecos River(Brewster and El Paso Counties) ; and thence to southern Chihuahua(vSanta Rosalia).Remarks.?The misstatements in regard to the type of Bufo dorsalisand the assumed type of Bufo woodhousii published by Cope in his
"Batrachia of North America" havebeen corrected by Doctor Stejneger.^'The actual type of Bujo dorsalis (U.S.N.M. No. 2531) is still extant andwas collected in October, 1851, on SanFrancisco Mountain, Coconino County,Ariz., by Dr. S. W. Woodhouse, whoaccompanied Capt. L. Sitgreaves onthe expedition down the Zuni and Col-orado Rivers. Girard found that thename Bvfo dorsalis was preoccupiedby an earlier application of dorsalis bySpix to a toad found in the provinceof Rio de Janeiro, Brazil, and thereforehe proposed Bvfo woodhousii as a sub-stitute name. Both of these specificnames are based on the same typespecimen.No specific type locality for Bufofrontosus is given in the original de-scription, except for a rather generalone?"Territory of Arizona"? wliichappears in the introductory remarks,reprinted by Dr. Elliott Coues in his "Synopsis of the Reptiles andBatracliians of Arizona," and the following is added: "One speci-men from Los Pinos, Rio Grande, N. Mex. [ = Valencia County, N.Mex.l; type of the species." This specimen was collected in 1864by Coues, whose collection and field notes were placed in the hands ofProfessor Cope. There is no record that indicates that the type ofBufo frontosus was ever sent to the National Museum by Coues, andit is possible that it may have been sent direct to Cope. At any event
? Stejneger, L., Annotated list of reptiles and batrachians collected by Dr. C. Hart Merriam and VernonBailey on the San Francisco Mountain Plateau and desert of the Little Colorado, Arizona, with descriptionsof new species. North Amer. Fauna No. 3, pt. 5, pp. 116, 117, Sept. 11, 1890.
? Ob'^ o
' -^ bFigure 16.?Head of Bujo woodhousii (U.S.N.M. No. 2531) from San Francisco Moun-tain, Ariz.: a, Lateral view; b, dorsal view.Type specimenThe original description is
74 BULLETIN 160, UNITED STATES NATIONAL MUSEUMthe type has never been recognized among the unlabeled specimensreturned to the National Museum by the executors of Cope's estate.The habitat of Bujo woodhousii is by no means restricted to moun-tainous regions, but includes surroundings as diverse as the sagebrushflats of eastern Montana, the prairie fields among the chalk cliffs ofwestern Kansas, the Hudsonian Zone mountain sides of eastern Colo-rado, the irrigation ditches that traverse the mesquite plains of NewMexico, and the bottomlands along the Colorado River near Yuma,Ariz. During May and June, according to locality, adults of thisspecies may be found breeding in shallow, sluggish creeks, in irriga-tion ditches, or in fresh-water pools in the canyons.The upperparts of this toad are generally grayish or dull yellowishbrown, conspicuously blotched with a darker color or without suchmarkings; vertebral stripe wliitish; limbs obscurely barred; under-parts light yellowish, with or without black markings on breast;largest warts red-tipped and encircled at base with narrow blackborder. The cranial crests are distinct, but not high; canthus rostra
-
lis generally distinct; interorbital space either concave or convex,depending upon the approximation of supraciliary crests, which mayeither parallel one another or diverge slightly posteriorly; parietalcrests generally lacking, but supracihary crests may be thickened attheir posterior extremities; parotoid glands often in contact with post-orbital crests, about twice as long as broad, their maximum diameterbeing almost twice the length of the orbit; inner metatarsal tuberclelarge, with free cutting edge.Specimens examined.?Two as follows
:
Bufo woodhousii
Museum
MEXICAN TAILLESS AMPHIBIANS 75
mining what is a family character. The phylogenetic importance ofthe various characters that have been selected from time to time toexpress relationships of supposedly allied groups of genera is in mostcases rather doubtful, for the grouping is based solely on recentgenera. In view of the admitted deficiency in the paleontologicalrecord of an undoubtedly old group like the Anura, a classificationbased upon structures that have been either remodeled or modifiedmay fail to eliminate all cases of convergent and parallel develop-ment. By definition the limits of families can be extended orrestricted as the taxonomist so desires. Difficulties familiar to allare encountered when one attempts to set up arbitrary limits orstructural features that will invariably characterize all genera of thefamily. If we grant that families and genera are derived from oldertypes, then we may expect to encounter overlapping structuralfeatures in related groups.It may be conceded that any classification based on dentitiondoes not adequately express true relationships. There seems to bean evolutionary tendency in some groups toward the elimination ofteeth and equivalent stages in this process do not necessarily indicaterelationship. Engystomops and Bufo lack maxillary and vomerineteeth. There are, however, important differences in the mode of lifehistory. Engystomops and Bufo are very much alike externally, batthe former lacks the characteristic protruding parotoid glands of thelatter, and furthermore their life histories are quite unlike. The toadsof the genus Bufo lay their eggs in long or short strings in the water,while the leptodactylids lay theirs in masses or clusters either on landor floating on the water's surface.According to Noble,^^ Syrrhophus "is a polyphyletic group of speciesof Eleutherodactylus which have independently lost the vomerineteeth." Granted that this is the case, a stage in this general processis represented by Eleutherodactylus mexicanus, in which all steps fromtoothless to a normal vomerine series may be encountered in a seriesof individuals. Both Eleutherodactylus and Syrrhophus have T-shaped terminal phalanges and the same type of sternal construction,including a cartilaginous style. In Leptodactylus the mesosternum hasossified. Tomodactylus seems to be related to Eleutherodactylus andSyrrhophus, since it has T-shaped terminal phalanges and a cartilagi-nous mesosternum, but lacks vomerine teeth. Engystomops is moredistantly related to the Mexican leptodactylids, for it lacks bothvomerine and maxillary teeth and possesses terminal phalanges -wathnodular extremity, as well as a straplike bony mesosternum witheach branch of distal bifurcation furnished with a cartilaginous plate.The egg-laying habits of these leptodactylids indicate that theyhave taken advantage of the limitations of their mode of life. Of
" Noble, G. K., Bull. Amer. Mus. Nat. Hist., vol. 46. art. 1, p. 18, 1922.
76 BULLETIN 160, UNITED STATES NATIONAL MUSEUMthose in which development is indirect, Engystomops makes floating,foamy nests for the eggs, while Leptodadylus forms froth nests incontact with water. Although no observations have been publishedon the life histories of the Mexican species of Eleutherodactylus, itis quite likely that some of the species have the direct development inwhich the young hatch full}^ formed from the egg capsules. The lifehistories of Tomodactylus and of the Mexican species of Syrrhophushave not been observed.KEY TO GENERA OF MEXICAN LEPTODACTYLIDAE
1. Vomerine and maxillary teeth present 3Vomerine teeth absent 22. Maxillary teeth low, projecting very little if at all bej^ond outerridge of this bone, and often barely penetrating the gum 5Maxillary teeth absent; a diamond-shaped area inclosed byglandular dermal ridges, with black outer border and brown,orange, or vermilion-colored center in interscapular region;omosternum cartilaginous, with terminal dilation;mesosternum ossified, forming a straplike bony style,which is bifurcated distally but not proximally, andeach of the branches is furnished with a separate car-tilaginous plate (xiphisternum) ; sacral diapophysesmoderately dilated; skin of upperparts decidedlytubercular and glandular; tympanum minute or hid-FiGURE 17? ^^^ ^y tubercles; a conical tubercle on middle of innerThe T-shaped edge of tarsus; fingers free; toes nearly free; ends ofterminal j3ha- fingers and toes slightly swollen, no distinct terminaldisks; outer metatarsals united; terminal phalangeswith peculiar nodular extremity and rarely withpseudo-T-shaped distal end Engystomops (p. 77)3. Mesosternum ossified, forming a bony style; fingers and toes free,without distinct distal dila,tions or disks; male of Leptodactylnsmelanonotus with two spine-bearing tubercles on inner side offirst finger; outer metatarsals united; tympanum distinct;pupil horizontal Leptodactylus (p. 81)Mesosternum not forming a bony style, but cartilaginous, withbisagittate (double arrow-shaped) extremity or xiphisternum 44. An external tj^mpanum, more or less distinctly outlined; fingersfree; toes either free or slightly webbed; terminal phalangesT-shaped (fig. 17); digital dilations or disks present, althoughgenerally small, and rarely as large as tympanum; outer meta-tarsals united; pupil horizontal Eleutherodactylus (p. 90)No external indication of a tj-mpanvmi; fingers with or withoutvestige of web at base; toes nearly fully webbed; terminalphalanges T-shaped; apical disks of fingers and toes welldeveloped; body stout; hind limbs rather short; male ofCawphias guatemalensis with well-developed clawlike vestigeof pollex buried in fleshy apophysis on side of first finger. Cauphias (p. 118)5. Large lumbar gland present, barely visible above the skin, itsposition marked by an irregular dark brown or black area,with two or three white spots, or with white vermiculations;outer metatarsals united; fingers and toes free, with smalldistally truncated dilations or disks; terminal phalanges
lanx of FAeu-therodaclylus
MEXICAN TAILLESS AMPHIBIANS 77T-shaped; tympanum distinct; cartilaginous omosternum andmesosternum; mesosternum sometimes forms a straplike bonystyle, possibly an age character Tomodactylus (p. 1 20)No lumbar gland present; mesosternum not forming a bonystyle but cartilaginous, straplike, and often spatulate; toesfree; apical disks, if developed, generally small; terminalphalanges T-shaped; subarticular tubercles generally extend-ing backward on plantar surface; outer metatarsals united;tympanum distinct; pupil horizontal Syrrhophus (p. 123)Genus ENGYSTOMOPS Espada1872. EngTjstomops Jimenez de la Espada, Anales Soc. Espan. Hist. Nat., vol. 1,p. 86. [Genotype, Engysiomops petersi Jimenez de la Espada, idem, p. 86(Provinciade Oriente, Ecuador).]
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Jimenez de la Espada, 1875, Vertebradosdel Viaje al Pacifico, etc., Batracios, pi. 2, figs. 3-3d.1873. Microphryne Peters, Monatsber. k. preuss. Akad. Wists. Berlin, p. 616,Oct. [Genotype, Paludicola pusiulosa Cope, 1864, Proc. Acad. Nat. Sci.Philadelphia, vol. 16, p. 180, Aug. (Truando River, New Granada,Colombia).]1875. Peralaimos Jimenez de la Espada, Vertebrados del Viaje al Pacifico, etc.,Batracios, p. 163. [Genotype, Peralaimos stentor Jimenez de la Espada,idem, p. 163, pi. 2, figs. 4, 4b (Isla de Tobaga) = B?/o stentor Jimenez de laEspada, 1875, Anales Soc. Espan. Hist. Nat., vol. 1, p. 85.]In 1882, Boulenger ^^ recognized Engystomops as a valid genus thatincluded the following species: petersii, pustulosus, and stentor.Boulenger stated in his catalogue that Eupemphix nattereri seemedto agree with Paludicola Juscomaculata. Six j^ears later he ^^ revisedhis views on this group and published the following explanatorystatement
:
The genus Eupemphix, Stdr. {= Engystornops, in part), must, on account ofthe absence of teeth, be referred to the family Bufonidae, although it is in everyother respect identical with Paludicola, to which it stands in the same relationas Pseudophryne to Crinia; this shows that frog-families founded upon thepresence or absence of teeth are artificial associations.A key to four species of Eupemphix follows after this statement,two of which, nattereri (the genotype) and nana, were formerly includedin Eupemphix, but pustulosa and stentor were allocated to Engystomops.No reference is made to the allocation of Engystomops petersii.Professor M6hely in 1904 did not concur with the allocation of Eupem-phix to the family Bufonidae, and held that the structural featuresdid not warrant this classification. Hence Eupemphix and Paludicolawere referred to the Cystignatliidae [ = Leptodactylidae].A small group of species, including petersii, stentor, and pustulosus,seems to form a natural assemblage somewhat closely related toPleurodema and Eupemphix. Southern Mexico on the north and
2< Boulenger, G. A., Catalogue of the Batrachia Salientia s. Ecaudata in the collection of the BritishMuseum, 2d edit., pp. 27,5-277, 1882.25 Boulenger, O. A., Descriptions of new Brazilian batrachians. Ann. and Mag. Nat Hist., ser. 6, vol.1, no. 3, p. 188, Mar., 1888.t)G7S5?32 6
78 BULLETIN 160, UNITED STATES NATIONAL MUSEUMEcuador on the south are the known limits of the range of this group.These small species have much more slender bodies than those of eitherPleurodema or Eupemphix, and the sldn on the upperparts is decidedlyglandular. A diamond-shaped mark on the back bounded by glan-dular ridges seems to be a constant character.The sternal apparatus of Engystomops is quite similar to that of thePaludicola signifera figared by M^hely.^^ The omosternum, however,is cartilaginous and not semiossified as in P. signifera, and the bonystyle is bifurcated distally but not proximally also, as in the latter.Each of the branches of the bony style are furnished with a separatecartilaginous plate. The terminal phalanges are rarely simple, butgenerally have a peculiar nodular distal end, and occasionally apseudo-T-shaped distal end.ENGYSTOMOPS PUSTULOSUS (Cope)Figure 181864. Paludicola pustulosa Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 16,p. 180, Aug.1876. Microphryne pustulosa Cope, Journ. Acad. Nat. Sci. Philadelphia, n. 8.,vol. 8, pt. 2, p. 155.1882. Engystomops pustulosus Boulenger, Catalogue of the Batrachia Salientia s.Ecaudata in the collection of the British Museum, 2d edit., p. 275.
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GiJNTHER, 1901, Biologia Centrali-Americana, Reptilia and Batrachia, p.242, Feb.1882. Bufo {Microphryne) pustulosus Brocchi, Mission scientifique au Mexiqueet dans TAm^rique Centrale, recherches zoologiques, pt. 3, sect. 2, p. 86,pi. 8, figs. 1, la-d.1923. Eupemphix pustulosa Nieden, Das Tierreich, Lief. 46, Amphibia, Anura I,p. 165.Type locality.?New Granada, on the River Truando, Colombia.Range.?Interior valleys on the southern side of the Isthmusof Tehuantepec in southern Oaxaca, southward to Panama andVenezuela.Remarks.?Cope designated a specimen in the National Museum(No. 4339) as the type and stated that it was collected by W. S. Woodof Lieutenant Micliler's surveying expedition. The original entryin the museum catalogue for this number gives Darien [Panama] asthe locality and [Arthur] Schott as the collector. The type can notnow be found.Surprisingly little is known in regard to the natural habitat of thiswidely distributed species. According to an annotation in the fieldcatalogue of Nelson and Goldman, this small leptodactylid was foundto be rather common during wet weather in the vicinity of SantoDomingo in Oaxaca at an elevation of 900 feet. It was observedthat they were especially active on rainy nights and that they oftencame into the house where Nelson and Goldman were staying.M M6hely, L. v., Ann. Hist. Nat. Mus. Nat. Himgarici, Budapest, vol. 2, pi. 13, fig. 6, 1904.
MEXICAN TAILLESS AMPHIBIANS 79The following account of the life history of this species has beenpublished by Doctor Noble : ^^The species of Eupemphix which both Mr. Breder and Doctor Chapin have stud-ied, is the wide-spread pustulosus. It has the peculiar shoulder girdle characteristicof the genus and of Paludicola. Breder made his observations in Darien, Panama,during February, March, and April, 1924. Most of his breeding pairs he foundduring the second and third week of March. E. pustulosus lays its eggs in afoamy mass which floats so lightly on the water that it does not break the surfacefilm. No less than thirty-six nests were studied criticallj' by Breder. The nestswere usually at the water's edge, generally floating but adhering to any adjacentobject. The few "nests" that were not in contact with the water had probablybeen left stranded as the water in the pool receded during a drought. Sometimesthe "nests" were partly and, rarely, completely hidden by leaves. The eggsdeveloped normally whether or not the mass was in contact with the water. Doc-tor Chapin found E. pustulosus breeding at Tapia, Canal Zone, August 7, 1923, incircular basins of cement placed around the bases of trees to protect them from leaf-cutting ants and kept filled by rain water. The egg masses werelaid on the edges of this basin exactly as in the case when largerbodies of water were utilized. Other egg masses were found insmall stagnant pools near by or even in hoof marks which hadbecome filled with water.The eggs of E. pustulosus are practically identical with thoseof P. fuscomaculata, being only 1.4 mm. in diameter, unpigmentedand laid in large numbers. Development, however, is slower thanin the latter species and many details which Budgett tells us are
"blurred and, as it were, hurried over" in P. fuscomaculata are ' ,'accentuated in E. pustulosus. In both species the yolk is small, ^but obviously dense, for the embryo soon curves around it in- Figure 18.?Headstead of incorporating it at once into the body as in our common ? , "''^* omopapustulosus (U.frogs and toads. The adhesive organs, which Bles found to arise g j^ j^ -^^separately in Paludicola, differentiate in Eupemphix from a cres- 47124) fromcent as in less specialized frogs {Scaphiopus, etc.). The external Santo Domingo,gills, which remain small in the first genus, soon develop extraor- Oa^aca, Mexico:dinary proportions in the latter. The nasal pits differentiate j' dorsal viewin the latter at the apex of the head and are not visible from theventral surface as in the case of the former. In short, practically every larvalfeature other than general body form differs in Paludicola and Eupemphix duringthe early stages.Here again as in the pelobatids and ranids mentioned above the early larvalstages may show great differences when the other life history features are verysimilar. In the case of Eupemphix and Paludicola the differences, however, arenot due merely to the different stage at which the frontal organ cells begin tosecrete their liquifying fluids, a change in the growth rate of the whole or a partof the Eupemphix embryo must have occurred to produce the large gills and otherdistinctive features of the early larva. Nevertheless, the place of egg deposition,the form of the egg mass and egg-capsules, the structure of the mature larvae andthe general breeding habits are much aUke and conform to our conclusions thatthe genera are closely related.
" Noble, G. K., Ttie value of life history data in the study of the evolution of the Amphibia. Ann.New York Acad. Sci., vol 30, pp. 87, 88, Oct. 31, 1927,
80 BULLETIN 160, UNITED STATES NATIONAL MUSEUMAccording to Ruthven ^^
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The colors of this species have not been accurately described, probably becausethe red fades rapidly in alcohol. The ground color above is gray, relieved byirregular black blotches. Most of the warts on the head and nape and a few onthe body are dull orange (142) 29 or red orange (78c or 78d), as are also the lightcoccygeal line, the broad light area on the fore limbs, and more or less of the lightcross bands on the hind limbs, particularly on the feet. The light spot in theinterscapular region and the one on either side of the back are present in all speci-mens, and, with the line of enlarged warts behind the angle of the mouth, areusually very pale 3'ellow, but occasional!}^ red. The line in the coccygeal regionis usually well defined only to the pair of large blotches in the lumbar region. Itis occasionally continued narrowly to the broad black margin of the interscapularspot, and occasionally bej'ond this to the snout, in which case it is interrupted bythe supraocular band. In the single very young example these three light ver-tebral marks form a narrow line from snout to anus, that is narrowly interruptedin the same places as in the adults. With the exception of a median line, thechin, throat, and breast are gray, generally profusely spotted with black or brownand generally somewhat speckled with pale yellow. The median line, which ispresent even in very young specimens, is whitish, or pale yellow, which is the dom-inant color of tlie abdomen, the dark color of the chin, throat, and breast beingbroken up into large spots on the posterior part of the ventral surface.This little toad was found in about the same places as B. valUceps, but seemedto prefer the wetter places and was more secretive. It was most numerous underlogs and decaying vegetation in a low partially cleared area along La Laja Creek,but it was also found under logs on the shore of the prairie ponds, and in the bananagrove at San Juan. During the rains it was found at night in the pools, and animmature specimen that had but recently emerged from the tadpole stage wasfound on the shores of a temporary pool on La Laja Creek, August 14. The noteis a low chirp that is repeated regularly at intervals of one second.The species that occurs in Mexico may be recognized by the fol-lowing combination of characters : A diamond-shaped area inclosed byglandular dermal ridges, with black outer border and brown, orange,or vermilion center in interscapular region ; a short light-colored sacralstripe; coloration of remainder of upperparts grayish olive or blackishbrown with scattered darker markings; fore and hind limbs and feetbroadly banded with pinkish gray or brownish; under surfaces oflimbs yellowish; underparts yellowish or white, marbled or spottedwith black; throat and breast heavily mottled with grayish or brown-ish, frequently bisected by longitudinal pin stripe and occasionally bytransverse stripe of same color; neither vomerine nor maxillary teeth;tympanum minute and distinct, or for the most part hidden bytubercles; fingers slender and free; first finger not extending beyondsecond; no subarticular tubercle on penultimate finger joint; smallsupernumerary tubercles continued backward on palmar surface; twopalmar callosities; no prominent terminal disks; toes not unusuallylong, free, and with lateral dermal ridges extending to distal phalanx
;
28 Ruthven, A. G., The amphibians and reptiles collected by the University of Michigan-Walker Expe-dition in southern Vera Cruz, Mexico. Zool. Jahrb. (Syst. Abt.), vol. 32, pt. 4, pp. 308, 309, 1912." The numbers refer to the "Code des Couleurs," by Klincksiek and Vallete.
MEXICAN TAILLESS AMPHIBIANS 81third toe slightly longer than fifth; prominent subarticular tubercles;supernumerary tubercles continued backward on plantar surface; aconical tubercle on the middle oj inner edge of tarsus marks the end of thetarsal fold; two small metatarsal tubercles, the inner larger than theouter; the hind limb being carried forward along the body, the tibio-tarsal joint barely reaches beyond insertion of fore limb; head small;snout prominent, acuminate; width of interorbital space greater thanthat of upper eyelid; canthus rostralis rounded; loreal region sub-vertical; large vocal sac; skin of upperparts, including top of head andupper surfaces of fore and hind limbs, pustulose or tubercular andexceedingly glandular; throat smooth; belly areolate; under surfacesof thighs granular; head-and-body length, 27 to 30 mm.Specimens examined.?Thirty-four, as follows
:
Engystomops pustulosvs
Museum
C.A.SU.S.N.M...DoDo
CatalogueNo.
5108-51234712410023-2830247-57
Num-ber ofspeci-mens Locality collectedOaxapa:CosolapaSanto Domingo (900 feetaltitude).Tehuantepee
Date collected
July 14, 1925June 10, 1895
By whom collected orfrom whom received
Joseph R. Slevin.Nelson and Goldman.Francis Sumichrast.Do.Genus LEPTODACTYLUS Fitzinger1826. Leptodadylus Fitzinger, Neue Classification der Reptilien, p. 38. [Geno-type, Lepiodactylus typhonia Fitzinger= /?aMa typhonia (part) Daudin, 1803,Histoire naturelle des rainettes, des grenouilles et des crapauds, p. 36, pi. 17,figs. 3, 4 (Surinam).] (Type fixed by Fitzinger, 1843, Systema reptilium,fasc. 1, p. 31; nee Rana typhonia Linnaeus, 1758.)1830. Cystignathus Wagler, Naturliches System der Amphibien, p. 202. [Geno-type, Cystignathus pachypics Wagler= ^ana pachypus Spix, 1824, Animalianova, sive species novae testudinum et ranarum quas in itinere per Brasiliam,etc., p. 26, pi. 2, figs. 1, 2 (Brazil).] (Type fixed by Fitzinger, 1843,Systema reptilium, fasc. 1, p. 31.)1843. Gnathophysa Fitzinger, Systema reptilium, fasc. 1, p. 31. [Genotype,Cystignathus labyrinthicus Dumeril and Bibron, 1841, Erp^tologie generate,vol. 8, p. ^07= Rana labyrinthica Spix, 1824, Animalia nova, sive speciesnovae testudinum et ranarum quas in itinere per Brasiliam, etc., p. 31, pi. 7,figs. 1, 2 (Brazil) = /?ana pentadadyla Laurenti, 1768, Synopsin reptilium,p. 32, "in Indiis" (erroneous).]1870. Entomoglossus Peters, Monatsber. k. preuss. Akad. Wiss. Berlin, p. 647.Aug. [Genotype, Enio7noglossus pustulatus Peters, idem, p. 647, pi. 2, fig. 1(Ceara, Maranhao, northern Brazil).]Various devices have been tried out by amphibians to insure thedevelopment of their eggs. Leptodadylus seems to have adoptedfroth nests for this purpose. These are generally made in water ornear water in damp hollows. One South American species lays itseggs on the surface of pools, while another deposits them in holes near
82 BULLETIN 160, UNITED STATES NATIONAL MUSEUMthe water's edge. No observations on the breeding habits of Lepto-dadylus in Mexico have been published, but it is quite probable thatthe breeding habits of L. albilabris in southern Mexico agree withthose of their West Indian relatives. Doctor Stejneger has reportedthe finding of a remarkable foamy *'nest" of L. albilabris, "withnumerous small black larvae just hatched," on April 7 in a smallstream near Utuado, Porto Rico.KEY TO MEXICAN SPECIES OF LEPTODACTYLUS1. A white stripe on upper lip, most conspicuous in front of eye,commencing on snout and extending backward to insertionof fore limb, and touching inferior margin of eye and tym-panum; a dark stripe running backward along canthusrostralis from tip of snout to eye and from eye to abovetympanum and then downward to above insertion of forelimb; upperparts clay-colored, reddish brown, or a pale orange-brown with four longitudinal rows of more or less irregularlyplaced dark brown or black spots; a W-shaped mark extendinglaterally upon upper eyelids in posterior interorbital region;underparts white; distal phalanges of fingers without distinctdilations or disks; first finger longer than second; fingers notwebbed at base; two palmar callosities; subarticular tuberclescontinued backward on palmar surface; toes in order ofdecreasing length, 4, 3, 5, 2, and 1, with vestigial well at base;subarticular tubercles continued backward on plantar sur-face; two metatarsal tubercles, the inner connected with aslight but distinct tarsal fold; the hind limb being carriedforward along the body, the tibio-tarsal joint reaches at mostto anterior margin of eye; vomerine teeth in two transverseslightly curved series behind and generally within the levelof inner margins of the choanae; tongue large, ovoidal, andslightly notched behind; canthus rostralis rounded; lorealregion oblique; snout pointed and projecting beyond upperlip; interorbital space equals width of upper eyelid; tympanumgenerally about two-thirds and always more than one-halfthe diameter of the eye, overhung by a dermal fold; skin ofupperparts smooth; numerous small pointed tubercles onouter surface of tibia; a narrow interrupted dorso-lateraldermal fold and another similar one, though less pronounced,on side from shoulder to groin; abdominal disk distinct;ventral surfaces of thighs granulated; head-and-body length,33 to 36 mm albilabris (p. 83>No white stripe on upper lip anterior to eye; adults with shortwhite line below eye and often with short posteriorly directedoblique continuation of the same; coloration of upperpartsdark brown or brownish orange, with as many as four longi-tudinal rows of elongate black spots; a black, light-margined,
"butterfly-shaped" mark extending laterally upon uppereyelids in posterior interorbital region; hind limbs distinctlycross banded with a darker color; fore limbs with dark spots;a dark stripe extending from nostril backward along canthusrostralis to eye and from eye to above tympanum and thendownward; sides, throat, belly, and under surface of hind
MEXICAN TAILLESS AMPHIBIANS 83limbs reticulated, marbled, or spotted with brown; distalphalanges of fingers very slightly if at all dilated; first fingeras long as or longer than second; fingers not webbed at base;palmar callosities fairly distinct; subarticular tubercles con-tinued backward upon palmar surface; male with two spine-bearing tubercles on inner side of first digit; toes in order ofdecreasing length, 4, 3, 5, 2, and 1, with vestigial web at base;subarticular tubercles not continued backward upon plantarsurface; two metatarsal tubercles, the inner connected withdistinct tarsal fold; legs stout; the hind limb being carriedforward along the body, the tibio-tarsal joint reaches at leastto the tympanum and often to eye; vomerine teeth in twotransverse series behind the choanae; tongue small, ovoidal,and slightly notched behind; canthus rostralis indistinct; lorealregion oblique; tympanum one-half to two-thirds the diameterof the eye, overhung by a dermal fold; skin of upperparts,sides of body, and upper surfaces of hind limbs with longitu-dinal series of small tubercles; ventral surfaces of thighsgranulated; head-and-body length, 35 to 46 mm melanonotus (p. 88)LEPTODACTYLUS ALBILABRIS (Gunther)1859. Cystignathus albilabris Gunther, Ann. and Mag. Nat. Hist., ser. 3, vol. 4,no 21. p. 217.1877. Cystignathus labialis Cope, Proc. Amer. Philos. Soc, vol. 17, no. 100, p. 90,July 20 (type locality uncertain, probably Mexico).1877. Cystignathus fragilis Brocchi, Bull. Soc. Philom. Paris, ser. 7, vol. 1, no. 4,p. 182 (Tehuantepec, Mexico).1879. C[ystignathus] gracilis Cope, Proc. Amer. Philos. Soc, vol. 18, no. 104,pp. 269, 270. June 20. [Nee Cystignathus gracilis Dumeril and Bibbon,1841, Erp6tologie g^n^rale, vol. 8, pp. 406, 407 (Montevideo).]1882. Leptodactylus fragilis Brocchi, Mission scientifique au Mexique et dansI'Am^rique Centrale, recherches zoologiques, pt. 3, sect. 2, p. 19, pi. 5, figs.2-2b.1900. Leptodactylus albilabris GtJXTHER, Biologia Centrali-Americana, Reptiliaand Batrachia, p. 213, Apr.Type locality.?St. Thomas, West Indies.Range.?From Cordoba in Vera Cruz south to Tekanto in Yucatanand southward through Central America to Nicaragua; apparentlyrestricted to the State of Oaxaca on the Pacific coast of Mexico.Remarks.?The occurrence of this frog on West Indian islands, aswell as on the mainland of tropical America, still remains one of theunexplained problems of distributional zoology. Some herpetolo-gists have advocated the recognition of the mainland form as a spe-cies distinct from the one that occurs on a number of the islands.None of the diagnostic criteria thus far suggested will even in themajority of cases serve as a means for the identification of specimensmthout reference to the locality. A reexamination of the types ofthe species included above in the synonymy and a critical comparisonof specimens from the mainland with those from the islands failed toindicate any reliable criteria for such a distinction.
84 BULLETIN 160, UNITED STATES NATIONAL MUSEUMThe following notes were made on the cotypes of Cystignathusalbilabris :B.M. Nos. 60. 4. 18. 61-64, St. Thomas, Riises Coll.: One adultand three young (head-and-body length of largest being 17.2 mm.).Adult individual : Head-and-body length, 41.2 mm.; transverse diam-eter of tympanum, 2.5 mm.; transverse diameter of eye, 4.7 mm.;anterior edge of eye to nostril, 3.3 mm.; the hind limb being carriedforward along the body, the tibio-tarsal joint reaches to center oftympanum; a vertical white line on lower eyelid; white stripe onupper lip indistinct, although it is present on the three young; adistinct tarsal fold; large inner and small outer metatarsal tubercles;long longitudinal dermal folds on sides, with intercalated linear seriesof smaller dermal ridges, in addition to others irregularly placed;white reticulations appear through brown on the sides; underside oftibia spotted; vomerine teeth in transverse rows far behind level ofchoanae.B.M. Nos. 60. 4. 18. 65-67, St. Thomas, Ruses Coll.: One adultand three young (the head-and-body length of largest being 17.2mm.). Adult individual: Head-and-body length, 42 mm.; trans-verse diameter of tympanum, 2.8 mm.; transverse diameter of eye,4.8 mm. ; anterior edge of eye to nostril, 3.8 mm. ; the hind limb beingcarried forward along the body, the tibio-tarsal joint reaches to poste-rior margin of orbit; white stripe on upper lip indistinct, although it ispresent on the three young; other characters the same as in the pre-ceding cotype, except that the sides are more nearly smooth, therebeing a few scattered dermal nodular elevations in place of dermalridges; throat, sides of chest, and concealed surfaces of tibia thicklyspotted with small brown specks; vomerine teeth in slightly archedtransverse rows far behind level of choanae.B.M. No. 60. 4. 18. 68, St. Thomas, Riises Coll.: One adult andtwo young (the head-and-body length of largest being 17 mm.).Adult individual: Head-and-body length, 36.8 mm.; transverse diam-eter of tympanum, 2.2 mm.; transverse diameter of eye, 4.5 mm.;anterior edge of eye to nostril, 3.4 mm.; the hind limb being carriedforward along the body, the tibio-tarsal joint reaches to center of eye;white stripe on upper lip indistinct, although it is present on the twoyoung; a sharp tarsal fold; toes with vestigial web at base; throatand sides of chest with closely approximated small brown spots; skinon sides fairly smooth, with no distinct longitudinal dermal folds orridges, though there are a few scattered nodular elevations; minutesupernumerary tubercles on plantar surface of foot; vomerine teethin transverse rows far behind level of choanae.Cope states that his supposedly new species, Cystignathus labialis(U.S.N.M. Nos. 31300-05), was based on young specimens in which
MEXICAN TAILLESS AMPHIBIANS 85the posterior limbs are not so long as in adults. In 1879 Cope ^?referred to "numerous specimens from Tehuantepee, wliich fix thecharacters and the locality." One of the six cotypes (U.S.N.M.No. 31302) agrees with the measurements given by Cope in theoriginal description of C. labialis, but inasmuch as all of them arequite young individuals the designation of an electotype will notsimplify taxonomic procedure. The color markings of these youngindividuals are either indistinct or obliterated by action of the pre-servative. The rows of tiny asperities or tubercles on back and onupper surfaces of the hind limbs are quite distinct.The specimens identified as C. gracilis Dum^ril and Bibron, byCope (U.S.N.M. Nos. 10018-19), were collected in Tehuantepec,Mexico, by Francis Sumichrast, who wrote that the specimens for-warded to the National Museum, as well as those sent to the ParisMuseum, were found under old logs and stones near the water.The type of Cystignathus fragilis (M.H.N.P. No. 195a, parchmentlabel No. 6316) is labeled as having been collected by Bocourt inTehuantepec, Mexico. The following notes were made on thisspecimen: Head and body length, 35.2 mm.; transverse diameterof tympanum, 2.6 mm.; tympanum two-thirds the diameter of theeye; the hind limb being carried forward along the body, the tibio-tarsal joint reaches to center of eye; a tarsal fold present; vomerineteeth in two transverse rows behind the level of the choanae.Ives ^^ reports the finding of four specimens of Leptodactyluslahixilis near a water trough on a hacienda near Tekanto, Yucatan,and Noble ^^ has recorded the accession of a large series from severallocalities in Nicaragua.Schmidt ^^ dissents from the prevailing opinion that the MexicanL. labialis is identical with the Porto Rican L. albilabris, which alsooccurs on several of the Virgin Islands, and says:I am unable to agree with Stejneger and Noble, who have compared theCentral American L. labialis with this species, that they are identical. In aseries of many hundreds of specimens from Nicaragua in the American Museumof Natural History, there is no approach to the adult size of the Porto Ricanspecimens; other differences have been pointed out by Noble (1918, Bull. Amer.Mus. Nat. Hist., Vol. XXXVIII, p. 323); and a comparison of the voices andbreeding habits of the two species remains to be made. They are so closelyrelated, however, that the problem of distribution is scarcely altered by thisview.Noble, in 1918, after handling a large series of these frogs fromNicaragua, called attention to three characters that he thought
aoCope, E.D., Eleventh contribution to the heipetolog>' of tropical .\merica. Proc. Amer. Philos.Sec, vol. 18, no. 104, p. 270, June 20, 1879.31 Ives, J. E., Reptiles and batrachians from northern Yucatan and Mexico. I'roc. Acad. Nat. Sci.Philadelphia, 1891, p. 461.22 Noble, G. K., Bull. Amer. Mus. Nat. Hist., vol. 38, art. 10, p. 323, 1918.33 Schmidt, K. P., Scientific survey of Porto Rico and the Virgin Islands. New York Acad. Sc:., vol.10, pt. 1, Amphibians and land reptiles, pp. 38, 39, 1928.
86 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
would separate average individuals on the mainland from those onthe islands. These distinctions, however, are qualified by the remarkthat there are individuals that are so similar that no distinguishingcharacters are apparent. The series of specimens from St. ThomasIsland, West Indies, and from the mainland of Mexico available forcomparison is not very large, but nevertheless the distinctions madeby Noble fail to separate these lots into two categories. The speci-mens from St. Thomas Island used in these comparisons do not havethe skin on the tibia any more noticeably studded with numeroussharp-pointed tubercles than those from the mainland, and the throatis stippled with brown around the edges. Furthermore, many ofthe mainland specimens have light-margined brown spots on the upper-parts, and the color pattern is quite if not almost identical with theisland specimens. It is true, however, that specimens from St.Thomas Island are somewhat larger than any of those collected inMexico, but otherwise they are inseparable from the latter.In commenting upon the relations and origin of the Porto Ricanherpetological fauna. Doctor Stejneger ^^ says:The status of the frog, Leptodadylus albilabris, is quite different from that ofthe other species of the fauna. In the Antilles it is restricted to the VirginIslands, St. Croix, Vieques, and Porto Rico. There is apparently no indigenousspecies of the genus in the Caribbean chain, the other two which occur in variousislands being evidently introduced by man and belonging to widely differentSouth American species. No Leptodadylus or related form is found in Haiti,Cuba, or Jamaica. But the most curious feature is that a frog w^hich neitherDr. Boulenger nor I can distinguish from L. albilabris, is a native of southernMexico, State of Vera Cruz, and the Isthmus of Tehuantepec. I know of abso-lutely no parallel to this extraordinary range, which is inexplicable on ordinarydistributional grounds, for certainly it would transgress the boundaries of theprobable to suppose either that this species had once covered the whole countrybetween Tehuantepec and the Virgin Islands and become extinct in the interme-diate territory, or that there had at any time been a direct connection betweenthe localities mentioned to the exclusion of the large Antilles. Nor can it for amoment be supposed that the species exists in the latter without having attractedattention. No doubt there are many species yet to be discovered in these islands,but L. albilabris is not likely to be one of them, for it is one of the commonest,most obtrusive, and most easily caught batrachians wherever it occurs. Insuggesting accidental introduction by man I am fully aware that this explanationdoes not at first appear plausible, as there does not seem to be or to have beenany direct route of communication between southern Mexico or Yucatan andPorto Rico or the Virgin Islands, but I offer it as the only possibility I canthink of. The wrecking of a vessel with a cargo of logw'ood or mahogany a hun-dred years or more ago might account for this remarkable distribution.Doctor Barbour '^^ is inclined to question this assumption andcontends that the occurrence of this species in the West Indies is nottraceable to any agency of man. He says:
s< stejneger, L., The herpetology of Porto Rico. Ann. Kep. U. S. Nat. Mus. for 1902, pp. 661, 662, 1904.8' Barbour, T., A contribution to the zoogeography of the West Indies, with special reference to amphib-ians and reptiles. Mem. Mus. Comp. Zool., vol. 44, no. 2, p. 253, Mar., 1914.
MEXICAN TAILLESS AMPHIBIANS 87Now, considering that L. albilabris occurs upon more than one island, it isnecessary to postulate a series of unlikely events to explain its accidental occur-rence. It is more probable that the Porto Rican and maniland individualshave by some fortuitous parallelism become indistinguishable if they originallybelonged to different stocks or if both were derived from the same stock thenthey have preserved their identity in parts of their range and varied with isolationin other parts. The occurrence in the East Indies of Rana rnacrodon in thegreater Sunda Islands and New Guinea, and of Rana modesta, a deiivative ofR. rnacrodon in Celebes is a case of similar distribution. The Lesser Antilleanspecies are almost certainly not introduced, since Labat informs us that frogs(L. pentadadylus) were a common article of food in Martinique and other islandsas early as 1724. That they probabl}"^ belonged to this species is shown by hisstating that they lived in the woods and not in the water and by his descriptionof their colors. The occurrence of a peculiar Leptodactylus upon Haiti, andthere is evidence that a species occiu's upon Cuba also, is not as astonishing asappears at first.Schmidt ^^ contributes the following notes on the life history ofthis species.At Coamo Springs, on the terrace behind the bathhouses of the hotel, the waterof some of the springs forms a permanent rivulet at the base of the cliff. Lep-todactylus albilabris was abundant on the terrace, beneath loose stones, and undera large stone at the edge of the creek, the writer found, on August 27, 1919, ashallow, rounded excavation, 6 or 7 cm. in diameter and about 3 cm. deep, filledwith a mass of white foam, in which were the small tadpoles of this species (12mm. in length, bodj' 3-4 mm.). There were between 75 and 100 tadpoles in themass, by no means confined to the central hollow, which was present, as in thefoam mass described by Stejneger. The bottom of the excavation was about3 cm. above the water level. Two similar excavations were discovered in theimmediate vicinity, in the same relative position with reference to the water,but empty. On August 29, near Bayamon, a small mass of foam, between 3and 4 cm. in diameter, containing similar tadpoles, was noted under a stone on ahilltop, with no water whatever in the neighborhood. On October 1, near theforester's cabin on El Yunque, at about 1,200 feet, a nest of this species wasobserved beside a pool of standing water (also at a slightly higher level than thatof the water) under a rotten log. This nest contained between 150 and 200 eggs,uniformly distributed through the foam, and with no central hollow. It wassomewhat larger than those previously observed, measuring 8 cm. in diameter.The eggs are light yellow and measure 2.5 mm. to 3 mm. in diameter. Thesmallest tadpoles taken swimming at large measured 6 mm. in body length, whichprobably represents their size at the time they escape from the foam. It isevident that the tadpoles usually will be washed from the nest into the adjacentwater by a flood or heavy rain. The location of the small nest away from waterwas probably a mistake on the part of the frog, and the nest described by Stej-neger under water probably had been covered by a rise in the creek after thedeposition of the eggs. The largest larvae, nearly ready to transform, measure13 mm. from snout to vent. The V-shaped dorsal markings are already evidentin the tadpoles at this stage. The median dorsal white line is probably an adultcharacter.
35 Schmidt, K. P., Scientific survey of Porto Rico and the Virgin Islands. New York Acad. Sci? vol. 10,pt. 1, Amphibians and land reptiles, pp. 41, 42, 1928.
88 BULLETIN 160, UNITED STATES NATIONAL MUSEUMSpecimens examined.?Twenty-four, as follows
:
Lcptodactylus albilabris
Museum
MEXICAN TAILLESS AMPHIBIANS 89The type of Cope's Cystignathus perlaevis (U.S.N.M. No. 10041),which was found in a well near Japana, in western Tehuantepec, issimply an emaciated and somewhat faded specimen of an adult Lep-todactylus melanonotus.Ruthven ^^ writes that Leptodadylus caliginosus is not uncommonat Cuatotolapam and Lake Catemaco, in southern Vera Cruz. Hesays:In those in which the color was examined in life the ground color above wasbrownish orange (138) ,^8 the margin of the occipital spot a pale dull j-ellow (153d),and the ventral parts black and white or gray and white. The colors of thedorsal surface are frequently darker, however, and the pale margin of the occipitalspot is not always well defined. The dark head markings are quite regular inarrangement, consisting of a band along the canthus rostralis, one from the lowerpart of the anterior margin of the orbit to the mouth and two from the posteriormargin of the orbit on the neck, the lower involving the tympanum. These darkbands with the occipital spot are usually' margined with pale yellow, and the palelines that margin the dark bands on the canthus rostralis are often connectedwith the anterior margin of the occipital spot to form a triangle. These lightbands are frequently more or less obsolete, occasionally entirely so.This little amphibian, while not uncommon in the region studied, is so secretiveas to be little in evidence. We only found it under logs and other decaying vege-table debris in the vicinity of water. It is quite generally distributed on lowground, however, occurring both in the woods and on the prairie. It breeds inthe lowland forest pools and in swampy places on the shores of Lake Catemaco,in July and August, and immature individuals were taken in both places.Specimens examined.?Sixty-three, as follows:Leptodadylus melanonohis
Museum
U.S.N.M.Do...Do.Do.F.M.N.H.M.C.Z....U.S.N.M.Do....F.M.N.H-U.S.N.M..Do....Do....F.M.N.H.U.S.N.M..Do....Do....Do....
CatalogueNo.
25154-5914080
14838369-7316552-5657765163117439-4030523-2410041136630300-0130437-5270402-0330116-25
Nnin-|bar ofsped-
!
mens I
21
Locality collected
Sinaloa:MazatlanPresidio, .50 miles from Maz-atlan.Guanajuato:GuanajuatodoVera Cruz:AchotolAlvaradoVicinity of Orizaba.Colima:ColimadoJalisco: GuadalajaraOaxaca:Barrio in TehuantepecJapana in Tehuantepec.Rio Obispo...Tehuantepecdo.doTehuantepec (west side)
.
Date collected
Apr. 3, 1884
187
-do.
By whom collected orfrom whom received
Ferdinand Bischoff.jVlphonso Forrer.
Alfred Duges.Do./Dec. ? , 19041 IUan. ? , 1905/ [1921
July 8. 1902
Apr. 24,1903
E. Heller and CBarber.E. R. Dunn.Francis Sumichrast.J. Hurter.C. H. T. Townsend.
.\lfred Dugfs.Francis Sumichrast.Do.S. E. Meek.Francis Sumichrast.Do.Do.Do.
M.
" Type Cystignathw microixs. ^ Type Cystignathus perlaevis.Ruthven, A. Q., Zool. Jahrb. (Syst. Abt.), vol. 32, pt. 4, pp. 306, 307, 1912.The numbers refer to the "Code des Couleurs," by Elincksiek and Vallete.
90 BULLETIN 160, UNITED STATES NATIONAL MUSEUMGenus ELEUTHERODACTYLUS Dumeril and Bibron1841. Eleutherodactylus Dumeril and Bibron, Erp^tologie g^n^rale, vol. 8, p. 620.[Genotype, Eleutherodactylus martinicensis= Hylodes martinicensis Tschudi,1838, M^m. Soc. Sci. Nat. de NeuchStel, vol. 2, p. 77 (no locality designated).]1843. Batrachyla Bell, Voyage of H. M. S. Beagle, Zoology, pt. 5, Reptiles, p. 43,Oct. [Genotype, Batrachyla leptopus Bell, idem, p. 43, pi. 18, fig. 5 (Valdivia,Chile).]1843. Euhyas Fitzinger, Systema reptilium, fasc. 1, p. 31. [Genotype, Hylodesricordii Dumeril and Bibron, 1841, Erp^tologie g6n6rale, vol. 8, p. 623(Cuba).]1843. Lithodytes Fitzinger, Systema reptilium, fasc. 1, p. 31. [Genotype,Hylodes lineatus Dumeril and Bibron=(i2ano) lineata Schneider, 1799,Historiae amphibiorum, naturalis et literariae, fasc. 1, p. 138 (no localitygiven) = /2. (ana) lineata Gravenhorst, 1829, Deliciae musei zoologiciVratislaviensis, fasc. 1, p. 44, pi. 8, fig. 2.]1862. Craugastor Cope, Proc. Acad. Nat. Sci. Philadelphia, vol 14, p. 153, Mar.[Included species: Hylodes hallowellii, idem, p. 153 (near Carthagena, NewGranada), and Hylodes fitzingeri Schmidt, 1858, Denkschr. k. Akad. Wiss.math.-naturw. Classe, Wien, vol. 14, pt. 2, p. 248, pi. 1, fig. 10 (Cordillerenvon Neu Granada, elevation 4,000 feet).]1863. Strabomantis Peters, Mouatsber. k. preuss. Akad. Wiss. Berlin, p. 405,Oct. [Genotype, St{rabomantis) biporcatus Peters, idem, p. 405 (Veragua).]1866. Epirhexis Cope, Journ. Acad. Nat. Sci. Philadelphia, n. s., vol. 6, pt. 1,p. 96, July. [Genotype, Batrachyla longipes Baird, 1859, Report on theUnited States and Mexican Boundary Survey, vol. 2, Reptiles, p. 35, pi. 37,figs. 1-3 (no description) (Mexico).]1868. Leiyla Keferstein, Archiv. f. Naturg., Berlin, vol. 1, pt. 3, p. 296.[Genotype, Leiyla giintheri Keferstein (Costa Rica).]1869. Liyla Cope, Proc. Amer. Philos. Soc, vol. 11, no. 82, p. 160, July 16.[Errore typogr. for Leixjla Keferstein, 1868.]1870. Limnophys Jimenez db la Espada, Jorn. Sci. Math. Phys. e Nat., Lisboa,vol. 3, no. 9, p. 59, June. [Included species: Livmophys cornutus and L. na-paeus Espada, idem., p. 60 (Ecuador).]1870. Pnstimantis Jimenez de la Espada, Jorn. Sci. Math. Phys. e Nat., Lisboa,vol. 3, no. 9, p. 61, June. [Genotype, Pristimantis galdi Espada, idem, p. 61(Ecuador).]1893. Liohyla Cope, Proc. Amer. Philos. Soc, vol. 31, no. 142, p. 335, Nov. 17.[Genotype, Liohyla ranoides Cope =^ Lithodytes ranoides Cope, 1885, Proc.Amer. Philos. Soc, vol. 23, no. 122, p. 275, Nov. 20 (Nicaragua).]A critical review of the status of Fitzinger's Hylodes lias beenpublished by Dr. L. Stejneger,^^ and is summarized as follows:1826. Hylodes Fitzinger, Neue Classification der Reptilien, pp. 38, 64. [Geno-type, Hyla ranoides Spix, 1824, Animalia nova, sive species novae testu-dinum et ranarum quas in itinere per Brasiliam, etc., p. 32, pi. 6, fig. 3(Brazil)] =Elosia Tschudi, 1838, M^m. Soc. Sci. Nat. de Neuchatel, vol.2, p. 77 [Genotype, Hyla nasus Lichtenstein, 1823, Verzeichniss d. Doublet-ten Zool. Mus. k. Univ. Berlin, etc., p. 106 (Brazil).] [Hyla ranoides Spixis not equivalent to Hyla nasus Lichtenstein; for illustrations of the pec-toral girdles of these two species see Miranda-Ribeiro, 1926, Archiv. Mus.Nac. Rio de Janeiro, vol. 27, p. 54, fig. 31, and p. 32, fig. 17.]
3? .Stejneger, L., The herpetology of Porto Rico. Ann. Rep. U. S. Nat. Mus. tor 1902, pp 682, 583, 1904-
MEXICAN TAILLESS AMPHIBIANS 911830. Enydrobius Wagler, Natiirliches System der Amphibien, p. 202. [Sub-stitute name for Hylodes Fitzinger, 1826.]In this review ten species of the genus Eleutherodadylus are recog-nized within the borders of Mexico. Three of these, mexicanus,rhodopis, and rugidosus, have been taken much more frequently thanany of the others. Larger series of dunnii and venustus, obviouslyrelated to rhodopis, are necessary for an accurate determination oftheir status. The robber frog, E. augusti, is quite closely related toE. laticeps. The species laticeps and longipes seem to be the rarest ofall Mexican eleutherodactylids, since at present they are known fromone and two individuals, respectively. The specimen taken north ofVera Cruz by Duges and referred to E. melanostidus by Giinther inhis Biologia Centrali-Americana, 1900, p. 236, is the only reportedoccurrence of this species in Mexico, and as this determination hasbeen questioned it has not been listed herein.In contrast to many other well-known kinds of frogs, some of thespecies referable to Eleutherodadylus hatch fully formed from theegg capsules. Their larvae have a peculiar hatching mechanism, theso-called egg-tooth, which consists of a horny structure derived frommodified epidermis. There are, however, many species of Eleuthero-dadylus whose life histories are entirely unknown. Although directdevelopment is definitely known to take place in a few species, thismay not necessarily be the invariable process. The Texan cliff frog,or robber frog, E. latrans, is reported by Strecker to pass through atadpole stage in the water.According to Noble ^?
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The diagnostic characters of the life history of Eleutherodadylus are the ter-restrial mode of egg-laying, the large eggs with two or more resistent capsulesdevoid of peduncles or "foam," the absence of tadpole mouth parts, the presenceof a premaxillary egg tooth, and a broad highly vasculated tail.The life histories of the two species studied by Noble " are suppos-edly typical for most of the members of this genus; the followingquotation summarizes the essential steps in their development
:
The eggs of the various species of Eleutherodactylus may be laid between leavesof epiphytes, palms, bananas, etc., or on the ground under rocks, logs, vegetation,or merely in a hole in the bank. The breeding site may be near water or faraway from it. As a rule, each species has a definite habitat preference for itseggs, but this is not constant. I have found in Santo Domingo that two species,E. abbotti, and E. flavescens, may lay their eggs among leaves above the groundor on the ground at the base of reeds. Development is in these species direct.All species of Eleutherodactylus fully investigated have been found to have an eggtooth lying directly over the symphysis of the premaxillae. This may vary insize and shape in the different species. I have found that E. abbotti on hatching
? Noble, G. K., The value of life history data in the study of the evolution of the Amphibia. Ann. MewYork Acad. Sci., vol. 30, p. 90, Oct. 31, 1927." Noble, O. K., The hatching process in Alytes, Eleutherodadvlus, and other amphibians. Amer. Mus.Nov., no. 229, pp. 4, 5, Oct. 14, 1926.
92 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
rips first the inner capsule and then the outer with its egg tooth. E. inoptatus,on the other hand, has a very small egg tooth and one bunch of eggs on which Iexperimented could not be induced to hatch without the application of water.Then the eggs elongated, the outer capsule broke first and the inner only sometime later. It would therefore seem that forms such as E. abbotti may hatch anj-time the encapsuled larva has the strength to rip the capsules, while in nature,E. inoptatus must Avait for the rains. The first species sometimes lays its eggsamong reeds in such a position as to be well protected from the rains. The latteralways (three observations) lays its eggs in shallow holes in the bank. Suchsituations become soaking wet with every rain. The eggs of E. inoptatus wouldtherefore become wet long before they are ready to hatch. Thus there is probablya time factor which prevents the recently laid eggs from splitting open at the firstsoaking. In other Salientia the egg capsules after a long soaking are much lessresistant than recently laid ones.The habitats of most of the Mexican species of Eleutherodadylusare very imperfectly known. In March, 1895, Nelson and Goldmancollected a specimen of E. rugulosus in a small stream in the woods at3,000 feet altitude near Pluma in Oaxaca. On August 30, 1894, theytook specimens of mexicanus at an altitude of 10,000 feet in densegrowths of grass about springs and little streams in the vicinity ofCerro San Felipe, and during September, 1894, found this species ataltitudes varying from 4,000 to 9,500 feet in oak woods on the moun-tains west of Oaxaca. Gadow found rhodopis living on the ground inmasses of rotten leaves, in tillandsia clusters, and on green shrubs, ashigh as 10,000 feet on Citlaltepetl in Vera Cruz.KEY TO MEXICAN SPECIES OF ELEUTHERODACTYLUS1. A white or silvery white streak edged with black or with darkiron gray extends along upper lip from snout, below eye andtympanum, to fore limb; a light or dusky band across inter-orbital region; upperparts dark brown or dark mahoganybrown; limbs with dark, light-edged crossbars; lower partswhite, more or less speckled with brown; male with two vocalsacs, indicated externally bj' a fold on each side of throat;first and second fingers equal; fingers and toes slender, withvery small terminal disks and fairly strong subarticular tu-bercles; a very distinct vestige of web between the toes; twometatarsal tubercles, the inner oval, the outer round andsmaller; plantar surface with small scattered tubercles; a dis-tinct crested tarsal tubercle or ridge about midway but notarsal fold; the hind limb being carried forward along the body,the tibio-tarsal joint reaches the tip of snout or a little beyond;vomerine teeth in two oblique series behind the level of thechoanae; tongue oval, or oblong; snout pointed, projectingbeyond mouth; canthus rostralis distinct; loreal region slightlyconcave; nostril near end of snout; interorbital space a littlebroader than upper eyelid; tympanum round, very large anddistinct, as large as or larger than the eye; skin of upperpartswith many small tubercles and plicate rugosities; a faint ver-tebral linear dermal ridge; a pair of converging dorsal dermalridges present or absent; several other irregular lateral glan-
MEXICAN TAILLESS AMPHIBIANS 93dular folds present; upper surfaces of thighs with scatteredtubercles like back; underparts smooth; inferior surfaces ofthighs granular; discoidal abdominal fold present; head-and-body length, 25 mm beatae (p. 104)No white streak edged with black along upper lip 22. Loreal region oblique, shelving outward to edge of upper jaw;either an intratympanic or postcephalic dermal fold; a largeabdominal disk 3Loreal region not shelving outward to edge of upper jaw; neitheran intratympanic nor a postcephalic dermal fold 43. A distinct sharp-edged tarsal fold; tarsal-metatarsal joint to tipof fourth toe, 36 mm.; a large protuberant inner and a verysmall outer metatarsal tubercle; subarticular tubercles at endsof metapodials fairly large; a few small supernumerary tuber-cles on metatarsals; toes webbed at base; the hind limb beingcarried forward along the body, the tibio-tarsal joint reachesbetween eye and nostril; first finger exceeds second by lengthof its apical disk; large subarticular tubercles and palmer cal-losities; fingers and toes with blunt apical disks; fingers with-out web at base; head thick, remarkably wide, its transversediameter being one-fifth greater than its length; muzzle obtuseand rounded; canthus rostralis rounded; tympanum large,higher than wide, overhung by prominent dermal fold extendingfrom eye to angle of lower jaw, where one of its two branchesgoes to shoulder and the other to insertion of fore limb;vomerine teeth in two short oblique clusters, considerably be-hind but within level of inner margins of choanae; tongue broadand somewhat oval in outline; skin on throat, abdomen, andunder surfaces of thighs smooth; posterior surfaces of thighsgranulated; a transverse post-tympanic dorsal dermal fold;skin of upperparts somewhat shagreen; general tint of upper-parts dark greenish brown; loreal region and tympanum black;hind limbs with black transverse bars; soles of feet black; un-der surface of forearm black; underparts yellowish white; outermargins of throat and under surfaces of thighs and tibiae mar-bled with brown specks; sides finely marbled with brown;head-and-body length, 74 mm laticeps (p. 106)No tarsal fold; tarsal-metatarsal joint to tip of fourth toe, 28mm.; inner and outer metatarsal tubercles large, the innerelongated and swollen, the outer broad and flattened, and inaddition the outer metatarsal tubercle is nearly as large as theinner; large subarticular tubercles at ends of metapodials; feetwith numerous supernumerary tubercles on plantar surface;toes free from web at base and without marginal dermalfringes; the hind limb being carried forward along the body,the tibio-tarsal joint reaches to center or to anterior margin ofeye; heels touch but do not overlap when legs are placed trans-verse to axis of body; first finger exceeds the second by lengthof its apical disk; fingers long and slender, with terminalT-shaped phalanges; apical disks blunt, swollen inferiorly; twolarge palmar callosities, the external one divided by a longi-tudinal groove; head much wider than long, with large eyeswidely separated; canthus rostralis fairly distinct; muzzletruncate; width of upper eyelid one-half to three-fifths that of66785?32 7
94 BULLETIN 160, UNITED STATES NATIONAL MUSEUMinterorbital space; transverse diameter of tympanum occasion-ally less than one-half, always less than two-thirds, that of theeye, and overhung by a dermal fold that extends from eyebackward over tympanum to angle of mouth; vomerine teethin adults in two essentially transverse clusters between ornearly on a line with posterior margins of choanae [teethbehind but within level of choanae in young]; tongue narrow,elongate; skin on abdomen smooth; circular abdominal diskpresent; under surfaces of thighs granulated; a transverseintratympanic dorsal dermal fold; skin of upperparts and topof head either shagreen or with small scattered tubercles; colorof upperparts brown or gray, with dark brown spots; sideswith light-margined black spots; hinder surface of thighs withclosely approximated or with large coalesced black markings;upper surface of tarsus and foot with black blotches; lowerfore limb with transverse black bars; abdominal disk yellowish,with faint vermiculations; throat black [in young or immatureindividuals, the color surrounding elongated spots on top ofhead and middle of back is bright salmon pink or pale yellow;a broad transverse pink or pale yellow band with several darkspots crosses back behind level of origin of fore limbs; anaggregation of smaller dark brown spots in sacral region;small irregular dark spots on top and sides of head; brownspots below tympanum, eye, and loreal region]; head-and-bodylength, 64.5 to 75 mm augusti (p. 100)4. Terminal disks distinctly enlarged or dilated, those of fingers 2and 3 about twice the width of corresponding digit 5Terminal disks less noticeablj' dilated, generally rounded or ball-like, rarely flattened, and those of fingers scarcely larger thanthose of toes 66. Terminal disks of fingers 2, 3, and 4 fully twice the width ofcorresponding digit, subtriangular in outline, flattened, andtruncated distaUy, their diameter nearly equaling that of thetympanum; first finger shorter than second; large subarticulartubercles at ends of metapodials; toes free or with vestiges ofweb at base; third toe as long as or only reaching to disk offifth; terminal disks of toes flattened and dilated, but notice-ably smaller than those of fingers; subarticular tuberclesmoderately large, very prominent, and not continued back-ward on plantar surface; two small metatarsal tubercles, theinner larger than outer; tarsal fold indistinct; the hind limbbeing carried forward along the body, the tibio-tarsal jointreaches to or beyond the tip of the snout; vomerine teeth intwo short transverse groups behind and well within the level ofinner margins of choanae, the distance between their outeredges being equivalent to slightly more than one-half theinterval between the choanae; tongue subcordiform ; snoutdepressed, rather short, and acuminate; canthus rostralis fairlydistinct; loreal region rather oblique, very slightly concave;upper eyelid wider than interorbital space; tympanum verydistinct, one-half to two-thirds diameter of eye, dependingupon the sex; a dermal fold above tympanum; skin of
. upperparts either smooth or sparsely studded with small tuber-cles and glandular dermal folds; an indistinct fine vertebraldermal ridge from muzzle to vent; sides areolate; underparts
MEXICAN TAILLESS AMPHIBIANS 95generally smooth, but hinder half of belly may be faintly areo-late; under surface of thighs granular; color of upperparts andsides grayish or brownish, mottled and speckled with blackishor grayish; occasionallj' with a narrow interorbital black bandand a narrow preorbital transverse black bar on light-coloredsnout; a narrow dark chevronlike mark with apex pointedinward on each shoulder; one or more pairs of narrow longitu-dinal mid-dorsal markings; a narrow black stripe from posterioredge of eye to above tympanum and downward behind latternearly to rictus oris; upper lip and loreal region with continuousor discontinuous oblique dark bars; fore and hind limbs cross-barred; under surfaces either uniformly white or throat mar-bled with brownish; head-and-body length, 50 mm alfredi (p. 99)Terminal disks of fingers 2, 3, and 4 almost twice the width ofcorresponding digit, rounded, but their diameter is less thandiameter of tympanum; first finger with tip scarcely if at alldilated; subarticular tubercles at ends of metapodials; toeswith vestigial web at base; third toe as long as fifth; terminaldisks of toes smaller than those of fingers and truncateddistally; no supernumerary tubercles on plantar surface; innerand outer metatarsal tubercles rather small and flat; tarsal foldpresent; the hind limb being carried forward along the body,the tibio-tarsal joint reaches to or beyond the tip of the snout;vomerine teeth in two small round groups well behind andwithin level of inner margins of choanae; tongue large, sub-circular; canthus rostralis distinct; loreal region oblique andslightly concave; diameter of tympanum less than half thatof eye; skin of upperparts smooth; underparts, sides, andlower surfaces of thighs coarsely granular or areolate; upper-parts brownish with triangular interorbital black spot extend-ing laterally on eyelids, a transverse black band across shoul-ders, and six or more large irregular black spots on back; foreand hind limbs with broad dark cross bands, or else with orna-mental markings obliterated or merging into the dark brownor almost black ground color; underparts lighter brown oryellov/, with fine black specks on throat and belly; head-and-body length, 38 mm longipes (p. 107)Toes united at base by vestigial web, with obvious marginal der-mal fringes extending to terminal disks; web at base of toesextends to or a little beyond basal subarticular tubercle;marginal dermal fringes extend to terminal disks; toes inorder of decreasing length, 4, 3, 5, 2, and 1; fifth toe reach-ing to terminal disk of third; two metatarsal tubercles, theinner large and elongated, the outer small and indistinct;a distinct tarsal fold; no supernumerary tubercles on plantarsurface; disks of toes distinct, those of 2, 3, and 4 mostnoticeably dilated; first finger either shorter than or slightlylonger than second; terminal disks of fingers distinct, butnot noticeably enlarged; subarticular tubercles not contin-ued backward on palmar surface; two palmar callosities, theinner large and elongate, the outer flattened and usuallydivided by longitudinal groove; the hind limb being carriedforward along the body, the tibio-tarsal joint reaches at leastto between eye and nostril and at most beyond end of snout;vomerine teeth in two nearly transverse groups, behind and
96 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
almost within level of inner margins of choanae, and separatedmedially by very narrow interval; tympanum very distinct,its horizontal diameter generally about two-thirds diameterof eye, rarely in very large individuals more than one-thirdbut less than one-half diameter of eye; tympanum overhungby a dermal fold; snout acuminate; canthus rostralis distinct;loreal region oblique and concave; upper eyelid wider thaninterorbital space; nostrils lateral and near end of snout,their distance from eye equal to or less than diameter of latter;skin of upperparts generally rugose and tuberculate, rarelysmooth, but always with short interrupted glandular dermalfolds along lateral borders; folds are most conspicuous inintratympanic region and on back; a distinct abdominal disk;hinder border of belly areolate; under surfaces of thighsstrongly granular; a more or less distinct continuous orinterrupted lateral dermal fold; a continuous or discontin-uous oblique dermal ridge at level of quadrate cartilage;ground color of upperparts brown, with scattered spots ofyellowish, and mottled or blotched with dark brown or black;white or yellowish ground color of sides vermiculated or vari-egated with blackish brown; darker brown ground color ofposterior surfaces of thighs vermiculated with white or yel-low; limbs with more or less distinct cross bands, the band-ing extending upon fingers and toes; oblique dark brownbands, separated by narrower light spaces, extend upwardacross lower and upper lips to eye and loreal region; the twobelow eye widest at lip margin, another one nearer the nos-tril than the ej^e, and a somewhat larger dark spot bisectedby two narrow vertical light lines on end of snout; anteriorrim of orbit margined with black; a light brown or yellowloreal spot; an interorbital dark band behind light-coloredsnout; a black streak extends backward from eye over tym-panum and downward to insertion of fore limb; dark streakalong canthus rostralis present or absent; underparts white;throat marbled with brown; head-and-body length, 43 to65 mm rugulosus (p. 116)Toes free, not united at base by vestigial web, and withoutobvious marginal dermal fringes 77. A broad longitudinal mid-dorsal purplish band extending fromsnout to vent between a pair of undulating medially converg-ing linear dermal ridges, each of which extends from supra-ciliary to end of sacral diapophysis; remainder of upperpartslight mauve; linear dermal ridges, stripe along canthus ros-tralis extending backward above and behind the tympanum,an oval spot on each side of pelvis, and some small irregularspots on sides of body, black; upper lip and posterior surfacesof thighs powdered with gray; fore and hind limbs cross-banded; underparts white, immaculate; first finger longerthan or subequal with second; subarticular tubercles con-tinued backward on inferior surfaces of metacarpals; twopalmar callosities; disks of fingers small; toes not united byweb at base; subarticular or supernumerary tubercles con-tinued backward on plantar surface; two metatarsal tubercles,the inner one the largest; tarsal fold indistinct; the hindlimb being carried forward along the body, the tibio-tarsal
MEXICAN TAILLESS AMPHIBIANS 97joint reaches beyond the end of the snout; vomerine teethin two small rounded groups behind the choanae, well sepa-rated from each other, and not extending outward beyondlevel of inner margins of choanae; tongue subround, scarcelyor not notched behind; canthus rostralis distinct; loreal regionsubvertical; head with flat crown and subacuminate snout;tympanum round, one-half, three-fourths, or even equivalentto diameter of eye; skin of upperparts smooth, with excep-tion of a pair of undulating, anteriorly bifurcating linear dermalfolds; upper surface of head smooth, with a few indistincttubercles on upper eyelid; legs and underparts smooth; nodermal fold across chest; head-and-body length, 35 mm-_venustus (p. 117)No broad longitudinal dark-colored band bounded laterally byconverging dermal folds; a black stripe runs from nostril alongcanthus rostralis through eye to above tympanum, decurvingbehind the latter; plantar surface often black or dark colored 8Skin of upperparts finely granular, with scattered tubercles; afaintly defined granular vertebral dermal ridge; skin of under-parts smooth or with hinder half of belly areolate; thighsheavily granulate; canthus rostralis fairly distinct; lorealregion subvertical; upper eyelid narrower than interorbitalspace; tympanum large, round, considerably more than halfdiameter of eye, its distance from eye being just over one-halfof its own diameter; vomerine teeth in two short heavyoblique series some distance behind the choanae, the intervalbetween them being less than distance from outer end of eachone and corresponding choana; first finger slightly longerthan or subequal with second; fingers with small ball-liketerminal disks; subarticular tubercles on palmar surface, aswell as on digits; toes with more flattened and less ball-liketerminal disks; two metatarsal tubercles, the outer one small,round, and conical, the inner one oblong and large; a ratherindistinct tarsal fold; the hind limb being carried forwardalong the body, the tibio-tarsal joint reaches beyond the tipof the snout; upperparts lichen gray, rather smoky, withminute black spots scattered over head and back; grayish foreand hind limbs with indistinct darker cross bands; a blackstreak curves around tympanum from eye to above insertion offore limb; three subvertical dark brown bars, separated bynarrower light spaces, extend across lower and upper lips toeye and loreal region; a pair of dark spots on end of snout infront of nostrils; head-and-body length, 35 mm dunnii (p. 105)Skin of upperparts either areolate or smooth, occasionally withscattered or sparsely placed tubercles; 9A tarsal fold or a crested tubercle 2 or 3 mm. above inner meta-tarsal tubercle generally present; skin of belly either areolateor smooth; throat smooth; under surface of thighs granular;skin of upperparts areolate or apparently smooth, but with tinyasperities; several exceedingly fine dermal ridges generallypresent, the longest of which extends along vertebral columnfrom snout to vent; one from eye above tympanum to shoulderand thence along side; another oblique one extending from eyetoward center of back and thence backward and outward topoint where sacral diapophysis juts out, which together withcorresponding one on opposite side incloses an hourglass-shaped
98 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
area; and one between the former and the latter on hinder halfof back; canthus rostralis fairly distinct; loreal region oblique,flattened; snout acuminate; interorbital space much wider thanupper eyelid; tongue subovate; vomerine teeth in two slightlyoblique series well behind the choanae, well separated fromeach other, and each series not extending outward beyond levelof inner margin of corresponding choana; horizontal diameterof tympanum two-thirds, three-fourths, or almost equal todiameter of eye, and overhung by a dermal fold; first fingergenerally a little longer than second, rarely shorter; two palmarcallosities, the outer with longitudinal groove; subarticulartubercles continued backward on palmar surface; terminal disksof fingers and toes very small; third and fifth toes equal inlength; supernumerary tubercles may or may not be continuedbackward on plantar surface; two small metatarsal tubercles,the inner larger than the outer; the hind limb being carriedforward along the body, the tibio-tarsal joint reaches beyondanterior margin of eye and rarely beyond end of snout; upperparts dark brown, reddish brown, or green (actually grayisholive or brownish in alcohol) ; a narrow indistinct dark interor-bital band behind light colored snout; a black streak, marginedabove with a lighter color, commences at nostril and extendsbackward along canthus rostralis to eye, and from eye back-ward above tympanum to side or curving downward behindtympanum; vent black; a pair of short black glandular ridgesin center of back, present or absent; forearm with at least onedark cross band; posterior limbs obscurely cross-banded; un-derparts white; head-and-body length, 28 to 36 mm
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rhodopis (p. 112)Tarsal fold normally absent; skin of hinder half of belly generallyareolate, occasionally smooth; under surface of thighs granular;skin of upperparts either smooth or else roughened by dermalridges; exceedingly fine or inconspicuous dermal folds inclosingan hourglass-shaped area on back may or may not be present;a short dermal ridge along vertebral edge of scapula generallypresent; discoidal fold normally present; canthus rostralisfairly distinct and curved; loreal region oblique; snout ratherbroad and acuminate; width of upper eyelid equal to or lessthan interorbital space; vomerine teeth absent, hidden in thegum, or represented by two short oblique groups well behindand within level of inner margins of choanae; horizontal di-ameter of tympanum one-half to two-thirds diameter of eye;first finger generally shorter and rarely longer than second; twopalmar callosities, the outer generally unequally divided by alongitudinal groove; subarticular tubercles continued backwardon palmar surface; terminal disks of fingers and toes slightlydeveloped, scarcely dilated; toes almost free, but with vestigesof web at base; supernumerary tubercles may or may not becontinued backward on plantar surface; two metatarsal tu-bercles, the inner one usually large, prominent, and com-pressed, the outer one small and rounded; the hind limbbeing carried forward along the body, the tibio-tarsal joint gen-erally reaches beyond end of snout and rarely only to anteriormargin of eye; color of upperparts variable, grayish rose,brownish gray, or grayish, marbled or mottled with some darkertint, and separated from light-colored snout by a dark inter-
MEXICAN TAILLESS AMPHIBIANS 99
orbital band; darker spots on back sometimes confluent andform a pair of longitudinal bands; a sharply defined vertebralwhite stripe extending from muzzle to vent may or may not bepresent; limbs with narrow crossbars, which on hind limb occa-sionally encroach upon concealed hinder surface; throat brown-ish or speckled with brown; remainder of underparts whitish oryellowish; a blackish streak along canthus rostralis sometimesextends over whole side of head and may be continued backwardon side of body; lower and upper lips with dark spots or barsthat generally have light edges; head-and-body length, 36 to40 mm. mexicanus (p. 108)ELEUTHERODACTYLUS ALFREDI (Boulenger)1898. Hylodes alfredi Boulenger, Proc. Zool. Soc. London, p. 480, pi. 39, fig. 1,June 7.1882. Hylodes conspicillatus Brocchi, Mission scientifique au Mexique et dansI'Am^rique Centrale, recherches zoologiques, pt. 3, sect. 2, p. 59 (Andes,Orizaba). [Nee Hylodes conspicillatus Gunther, 1858.]Type locality.?Atoyac, State of Vera Cruz, Mexico.Range.?So far as known, found only at Orizaba and at Atoyac inVera Cruz.Remarks.?The leptodactylid from the Andes of Orizaba, whichBrocchi identified as Hylodes conspicillatus Giinther, is certainly notthat species. Brocchi states that the adhesive disks are very distinct,and since the external characters and general coloration are describedrather briefly, the relationships of this frog are estabUshed mainly bythe process of elimination. Distinctive characters of constant valueare hard to find in this genus, but the species alfredi fortunately hasespecially large subtriangular and rather squarely truncated adhesivedisks on the two outer fingers.This frog was named in honor of the donor. Prof. Alfred Dug^s, ofGuanajuato, Mexico, and there are two cotypes in the collection of theBritish Museum of Natural History. The following notes were madeon these specimens:B.M. No. 98. 2. 19. 1 : Head-and-body length, 37.2 mm.; transversediameter of tympanum, 2.8 mm. ; transverse diameter of eye, 4.2 mm. ;anterior edge of eye to nostril, 4.8 mm.; the hind Hmb being carriedforward along the body, the tibio-tarsal joint reaches considerablybeyond the tip of snout; apical disks of fingers very large; vomerineteeth in two small oblique posteriorly converging rows behind andwithin level of the choanae; skiu of abdomen smooth; large subarticu-lar tubercles; soles of feet with supernumerary tubercles ; large elongateinner and small conical outer metatarsal tubercles; tarsal fold indis-tinct, if present; apical disks of toes considerably smaller than thoseof fingers.B.M. No. 98. 4. 7. 1: Head-and-body length, 30.8 mm.; transversediameter of tympanum, 2 mm.; transverse diameter of eye, 3.6 mm.;anterior edge of eye to nostril, 4.1 mm.; the hind limb being carried
100 BULLETIN 160, UNITED STATES NATIONAL MUSEUMforward along the body, the tibio-tarsal joint reaches the tip of thesnout; apical disks of fingers large and subtriangular; vomerine teethas in the preceding; large subarticular tubercles; soles of feet withoutsmall supernumerary tubercles; apparently no tarsal fold.Specimens examined.?Four, as follows:
Eleutherodactylus alfredi
Museum
MEXICAN TAILLESS AMPHIBIANS 101
[ = Nayarit], form the basis for the following notes: Tarsal fold absent;tarsal-metatarsal joint to tip of fourth toe, 28 mm.; inner and outermetatarsal tubercles large, the inner elongated and swollen, the outerbroad and flattened; large subarticular tubercles at ends of metapo-dials; feet with numerous supernumerary tubercles on plantar sur-face; the hind limb being carried forward along the body, the tibio-tarsal joint reaches to center or to anterior margin of eye; heels touchbut do not overlap when legs are placed transverse to axis of body;first finger exceeds the second by length of its apical disk; canthusrostralis fairly distinct; transverse diameter of tympanum occasion-ally less than one-half, always less than two-thirds, diameter of eye,and overhung by a dermal fold that extends from eye backward overand behind tympanum to angle of mouth (transverse diameter oftympanum, 3.7 mm.; transverse diameter of eye, 8.2 mmx.); distancefrom eye to nostril equal to or less than diameter of eye ; tongue nar-row, elongate; skin on abdomen smooth; circular abdominal diskpresent; under surfaces of thighs granulated; a transverse intratym-panic dorsal dermal fold; skin of upperparts and top of head eithershagreened or with small scattered tubercles; sides with light marginedlarge black blotches; hinder surface of thighs with closely approxi-mated or with large coalesced black marldngs ; upper surface of tarsusand foot with black blotches; lower fore limb with transverse blackbars; abdominal disk yellowish, with faint vermiculations; throatblack; head and body length, snout to vent, 75, 65.5, 65.5, 64.5, 67.5,and 66.2 mm., respectively.Mocquard concluded that E. augusti was identical with Cope'sE. latrans from central Texas. Direct comparison of Mexican speci-mens with the cotypes of E. latrans does not entirely confirm thisassumption. Though there are no constant structural features thatwill distinguish specimens from these two areas, it was observed thatin E. latrans the fourth toe is relatively longer, the color pattern con-sists of fairly closely aggregated large black blotches, the sides andhinder half of the abdomen are faintly areolate, and the skin on theupperparts of old adults is stiff, coarse, and areolate. These twoforms are unquestionably rather closely related. An immature indi-vidual from Jalisco and an adult individual (with a body length of 75mm.) collected by Ruthling, which unfortunately is without anydefinite locality, were used in these direct comparisons. The skin onthe upperparts of the immature individual from Jalisco is muchmore tubercular and warty than that on the Texas specimens.Juvenile characters, such as vestigial postcephalic intratympanicdermal fold and vomerine teeth in minute clusters, are not unusual,but the presence of an abdominal disk seems rather remarkable forso young an individual. Mocquard ^^ has pubHshed some interesting
" Mocquard, F., Reptiles et batraciens recueillis au Mexique par M. TAon Dlguet en 1896 et 1897. Bull.Soc. Philom. Paris, ser. 9, vol. 1, no. 4, pp. 160, 161, 1899.
102 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
notes on E. augusti, which are taken from the field notes of L^onDiguet and from which the following is quoted (translation)
:
H. augusti appears to be essentially nocturnal, which explains its rarity incollections. Cope had already made known some facts regarding their habits,after the data furnished to him by Mr. G. W. Marnock, who discovered it inTexas.In confirming these details for me, Mr. Diguet has been good enough to add tothem the following information: "This batrachian is encountered in the territoryof Tepic in damp ravines at the commencement of the rainy season; that is, theend of June and July. Its voice is resounding and can be heard at a distanceafter sunset. At this time, it is found attached upon certain smooth-barkedtrees, such as the Burseras, the color of which is the same as that of the animal."This last observation of Mr. Diguet has its importance, in that it establishes forus the significance of the ventral disk: It is, undoubtedly, an adhesive disk.The fingers, in fact, lack terminal disks and are only feebly swollen at theirextremities; the animal therefore can not maintain itself along vertical surfacesexcept by means of this ventral disk. Furthermore as this disk can not be otherthan an organ of attachment, the progression of this batrachian upon the trunksof trees or in the vertical cracks of rocks, where it has been found by Mr. G. W.Marnock in Texas, must be effected by leaps: This is actually the case and Mr.Diguet has seen it constantly leaping from one branch to another when it tried tohold on.Mr. Diguet has never encountered H. augusti during the day, but [only] atnight with the aid of a light. He was able to capture eight specimens, seven malesand one female, in the vicinity of small puddles of water on the western slope ofthe Cerro San Juan, territory of Tepic. He had at first grasped a male, next afemale, whose sex he identified by the presence of ovaries loaded with eggs, whichthe transparency of the abdominal wall let be seen; he held this female uponthe ground and saw arriving successively 6 other males which followed her andwhich he seized.Inasmuch as Mocquard makes reference to the field observationsof Marnock published by Cope and since E. augusti and E. latransseem to be quite closely related, quotations from accounts dealingwith the habits and life histories of E. latrans are here given.In his preliminary notice of the discovery of the Texas cliff frog,Cope writes ^* that
?
It lives in fissures in the limestone cliflFs that stretch across that section of theState. According to Mr. Marnock the eggs hatch out in the winter, and thetadpoles live in the rainwater which is caught in the shallow holes in the rocks,far from the creeks. During the winter the adults are very noisy, the rocksresounding in the evening with their dog-like bark.Miss Dickerson ^^ has published a very interesting account of thisspecies, from which the following is taken:This smooth frog-like batrachian attains a surprisingly large size for one sodelicately built. The arms and legs are peculiarly slender, and look out of pro-portion when seen on a frog three and a half inches long.In fact, Lithodytes latrans is a very curious-looking creature. It rests on handsand feet only, the tarsus and other parts of the legs as well as the body being kept
" Cope, E. D., Amer. Nat., vol. 12, no. 3, p. 186, Mar., 1878.?? Dickerson, M. C, The frog book, p. 164, 1906.
MEXICAN TAILLESS AMPHIBIANS 103
elevated some distance above the ground. If its tracks could be seen, they wouldshow impressions of the soles and long toes, of the palms and long fingers only.It moves about slowly and seriously in this stilted fashion, a grotesque littlecreature indeed. Its grotesqueness is enhanced bj'^ the transversely elongatedspots of the back, set in their light pink background like two staring eyes.This species lives in fissures of the limestone cliffs along the borders of the firstplateau region of Texas. The method of proceeding with body elevated, insteadof dragged on the ground, is perhaps correlated with its habit of living among thelimestone rocks.Some additional observations on the breeding habits of the Texanrobber frog have been piibhshed by Strecker : ^^Lilhodytes latrans has in all probability an extensive range, but, on account ofits peculiarly secretive and nocturnal habits, has been overlooked by the most emi-nent herpetologists who have visited Texas. Its distribution is entirely depend-ent on the presence of the exposures of white limestone which enclose many of thestreams of the central and southern sections of the State.It is a land animal, hiding in caves and fissures during the daytime, and, except-ing during the brief breeding period, venturing abroad only at night. Breedingin water-filled pockets and hollows in the rocks and in the rocky beds of smallstreams, it does not appear to be perfectly at home in the water at any time andspecimens observed by me made no attempt to conceal themselves by divingbut swam clumsily across small pools and sought to escape by leaping up the bankon the opposite side. A breeding pair remain in copula close in to the bank.The masses of water-soaked leaves which line the edges of the pools and hollowsserve them for the purpose of floating their fertilized eggs. * * *This species breeds unusually early in the year. Marnock informed Cope thatthe eggs were hatched in winter. Here in central Texas the breeding season islater than it is in Bexar County and the eggs are deposited early in February.If the eggs were deposited before the 9th of that month in the present year, theywere subjected to some of the hardest freezes we have had in years. On the 9thand 10th the ground was covered with two inches of water. A few days laterthe weather was warm and clear and melted snow filled the hollows in many ofthe gulches that are usually dry at this season.On March 5 a number of tadpoles were found in small pools in the gully 3 milesnorth of town. They were in two stages, the larger ones having the hind limbswell developed. In form these larvae were short and round bodied, with slender,but rather short, tails. In a specimen 36 mm. in total length, the distance frommuzzle to anus was 14 mm. In a smaller example, the tail was only 4 mm. longerthan head and body. * * *These little poUywogs are very active and on being disturbed conceal themselvesamong leaves in the bottoms and on the sides of the pools. The larger ones areunusually wary and it is a difficult matter to capture them even with a dip net.* * * By the 19th of March the larger tadpoles had become fuUy developedfrogs and left the water with their short tails still in evidence. They were slightlyover a third as large as full-grown adults. The complete metamorphosis mustnot take over six weeks, if we are to judge by the length of time required for otherfrogs to transform after the first appearance of the hind limbs.The indirect development described by Strecker and the presenceof narrow toes have led Noble *^ to infer that this species does not
" strecker, J. K., jr., Notes on the robber frog (Lithodytes latrans Cope). Trans. Acad. Sci. St. Louis,vol. 19, no. 5, pp. 73-79, June 14, 1910.? Noble, Q. K., An outline of the relation of ontogeny to phylogeny within the Amphibia, I. Amer,Mus. Nov., no. 16.5, pp. 14, 15, Apr. 16, 1925.
104 BULLETIN 160, UNITED STATES NATIONAL MUSEUMbelong in the genus Eleutherodadylus. Contrary to Noble's assump-tion, at least one of the cotypes of Lithodytes latrans (U.S.N.M. No.10058) has typical, well-developed, T-shaped terminal phalanges,and there is no reason to suspect that this modification does not occurin the remaining cotypes. A dissected specimen of this species wasfound to have the usual Eleutherodadylus type of sternum, withouta bony style and with a cartilaginous mesosternum with bisagittateextremity.Specimens examined.?Nine, as follows:Eleutherodadylus augusti
Museum
MEXICAN TAILLESS AMPHIBIANS 105supernumerary tubercles on plantar surface of foot; first finger longerthan second; a broad white streak extends backward from tip ofsnout above brown upper lip and below eye and tympanum to forelimb; lower lip variegated with light and dark; fore limb with darkcross bands; upper surface of thigh and tibia with indistinct dark crossbands; an exceedingly fine vertebral dermal ridge; skin of upper-parts with tiny asperities and small glandular elevations; a shortcurved dermal ridge along vertebral margin of scapula; canthus ros-tralis rounded; vomerine teeth in two oblique posteriorly convergingrows behind and within level of choanae.B.M. No. 1903. 9. 30. 237: Head-and-body length, 23.2 mm.; trans-verse diameter of tympanum, 2.7 mm.; transverse diameter of eye,2.8 mm. ; anterior edge of eye to nostril, 2.8 mm. ; the hind limb beingcarried forward along the body, the tibio-tarsal joint reaches beyondend of snout; a distinct crested tarsal tubercle about midway; super-numerary tubercles on plantar surface of foot; coloration the same asin the preceding, except that lower lip is brown and not variegatedwith a light color; canthus rostralis rounded; vomerine teeth similarto the other cotype.Specimens examined.?Three, as follows:Eleutherodactylus beatae
Museum
106 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
of these species may possibly show that E. dunnii is merely a variantof the widely distributed E. rhodopis.Doctor Dunn writes to me from London as follows
:
There are specimens in the British Museum which fit the description of E.dunnii more closely than any other. Ten specimens from Xometla (1903. 9. 30.238-247, Brit. Mus.) and four from Ste. Barbara in Vera Cruz (1903. 9. 30.248-251, Brit. Mus.) are referred to this species. These are differently preserved,but I think that they are the same. Tympanum black; considerable black aroundeye and on lip in spots. Glandular ridges do not show up. Apparently aW-shaped scapular fold. In these specimens the converging dorsal dermal ridgesare lacking.Specimens examined.?Two, as follows:
Eleutherodactylus dunnii
Museum
MEXICAN TAILLESS AMPHIBIANS 107and palmar callosities; head broad, 33.3 mm, wide at angle of jaws;loreal region somewhat shelving; canthus rostralis rounded; vomerineteeth in two short oblique clusters, considerably behind but witliinlevel of inner margins of choanae; tongue broad and somewhat ovalin outline; skin on throat, abdomen, and under surfaces of thighssmooth; a distinct abdominal disk; posterior surfaces of thighs granu-lated; a transverse post-tympanic dorsal dermal fold; skin of upper-parts somewhat shagreened; hind limbs with black transverse bars;soles of feet black; under surface of forearm black; outer margins ofthroat and under surfaces of thighs and tibiae marbled with brownspecks; sides with small anastomosing brown specks; head-and-bodylength, 74 mm. ; transverse diameter of tympanum, 4.7 mm. ; transversediameter of eye, 8.8 mm.; anterior edge of eye to nostril, 9.7 mm.Specimens examined.?One, the type.ELEUTHERODACTYLUS LONGIPES (Baird)1859. Bairachyla longipes Baird, Report on the United States and MexicanBoundary Survey, vol. 2, Reptiles, p. 35, pi. 37, figs. 1-3 (no description).
?
Barbour, 1923, Proc. New England Zool. Club, vol. 8, pp. 81-83 (80 milesS. W. Victoria, Tamaulipas, Mexico).1866. Epirhexis longipes Cope, Journ. Acad. Nat. Sci. Philadelphia, n. s., vol. 6,pt. 1, July, p. 96.?Cope, 1S75, U. S. Nat. Mus. Bull. 1, p. 31 (Lower RioGrande) .?Cope, 1887, U. S. Nat. Mus. Bull. 32, p. 16.Type locality.?Mexico.Range.?Exact limits of range unknown; probably occurs on theMexican Plateau from southern Tamaulipas to Puebla.Remarks.?Under the caption for the explanation of the plate withthe original illustration of this frog, Baird simply stated that it wascollected in Mexico. The original drawing of the type, which wasused by Professor Baird in his report on the reptiles of the UnitedStates and Mexican Boundary Survey, was made by J. H. Kichardand is dated 1856. At the right lower margin, "City of Mexico" iswritten in pencil and on the left lower margin "Batrachyla." In1875, Cope gave "Lower Rio Grande" as the locahty, but in 1887 hestated that the type was taken 40 leagues north of the City of Mexicoand that it bore the number 3207 in the collection of the NationalMuseum. An unlocated specimen from Camp Yuma, Calif., col-lected by Dr. R. O. Abbott, bears this number in the catalogue withthe original entries. A missing "Hyla" collected by John Potts "40leagues S. of City of Mex.," and catalogued under the number 3237,may be the original entry for the type of Bairachyla longipes. Thetype is now lost.Specimens examined.?One (M.C.Z. No. 9308), collected at Miqui-huana, 80 miles southwest of Victoria, Tamaulipas, in 1922 by W.W. Brown.
108 BULLETIN 160, UNITED STATES NATIONAL MUSEUMELEUTHERODACTYLUS MEXICANUS (Brocchi)1877. Leuiperus [sic] mexicanus Brocchi, Bull. Soc. Philom. Paris, ser. 7, vol. 1,no. 4, p. 184.
?
Brocchi, 1882, Mission scientifique au Mexique et dansFAm^rique Centrale, recherches zoologiques, pt. 3, sect. 2, p. 21, pi. 5, figs. 3,3a-c.1879. Hylodes lineatus Brocchi, Bull. Soc. Philom. Paris, ser. 7, vol. 3, no. 1, p. 22,(Attitlan, probably Guatemala). [Preoccupied by Hylodes lineatus GiJN-ther, 1858= Rana lineata Schneider, 1799.]1898. Borborocoeies mexicanus Boulenger, Proc. Zool. Soc. London, p. 481, pi.39, fig. 2, June 7 (Hacienda el Florencio, State of Zacatecas, Mexico)
.
1900. Syrrhaphus omiltemanus Gunther, Biologia Centrali-Americana, Reptiliaand Batrachia, p. 216, Apr. (Omilteme, State of Guerrero, Mexico).1900. Hylodes calcitrans Gxjnther, Biologia Centrali-Americana, Reptilia andBatrachia, p. 230, pi. 67, fig. B, Aug. (Omilteme, State of Guerrero, andJalisco, Mexico).1927. Pleurodema mexicana Parker, Ann. and Mag. Nat. Hist., ser. 9, vol. 20,no. 118, p. 475, Oct.Type locality.?Mexico.Range.?Western side of the Mexican Plateau from Zacatecas south-ward through Jalisco and Oaxaca to the Isthmus of Tehuantepec.Remarks.?An examination of the type of Leuiperus mexicanus(M.H.N. P. No. 226a, parchment label No. 6318) convinced Dr. E.R. Dunn that Parker had erred in referring it to the genus Pleurodemaand that this species was the same as Giinther's Hylodes calcitransfrom western Mexico. In commenting upon this species Parkerstated that:No specimens of this species have been examined, but it has been referred tothis genus rather than to Physalaemus on account of the absence of any tarsaltubercle; its real position is still uncertain. Known only from the type speci-mens from Mexico.Adolphe Boucard is listed as the collector of the type of Leuiperusmexicanus in the catalogue of the Museum National d'Histoire Natu-relle, Paris. The following notes were made on this specimen:Head-and-body length, 39 mm.; the hind limb being carried forwardalong the body, the tibio- tarsal joint reaches to or slightly beyondtip of snout; no tarsal fold; plantar surface of foot without super-numerary tubercles; small inner and outer metatarsal tubercles;faint black crossbars on hind limbs; vomerine teeth in two minuteclusters between the choanae; first finger longer than the second.It is clearly a member of the genus Eleutherodactylus.Brocchi's Hylodes lineatus seems to have been a rose-gray individualwith very narrow vertebral stripe, and in so far as color is concernedit is not unlike other specimens taken in the State of Oaxaca. Al-though a rather thorough search was made in the catalogues of thelaboratory of herpetology of the Paris Museum, no entry of a Hylodesfrom Attitlan was found.
MEXICAN TAILLESS AMPHIBIANS 109In response to an inquiry in regard to the type of Borborocoetesmexicanus, H. W. Parker, of the British Museum of Natural History,repHed as follows:As you suspect, this is probably an Eleutherodactylus. The terminal phalangesarc T-shaped, but the digits are scarcely dilated; fingers and toes subequal dis-tally. No tarsal fold seems to be present.After having examined the cotypes (B.M. Nos. 1892. 2. 8. 66-67),Doctor Dunn corroborated the assumption that Borborocoetes mexi-canus and Eleutherodactylus calcitrans did not represent distinctspecies and affirmed that they were the same. The cotypes of B.mexicanus were collected by Dr. A. C. Buller on the Hacienda elFlorencio, and according to his route map they were taken on August15, 1891, at a locality nearly directly north of San Cristobal on theeastern slope of Sierra de Florencio in Zacatecas. The followingnotes were made on these cotypes
:
B.M. No. 1892. 2. 8. 66: Head-and-body length, 37.4 mm.; trans-verse diameter of tympanum, 2 mm.; transverse diameter of eye,4.2 mm.; anterior edge of eye to nostril, 3.8 mm.; the hind limbbeing carried forward along the body, the tibio-tarsal joint reachesbetween the eye and tip of snout; vomerine teeth in two small clusters,each about the diameter of a choana, widely separated medially,within and considerably behind level of choanae ; no tarsal fold ; innermetatarsal tubercle very large, outer one quite small; large subar-ticular tubercles; supernumerary tubercles on plantar surface of foot;side of head black; a black band from eye to about middle of side;lower hp marbled with black; upperparts light gray; upper surfaceof thigh and tibia with dark cross bands; forearm with two dark crossbands.B.M. No. 1892. 2. 8. 67: Head-and-body length, 28.8 mm.; a darkinterorbital band ; upper surface of snout light colored ; lower and upperlips with light vertical bars; vomerine teeth in same position as in theother cotype; other characters likewise essentially the same as in theother cotype.Doctor Dunn, writing from London, says that Syrrhaphus omilte-manus (cotypes, B.M. Nos. 1901. 12. 19. 7-8) belongs unquestionablyin the genus Eleutherodactylus. Dunn and Parker after examiningthe cotypes reported that they have vomerine teeth but that they arevery much reduced, and that a backwardly projecting vomer is present.The following notes were made on these specunens: Head-and-bodylength of largest individual, 21.5 mm.; transverse diameter of tym-panum, 2.1 mm.; transverse diameter of eye, 2.4 mm.; anterior marginof eye to nostril, 2 mm.; vomerine teeth rather minute; the hind limbbeing carried forward along the body, the tibio-tarsal joint reachesto between the eye and end of snout; apical disks of fingers and toes66785?32 8
110 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
not grooved or divided; a pair of converging linear dorsal dermalridges inclosing an hourglass-shaped area, and a pair of dark spots,one on each side, external to ridge; abdomen and under surface ofthighs areolate; both specimens are young individuals. Syrrhaphusomiltemanus is a strict synonym of E. calcitrans = E. mexicanus.One of the cotypes of Hylodes calcitrans from Jalisco (B. M. Nos.1901 . 12. 19. 35-36) could not be found, and the other one does not haveconverging linear dermal ridges on the upperparts such as are present onsome of the specimens in the United States National Museum. Thefollowing notes were made on this cotype: Head-and-body length,34.5 mm.; transverse diameter of tympanum, 3.6 mm.; transversediameter of eye, 4.3 mm.; anterior edge of eye to nostril, 3.5 mm.;lateral dermal fold extends backward over tympanum and along sideof body; dermal asperities present on sides of body and on back insacral region; large subarticular tubercles; small supernumerarytubercles on plantar surface of foot; unusually large inner metatarsaltubercle and minute conical outer one; no tarsal fold; vomerine teethin two small clusters, each about the diameter of a choana, widelyseparated medially, within and considerably behind level of choanae;skin on abdomen areolate, but smoother than on Omilteme cotypes;light vertical bars on upper lip ; a black canthal stripe, but side of headis not so dark as in Borborocoetes mexicanus; upperparts gray.Nineteen cotypes (B.M. Nos. 1901. 12. 19. 25-43) from Omiltemein Guerrero were received from F. D. Godman. All these haveareolate skin on the abdomen, but this seems to have resulted in partat least from partial desiccation at some time. Only two have smallsupernumerary tubercles on the plantar surface of foot. On all theupper surface of the snout is light colored, the thighs and tibiaehave transverse dark bands, and vomerine teeth are present. Elevenspecimens have distinct medially converging dorsal dermal ridgesthat inclose an hourglass-shaped area. Nine have the anterior endof each of these dermal ridges ending in a black dermal elevation,lateral and posterior to which is a second short black dermal elevationthat is not connected with either of above ridges. Thus on eachside of the vertebral line in the suprascapular region there is a pairof short black dermal ridges. One cotype measured by Giintherhas: Head-and-body length, 36.6 mm.; transverse diameter of tym-panum, 2.5 mm.; transverse diameter of eye, 3.9 mm.; anterior edgeof eye to nostril, 3.3 mm.; no dark band on side of head, though it ispresent behind tympanum to a point about halfway length of side;skin of upperparts fairly smooth; converging linear dorsal dermalridges absent; large inner and small outer metatarsal tubercle; notarsal fold; undersurface of thighs coarsely granular; skin of abdomenareolate; large subarticular tubercles; no supernumerary tubercleson plantar surface of foot; vomerine teeth in short, oblique, posteriorly
MEXICAN TAILLESS AMPHIBIANS 111
converging rows behind and within level of choanae. In case of twoother cotypes, having head-and-body lengths of 38 and 41.7 mm.,respectively, the tibio-tarsal joint reaches to between eye and tipof snout when the hind limb is carried forward along the body.It is apparent from the synonymy of E. mexicanus that some ofthe more pronounced variants of this species have been incorrectlyallocated generically. Some of the confusion that has prevailed inthe past may possibly be explained by the occurrence of occasionalindividuals that lack vomerine teeth. Of the 13 specimens of thisfrog in the National Museum, all from the State of Oaxaca, there aretwo with a white vertebral stripe at least 1 mm. wide, one with awhite pin stripe, and the remainder show no trace of a vertebral line.Most of them have a short dermal ridge above the edge of the scapula.Eight have more or less distinct converging dorsal dermal folds,while five show no trace of such folds. On some the skin of theupperparts is roughened by dermal ridges, and on others it is quitesmooth. The skin on the posterior half of the abdomen is generallyareolate, but occasionally is smooth. No trace of vomerine teethcan be observed in 3 of these specimens, and in the remaining10 these teeth vary not only in the width of the series, but also in theextent to which they project beyond the gum.In general E. mexicanus has a much more variegated appearancethan E. rhodopis. Darker colors predominate, and the black andbrown mottling of the upperparts is especially noticeable. The darkbars on the hind limbs are quite distinct. The tarsal fold is normallyabsent, though this distinction will not hold invariably.Certain specimens from the west coast of Mexico are rather puz-zling, for they approach rhodopis in some details and mexicanus inothers. They are tentatively assigned to this species until a largerseries is available to determine their precise relationships. A short-legged individual (U.S.N.M. No. 47433) collected by Nelson andGoldman at Plomosas in Sinaloa is referred to E. mexicanus partly ongeographical grounds and partly because of the absence of a tarsalfold, although in general appearance it bears some resemblance toE. rhodopis. When the hind limb is carried forward along the body,the tibio-tarsal joint barely reaches the anterior margin of the eye.This dull-colored individual has a dark interorbital streak behindthe light-colored snout, and a pair of short black glandular ridges onthe mid-dorsum. Distinctive markings on the side of the headconsisting of subvertical dark brown bars, separated by narrowerlight spaces, that extend across lower and upper lips to eye and lorealregion are also present.Three specimens (U.S.N.M. Nos. 47905-07) were found aboutsprings and little streams in dense growths of grass and other plantsat 10,000 feet altitude at Cerro San Felipe. One (U.S.N.M. No.
112 BULLETIN 160, UNITED STATES NATIONAL MUSEUM47911) was found in a damp gulch in the shade of bushes and treesat 7,800 feet altitude at La Parada.Dr. Hans Gadow ^^ in reporting his experiences at Omilteme inGuerrero says:An insignificant and diminutive toad {Borborocoetes mexicanus) lived in theirrigated patches of Indian corn, and emitted a sharp, double, piping note. Oneof these specimens was inside an orange-headed grass snake {Tropidonotuschrysocephalus) caught by Rafael, the "arriero," who was much astonished whenhe received the promised pay for two instead of one creature caught.Specimens examined.?Forty-eight, as follows:Eleutherodactylus mexicanus
Museum Num-Catalogue ber ofNo. , speci-mens Locality collected Date collected By whom collected orfrom whom received
r.S.N.MB.MDoU.S.N.MB.M... --DoU.S.N.MC.A.SU.S.N.MDoDoDoDoDo.M.H.N.P
4743392. 2. 8.66-671901. 12.19. 35-3647013-141901. 12.19. 7-81901. 12.19. 25-4347905-0750424791146964-65
47143-50
1002030325-126, -28/47913226a
22
2 19
July 19, 1897Sinaloa: PlomosasZacatecas: Hacienda el Flor-encio.Jalisco: No definite locality..Guerrero: IOmilteme.. 1 May 20, 1903do
.do.Oaxaca:Cerro San Felipe, 10,000 feet Aug. 30, 1894altitude.Cosolapa ! Mar. ? , 1925La Parada, 7,800 feet altitude.! Aug. 19, 1894Mountains west of Oaxaca, Sept. 18, 1894oak woods at 9,400 feetaltitude.Mountains west of Oaxaca, | Sept. 19, 1894oak woods at 6,000-9,500 Ifeet altitude.Tehuantepec 'do .-I-Totontepec ..! July 26, 1894? State: No definite locality.. I
Nelson and Goldman.A. C. Buller.F. D. Godman.Nelson and Goldman.F. D. Godman.Do.Nelson and Goldman.Paul D. R. Ruthling.Nelson and Goldman.Do.Do.
Francis Sumichrast.Do.Nelson and Goldman.Adolphe Boueard.
' Cotypes of Borborocoetes mexicanus.2 Cotypes of Uylodes calcitrans.3 Cotypes of Syrrhophus omiltemanus. * Young.' Type of Leuiperus mexicanus.ELEUTHERODACTYLUS RHODOPIS (Cope)1866. Liihodytes rhodopis Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 18,p. 323, Nov.?Cope, 1869, Proc. Amer. Philos. Soc, vol. 11, no. 82, p. 160,footnote, July 16.
?
Cope, 1893, Proc. Amer. Philos. Soc, vol. 31, no. 142,p. 337, Nov. 17.1868. Hylodes sallaei Gunther, Proc. Zool. Soc. London, p. 487, pi. 38, fig. 3,June 25 (Mexico). Boxjlenger, 1882, Catalogue of the Batrachia Salientias. Ecaudata in the collection of the British Museum, 2d edit., p. 204.1900. Hylodes plicatus GtJNTHER, Biologia Centrali-Americana, Reptilia andBatrachia, p. 228, pi. 66, fig. B, June (Jalapa, Vera Cruz, Mexico).1900. Hylodes rhodopis Gunther, Biologia Centrali-Americana, Reptilia andBatrachia, p. 232, pi. 67, figs. C, C, Aug.
" Gadow, H., Through southern Mexico: Being an account of the travels of a naturalist, London, p. 380,1908.
MEXICAN TAILLESS AMPHIBIANS 113Type locality.?Orizaba and Cordoba, State of Vera Cruz, Mexico.Range.?From Orizaba in Vera Cruz southward through Chiapasto Costa Rica; apparently restricted to the eastern side of the MexicanPlateau.Remarks.?The type of Lithodyies rhodopis was not designated byCope. A specimen (U.S.N.M. No. 16558) from the vicinity of Orizabaagrees with the measurements published by Cope and may be des-ignated as the electotype. Some of the other specimens (U.S.N.M.Nos. 16557, 16559-61) mentioned by Cope in his original descriptionhave been located, and they should be considered as paratypes. In1869, Cope remarked that his L. rhodopis was the same as Giinther'sHylodes sallaei. According to Dr. E. R. Dunn, who has recentlyexamined the type (B.M. No. 57. 7. 31. 27), H. sallaei is unquestion-ably identical with E. rhodopis. This assumption is corroborated bythe following notes made on the type specimen:Head-and-body length, 27.7 mm. ; transverse diameter of tympanum2 mm.; transverse diameter of eye, 3.3 mm.; anterior edge of eye tonostril, 3.3 mm.; the hind limb being carried forward along the body,the tibio-tarsal joint reaches to the end of the snout; inner metatarsaltubercle of medium size, outer one minute; a short tarsal fold endingin a crestlike tarsal tubercle; rows of minute supernumerary tubercleson plantar surface of foot; first finger slightly longer than second;vomerine teeth in two short obHque rows, widely separated medially,well behind, and within level of the choanae; under surface of thighsgranular, abdomen areolate; an exceedingly fine vertebral dermalridge from occiput to vent; an oblique dermal ridge extending frompostero-external angle of eye to short glandular ridge in a black spotat about middle of back, forms inverted chevron with opposite dermalridge; a lateral linear dermal ridge from shoulder backward to andalong sacrum ; and lateral to this another linear dermal ridge extendsfrom posterior edge of eye, above tympanum, and backward alongside to a point slightly beyond middle of body; skin of upperpartswith rather scattered minute glandular asperities; glandular asperitieson sides are rather closely approximated and somewhat linear inarrangement; black streak from tip of snout to nostril and alongcanthus rostralis to eye, and from eye backward above tympanumand downward behind to below level of angle of jaw.The type of Hylodes sallaei was received from Auguste Sall6 andpresumably came from the State of Vera Cruz. Giinther laidconsiderable emphasis on the fact that the skin on the posterior halfof the abdomen was granular, though he admits that in consequenceof imperfect preservation this granulation may be obliterated. Onsome specimens the skin of the underparts seems to be smooth,though an areolated condition may be seen on closer inspection witha lens, and on others the skin is distinctly granulated. This criterion
114 BULLETIN 160, UNITED STATES NATIONAL MUSEUMwas used by Cope as a basis for generic discrimination of Lithodyiesand Hylodes.The type of Hylodes plicatus (B.M. No. 1901. 12. 19. 38) is hereidentified as an immature individual of Eleutherodadylus rhodopis.When direct comparisons were made between the types of H. plicatusand H. sallaei, it was observed that the existing differences are toosUght to have any taxonomic significance. For purposes of compar-ison, the following notes on the type of H. plicatus are here recorded
:
Head-and-body length, 19.8 mm.; transverse diameter of tympanum,1.8 mm.; transverse diameter of eye, 2.8 mm.; anterior edge of eyeto nostril, 2.2 mm.; the hind hmb being carried forward along thebody, the tibio-tarsal joint reaches to end of snout; a long narrowtarsal fold; inner metatarsal tubercle large and elongate, outer quitesmall and conical; minute supernumerary tubercles on plantar surfaceof foot; apical disks not very large; canthus rostralis fairly sharp;vomerine teeth in two small widely separated clusters far behind, andwell within level of, the choanae; a vertebral linear dermal ridge fromsnout to vent; a pair of medially converging dorsal dermal ridges thatinclose an hourglass-shaped area; skin of abdomen apparently areolate;a black streak with light upper margin from nostril to eye and fromposterior edge of eye backward above tympanum, and downwardbehind to above its lower edge; faint transverse dark bars on uppersurfaces of thigh and tibia.Some variation in the development of the tarsal fold may beexpected. Four of the specimens referred to this species have adistinct tarsal fold, three have a low crested tubercle 2 or 3 mm. abovethe base of the inner metatarsal tubercle, but the connecting tarsalridge is barely discernible, and on one the tarsal fold seems to be absent.Exceedingly fine medially converging dorsal dermal folds are generallypresent. Similarly the short black glandular ridges in the mid-dorsal region are present in some and absent in others. With theexception of the vomerine teeth, which are normally behind thelevel of the choanae, no constant character that will invariably dis-tinguish this species is apparent in the series studied.In life this little frog may be dark brown, reddish yellow, orgreen, depending to some extent upon the surroundings. The ventralsurface of the tarsus and of the foot is either dark brown or black.On the forearm there is either a dark-brown spot or a transversedark-brown bar. A dark-brown or black stripe with a light-coloredor white edge extends along the canthus rostralis from snout to eye;beginning again at posterior margin of eye it extends backward abovethe tympanum and along the lateral line for 6 or 8 mm,, or elsecurves downward behind the tympanum.
MEXICAN TAILLESS AMPHIBIANS 115Dr. Hans Gadow ^ contributes the following notes on this species:Of tailless amphibians only one kind exists, but this is rather plentiful, Hylodesrhodopis, a small Cystignalhid, which leads the life of a tree frog. It seems tobe a southerner, which, although not extending onto the plateau itself, ascendsthe high mountains on its eastern, southern and western borders. On Citlalte-petl, for instance, it occurs well up to 10,000 feet, whilst it also inhabits the"hot lands" in the State of Vera Cruz. Most of the specimens were darkbrown, with reddish tints, and lived on or near the ground amongst the darkmasses of rotten leaves; others had made their home in the tillandsia clusters,or on the green shrubs at the edge of the forest; these frogs were quite green,but when caught soon changed to reddish yellow, and ultimately assumed thenatural coloration, which is also that of specimens preserved in spirit. None ofthese frogs in the forest itself, upon the ground, showed a trace of green. Thegenus Hylodes comprises the H. mariinicensis, the "coqui" of the West Indies,which has become famous as the first frog known to lay a few large eggs only,from which within a few days the young are hatched, as almost perfect littlefrogs, they having hurried through the gilled and tailed tadpole stage in a dia-grammatically precipitate fashion. Most kinds of Hylodes seem to go throughsuch a "condensed" process of babyhood; the eggs are laid in a foaming latheror spume wrapped between leaves. Suitable places upon the mountain, shadyplaces with decaying leaves, were swarming with these little baby frogs, but oursearch for nests was without result, 'ince the proper breeding season was alreadypassed, and the adults kept quite mute. In any case, the presence of thesepeculiar frogs was significant, they, with the newts, being the sole representativesof amphibia. There were no other frogs or toads whatever, nor were any knownto occur. Both toads and frogs require standing water in which to deposittheir eggs, and there is no standing water anywhere near the Xometla level.The streams were quick running and very cold; some were fed by melting snow,or became rapid torrents when it rained, and for these reasons they containedno fish either.Specimens examined.?Eleven, as follows:Eleutherodadylus rhodopis
Museum
B.MU.S.N.MDo---.Do---.Do? .B.M
CatalogueNo.
1901. 12. 19.38709116558/ 16557I 59-6147199{57^7.31.}
Num-ber ofspeci-mens Locality collectedVera Cruz:JalapaOrizaba, alpine region
-
Vicinity of Orizaba
?
Date collected
1864 (7)
-do-Chiapas: Chicharras, 3,400 ft.altitude. Feb. 17, 1896?State: No definite locality...! 1855
By whom collected orfrom whom received
F. D. Godman.Francis Sumichrast.Do.Do.Nelson and Goldman.Auguste Sall6.
1 Type of Hylodes plicatus.* Type of LUhodytes rhodopis. ' Paratypes of LUhodytes rhodopis.* Type of Hylodes sallaei.
*? Gadow, H., Through southern Mexico: Being an account of the travels of a naturalist, London, pp. 55,66. 1903.
116 BULLETIN 160, UNITED STATES NATIONAL MUSEUMELEUTHERODACTYLUS RUGULOSUS (Cope)1869. Liylarugulosa Cope, Proc. Amer. Philos. Soc, vol. 11, no. 82, p. 160, July 16.1869. Hylodes berkenbuschii Peters, Monatsber. k. preuss. Akad. Wiss. Berlin,p. 879, Dec. (probably vicinity of Matamoros, Puebla).
?
Boulenger, 1882,Catalogue of the Batrachia Salientia s. Ecaudata in the collection of theBritish Museum, 2d edit., p. 202.?Nieden, 1923, Das Tierreich, Lief. 46,Amphibia, Anura I, p. 424.Type locality. ?Pacific region of the Isthmus of Tehuantepee,Mexico.Range.?Central Vera Cruz southward through the Isthmus ofTehaantepec to San Salvador.Remarks.?The type of Hylodes berkenbuschii (M.N.B. No. 6666)has been examined, independently, by Dr. E. R. Dunn and myself,and both of us agree that it is identical with Cope's Liyla rugulosa.The folloAving notes were made on this specimen
:
Head-and-body length, 42.5 mm.; transverse diameter of tym-panum, 3.9 mm.; transverse diameter of eye, 5.8 mm.; the hind limbbeing carried forward along the body, the tibio-tarsal joint reaches toend of snout; web at base of toes extends to or a little beyond proximalsubarticular tubercle; tarsal fold sharp and quite distinct; inner meta-tarsal tubercle quite large, outer tubercle minute; no sapernumerarytubercles on plantar surface of foot; apical disks of toes smaller thanthose of fingers; canthus rostralis sharp; loreal region oblique; tym-panum overhung by dermal fold; sldn of upperparts rather rough,with a pair of oblique dermal plicae on each shoulder and a linearseries of short interrupted glandular ridges along sides of back; skinof abdomen areolate; under surface of thighs areolate, with a fewwarts below vent; upperparts brown; a dark interorbital transverseband, in front of which the top of the head is suffused with white,extends laterally upon posterior edges of upper eyelids; narrow verticallight yellow stripes on upper lip alternating with subtriangular darkspots; dark stripe above and downward behind tympanum; sides ofbody mottled with a darker color; hind limbs with indistinct darkcross bands; a narrow wliite throat line; underparts brownish yellow,with darker mottling; posterior surfaces of thighs with small whitespots.The two cotypes of Liyla rugulosa (U.S.N.M. Nos. 29771-72)mentioned by Cope are now poorly preserved and rather soft. Por-tions of both specimens are bleached nearly white, and the colormarldngs are obliterated. The skin of the upperparts is somewhatwrinkled, but the dermal glandular ridges are still visible. One ofthe cotypes (U.S.N.M. No. 29772) has the sldn freed from the head,the sternum dissected out, and the extremity of the right sacraldiapophysis exposed. This specimen has a head-and-body length of33 mm., while the other cotype measures 36 mm. in length. A large
MEXICAN TAILLESS AMPHIBIANS 117individual with a head-and-body length of 65 mm., which was col-lected at Achotal in Vera Cruz, is referred to Cope's species.Specimens examined.?Twenty-eight, as follows
:
Eleutherodactylns rugulosus
Museum CatalogueNo. Num-ber ofspeci-mens Locality collected Date collected By whom collected orfrom whom received
F.M.N.H.U.S.N.M..Do....M.N.B..-.U.S.N.M.Do...Do...Do...Do...
71159165676666
479101003829971-723032746850
Vera Cruz:AchotalOrizaba.Vicinity of Orizaba...PUEBLA (?): No definite lo-cality.Oaxaca:Pluma, 3,000 feet altitude...Tehuantepec
....dodoChiapas: Tumbala, 4,000 feetaltitude.
/Dec. ?, 1904\Jan. ?, 1905
Mar. 19, 1895
NovVie.'isM'
E. Heller and C. M.Barber.Francis Sumichrast.Do.Herr Berkenbusch.Nelson and Goldman.Francis Sumichrast.Do.Do.Nelson and Goldman.
' Type of Hylodes berkenbuschii. 2 Cotypes of Liyla rugulosa.ELEUTHERODACTYLUS VENUSTUS (Gunther)1900. Hylodes venustus Gunther, Biologia Centrali-Americana, Reptilia andBatrachia, p. 234, pi. 68, fig. C, Aug.
?
Nieden, 1923, Das Tierreich, Lief.46, Amphibia, Anura I, pp. 409, 425.Type locality.?Jalapa, State of Vera Cruz, Alexico.Range.?Limits of range as yet undetermined.Remarks.?This variety was recognized as a distinct species byGunther chiefly on the account of the presence of a broad purplishband from snout to vent bounded laterally on the back by convergingdermal ridges. These converging dermal ridges are much more widelyseparated than on any of the specimens of E. rhodopis that are avail-able for comparison. A similar color variant has been observed inundoubted specimens of rhodopis. The chevronlike glandular ridgeover the vertebral edge of the scapula is merely the most conspicuousportion of the converging dermal ridge that extends backward fromupper eyelid to the level of the end of the sacral diapophysis. Theskin of the upperparts is granular, that on the throat is smooth, andthat on the abdomen is granular.The type of E. venustus (B.M. No. 1901. 12. 19. 37) has been com-pared with specimens of E. rhodopis, and it unquestionably belongsin the same group as the latter. The species venustus, however,^istentatively retained until a series sufficient to determine its status isavailable for examination. The following notes were made on thetype specimen: Head-and-body length, 36.9 mm.; transverse diam-eter of tympanum, 2.5 mm.; transverse diameter of eye, 3.9 mm.;anterior edge of eye to nostril, 3.8 mm.; the hind limb being carried
118 BULLETIN 160, UNITED STATES NATIONAL MUSEUMforward along the body, the tibio-tarsal joint reaches beyond tip ofsnout; tarsal fold with fairly prominent crestlike tubercle about half-way length of tarsus; inner metatarsal tubercle fairly large, a some-what smaller conical outer one; large subarticular tubercles; a fewsupernumerary tubercles on plantar surface of foot; vomerine teethin two small clusters, each slightly wider than choana, behind, andwithin level of, the choanae; medially converging dorsal dermalridges inclose a semi-hourglass-shaped area; a fine vertebral dermalridge; short dermal ridges and glandular elevations in sacral region;skin of abdomen smooth; a black canthal stripe with light uppermargin from nostril to eye, along outer edge of eyelid to above tym-panum and downward behind to level of its lower margin; a smallblack spot on shoulder behind this stripe; a purple median longitu-dinal stripe from tip of snout to vent extends over top of head andbehind the eyes is bounded laterally by black streaks, which arecoextensive with the converging dorsal dermal ridges; external to thesacrum is a Unear series of indistinctly outUned and irregular darkblotches; outer surface of thigh, tibia, and tarsus with indistinct darkor dusky transverse bars; forearm with black spots; fine dark specksscattered over skin of underparts and concealed surfaces of Umbs.Specimens examined.?Two, as follows:Eleutherodactylus venustus
Museum
MEXICAN TAILLESS AMPHIBIANS 1191882. Hyla crassa Boulenger, Catalogue of the Batrachia Salientia s. Ecaudatain the collection of the British Museum, 2d edit., p. 396.?GtJNTHER, 1901,Biologia Centrali-Americana, Reptilia and Batrachia, p. 281, Sept.Type locality.?Mexico.Range.?Unknown.Remarks.?There is no certainty that this amphibian was actuallycollected in Mexico. As originally described this species was placedin the genus Cauphias, which has for its genotype Cauphias guatema-lensis. Brocchi evidently considered that Cauphias guatemalensis andC. crassum were related to Hylodes [=Eleutherodactylus], since they areplaced after Hylodes in the family Hylodidae. A specimen ofCauphias guatemalensis ^'^ recently taken at Panajachel, near Solola,Lake Atitlan, at an altitude of 4,500 feet in the mountains of Guatemala,conclusively shows that this species is related to Eleutherodactylus.The webbing of the hind feet, however, of both Cauphias guatemalensisand C. crassum is quite unlike any known Eleutherodactylus.The two cotypes of Cauphias guatemalensis (M.H.N.P. No. 509^,parchment label No. 6332) were collected at Pacicilla, Guatemala, byAdolphe Boucard in 1873. They are, respectively, one adult and oneimmature individual. The head-and-body length of the largest indi-vidual is 54,5 mm. The diagnostic characters of this species may bedescribed briefly as follows : The hind limb being carried forward alongthe body, the tibio-tarsal joint reaches to posterior margin or to centerof eye; a well-developed tarsal fold extends full length of tarsus; alarge inner, but no outer metatarsal tubercle; toes fully webbed,slightly scalloped; first finger bifurcated at base; fingers 2, 3, and 4slightly webbed at base; well-developed and rounded apical disks; noexternal indication of tympanum; a thick dermal fold from posteriormargin of eye curves backward and downward to origin of fore limb;vomerine teeth in two transverse rows behind but within level ofinner margins of choanae; canthus rostralis distinct; loreal region sub-vertical; skin of upperparts, top of head, upper surfaces of fore andhind limbs with scattered tubercles; skin of throat, abdomen, andunder surface of thighs coarsely granular; coloration of upperparts(in alcohol) uniformly bluish slate.The first finger of Cauphias guatemalensis has a well-developedclawlike vestige of the pollex buried in a fleshy apophysis, whichseemingly arises from the side of the thumb, giving a pecuHar bifidappearance to that digit. This same structure is present on theimmature individual, but is not developed on the first finger of Cauphiascrassum. There are striking similarities in the general build of thesetwo species that are apparent when they are compared side by side.Boulenger referred this short-toed and nearly fully webbed crassumto the genus Hyla, but this is incorrect since the type has T-shapedM Barbour, T., Cauphias rediscovered. Copeia, no. 165, pp. 96-98, Dec. 23, 1927.
120 BULLETIN 160, UNITED STATES NATIONAL MUSEUMterminal phalanges. Cauphias crassum with its hidden tympanum istentatively recognized as a distinct species, though there are someindications that subsequent collecting may show that it is merely afemale of Cauphias guatemalensis.The type of Cauphias crassum (M.H.N.P. No. 509a, parchmentlabel No. 6331) was likewise collected by Adolphe Boucard in 1873,according to the catalogue somewhere in Mexico, though it is quitelikely that this locality may be incorrect. The following notes weremade on the type of this species: Head-and-body length, 54.8 mm.;the hind limb being carried forward along the body, the tibio-tarsaljoint reaches to center of eye; tarsal fold distinct and strong; a smallinner but no outer metatarsal tubercle ; toes fully webbed, with excep-tion of fourth toe on which the penultimate joint has a dermal fringe;terminal phalanges T-shaped; first finger without basal bifurcation;fingers without web at base; no external indication of tympanum; adermal fold above and behind normal position of tympanum; distancefrom eye to nostril less than transverse diameter of eye; vomerineteeth in two rounded clusters between choanae; loreal region oblique;sldn on abdomen coarsely granular; throat with scattered granula-tions; under surface of tliighs areolated; coloration of upperparts gray,without darker markings; slight grayish marbling on upper lip; sideslight with gray vermiculation ; throat with gray marbling; abdomenand under surfaces of thighs yellowish.Specimens examined.?One, the type.Genus TOMODACTYLUS Giinther1900. Tomodactylus Gunther, Biologia Centrali-Americana, Reptilia and Batra-chia, p. 219, Apr. [Genotype, Tomodactylus amulae Gunther, idem, p. 219,pi. 64, fig. C (Amula, Guerrero, Mexico).]For many years this unique little frog was confused with Physalae-mus and Pleurodema, chiefly on account of the occurrence of a blacklumbar spot on some of the species referred to these genera. Thereare species of Pleurodema that have a lumbar gland, but they havesimple terminal phalanges and a sternum with a bony style. Tomo-dactylus, however, has T-shaped termmal phalanges and a cartilaginoussternum and omosternum, but lacks vomerine teeth. It is here recog-nized as a vahd genus related to Eleutherodactylus and Pleurodema.TOMODACTYLUS NITIDUS (Peters)1869. Liuperus [sic] nitidus Peters, Monatsber. k. preuss. Akad. Wiss. Berlin,p. 878, Dec.1900. Tomodactylus amulae Gunther, Biologia Centrali-Americana, Reptilia andBatrachia, p. 219, pi. 64, fig. C, Apr. (Amula, State of Guerrero).1927. Paludicola [Liuperus] nitidus Parker, Ann. and Mag. Nat. Hist., ser. 9,vol. 20, no. 118, p. 451, Oct.
MEXICAN TAILLESS AMPHIBIANS 121Type locality.?State of Puebla, Mexico, possibly from the vicinityof Matamoros.Range.?Exact limits of range unknown; records of capture indicatethat it occurs from southern Nayarit (El Ocotillo) and Guanajuato,southward through Paebla and Michoacan to central Guerrero(Chilpancingo).Remarks.?The earliest available specific name for this small frogseems to be Liuperus [sic] nitidus Peters. This name was based on aspecimen (M.N.B. No. 6669) that was included in a small collectionmade by Herr Berkenbusch in Mexico. Inasmuch as the diagnosticfeatures of this species have been ignored for so many years, a shortdescription of the type of nitidus is herewith given: Head-and-bodylength, 20 mm.; transverse diameter of tympanum, 1 mm.; transversediameter of eye, 1.8 mm.; the hind limb being carried forward alongthe body, the tibio-tarsal joint reaches to anterior margin of tympan-um; tarsal fold very indistinct, if present; large subarticular tubercles;minute supernumerary tubercles on sole of foot; fore limbs slender;toes slender; fourth toe considerably longer than third or fifth; skin ofabdomen smooth; under surface of thighs areolate; no visible vomerineteeth; coloration, seemingly dark brown with light marbling; lumbargland white with wavy or zigzag dark bands; transverse dark bands onhind limbs rather indistinct.Brocchi likewise referred nitidus to Leiuperus, while Boulenger,Giinther, and Nieden interpreted the original description as indicat-ing that this species should be allocated in the genus Paludicola.Giinther seemingly ignored the resemblances to his Tomodadylusamulae that are clearly disclosed in the pubUshed description ofnitidus. The obvious similarity of the specimens dealt with in thesetwo descriptions is at once apparent when they are compared side byside. Parker seems to have been the first to suggest that the correctrelationships of nitidus are with Tomodadylus and cites as follows hisreasons for this allocation:A specimen in the British Museum which agrees with the type in all externalcharacters is scarcely distinguishable from the type-specimen of Tomodadylusamulae. Like the latter it also has T-shaped terminal phalanges, cartilaginoussternum and omosternum, and a lumbar gland, and so must be referred to thegenus Tomodadylus, at least until the position of this genus has been satisfactorilydetermined.Inasmuch as it seemed improbable that nitidus and amulae weredistinct species, the cotypes of the latter (B.M. Nos. 1901. 12. 19.9-12) have been reexamined. They comprise one adult and threeyoung. The following notes were made on the adult specimen : Head-and-body length, 23.8 mm.; transverse diameter of tympanum, 0.8mm.; transverse diameter of eye, 2.9 mm.; anterior edge of eye tonostril, 2.4 mm.; the hind limb being carried forward along the body,the tibio-tarsal joint reaches to center of tympanum; no tarsal fold;
122 BULLETIN 160, UNITED STATES NATIONAL MUSEUMinner metatarsal tubercle slightly larger than outer one; large sub-articular tubercles; small supernumerary tubercles on plantar surfaceof foot; ball-like apical disks on fingers; first finger shorter than second;no vomerine teeth; tongue subpyriform in outline, indistinctly notchedbehind; abdomen and under surface of thighs coarsely areolate.Some individual variation in the general color pattern was observedin the small series of specimens examined. The darker markings aremuch more conspicuous on some than on others, and the ground colorvaries from olive-gray to reddish brown. Some of the specimens havea light-colored lumbar gland with wavy or zigzag dark streaks. Theothers have the lumbar gland largely black wdth branching lightstreaks or with Hght centers. A surprising degree of variation existsin the relative measurements of the body, tympanum, and eye. Fourspecimens, one of which is from Guerrero (U.S.N.M. No. 47012), twofrom Nayarit (A.M.N.H. Nos. 12552-53), and one from Puebla(A.M.N.H. No. 13867) measure, respectively, as follows: Head-and-body length, 26.8, 22.4, 25.2, and 26 mm.; transverse diameter oftympanum, 1.3, 0.9, 1.2, and 1 mm.; transverse diameter of eye, 3.1,2.9, 3, and 3.7 mm.; anterior edge of eye to nostril, 2.8, 2.3, 2.5, and2.6 mm.Examination of specimens in the British Museum of Natural His-tory by Dr. E. R. Dunn disclosed three additional records for this frog.These locahties are: Guanajuato (No. 1898. 7. 19. 10); Tancitaro inGuerrero (Nos. 1914. 1. 28. 176-180); and Chilpancingo in Guerrero(No. 1906. 6. 1. 1899).The single known species may be recognized by the folloAving com-bination of characters: Coloration of upperparts brownish, grayish,or olive gray, spotted or splotched with some darker color or withblack; a dark triangular interorbital spot sometimes present; snoutin front of transverse interorbital light line rarely light colored; lipborder spotted with brown or green; lumbar gland coextensive withan oval partly brown or partly black spot, with two or three whitedots, or with hght background and zigzag dark streaks; fore and hindlimbs with brown or black cross bands; a conspicuous light-coloredspot on anterior surface of upper end of thigh; remainder of thighbroadly marbled with brown and white; coloration of underparts uni-form white, or spotted with black, or brownish with obscure hghtstipphng; canthus rostralis rounded; snout subacuminate and rounded;loreal region subvertical ; tympanum vertically oval or circular, aboutone-third diameter of eye ; diameter of eye a little more than or equiv-lent to its distance from nostril; body slender; tongue entire or indis-tinctly notched behind; maxillary teeth present; no vomerine teeth;sacral diapophysis not widened; skin of upperparts fairly smooth, butwith scattered rugosities and small tubercles; skin of abdomen smoothor faintly granulated; fore Umb slender; first finger slightly shorter
MEXICAN TAILLESS AMPHIBIANS 123than or subequal with second; fingers and toes quite free, with smallbut distinct terminal disks; subarticular tubercles very prominent;supernumerary tubercles continued backward on palmar and plantarsurfaces; two distinct metatarsal tubercles; the hind limb being carriedforward along the body, the tibio-tarsal joint reaches the tympanumor at most to the posterior margin of the eye; head-and-body length,20 to 26.5 mm.; adults probably 25 to 26.8 mm.Specimens examined.?Nine, as follows
:
Tomodactylus nitidus
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124 BULLETIN 160, UNITED STATES NATIONAL MUSEUMA taxonomist should act as an impartial judge in matters relating tothe validity of species, for it is his business to ascertain how manyactually exist and, having done so, to designate the characters bywhich they may be recognized. Unfortunately, the lack of a uniformyardstick for measurement of distinctness makes the recognition ofspecies a matter of interpretation, since different sets of charactersmust unavoidably be used in the discrimination of species in unrelatedand occasionally in related genera.Specimens collected half a century or more ago are generally sopoorly preserved that any precise determination of their structuralpeculiarities is out of the question. Freshly preserved specimenswould be much better to work with than some of the types of Mexi-can species of Syrrhophus described by Cope. A satisfactory eluci-dation of the Mexican species of Syrrhophus will necessarily awaitthe accumulation of an adequate series of well-preserved or of livingindividuals, preferably topotypes. For the present the toothlesseleutherodactylids to which names have been applied are listedmerely as nominal species without regard to their possible relation-ships, since in most instances they are known solely from the materialavailable to the original describer.According to H. W. Parker ^' the large eggs of Syrrhophus suggesta shortened life history.KEY TO MEXICAN SPECIES OP SYRRHOPHUS1. Tympanum very small, one-third or at most less than one-halfthe diameter of the eye 2Tympanum larger, one-half to two-thirds the diameter of theeye2. Smooth skin on hinder half of abdomen; tympanic membraneone-third the diameter of the eye; canthal region rounded,with no discernible ridge; loreal region subvertical; muzzlebroadly acuminate and obtuse; head flat above, the maxi-mum width of upper eyelid two-thirds the width of interor-bital region; nares lateral and terminal, and their distancefrom eye less than one-half the diameter of the latter; tonguesubpyriform; skin of upperparts, sides, throat, chest, andbelly smooth; underside of thighs coarsely granulated; fingersfree, without any vestige of membrane at base; first fingershorter than second; large subarticular or supernumerary tuber-cles at ends of metapodials of fingers; apical disks of fingersand toes slightly wider than corresponding digit, but notespecially enlarged, truncated at extremity, and flattened;a large palmar tubercle; toes with slight vestige of web atbase; free portion of fourth toe more than twice the length offree portion of fifth; small subarticular or supernumerarytubercles extended backward on inferior surfaces of metatar-sals; inner metatarsal tubercle larger than outer; the hind limbbeing carried forward along the body, the tibio-tarsal jointreaches about the middle of the eye; the hind limbs being
? Parker, H. W., Ann. and Mag. Nat. Hist., ser. 9, vol. 20, no. 118, p. 452, Oct., 1927.
3
MEXICAN TAILLESS AMPHIBIANS 125placed at right angles to the long axis of the body, the heelstouch but do not overlap; upperparts dark mulberry brown,dotted with moderately large gray spots (yellow in life) ; loresand upper lip black; underparts pale pinkish brown, withoutmarkings; head-and-body length, 24 mm lepras (p. 128)Areolate skin on hinder half of abdomen; tympanic membranemore than one-third but less than one-half the diameter of theeye; canthal region rounded, with no discernible ridge, lorealregion subvertical; head fiat above; maximum width of up-per eyelid one-half to two-thirds the width of interorbitalregion; external nares small, rather widely separated, and theirdistance from the eye slightly less than diameter of the latter;tongue subpyriform; skin of upperparts and sides apparentlysmooth, but actually areolated as seen under the lens;areolation on lower half of sides visible without aid of lens;skin of underparts smooth, except posterior half of abdomen,which is areolated; inferior surfaces of thighs coarsely gran-ular; fingers free, without vestige of membrane at base; firstfinger shorter than second; large subarticular or supernum-erary tubercles continued backward on metacarpals; a largepalmar callosity; apical disks slightly wider than finger,truncated at extremity, and apparently flattened; toes withslight vestige of web at base; free portion of fourth toe twiceas long as free portion of fifth; subarticular or supernumerarytubercles especially large at ends of metapodial bones; plantarsurface of foot covered with numerous minute tubercles;inner metatarsal tubercle larger than outer; no tarsal fold;apical disks of toes smaller than those of fingers; the hindlimb being carried forward along the body, the tibio-tarsaljoint reaches to posterior edge or to middle of eye; the hindlimbs being placed at right angles to the long axis of the body,the heels overlap slightly; upperparts (in alcohol) rangingfrom dark vinaceous-drab to brownish drab, marked withnumerous blue-black spots without sharply defined borders,but tending to merge with ground color of upperparts; ablue-black mark curving from above downward behind tym-panum; fore and hind limbs with dark cross bands; externaldigit of hand and foot with faint cross bands; upper andposterior surfaces of femur with white or light-colored spotson a darker ground color; edge of upper lip with a few lightsplotches; darker streak along canthal region from externalnarial opening to eye; ground color much lighter on sides,with distinct light spots; inferior surfaces light colored, un-spotted; head-and-body length, 22 mm guttilatus (p. 127)Skin of upperparts distinctly roughened, with wartlike tuber-cles; sides areolated; tympanum very distinct, in diameterequal to one-half to two-thirds of that of eye; canthal regionrounded, no discernible ridge; loreal region vertical; snouta little longer than diameter of eye; skin on throat and breastsmooth; hinderpart of belly areolated; underside of thighsclosely granulated; first finger shorter than second, fourth alittle longer than second; distinct subarticular tubercles;apical disks moderately large, but those of two outer fingersappreciably larger than the tolerably small disks of the toes;66785?32 9
126 BULLETIN 160, UNITED STATES NATIONAL MUSEUMtoes free, progressively increasing in length from first to fourth,the fifth being a little shorter than third; large subarticulartubercles extended backward upon inferior surfaces of metatar-sals; plantar surface covered with small supernumerary tuber-cles; inner and outer metatarsal tubercles slightly if at alllarger than the subarticular tubercles; upperside of body andlimbs yellow-brown, spotted and stippled with black; a darkstreak along canthal region and above tympanum; large tri-angular black mark with apex pointing backward occasionallypresent in interorbital region; loreal and temporal region black-ish brown; underside brownish yellow, sprinkled with dark;a black mark over large tympanum; head-and-body length,20 mm verruculatus (p. 129)Skin of upperparts generally apparently smooth, but oftenareolate, and rarely shagreen 44. Tympanum subround, about one-half, occasionally less, diameterof eye; canthal region rounded, with no discernible ridge;loreal region subvertical; muzzle rather elongate, with nar-rowed and rounded extremity; nares lateral and their distancefrom eye less than diameter of the latter; tongue pyriform,much narrowed in front; skin of upperparts and sides appar-ently smooth, but actually areolated under the lens; skin ofthroat and chest smooth; hinder half of belly smooth or areo-late; inferior surfaces of thighs coarsely granulated; fingersfree, without any vestige of membrane at base; first fingershorter than second; large subarticular tubercles at ends ofmetapodials of fingers; apical disks somewhat wider than cor-responding finger and appreciably larger than small disks onthe toes; a large palmar callosity; toes with slight vestige ofweb at base; free portion of fourth toe more than twice thelength of free portion of fifth; small subarticular tuberclescontinued backward on inferior surfaces of metatarsals; innermetatarsal tubercle slightly larger than outer; the hind limbbeing carried forward along the body, the tibio-tarsal jointreaches either to the middle or to the anterior margin of theeye; the heels being placed at right angles to the long axis ofthe body, the heels overlap; upperparts reddish or dark mul-berry brown, thickly spotted or speckled with large darkerbrown or blackish blotches, the largest of which reaches tobetween the eyes; limbs obscurely cross-banded; a black ordark mark extends from above to behind tympanum; upper lipwith a few pale spots; loreal region dark colored or with darkstreak along canthal region from external narial opening toeye; sides paler, with distinct spots; underparts uniformlyleather brown or yellowish; head-and-body length, 22.5 mm.cystignathoides (p. 127)Tympanum more than two-thirds the diameter of the eye; can-thus rostralis obtuse or blunt; loreal region vertical; head flatabove, moderately wide, with muzzle slightly prominent;external nares almost at extremity of muzzle; skin every-where smooth except on the sides, which are tubercular areo-late; no abdominal or gular fold; apical disks of fingers a littlelarger than those of toes; one large palmar tubercle; toes ofmoderate length, with small disks, and with prominent sub-articular tubercles, which are especially large at ends of meta-
MEXICAN TAILLESS AMPHIBIANS 127podial bones; two small metatarsal tubercles; the hind limbbeing carried forward along the body, the tibio-tarsal Jointreaches the anterior edge of the eye; color above, dark bluishgray, marked with numerous black spots with ill-definedborders; limbs similar, with black cross bands, which arecontinued on external digit of each foot; closely spaced whitespots on dark ground color of upper surface of femur; a fewsmall white spots on the dark posterior face of femur; edge oflips with a few light spots; underparts, white, unspotted;head-and-body length, 23.5 mm verrucipea (p. 128)SYRRHOPHUS CYSTIGNATHOIDES (Cope)1877. Phyllobates cystignathoides Cope, Proc. Amer. Philos. Soc, vol. 17, no. 100,p. 89, July 20.Type locality.?Potrero, near Cordoba, State of Vera Cruz, Mexico.Range.?Not known to occur elsewhere than at the type locality.Remarks.?Cope stated that "numerous specimens of this specieswere found by Francis Sumichrast at Potrero, near Cordoba, VeraCruz, under the decayed trunks of trees." No other observations onits habits have been published. There are now eight specimens inthe jar containing the cotypes of this species, although it is possiblethat a larger series was loaned Cope. At that time Cope was activelyengaged in extending our knowledge of the herpetology of North andSouth America, and incoming material apparently was often shippedby the Smithsonian Institution directly to Cope without being un-packed. Cope managed to publish reports on these collections aboutas fast as they were received. The cotypes of cystignathoides havethe fronto-parietals meeting medially, eliminating the fontanelle. Notype is designated, and only one specimen (U.S.N.M. No. 32402)has the skin dissected away from the cranium to expose the fronto-parietal region. The following measurements were taken from acotype (U.S.N.M. No. 32404): Head-and-body length, 22.5 mm.;transverse diameter of tympanum, 1.2 mm.; transverse diameter ofeye, 3 mm.; anterior edge of eye to nostril, 2.3 mm. It is quite pos-sible that these cotypes may be the young of Leptodactylus melano-notus, as all of them have a dark occipital spot that extends laterallyupon the upper eyelids in the posterior interorbital region.Specimens examined.?The eight cotypes (U.S.N.M. Nos. 32402-09) collected by Francis Sumichrast at Potrero, near Cordoba, VeraCruz. SYRRHOPHUS GUTITLATUS (Cope)1879. Malachylod.es guttilatus Cope, Proc. Amer. Philos. Soc, vol. 18, no. 104,p. 264, June 20.Type locality.?Guanajuato, State of Guanajuato, Mexico.Range.?So far known to occur only in the sierras of southern SanLuis Potosi and in the vicinity of Guanajuato.
128 BULLETIN 160, UNITED STATES NATIONAL MUSEUMRemarks.?Cope considered that the presence of a fronto-parietalfontanelle was of sufficient importance to warrant a generic separationof this form from related species of Syrrhophus. The type of thisspecies (U.S.N.AI. No. 9888) should be compared with the young ofLeptodactylus melanonotus . It was collected by Alfred Duges at Guana-juato and is now somewhat shrunken and faded. It measures asfollows: Head-and-body length, 21.7 mm.; transverse diameter oftympanum, 1.2 mm.; transverse diameter of eye, 3 mm.; anterioredge of eye to nostril, 2.2 mm. The terminal phalanges of the toeson left hind foot are missing.Specimens exajnined.?Three, as follows:Syrrhophus guttilatus
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MEXICAN TAILLESS AMPHIBIANS 129Remarks.?The present location of the type of this species is un-known. It is possible that it may be found in the Academy ofNatural Sciences of Philadelphia along with other types from Cope'scollection, since it formed part of a small collection made by Copeand Dr. Santiago Bernad. No specimens referable to this specieswere found in any American institution.Specimens examined.?None.SYRRHOPHUS VERRUCULATUS (Peters)1870. PhyUobates verruculatus Peters, Monatsber. k. preuss. Akad. Wiss.Berlin, p. 650, Aug.Type locality.?Huanusco [==? Huatusco, State of Vera Cruz,Mexico].Range.?From the Rio Necaxa in northeastern Puebla southward tocentral Vera Cruz (? Huatusco).Remarks.?It seems quite probable that some mistake in the originallabel or in the printing of the description led Peters to publishHuanusco as the type locality. No place name with this spelling waslocated on any of the Mexican maps, and it would appear that Huat-usco in Vera Cruz was the intended locality.The genus Hypodictyon was proposed by Cope in 1885 for specieswith an areolated abdomen. Peters's verruculatus was one of the threespecies mentioned as possessing this character. A reexamination ofthe cotypes of cystignathoides shows that they likewise have an areo-lated abdomen and this condition also exists in guttilatus. Faultypreservation may account for some of the differences noted. On thebasis of available material it would seem that this areolated conditionof the skin is of little significance.The type of PhyUobates verruculatus (M.N.B. No. 6957), accordingto Doctor Dunn, has a large tympanum with a black mark over it,no distinct terminal disks, heavy subarticular tubercles, no vomerineteeth, and a parotoid gland. The lumbar gland is not present. Aspecimen in the British Museum (No. 1906. 6. 1. 100) from BuenaVista, Guerrero, is referred to this species by Dunn.Specimens examined.?One (U.M.U.M. No. 2), collected at Necaxa,Puebla, July 5, 1926, by H. B. Baker.Family HYLIDAE Gunther1858. Hylidae Gunther, Catalogue of the Batrachia Salientia in the collectionof the British Museum, p. 96.Tropical forests must have tree frogs, according to Dr. Hans Gadow,and in each region these are fashioned from the nearest availablematerial. In Mexico, most of the frogs that hve in the trees belongto the family HyHdae, but elsewhere in Central and South America,as well as in Mexico, there are a number of less closely related frogs
130 BULLETIN 160, UNITED STATES NATIONAL MUSEUMthat have been adapted to an arboreal life. The ends of the fingersand toes are provided with adhesive disks, by means of which thesetree frogs are able to obtain a firm foothold on relatively smoothsurfaces, though they are employed mainly for climbing and forclinging to foliage and limbs when jumping. Diverse ecological nichesare filled mth different kinds of animals, and it is therefore not sur-prising that Mexico with its varied climate and topography shouldbe endowed with a remarkable assemblage of hylids. Not less than7 genera and 20 species are recognized in this review.Most peculiar of all these hylids is the marsupial frog Gastrotheca,in which the whole brood is sheltered in a common pouch on the backof the female. Another interesting genus, Phyllomedusa, is repre-sented herein by three species. Some tree frogs of the genus Phyllo-medusa are known to lay their eggs in nests of froth attached to leaves,but unfortunately the life histories of those that occur in Mexico areunknown, and it is barely possible that all of them may not haveacquired this curious habit. The eggs of one species of the diminutivetoothless Hylella are laid in the sheathing leaf bases of a dust-feedingplant that grows on tropical trees.A curious adaptation to the habit of digging occurs in the genusPternohyla, which differs from other hylids in having a shovellikeinternal metatarsal tubercle. Tree frogs without a visible covering ofskin on their heads also occur in Mexico. Besides Gastrotheca andPternohyla, there are two other little-known genera, Diaglena and Tri-prion, with bony casques. The presence of palatine or parasphenoidteeth in Diaglena and their absence in Triprion are quite inexplicable.The gradation in the size of the adhesive disks possessed by theMexican hylas suggests that arboreal adaptation has not progressedwith equal rapidity and that some species of the genus Hyla are betteradapted for climbing than others. The hind limbs of most of thesehylas are elongated for leaping, though the ability to jump seems tovary with the different species. One of the larger species, H. venulosa,has been known to leap and alight without injury from a height of140 feet. The extent of the webbing between the toes varies in thedifferent species. In H. mioiympanum the toes are four-fifths webbed.Some individuals of H. eximia have the toes not more than halfwebbed.Within certain limits tree frogs can change their general colorationso that their bodies will blend more perfectly with the surroundings.These changes in color are more marked in some species than in others.One of the most widely distributed Mexican species, H. baudinii,seems to have an exceptional color range. Most of the Mexican hylashave their own distinctive color patterns or markings and, while thegeneral coloration is not necessarily dull, brilliant hues are nevercontiguous.
MEXICAN TAILLESS AMPHIBIANS 131KEY TO GENERA OF MEXICAN HTLIDAE1, Derm of head of adults completely involved in cranial ossifi-cation, forming a bony casque 2Derm of head of adults not involved in cranial ossification andnot forming a bony casque; head covered with soft loose skin;no parasphenoid teeth 52.^Posterior edge of bony casque studded with high, conical, erect,bony spines; top of head bony; canthus rostralis sharplydefined, granular; snout rounded; loreal region concave;upper labial borders smooth, not projecting conspicuouslybeyond lower jaws; supratympanic ridge strongly elevated,tubercular; supraciliary crest tubercular; interorbital spaceconcave; vomerine teeth in two nearly transverse rows on levelwith posterior margins of choanae; no parasphenoid teeth;tongue subcircular, not or slightly notched behind; pupil hor-izontal, circular when contracted; tympanum circular, two-thirds diameter of eye; fingers free; toes one-third webbed;adhesive disks of fingers and toes smaller than tympanum;tarsal fold distinct; large internal metatarsal tubercle; noouter metatarsal tubercle; female with dorsal pouch.-.Qastrotheca (p. 133)Posterior edge of bony casque more or less upturned, but withouthigh conical spines; upper labial borders projecting conspic-uously beyond lower jaws; snout extended beyond symphysisof lower jaws; females without dorsal pouch 33. Inner metatarsal tubercle greatly enlarged and compressed;outer metatarsal tubercle quite small; eye separated fromouter margin of flaring labial border of upper jaw by an intervalequivalent to more than one-third of its diameter; posterioredge of rugose bony casque not upturned; canthus rostralisraised, curved, separating external nares; snout rounded;loreal region very wide, concave; upper labial borders pro-jecting and slightly upturned; interorbital space broader thanupper eyelid, deeply concave; vomerine teeth in two groupsbetween choanae; no parasphenoid teeth; tongue circular,notched behind; pupil transversely oval; tympanum oval,one-half to two-thirds diameter of eye; fingers slender, freeor with vestigial web at base; no projecting rudiment of poUex;toes slender, one-third webbed; adhesive disks of fingers andtoes quite small, considerably smaller than tympanum; sub-articular tubercles small, prominent; tarsal fold absent orfaint; skin of back closely granulated; underparts and inferiorsurface of thighs coarsely granulated; upperparts brov/nish,with large elongate insuliform or anastomosing dark-brownblack-edged spots; thighs sulphur yellow, marbled with darkbrown; upper surfaces of fore and hind limbs with large darkbrown transverse spots; underparts white Pternohyla (p. 135)Inner metatarsal tubercle not greatly enlarged; eye almost incontact with outer margin of flaring labial border of upperjaw; tympanum distinct; transverse vomerine and longitudinalparasphenoid teeth present, the former in two small clustersand the latter in a single narrow row 44. Palatine teeth present, forming slightly curved row on each sideposterior to choanae; vomerine teeth in two narrowly separatedclusters between choanae; parasphenoid toeth small, closelv
132 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
set, and forming single longitudinal series; top of head bony;canthus rostralis or preorbital ridge very little elevated, ter-minating posteriorly in an obtuse tuberosity above anteriormargin of eye, meeting the opposite one at level of or in frontof the external nares, and continuing forward to muzzle asnarrow elevated ridge; crenelations on superior labial bordersextending inward at least halfway to canthus rostralis; upperlabial margin moderately thick, abruptly elevated in front ofeye, producing there a sharp angle from which it curves or slopestoward muzzle and tympanum; supratympanic ridge not verysharp; posterior margin of bony casque thick, rugose; inter-orbital space rather concave; tongue not or slightly notchedbehind; pupil transversely oval when expanded, subquadran-gular when contracted, but never vertical; tympanum oval orcircular, two-thirds diameter of eye; fingers free; toes one-half webbed; adhesive disks of fingers and toes much smallerthan tympanum; short tarsal fold; upperparts light olive,with dark spots Diaglena (p. 137)Palatine teeth absent; vomerine teeth in two narrowly separatedclusters between choanae; parasphenoid teeth small, closelyset, and forming single longitudinal series; top of head bony;canthus rostralis developed as high preorbital ridge, which iscontinuous or nearly so with the opposite one considerablybehind external nares, terminating posteriorly in strong pro-jection above anterior margin of eye, and not continuedforward in front of external nares as elevated ridge in case ofadults; upper labial margin thin, curved, but not abruptlyelevated in front of eye; prominent bony supratympanic ridge;posterior margin of bony casque thin, upturned, and nearlystraight; interorbital space deeply concave; tongue subcircular,slightly notched behind; pupil vertical; tympanum oval, one-half diameter of eye; fingers united by vestigial web at base;toes one-half webbed; adhesive disks of fingers and toessmaller than tympanum; indistinct tarsal fold; coloration ofupperparts brownish, with darker spots; head with lighterspots Triprion (p. 138)6. Pupil vertical; vomerine teeth generally present in two obliqueclusters between choanae; palatine and parasphenoid teethabsent; tympanum distinct, more than one-half diameter ofeye; canthus rostralis rounded; upper labial margin not pro-jecting; tongue entire or slightly notched behind; interorbitalspace broader than upper eyelid, imperceptibly convex;webbing between fingers and toes variable in extent; adhesivedisks of fingers and toes varying from less than half to morethan the diameter of the tympanum; subarticular tubercles offingers and toes distinct; first finger shorter than second;first and second fingers opposable; a projecting rudiment ofthe pollex; a faint tarsal fold present; outer metatarsalsunited; diapophyses of sacral vertebrae strongly dilated;mesosternum cartilaginous, posteriorly notched and rounded.Phyllomedusa (p. 139)Pupil horizontal; tympanum distinct or indistinct 66. Vomerine teeth present; tongue not or slightly notched behind;fingers and toes with adhesive disks, the former free or webbed,the latter always webbed Hyla (p. 147)
MEXICAN TAILLESS AMPHIBIANS 133Vomerine teeth absent; tongue circular or suboval, slightlynotched behind; fingers and toes with adhesive disks, theformer webbed at the base and the latter extensively webbed.Hylella (p. 179)Genus GASTROTHECA Fitzinger1843. Gastrotheca Fitzinger, Systcma reptilium, fasc. 1, p. 30. [Genotype, Hylamarsupiata Dum^ril and Bibron, 1841, Erp^tologie gen^rale, vol. 8, p. 598,pi. 98 (Cuzco, Peru).]1854. Notodelphys Lichtenstein and Weinland, Monatsber. k. preuss. Akad.Wiss. Berlin, p. 373, July. [Genotype, Notodelphys ovifera Lichtensteinand Weinland, idem, p. 373 (Puerto Cabello, Venezuela).] [Preoccupiedby a genus of Entomostraca, Notodelphys Allman, 1847, Ann. and Mag.Nat. Hist., ser. 1, vol. 20, no. 130, p. 2, July.]1858. Nototrema Gunthek, Catalogue of the Batrachia Salientia in the coUec-V. , tion of the British Museum, p. 115. [Genotype, Hyla marsupiata Dumeril
..' and BiBRON, 1841, Erp^tologie gen(5rale, vol. 8, p. 598, pi. 98 (Cuzco, Peru).]1858. Opisthodelphys Gunther, Catalogue of the Batrachia Salientia in thecollection of the British Museum, p. 117. [New name for NotodelphysLichtenstein and Weinland, preoccupied.]One of the most extraordinary adaptations for protecting the devel-oping eggs is found in the marsupial frog Gastrotheca. The female hasa dorsal pouch into which the eggs are probably introduced by the male.The whole development of the embryo, including the gilled stage, ispassed within the gelatinous capsule of the egg in some of the species.In some of the South American species, particularly G. marsupiata,about 100 large eggs are deposited in the dorsal pouch and only a partof the metamorphosis is undergone within the egg. In this speciesthe young escape from the common brood pouch in the tadpole stage.In other species, especially G. ovifera, a few unusually large eggs areintroduced into the dorsal brood pouch, and there the young remainuntil the metamorphosis is completed. The life history of the Mexicanspecies, G. coronata, is unlaiown.GASTROTHECA CORONATA Stejneger1858. Opisthodelphys ovifera Gunther, Catalogue of the Batrachia Salientia inthe collection of the British Museum, p. 117 (Cordoba, Mexico). [NeeNotodelphys ovifera Lichtenstein and Weinland, 1854, Monatsber. k.preuss. Akad. Wiss. Berlin, p. 373, July; and Weinland, 1854,in Joh. Miiller'sArchiv. f. Anat. u. Physiol., Berlin, pp. 449-477, pis. 17-19 (Puerto Cabello,Venezuela) .]1882. Nototrema oviferum Boulenger, Catalogue of the Batrachia Salientia s.Ecaudata in the collection of the British Museum, 2d edit., p. 418.
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Gun-ther, 1901, Biologia Centrali-Americana, Reptilia and Batrachia, p. 288, pL74, fig. A, Sept.1911. Gastrotheca coronata Stejneger, Proc. U. S. Nat. iMus., vol. 41, no. 1857,p. 287, Aug. 14.Type locality.?Palomo, Valle de Orosi, Cartago, Costa Rica.
134 BULLETIN 160, UNITED STATES NATIONAL MUSEUMRange.?Exact limits of range unknown; occurs at Cordoba, VeraCruz, Mexico, as well as in vicinity of type locality in Costa Rica.No specimens have been recorded from intermediate localities.Remarks.?In commenting upon Gastrotheca coronata, Dr. L.Stejneger concluded that it is
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probably the same as the young specimen from Carillo, Costa Rica, described andfigured by Guenther in the Biologia Centrali-Americana as Nototrema oviferum,and possibly also with the Mexican specimen from Cordova in British Museum,described by Boulenger under the same name. I feel confident, however, thatnone of these belong to the true Gastrotheca ovifera of Weinland, from Venezuela.Two marsupial frogs were listed by Giinther as having been collectedin Mexico. The one from the Sall^ collection is a half-grown malecollected at Cordoba in Vera Cruz. The adult male labeled as comingfrom Mexico was purchased from fimile Parzudaki, a Paris dealer innatural-history specimens. According to Gerrit S. Miller, jr., thelocahties given to specimens purchased from Parzudaki have thereputation in the British Museum of being quite unreliable.The following notes were made on the specimen purchased fromSaU^:B.M. No. 1930. 4. 10. 1; Cordoba, Vera Cruz, Mexico, Salle'scollection: Head-and-body length, 64.6 mm.; transverse diameter oftympanum, 5.1 mm.; transverse diameter of eye, 8.2 mm.; anterioredge of eye to nostril, 5.8 mm.; posterior margin of bony casque totip of snout, 20.6 mm.; width of head at level of posterior angles ofjaws, 21.3 mm.; the hind Umb being carried forward along the body,the tibio-tarsal joint reaches sUghtly beyond anterior margin of orbit;a thin tarsal fold reaches more than halfway from base of first toe totibio-tarsal joint; toes about one-third webbed, the membrane reachingto base of last phalanx on first, second, third, and fifth toes, and leavingfree the last two phalanges on fourth toe ; a prominent inner metatarsaltubercle, but no outer one; subarticular and supernumerary tubercleson plantar surface of foot; first finger barely reaches apical disk ofsecond; apical disks of fingers large; fingers not webbed at base;vomerine teeth in essentially transverse rows, between and slightlybehind level of choanae; tongue subcircular, scarcely free behind;canthus rostralis raised and surmounted by a ridge of minute tubercles;interorbital width 7.9 mm., considerably greater than width of uppereyehd; posterior edge of bony casque with high, conical, erect bonyspines; a similar series above tympanum, and a smaller series of spinestakes the place of the combined supraciliary and postorbital curvedcranial crest; upperparts apparently without darker color markings;sides with four light-edged dark pear or gourd shaped spots, obliquein position, and with the constricted end dorsal; the anterior spot isnearly longitudinal and extends forward over scapula; upper surfaceof thigh with three light-edged dark cross bands; under or concealedsurface of tibia with three pairs of large light-edged dark spots in
MEXICAN TAILLESS AMPHIBIANS 135place of transverse bars; four light-edged transverse dark bands onupper surfaces of tarsus and metatarsus.Specimens examined.?Two, as follows:Gastroiheca coronata
Museum
136 BULLETIN 160, UNITED STATES NATIONAL MUSEUMAfter I read Mocqiiard's description of Hyla rudis, it becameapparent that it was certainly not a Hyla and furthermore that itseemed to agree rather closely with specimens of this species. Areexamination of the type revealed that it was unquestionably ayoung individual of Pternohyla fodiens. The type of H. rudis(M.H.N.P. No. 373a, parchment label No. 97-217) was collected atGuadalajara in Jalisco by Leon Diguet. The following notes weremade on this specimen : Head-and-body length, 34.5 mm. ; tympanumone-half diameter of eye ; top of head flat and rugose ; the hind limbbeing carried forward along the body, the tibio-tarsal joint does notreach to edge of minute tym.panum; a very faint indication of a tarsalfold; toes scarcely webbed; fingers free; very small adhesive disks;large inner metatarsal tubercle; abdomen and under surface of thighscoarsely areolate or granular. The color pattern described by Moc-quard, the presence of a large internal metatarsal tubercle, as well asother peculiarities noted above are all characters that distinguishP. fodiens from other Mexican hylids.A series of 67 of these burrowing tree frogs (A.M.N.H. Nos.12415-12458 ; 12460-12482), all collected in Mexico by Paul D. K. Ruth-ling, are especially interesting, for they range in size from youngwith a body length of 17.5 mm. (No. 12482) to adults with ahead-and-body length of 45 mm. (No. 12457). The young generallyhave a single pair of black spots on the posterior sacral region in frontof the vent, and all except the very youngest have the characteristicsplotched upperparts. The shape of the head is about the same inyoung and in adults, but in the former the ornamentation is lessobvious because of the less advanced stage of dermal ossification, andin quite young indi\'iduals the loreal region is oblique and not shelving.Nelson and Goldman removed one of these frogs from the stomachof a snake {Leptophis diplotropis) killed at Acaponeta in Nayarit.No field notes on the habits of this frog are available, but Boulenger{op. cit.) has published some observations on a living specimenobtained at Presidio by A. Forrer. He says:It is a timid creature, getting very frightened when handled, whilst all otherHylidae 1 have seen alive are very indifferent under similar circumstances. It isslow in its movements, and not a good climber. Its habits are more burrowingthan arboreal. It is not able to climb up a glass, but burrows itself deeply in themoss by means of its metatarsal shovels, the movements executed in this pro-ceeding being exactly those of Pelobates. I believe this to be the first instance ofan adaptation to both burrowing and arboreal life.Specimens examined.?Seventy-eight, as follows
:
MEXICAN TAILLESS AMPHIBIANSPternohyla fodiens 137
Museum
B.MU.S.N.M....DoU.S.N.M...-A.M.N.H...DoM.n.N.P..A.M.N.H..Do..M.C.Z_.A.M.N.H....Do
CatalogueNo.
138 BULLETIN 160, UNITED STATES NATIONAL MUSEUMB.M. Nos. 82. 12. 5. 11-12, two cotypes: (a), Head-and-bodylength, 84 mm.; transverse diameter of tympanum, 4.4 mm.; trans-verse diameter of eye, 7.5 mm.; posterior margin of bony casque to tipof snout, 30.2 mm.; width of head at level of posterior angles of jaws,25.2 mm.; tip of snout spinose; skin on abdomen fairly smooth; bodysoft; (b), head-and-body length, 87.9 mm.; transverse diameter oftympanum, 4.2 mm.; transverse diameter of eye, 7.2 mm.; posteriormargin of bony casque to tip of snout, 30.9 mm.; width of head atlevel of posterior angles of jaws, 23.6 mm.; tip of snout fairly smooth;skin on abdomen strongly areolate; body somewhat dried.An entomological collector, J. Aug. Kusche, presented an adultspecimen of this species to the National Museum and noted on theaccompanying label that it was found in a white-ant nest in a tree atVenodio. In 1882, Doctor Forrer returned to London with threeliving specimens, which were described by Giinther as follows
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The coloration is a uniform light olive, without any spots, changing in intensityof shade only; the upper side of the head is sometimes of a yellowish bronze color* * * the pupil is transversely oval when expanded, and subquadrangularwhen more contracted, but never vertical. It can be shut entirely.Specimens examined.?Four, as follows
:
Diaglena spatulata
Museum
MEXICAN TAILLESS AMPHIBIANS 139Type locality.?Vicinity of Merida, State of Yucatan, Mexico.Range.?Not known to occur elsewhere than on the peninsula ofYucatan.Remarks.?The type of this species (U.S.N.M. No. 12287) wastaken from a hole in the rocky wall of the Cenote Pamanche nearMerida on the road to Progressa. It is fairly well preserved not-withstanding numerous dissections made by Cope. The sternum hasbeen dissected, the viscera are exposed by a longitudinal slit in theabdominal wall, and the shoulder girdle, sacrum, and sacral diapo-physes have been freed from their muscle attachments through alongitudinal incision in the skin on the back. The measurements ofthis specimen are as follows: Head-and-body length, 72.5 mm.;transverse diameter of tympanum, 4 mm.; transverse diameter of eye,6.8 mm.; anterior edge of eye to nostril, 10.3 mm; width of head atlevel of posterior angles of jaws, 22.7 mm.; posterior margin of bonycasque to tip of snout (extremity missing), 25 mm.The following field notes of Leon J. Cole ^^ made at Chichen Itzain Yucatan are of special interest
:
Note an unmusical, rather drawn-out quarr-quarr-quarr . Not guttural, butwith a rasping quality. Life colors as follows: Top of head fuscous, with silverygreenish gray dots; back silvery gray, with dark fuscous blotches and smallerspots; sides with yellowish green suffusion; arms and legs brown, with yellowishblotches on upper arms and legs; silvery gray on lower arms and legs. Undersides whitish. The gray has a decided greenish tinge, which became more markedin a short time while the creature was held in the hand. This frog was not heardduring the drier part of the season (February and most of March), but was heardquite frequently during the last part of March, when there was more rain.Specimens examined.?Four, as follows:Triprion pelasatus
Museum
140 BULLETIN 160, UNITED STATES NATIONAL MUSEUM1864. Agalychnis Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 16, p. 181, Aug.[Genotype, Hyla callidryas Cope, 1862, Proc. Acad. Nat. Sci. Philadelphia,vol. 14, p. 359, footnote, Sept. (Panama).]
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Cope, 1865, Proc. Acad. Nat.Sci. Philadelphia, vol. 17, p. 194, Oct.There are three species of Phyllomedusa in Mexico, and the rangesof two of these extend southward into Central America. Relatedspecies occur in Central and South America. Some of the specieshave rather bizarre and striking color schemes. The largest one,P. bicolor, which is found in Guiana and BrazU, has a body length of110 mm., and the smallest, P. perlata, a native of northeastern Peru,measures onlj'^ 23 mm. Large individuals of a Mexican species, P. dacni-color, may attain a body length of 90 mm. These frogs are supposedlynocturnal in habit, for all of them have eyes with vertical pupils, andpresumably spend most of their Uves in arboreal surroundings. Mostof them have an eye with a red iris, and the body color is some shadeof green. In one of the Mexican species, P. dacnicolor, the iris isblack, vermiculated with gold. The viscous under surfaces of thelarge adhesive disks enable these frogs to attach themselves to thesurfaces of leaves, limbs, and trunks of trees and shrubs. It is gen-erally thought that those that have the hind toes united by a web ormembrane are best fitted for swimming in water, and if this be trueP. moreletiiwith. its extensively webbed fingers and toes is exceptionallywell adapted for this purpose. One of the Mexican species, P. dacni-color, seems to be as much at home on the ground in the grass as it iswhen hidden in the foliage of trees and shrubs. Some species ofPhyllomedusa merely deposit their eggs on the upper surface of leavesoverhanging the water, while others seemingly succeed in curling leavesaround the egg mass. After several days in the egg mass the tadpoleslose their external gills and drop into the water, where the metamor-phosis is completed.The supposedly least specialized species of these neotropical treefrogs were allocated to the genus Agalychnis by Cope. In character-izing the genus Agalychnis he cited the elongation of the inner toes,which are more or less opposable, the deeply emarginate xiphisternum,and the fact that the inferior palpebra is latticed or reticulated withstrong white veins. In formulating the diagnoses for the generaPhyllomedusa and Agalychnis, Boulenger in 1882 made no referenceto some of the characters cited by Cope, and introduced other struc-tural characters overlooked by him. Although a number of structuralfeatures are hsted in each, the diagnosis of Agalychnis differs from thatof Phyllomedusa solely in having the " first [fingers and toes] opposite tothe others." It has been confirmed by direct observation of hvingPhyllomedusa that the first finger and the first toe usually oppose theother digits when some small branch is grasped. This faculty ofopposing the inner finger and the inner toe is by no means confinedto the limited group formerly segregated under the name of Agalych-
MEXICAN TAILLESS AMPHIBIANS 141
FlOUEE 19.?Fore feet of Central American hylids, illustrating the extent of the web between thefingers: a, Phyllomedusa morelelii (U.vS.N.M. No. 25370), two-thirds webbed; 6, Hyla gabbi(U.S.N.M. No. 30658), two-thirds webbed; c, H. venulosa (U.S.N.M. No. 13976), one-halfwebbed; d, H. mioiympanum (U.S.N.M. No. 71092), one-half webbed; e, H. baudinii (U.S.N.M.No. 25369), one-third webbed.
Figure 20.?Hind feet of Mexican hylids,! illustrating the extent of the web between the toes: a,Phyllomedusa moreletii (U.S.N.M. No. 46916), fully webbed; 6, P. moreletii (U.S.N.M. 25370),three-fourths webbed; c, Hyla baudinii (U.S.N.M. No. 25369), three-fourths webbed; d, H.venulosa (U.S.N.M. No. 13975), two-thirdsj webbed; e, H. arenicolor (U.S.N.M. No. 71183),one-half webbed.66785?32- -10
142 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
nis, but exists to a varying degree in most of the species of Phyllo-medusa. In P. dacnicolor, a species allocated to Agalychnis by Copein 1866, the ability to oppose the inner digits of the fore and hindfeet is less marked than in some of the typical species of Phyllomedusa.KEY TO MEXICAN SPECIES OF PHYLLOMEDUSA
1. A single linear series of large quadrangular black spots on sides,which are set off from upperparts by a continuous or inter-rupted yellow border, and separated from one another by nar-row oblique yellow stripes, but not extended ventrally uponthe belly; upperparts, upper surfaces of forearm, tibia, tarsus,fifth finger, fifth toe, and narrow band on upper surface offemur greenish (green or sky blue in life); humerus togetherwith concealed surfaces of limbs dark purplish; forearm, hand,tarsus, and foot margined with white; toes and under surfacesyellow, a deeper hue on belly; the hind limb being carriedforward along the body, the tibio-tarsal joint reaches beyondthe eyes and usually to the tip of the snout; tympanum indis-tinct, subvertical in direction, ovoidal, and two-thirds diameterof eye; tongue pyriform, and openly emarginate behind;vomerine teeth in short series, directed inward and backwardfrom antero-internal angles of large choanae; inferior palpebrawith oblique subparallel white veins; fingers one-third webbed;toes two-thirds webbed; digits of hands and feet furnishedwith large adhesive disks; skin of upperparts smooth; bellyand underside of thighs strongly granulated; dermal folds onouter margin of underside of forearm, and on inner and outermargins of upperside of tibio-fibular region; head-and-bodylength, 30 to 68 mm helenae (p. 145)No single linear row of large quadrangular black spots; sideseither unicolored or marbled with yellow and reddish brown 22. Fingers slightly webbed at base; toes one-third webbed; adhesivedisks of medium size; the hind limb being carried forwardalong the body, the tibio-tarsal joint does not reach beyondthe eye; tympanum distinct, two-thirds the diameter of theeye; tongue large, pyriform, openly emarginate behind; vo-merine teeth in two short or long, transverse or oblique rowsbetween anterior margins of choanae; inferior palpebra reticu-lated with white veins; skin of upperparts smooth; belly andunderside of thighs strongly granulated; superior surfacesgrass green or violet-blue (narrowly on femur), occasionallywith a few scattered yellow spots; gular region and posteriorfaces of femora immaculate; forearm, hand, and foot marginedwith white; sides marbled with yellow and reddish brown; afew small pustules on anterior part of sides, which are yellow,like the inferior surfaces; head-and-body length, 60 to 83 mm.dacnicolor (p. 1 48)Fingers united for one-half or two-thirds of their length by web(fig. 19, a) , the membrane often extending to disks of the secondand fourth digits; and last phalanx of third is bordered bydermal fringe; toes two-thirds to fully webbed (fig. 20, a and b),the membrane often reaching to disks of all digits; tarsal fold
MEXICAN TAILLESS AMPHIBIANS 143faint; adhesive disks of fingers slightly larger than those oftoes and often larger than tympanum; the hind limb beingcarried forward along the body, the tibio-tarsal joint reachesbetween eye and tip of the snout; vomerine teeth in twooblique series between anterior margins of choanae, almost incontact medially; fronto-parietal bones not in contact anteri-orly, leaving a small fontanelle; tympanum more than one-halfand often three-fourths diameter of eye; tongue broad, notchedbehind; head broad; canthus rostralis obtuse; loreal regionconcave, oblique; skin of back smooth, with a few scatteredsmall warts; top of head shagreened between tympani; under-parts, with exception of gular region, coarsely granulated; upper-parts (of specimens in alcohol or formalin) green, blue-green, orviolet, rarely with round yellow spots; sides often with lateralyellow bands; concealed surfaces of limbs, hands, and feetyellowish, with exception of outer finger and toe, which aresame color as back; a white line along outer edge of forearm,lower hind limb, and tarsus; under surfaces white; male withsingle subgular sac; head-and-body length, 58 to 77 mm.raoreletii (p. 146)PHYLLOMEDUSA DACNICOLOR Cope1864. Phyllomedusa dacnicolor Cope, Proc. Acad. Nat. Sci. Philadelphia, vol.16, p. 181, Aug.Type locality.?Near Colima, State of Colima, Mexico.Range.?West of the Mexican Plateau, from southern Sinaloa tothe southern boundary of Guerrero.Remarks.?The type of this species, which can not now be located,was evidently collected by John Xantus de Vesey, for Cope statesthat it was removed "from the large Xantusian Coll." Several livingindividuals of this species collected near the Presidio in Sinaloa werebrought to London by Alphonso Forrer. In describing these frogsBoulenger ^^ remarks that large individuals may have a head-and-body length of 83 mm. and in addition makes the following comments:The faculty of opposing the inner finger and toe is conspicuous, though less sothan in the typical species of Phyllomedusa. The colour of the upper parts isnormally bright green, but rapidly changes to olive or brown; white dots arescattered on the flanks; the lower parts are pure white; the two inner fingers, thethree inner toes, the lower surface of the hands and feet, and the sides of the limbsare yellowish pink. The iris is black, vermiculated with gold; a golden line bordersthe vertical pupil. The nictitating lid is veined with gold, and the lower eyelidcompletely opaque, green.Entries made in the field catalogues of Nelson and Goldman showthat two of these frogs (U.S.N.M. Nos. 46954-55), taken on June 30,1897, in green grass on a plain near Acaponeta in Nayarit, were brightgreen on the back, and another individual (No. 46950) caught in apublic garden in the middle of the city of Rosario, Sinaloa, on July 6,
?? Boulenger, Q. A., Description of a new genus and species of frogs of tlie famOy Hylidae. Ann. and Mag,Nat. Hist., ser. 5, vol. 10, no. 68, p. 328, October, 1882.
144 BtJLLETIN 160, UNITED STATES NATIONAL MUSEUM1897, was bright grass green on the back and yellowish white below.While collecting on February 13, 1895, in Guerrero, thej found one(No. 47904) in dense green grass on the river bottom near Ometepec.These observations would seem to indicate that this species spendsconsiderable time on the ground at periods not necessarily coincidentwith the breeding season.Very little is kno^vn in regard to the breeding habits of P. dacnicolor.The following observations on Phyllomedusas at Balsas in Guerreromade by Dr. Hans Gadow ^^ summarize what little is actually known
:
Lovely big tree-frogs, of a saturated green, dwelt during the heat of the day inthe thick thatch of the huts, whilst at night especially during the rain, they hoppedabout on the ground, to pair. As a rule these Phyllomedusas do not deposit theirspawn directly in the water; they wrap their eggs in a foamy lather, and suspendthe whole mass between leaves or grass, over water, in such a position that thenext heavy rain washes the rapidlj' developing eggs or tadpoles into it. Thosewhich we could observe behaved in a rather puzzling manner. The pairs sat ina tangle of herbs at the edge of a little ditch in the middle of the village, whence therain could wash the eggs only into a filthy lagoon some fifty yards off. In themorning the ditch was alwaj^s quite dry, and there were no traces of frogs oreggs left. This may have been an error of judgment of the tree-frogs. Themajority were barking, snarling, and entreating each other's attention on moresuitable ground, but we failed ignominiously in our observations during the darkof night, and in the tropical downpour. The most remarkable feature, so farapparently unique, is the colour of the numerous and very small eggs, these allbeing of a light green!Specimens examined.?Twenty-four, as follows
:
Phyllomedusa dacnicolor
Museum CatalogueNo. Num-ber ofspeci-mens Locality collected Date collected By whom collected orfrom whom received
U.S.N.M.M.C.Z...A.M.N.HU.S.N.M.Do...A.M.N.HU.S.N.M.Do...A.M.N.HDo...M.C.Z...A.M.N.HDo...M.C.Z...U.S.N.M.
68378631-321312614081469501256546954-55469531256312565-66242013266-6713265106047904
Sinaloa:Mazatlan.
.do.North of MazatlanPresidio, 50 miles from Ma-zatlan.Rosario.San FrancisquitoNatarit:AcaponetaSantiagoRosamocadadoColima:Colima2 miles from ColimaEast of Lo de VillaGuerrero:Acapulco -.Ometepec, 200 ft. altitude.
18681921Feb. 2, 1920Apr. 3, 1885July 6, 1895fDec. 2, 1919[Dec. 4, 1919June 30,1897June 21,1897Nov. 20, 1919Nov. 19, 1919July 14,1902Apr. 17,1919Apr. 19,1919
Feb. 13,1895
F. Bischofl.C. C. Torron.Paul D. R. Ruthling.Alphonso Forrer.Nelson and Goldman.}Paul D. R. Ruthling.Nelson and Goldman.Do.Paul D. R. Ruthling.Do.Thomas Barbour.Paul D. R. Ruthling.Do.Hassler expedition.Nelson and Goldman.M Gadow, H.329, 330, 1908. Through southern Mexico: Being an account of the travels of a naturalist, London, pp.
MEXICAN TAILLESS AMPHIBIANSPHYLLOMEDUSA HELENAE (Cope)Banana Frog 1451884. Agalychnis helenae Cope, Proc. Araer. Philos. Soc, vol. 22, pt. 2, no. 118,p. 182, Dec. 19.Type locality.?Nicaragua.Range.?From Costa Rica and Nicaragua northward throughTabasco to Piiebla.Remarks.?The type of this species (U.S.N.M. No. 13737) wascollected on December 4, 1883, by Lieut. J. F. Moser. Althoughthis frog is widely known in Central America as the banana frog,there are very few published observations on its habits. Clarence R.Halter kept a record in liis field notebook of the color changes observedon a captive frog taken from the upper side of a leaf on a banana treestanding near the edge of the jungle at Colorado Bar, northeasternCosta Rica. The following notes published by Noble ^^ were extractedfrom this notebook:During captivity, the dorsal coloration was subject to considerable variation.Generally the ground tone was a dark slaty green or a bright leaf-green. Anumber of pale green spots would often appear. These did not seem to bedependent upon the intensity of the light. They appeared irregularly scatteredover the back. Sometimes they would form an H-shaped mark just anterior tothe pectoral girdle. The changes in the color of the ground tone did not seem tobe correlated with changes in the lighting. The concealed portions of the legs,which appear purplish in alcohol, were a deep blue in life. The feet and handswere brilliant orange. The edge of the upper ej^elid was a deep yellow, in strikingcontrast to the orange-red iris. The ventral surface of the throat and bod}' wasyellowish white, of the anterior appendages a slaty color, of the posterior append-ages a yellowish white washed entirely, except for a narrow strip, with the deepblue of the concealed portions.The specimen was rather awkward in its movements. Before jumping, it wouldcrouch on all fours with appendages at right angles to the axis of the body. Itwould then laboriously raise the body about an inch from the ground. It wouldremain in that position for a few seconds, studying the nearby objects, and thenwould suddenly leap toward one.Halter suggests that the breeding season in eastern Nicaragua mayoccur about the end of August, inasmuch as a female with ovariesdistended \\dth eggs was taken during that month at Culvra.Specimens examined.?Two, as follows:
146 BULLETIN 160, UNITED STATES NATIONAL MUSEUMPHYLLOMEDUSA MORELETH (Diun6ril)Morelet's FrogFigures 19, a; 20, a, b1853. Hyla moreletii Dumeril, Ann. Sci. Nat., Paris, ser. 3, Zoology, vol. 19,p. 169.1861. Hyla holochlora Salvin, Proc. Zool. Soc. London, 1860, pt. 3, p. 460, pi. 32^fig. 2. Mar. (Coban, Guatemala.)1882. Hyla moreleti Brocchi, Mission scientifique au Mexique et dans I'Am^riqueCentrale, recherches zoologiques, pt. 3, sect. 2, p. 31, pl. 1, fig. 1; pl. 13, fig. 1.1901. Agalychnis moreletii Gunther, Biologia Centrali-Americana, Reptilia andBatrachia, p. 289, Sept.Type locality.?Coban, Vera Paz, Guatemala.Range.?From Volcan de Tuxtla in southern Vera Cruz and SanMiguel in northern Oaxaca southward through Central America toPanama.Remarks.?The two cotypes of Hyla moreletii (M.H.N.P. No. 428,parchment label No. 767) were collected at Coban in Vera Paz, Guate-mala, by Arthur Morelet. The bodies of these cotypes are quite soft,and the coloration has faded to a creamy white. The head-and-bodylengths are, respectively, 68.5 and 59 mm. It was further noted thatthe hind limb being carried forward along the body, the tibio-tarsaljoint reaches to between eye and tip of snout; toes fully webbed withexception of last phalanx of fourth toe, which has a dermal fringe;tarsal fold indistinct; fingers two-thirds webbed, the last phalanx ofthird finger bordered by a dermal fringe; an external callosity at baseof first finger; sldn on abdomen and under surface of tliigh granular;vomerine teeth in transverse rows between and at level of anteriormargins of choanao.Osbert Salvin states that he obtained three examples of his Hylaholochlora at Coban in Vera Paz, Guatemala, and that all of themwere caught by the Indians. One of these (B.M. No. 1864. 1. 26.142) is Hsted in the catalogue as the type of this species. The follow-ing notes were made on this specimen: Head-and-body length, 76.5mm.; transverse diameter of tympanimi, 4.6 mm.; transverse diameterof eye, 6.6 mm.; anterior edge of eye to nostril, 7.2 mm.; width ofhead at level of posterior angles of jaws, 25.5 mm.; the hind limb beingcarried forward along the body, the tibio-tarsal joint reaches to abouthalfway between eye and tip of snout; long tarsal ridge from base offirst toe to tibio-tarsal joint; toes fully webbed with exception of lastphalanx of fourth toe, which has a dermal fringe; inner metatarsaltubercle large and flat, no outer one; subarticular and supernumerarytubercles on plantar surface of foot; tip of first finger reaches to apicaldisk of second; fingers two-thirds webbed, the last phalanx of thirdfinger bordered by dermal fringe; vomerine teeth between and atlevel of anterior margins of choanae; abdomen and under surface of
MEXICAN TAILLESS AMPHIBIANS 147thighs coarsely granular; coloration of upperparts (in alcohol) blue;underparts yellowish brown.From Bocourt's field notes *^ on a living frog and Salvin's descrip-tion of Hyla holochlora and from the observed pigmentation of theskin on specimens preserved in alcohol, it would appear that theupperparts as well as the outer or upper surfaces of the forearm,fourth finger, tibia, tarsus, and fifth toe are grass green. The remain-der of the fore and hind limbs, thighs, fore feet, and hind feet areorange-yellow. The sides are likewise orange-yellow, and the throat,chest, and belly are suffused with a paler tint of the same color.This species can be recognized at once by the extensive webbing ofthe fore and hind feet. The terminal phalanges of the fingers andtoes are clawUke, very sharp, and recurved. One individual in aseries from Atitlan lacked vomerine teeth, according to Brocchi.Specimens examined.?Sixteen, as follows:Phyllomedusa moreletii
Museum CatalogueNo.
148 BULLETIN 160, UNITED STATES NATIONAL MUSEUMdes crapauds, p. 26, pi. 14 ('' Caroline et en Virginie"); fixed by Stejneger,1907, U. S. Nat. Mus. Bull. 58, p. 76.]1830. Auletris Wagler, Natiirliches System der Amphibien, p. 201. [Genotype,[Rana] boans Linnaeus, 1758, Systema naturae, ed. 10, vol. 1, p. 213(America,) = Hyla boans Daudin, 1803, Histoire naturelle des rainettes, desgrenouilles et des crapauds, p. 21, pi. 11 (Surinam); fixed by Stejneger,1907, U. S. Nat. Mus. Bull. 58, p. 76.]1830. Hyas Wagler, Natiirliches System der Amphibien, p. 201. [Genotype,[Rana] arborea Linnaeus, 1758, Systema naturae, ed. 10, vol. 1, p. 213(Europe); preoccupied by Hyas Leach, 1815, Crustacea.]1830. Hypsiboas Wagler, Natiirliches System der Amphibien, p. 200. [Geno-type, Hyla palmata Daudin, 1803, Histoire naturelle des rainettes, desgrenouilles et des crapauds, p. 26, pi. 14 (" Caroline et en Virginie").]1830. Scinax Wagler, Natiirliches System der Amphibien, p. 201. [Genotype,H[yla] aurata Wied, 1825, Beitrage zur Naturgeschichte von Brasilien, p. 531(Bahia, Brazil); fixed by Stejneger, 1907, U. S. Nat. Mus. Bull. 58, p. 76.]1830. Dendrohyas Wagler, Natiirliches System der Amphibien, p. 342. [Substi-tute name for Hyas, preoccupied, ^rfe Stejneger, 1907, U. S. Nat. Mus. Bull.58, p. 76.]1838. Lophopus Tschudi, M^m. Soc. Sci. Nat. de NeuchStel, vol. 2, pp. 32, 73.[Genotype, Lophopus marmoratus Tschudi, idem, pp. 32, 73= Bufa marmor-atus Laurenti, 1768, Synopsin reptilium, p. 29 (Surinam); preoccupied byLophopus DuMERiL, 1837, Polyzoan.]1838. Sphaenorhynchus Tschudi, M6m. Soc. Sci. Nat. de Neuchatel, vol. 2, pp.28, 71. [Genotype, Sphaenorhynchus lacteus Tschudi, idem, pp. 28, 71 = Hylalactea Daudin, 1803, Histoire naturelle des rainettes, des grenouilles et descrapauds, p. 21, pi. 10, fig. 2 (America).]1838. Trachycephalus Tschudi, Mem. Soc. Sci. Nat. de NeuchStel, vol. 2, pp.33, 74. [Genotype, Trachycephalus nigromaculatus Tschudi, idem,, pp. 33,74 (America meridionalis).]1843. Acrodytes Fitzinger, Systema reptilium, fasc. 1, p. 30. [Genotype, Hylavenulosa Daudin? /2ana venulosa Laurenti, 1768, Synopsin reptilium,p. 31 ("Indiis").]1843. Dendropsophus Fitzinger, Systema reptilium, fasc. 1, p. 31 [Genotype,Hyla frontalis Daudin, 1803, Histoire naturelle des rainettes, des grenouilleset des crapauds, p. 17, pi. 7, figs. 1, 2 (Surinam).]1843. Dryomelictes Fitzinger, Systema reptilium, fasc. 1, p. 31. [Genotype, Hylalactea Daudin, 1803, Histoire naturelle des rainettes, des grenouilles et descrapauds, p. 21, pi. 10, fig. 2 (America).]1843. Dryophytes Fitzinger, Systema reptilium, fasc. 1, p. 31. [Genotype, Hylaversicolor LeConte, 1825, Ann. Lye. Nat. Hist. New York, vol. 1, pt. 2, p.281 ("Northern States," U. S. A.).]1843. Hypsipsophus Fitzinger, Systema reptilium, fasc. 1, p. 30. [Genotype,Hyla xerophyllum= Hyla xerophylla Dumeril and Bibron, 1841, Erp(5tologieg6n6rale, vol. 8, p. 549 (Cayenne).]1843. Lobipes Fitzinger, Systema reptilium, fasc. 1, p. 30. [Genotype, Hylapalmata Daudin, 1803, Histoire naturelle des rainettes, des grenouilles etdes crapauds, p. 26, pi. 14 (' Caroline et en Virginie') ; preoccupied by Lobipes,Cuvier, 1817, Aves.]1843. Osteopilus Fitzinger, Systema reptilium, fasc. 1, p. 30. [Genotype,Trachycephalus marmoratus Bibron, 1842, in Ramon de la Sagra, Histoirephysique, politique et naturelle de I'lle de Cuba, Atlas, pi. 29 (Cuba); neeHyla marmorata (Laurenti), 17&S=^ Hyla septentrionalis Boulenger, 1882,Catalogue of the Batrachia Salientia s. Ecaudata in the collection of theBritish Museum, 2d edit., p. 368 (Cuba).]
MEXICAN TAILLESS AMPHIBIANS 1491843. Phrynohyas Fitzinger, Systema reptilium, fasc. 1, p. 30. [Genotype,Hyla zonula Spix, 1824, Animalia nova, sive species novae tesludinum etranarum quas in itinere per Brasiliam, etc., p. 41, pi. 12, fig. 1 (Brazil).]1843. PhylloUus Fitzinger, Systema reptilium, p. 30. [Genotype, Hyla albo-marginata Spix, 1824, Animalia nova, sive species novae testudiniim etranarum quas in itinere per Brasiliam, etc., p. 33, pi. 8, fig. 1 (Brazil); pre-occupied by Phyllobius, Schonherr, 1824, Coleoptera.]1856. Centrotelma Burmeister, Erlauterungen zur Fauna Brasiliens, enthaltendAbbildungen und ausftihrliche Beschreibungen neuer oder ungeniigendbekannter Thier-Arten, p. 97. [Genotype, Hyla (Centrotelma) infulata=Hyla infulata Wied, 1825, Beitrage zur Naturgeschichte von Brasiliens, vol.1, p. 533 (Brazil).]1856. Hylomedusa Burmeister, Erlauterungen zur Fauna Brasiliens, enthaltendAbbildungen und ausftihrliche Beschreibungen neuer oder ungeniigend be-kannter Thier-Arten, p. 102. [Genotype, Htjla (Hylomedusa) crepitansBurmeister, idem, p. 103? H[yla] crepitans Wied, .1824, Abbildungenzur Naturgeschichte Brasiliens, pi. (47), fig. 1 (Tamburil im Sertong vonBahia, Brazil).]1862. Osteocephalus Steindachner, Archiv. Zool. Anat. e la Fisiol. Genova, vol.2, fasc. 1, p. 77. [Genotype, Osteocephalus taurinus Steindachner, idem,p. 77, pi. 6, figs. 1-3 (Barra do Rio Negro in Brasilien) . Not of Fitzinger,1843, a nomen nudum.]1862. Scytopis Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 14, p. 354, Sept.[Genotype, Scytopis hebes Cope, idem, p. 354 (Paraguay).]1865. Smilisca Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 17, p. 194, Oct.[Genotype, Smilisca daulinia Cope, idem, footnote, p. Q5= Hyla baudiniiDum^ril and Bibron 1841, Erp^tologie g6n6rale, vol. 8, pp. 564, 565(Mexico).]1867. Cinclidium Cope, Journ. Acad. Nat. Sci. Philadelphia, n. s., vol. 6, pt. 2,p. 200, July. [Genotj'pe, Cinclidium granulatum Cope, idem, pp. 200, 202(Surinam) .]1870. Cincloscopus Cope, Proc. Amer. Philos. Soc, vol. 11, no. 84, p. 554, foot-note, Sept. 16. [Substitute name for Cinclidium Cope.]1870. Cophomantis Peters, Monatsber. k. preuss. Akad. Wiss. Berlin, p. 650,Aug. [Genotj'pe, C[oph omantis] punctillata Peters, idem, p. 651, pi. 2,fig. 4 (Sta.Catharina, Brazil).]1872. Hylomantis Peters, Monatsber. k. preuss. Akad. Wiss. Berlin, p. 772.[Genotype, Hylomantis aspera Peters, idern, p. 772, pi.-, fig. 2 (Bahia,Brazil).]1885. Epedaphus Cope, Proc. Amer. Philos. Soc, vol. 22, pt. 4, no. 120, p. 383,Apr. 17. [Genotype, Hyla gratiosa LeConte, 1856, Proc. Acad. Nat. Sci.Philadelphia, vol. 8, p. 146, pi. 4 (printed VI) Aug. ("lower country ofGeorgia").]1899. Hyliola Mocouard, Nouv. Archiv. Mus. Hist. Nat. Paris, ser. 4, vol. 1,p. 337. [Genotype, Hyliola regilla Mocquard= //j/Za regilla Baird andGiRARD, 1852, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, p. 174, Oct.("Sacramento River, in Oregon, and Puget Sound"); fixed by Stejneger,1907, U. S. Nat. Mus. Bull. 58, p. 76.]Eleven species of the genus Hyla are liere recognized as occurringwitliin the limits of continental Mexico. There is reason to believethat five of these
?
arenicolor, baudinii, eximia, miotympanum, andvenulosa?are fairly common tree frogs, for they are most frequentlyfound in collections made in IMexico. One, //. phlebodes, may yet be
150 BULLETIN 160, UNITED STATES NATIONAL MUSEUMfound to be fairly abundant in the lowland swamps of Yucatan andQiiintana Roo. H. gracilipes is a very close relative of H. eximia, butseems to be sufficiently distinct to be recognized as a full species,rather than as a variety of the latter. The habitat of the tree frogdescribed by Brocchi as H. plicata is unknown, and, although the posi-tion of the vomerine teeth is quite different from other Mexicanhylas, it is otherwise structurally quite similar to H. eximia. Noadditional specimens of H. bistinda have been reported since it wasfirst described by Cope. The type of the peculiar little H. staujferiremained unique for more than 60 years, though quite recently Dr.E. R. Dunn discovered a second individual in the British Museumthat was collected at Motzorongo in Vera Cruz, and there are threeadditional specimens in the United States National Museum fromPotrero, Vera Cruz. Some uncertainty still exists as to the validityof H. taeniopus, a species based on two immature individuals.The genus Hyla, in spite of its nearlj^ world-wide distribution andnumerous species, seems to have retained a fairly uniform type of lifehistory. There is no evidence that any species of the genus skips thetadpole stage, and with the exception of certain Jamaican species allof them evidently lay their eggs in the water, where the tadpolesfollow the usual aquatic development. The eggs are deposited inponds, in flooded marshes, in stagnant pools of rain-water in forests,along the edges of streams where there is little or no current, or inlittle basins or pools on the edges of mountain brooks. In Jamaica,Dunn found three species of Hyla that laid their eggs in water caughtbetween the leaves of bromeliads. Some of the large tree frogs ofSouth America build mud basins, in which the eggs are deposited andin which the tadpoles pass through the larval stage.According to Noble "
?
In Hyla, as in many large genera of Salientia, the tadpoles do not conform to asingle type. Those species which lay their eggs in ponds have either broad-finnedtadpoles of the "sunfish type" or bottom-wriggling larvae of the "polUwog type."The "sunfish type" has larger lungs, remains more frequently between surfaceand bottom and is usually a graceful swimmer. The "polliwog type" has a nar-row fin restricted to the tail or only to the tail and posterior part of the body.This type is prevailingly a bottom-wriggler but may dart very rapidly in theundulatory manner characteristic of our common Rana poUiwogs.KEY TO MEXICAN SPECIES OF HYLA1. Second to fourth fingers practically free, at most united at baseby a vestigial web or entirely free from such 2Second to fourth fingers united at base by distinct web, whichmay extend for one-fourth to two-thirds of their length 82. Vomerine teeth behind or at most at level of posterior edge ofchoanae 3Vomerine teeth between or near level of anterior edge of choanae 4
" Noble, Q. K., The value of life history data in the study of the evolution of the Amphibia. Ann,New York Acad. Sci., vol. 30, p. 95, Oct., 31, 1927.
MEXICAN TAILLESS AMPHIBIANS 151Vomerine teeth behind or barely at level of posterior edge ofchoanae, each group consisting of about eight teeth; body withlateral color band; fronto-parietal bones ossified and in con-tact anteriorly, separated posteriorly, leaving a fontanelle;tympanum about two-fifths diameter of eye; adhesive disksnearly as large as tympanum; the hind limb being carriedforward along the body, the tibio-tarsal joint reaches theposterior portion of the orbit; tongue wide, not or scarcelynotched behind; canthus rostralis indistinct; loreal regionconcave, obhque; skin of upperparts thickened, studded onback with large obtuse warts; upperparts Ught purplishbrown, separated from sides in some specimens by a broadblackish band, which extends from above tympanum to loin;pupil cruciform, interspaces golden, black spot on outermargin venulosa, young [= spilomma Cope] (p. 17b)Vomerine'teeth at level of posterior edge of choanae, each groupconsisting of four teeth; body without lateral longitudmalcolor band; tympanum one-third diameter of eye, overhungby thick dermal fold; frontal bones cartilaginous, thickenedor stiffened in front, possibly ossified; fingers united at baseby vestigial web; thumb of male with two corneous callosi-ties on inner surface during breeding period; toes webbedfor two-thirds of their length, the membrane being deeplyincised and continued beyond the bases of the penultimatephalanges of the third to fifth toes as a narrow fringe; footshorter than combined length of tibia and femur; tarsal foldindistinct; adhesive disks of approximately the same size astympanum: the hind limb being carried forward along thebody, the tibio-tarsal joint reaches the anterior margin of eye;tongue broad, oval, not notched behind; canthus rostralisrounded; skin of upperparts smooth; underparts granulated;a dermal fold across chest; coloration of upper surfaces,including femur and humerus, dark bluish plumbeous (inalcohol), without darker markings; sides with a network ofdark streaks on a yellow background; posterior face of thighbrown, with few yellow specks along superior border; inferiorsurfaces yellow; eye unspotted; head-and-body length, 43 5bistmcta (p. loo;
1. VomTr'ine teeth'situated in transverse rows near level of anteriormargins of choanae; tympanum distinct, more than one-halfdiameter of eye and overhung by a dermal fold; adhesivedisks small, less than one-half diameter of tympanum; fin-gers free from web at base; first finger shorter than second;toes webbed for one-half their length and membrane is con-tinned as a narrow fringe on distal phalanges; an ovoidalinner metatarsal tubercle and an indistinct outer one; nosupernumerary tubercles on plantar surface of foot; a tarsalfold- the hind limb being carried forward along the body, thetibio-tarsal joint reaches between eye and tip of snout;tongue ovoidal, notched behind, and free on its posteriorone-third; canthus rostraUs angular; skin of upperpartssmooth; a very distinct dermal fold across chest; throat,abdomen, and under surface of thighs granulated; colorationof upperparts apparently green with indistinct spots of adarker color; a narrow light-yellow stripe along canthus
152 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
rostralis from nostril to eye, and from posterior edge of uppereyelid backward above tj^mpanum, and then obliquely down-ward to middle of side; this stripe seems to be upper marginof broad dark band that extends obliquely downward behindtympanum; head-and-body length, 38.7 mm plicata (p. 173)Vomerine teeth situated between the choanae 55. A tarsal fold present in adults, indistinct in young or immatureindividuals 6No tarsal fold; second to fourth fingers free from web at base;adhesive disks nearly as large as tympanum; toes two-thirdswebbed, the membrane reaching more than halfway to diskon penultimate phalanx of third and fifth toes; the hind limbbeing carried forward along the body, the tibio-tarsal jointreaches to center or at most to anterior margin of eye; tym-panum less than or slightly more than one-half the diameterof the eye and overhung by an indistinct dermal fold; dis-tance from ej'e to nostril somewhat greater than transversediameter of eye; vomerine teeth in two elongate clusters,presenting an obtuse angle posteriorly, each group consistingof about four teeth, and situated between the choanae; tonguecircular, scarcely notched behind; canthus rostralis indis-tinct; loreal region fiat, oblique; snout prominent, rounded;skin of upperparts fairly smooth; sides rugulose; abdomenand under surface of thighs granulated; a dermal fold acrosschest; upperparts dark olive (in alcohol), with a short longi-tudinal or curved black streak over each scapula, and onefrom eye to eye; indistinct dark stripe along coccyx; a lateraldark streak from posterior edge of upper eyelid to about half-way length of body; sides minutely marbled with dorsal andventral tints; upper lip olive, sending a pale line to near axil-la; limbs maj^ or may not have dark crossbars; underpartsyellowish, grading from a deeper hue to brownish on the ex-tremities; male with single large vocal sac; head-and-bodylength, 25 to 30 mm staufferi (p. 173)6. Skin of upperparts rough, tubercular, or with small warts, thetubercles extending forward beyond the interorbital space;vomerine teeth in two clusters, each group consisting offive to six teeth, and between the choanae; fronto-parietalbones ossified and in contact anteriorly, leaving no fontanelle;tympanum distinct, generally one-half to three-fourths,rarely one-third, diameter of eye; adhesive disks of fingers alittle larger than those of toes and often nearly as large astympanum; the hind limb being carried forward along thebody, the tibio-tarsal joint usually reaches to anterior marginof eye, occasionally between eye and tip of snout, and rarelybeyond tip of snout; toes one-half webbed (fig. 20, e), withnarrow dermal fringe on terminal phalanges; a tarsal fold;tongue broad, notched behind; canthus rostralis indistinct;loreal region oblique; abdomen coarsely, throat generallyfinely granulated; a distinct dermal fold across chest; upper-parts sand color, light or dark gray, or olive, with grayish orblackish blotches; a curved baud, interrupted or continuous,between the eyes; a white spot generally present below eye;upper lip and upper surface of limbs crossbarred; pos-terior surface of thigh orange colored; throat brownish, with
MEXICAN TAILLESS AMPHIBIANS 153black specks; rest of underparts pale yellow or white; maleswith external gular sac; head-and-body length, 30 to 43 mm.
-- arenicolor (p. 156)Skin of upperparts smooth, rarely sprinkled with minutetubercles; side of head and fore part of body with a longi-tudinal dark-colored streak 7Dark markings on the greenish upperparts run more to longi-tudinal dark streaks than to distinct dorsal spots; hindersurface of thigh occasionally exhibits a distinct color patternconsisting of a darker area more or less marbled or spottedwith yellow, and often with light upper edge; coloration ofupperparts greenish or brownish (bluish in alcohol) , with fewor numerous blackish, and often light-edged, elongate spots;a black or brown band, narrow anteriorly and broad poste-riorly, with white dorsal edge, commences at nostril andextends backward below canthus rostralis to eye, and fromposterior edge of upper eyelid through tympanum and alongside to about level of middle of sacrum, where it is interruptedor disappears entirely; horn-colored margins of upper andlower lips merging posteriorly into broad band of same colorwhich terminates on fore limb; normal specimens have awhite or light-colored infraorbital stripe from end of snoutto axilla; a pair of elongate dark spots or streaks in sacralregion ; paired longitudinal dark streaks or sublinear arrange-ment of dark spots on back; dark spot on each upper eyelid;dark spots often present on each side of vent; upper surfacesof hind limbs with dark spots or complete crossbars; alight-edged dark streak along outer surface of tibia may ormay not be present; underparts and concealed surfaces oflimbs light colored; fingers practically free from web at base;toes one-half to two-thirds webbed, the membrane not reach-ing to adhesive disks of third and fifth; a tarsal fold; adhesivedisks generally smaller and rarely as large as the tympanum;the hind limb being carried forward along the body, thetibio-tarsal joint generally does not reach beyond anteriormargin of eye; vomerine teeth in two elongate clustersbetween the choanae, occasionally in two slightly arched oroblique rows, and each group consisting of three to seventeeth; tympanum usuallj'^ one-half transverse diameter of eye,overhung by a weak dermal fold; tongue pyriform or circular,notched behind; canthus rostralis fairly distinct; loreal regionconcave, oblique; gular region, abdomen, and under surfaceof thighs coarsely granulated; a dermal fold across chest;males with single external gular sac; head-and-body length,25 to 41 mm ezimia (p. 164)Dark markings on greenish upperparts tend to be restricted todistinct spots on hinder half of body; hinder surface of thighunmarbled and without color pattern, the dark flesh colgrshowing through a bluish suffusion; upperparts dark green,grass green, olive green, or brownish in life (dirty blue inalcohol) with paired oblong white-edged dark spots onhinder back; sides of snout green; canthal stripe blackish,edged above with white, bronze, or light brown; from eye totympanum and behind tympanum a dark lateral stripe ofregular or irregular width extends backward to about half-
154 BULLETIN 160, UNITED STATES NATIONAL MUSEUMway length of body where it either disappears or is brokenup on hinder half of fiank into a series of spots; upper lipwith dark gray stripe, edged above, or above and belowwith white; outer edge of tibia with lengthwise light-edgeddark streaks or with dark spots; cross bands on limbs indis-tinct, or incomplete (small spots) , or well developed; a darkspot on inner side of forearm; underparts whitish; lingersunited at base by dermal fringe or vestigial web; first fingershorter than second; adhesive disks of fingers somewhatlarger than those of toes and about one-half as large astympanum; toes two-thirds webbed; tarsal fold weakly orstrongly developed; inner metatarsal tubercle large; outerone small and conical, or flat, or absent; distinct subarticulartubercles; the hind limb being carried forward along thebody, the tibio-tarsal joint reaches slightly beyond or to tipof snout; vomerine teeth in two short, somewhat oblique ortransverse rows between the choanae; transverse diameter oftympanum generally slightly more than one-half diameterof eye, rarely less than one-half; a dermal fold extends aboveand downward behind tympanum to axilla; tongue large,broad, and feebly notched behind; canthus rostralis distinct;loreal region sloping more or less obliquely; throat, abdomen,and under surface of thighs granulated; a dermal fold acrosschest; head-and-body length, 30-50 mm gracilipes (p. 168)8. Vomerine teeth on a level with or slightly behind posterioredge of choanae ; tarsal fold absent or indistinct 9Vomerine teeth in elongate clusters between the choanae; a dis-tinct tarsal fold in adults, indistinct and possibly absent inyoung 109. ^Coloration of upperparts olive or brownish, generally with large,subsj^mmetrical, dark blotches, more or less confluent into apair of longitudinal bands, and sharply defined against alighter lateral band that extends from upper eyelid to loin;a large dark blotch extending from interorbital region back-ward upon shoulders generally present; a broad black bandextends obliquely downward behind tympanum to beyondlevel of fore limb; warts often black; limbs may or may notbe broadly crossbarred; underparts generally uniform whit-ish; transverse diameter of tympanum one-half (rarely one-third) diameter of eye, and overhung by a dermal fold;vomerine teeth on a level with hinder edge of choanae, form-ing either transverse or slightly arched series, or a very openchevron, the apex of which is directed posteriorly; fronto-parietal bones in contact medially, leaving no fontanelle;fingers one-fourth webbed (fig. 19, c) ; no projecting rudi-ment of poUex; adhesive disks of fingers slightly larger thanthose of toes, and equal in area to that of tympanum; thehind limb being carried forward along the body, the tibio-tarsal joint reaches to but not beyond eye; toes two-thirdswebbed (fig. 20, d), the membrane reaching to adhesive disksof third and fifth toes, but deeply incised, and leaving free thepenultimate phalanx of fourth; tarsal fold absent or indis-tinct; tongue broad, notched behind; canthus rostralisrounded; loreal region oblique; skin of upperparts and sidescovered with large round flat warts; brownish black minute
MEXICAN TAILLESS AMPHIBIANS 155pustules on exposed surfaces of fore and hind limbs; skin ofgular region, abdomen, and under surface of thighs coarselygranulated; male with two large external vocal vesicles behindangle of mouth; head-and-body length, 40 to 64 mm_..venulo8a (p. 176)Coloration of upperparts vinaceous-drab, with coarse networkof dark purplish reticulations; tibia with a number of crosslines of same color; lower surfaces uniform white; transversediameter of tympanum one-third the diameter of the eye,and overhung by a dermal fold: vomerine teeth in two smallclusters on a level with posterior edge of the choanae, eachgroup consisting of about four teeth, and the interval betweenthe clusters about equivalent to their diameter; fingers one-fourth webbed; no projecting rudiment of pollex; adhesivedisks as large as tympanum; the hind limb being carriedforward along the body, the tibio-tarsal joint reaches tobetween eye and tip of snout; toes three-fourths webbed, themembrane reaching to the adhesive disks of the third andfifth toes, leaving free the penultimate phalanx of the fourth;tarsal fold absent or indistinct; tongue nearly circular, scarcelynotched behind; canthus rostralis rounded; loreal regionoblique, concave; skin of upperparts finely shagreened, withscattered indistinct warts; gular region smooth; rest ofunderparts coarsely granulated; a dermal fold across chest;head-and-body length, 28.5 mm phlebodes (p. 172)10. Tympanum not more than one-half diameter of eye 11Tympanum two-thirds to three-fourths diameter of eye, over-hung by dermal fold; fronto-parietal bone often not in con-tact medially, leaving large fontanelle; fingers from one-fourthto one-third webbed (fig. 19, e) ; adhesive disks of fingers notnoticeably larger than those of the toes, but smaller than tym-panum; the hind limb being carried forward along the body,the tibio-tarsal joint reaches to the eye, or between the eye andtip of the snout; toes three-fourths webbed (fig. 20, c), themembrane reaching the disks of the third and fifth, leavingfree the last phalanx of the fourth; a distinct tarsal fold;tongue circular, usually notched behind; canthus rostralisangular; loreal region concave; skin of upperparts smooth orfinely shagreened; underparts, including gular region andunderside of thighs, granulated; upperparts sometimesuniform green, olive, or grayish, but generally brownish orgreenish, with subsynimetrical markings of dark brown ordark green on back and crown of head; a dark band com-mences at tip of snout and extends backward below canthusrostralis through eye and tympanum, ending in a black spotbehind fore limb; a vertical dark band below eye separatingtwo large white spots on labial border of upper jaw; sidesmarbled with dark brown and black; fore and hind limbs withmore or less distinct dark crossbars; lower surfaces whitish,immaculate; male with two large external vocal sacs behindangle of mouth; head-and-body length, 55 to 73 mm baudinii (p. 160)11. Adhesive disks of fingers not noticeably larger than those oftoes, but as large as or larger than tympanum; vomerineteeth in two elongate clusters between the choanae, present-ing an obtuse angle medialh' and posteriorly, but theirhinder ends do not quite reach the level of posterior margins
156 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
of choanae; fronto-parietal bones separated by large fon-tanelle covered by cartilaginous roof; tympanum usuallyone-third or less, and at most one-half the diameter of theeye, and overhung by a dermal fold; fingers one-fourth toone-half webbed (fig. 19, d); the hind limb being carried for-ward along the body, the tibio-tarsal joint reaches beyond theeye and at the most to the end of the snout; toes four-fifthswebbed, the membrane reaching the disks of the third andfifth toes, but leaving free the last phalanx of the fourth;subarticular and supernumerary tubercles extended backwardupon metatarsals; a tarsal fold; tongue notched behind;canthus rostralis distinct, short; loreal region oblique, con-cave; skin of upperparts smooth; under surfaces of thighs,underparts, and gular region coarsely granulated; dermalfold across chest present or absent; upper surfaces purplishash (? green in life) or pale olive brown, with or without afew lighter specks, indistinct darker reticulations, or blackspots; limbs paler, not crossbarred; a light external stripefrom elbow and heel to digits; superior labial border andlateral stripe occasionally bright yellow, there being noabrupt demarcation between the latter and the abdomen insome cases; anal spot with light superior border; undersurfaces yellow; head-and-body length, 35 to 40 mmraiotympanura (p. 170)Adhesive disks of fingers larger than those of toes, and nearlyas large as tympanum; vomerine teeth in oblique rowsbetween and extending back to level of hinder edge ofchoanae; tympanum less than or slightly more than one-half the diameter of the eye; fingers one-fourth webbed;the hind limb being carried forward along the body, the tibio-tarsal joint reaches to center of the eye; toes two-thirdswebbed, the membrane reaching the disks of the third andfifth toes, but leaving free the last phalanx of the fourth;a tarsal fold; tongue oval, not notched behind; canthusrostralis rounded; loreal region oblique; skin of upperpartssmooth; underparts granulated; no dermal fold across chest;upper surfaces light purplish, with or without darker mar-bling; no post-tympanic black mark; sides with a networkof blackish brown spots or blotches on creamy white back-ground; hind limbs with indistinct dark cross bands; under-parts white, with scattered black specks on throat andanterior part of abdomen, concealed parts unpigmented,white; head-and-body length, 30 mm taeniopus (p. 175)HYLA ARENICOLOR CopeCaSon Tree ToadFigure, 20, e1864. Hyla affinis Baird, Proc. Acad. Nat. Sci. Philadelphia, vol. 7, p. 61, Apr.(preoccupied by Hyla affinis Spix, 1824).
?
Baird, 1859, Report on the UnitedStates and Mexican Boundarj' Survey, vol. 2, Reptiles, p. 29, pi. 38, figs.4-7.
MEXICAN TAILLESS AMPHIBLA.NS 1571866. (Hyla) arenicolor Cope, Journ. Acad. Nat. Sci. Philadelphia, n. s., vol. 6,pt. 1, p. 84, July.?DiCKERSON, 1906, The frog book, pp. 122, 123, pi. 48,figs. 143-146.?Stoker, 1925, Univ. California Publ. Zool., vol. 27, pp. 203-215, pi. 2, fig. 4; pi. 13, figs. 37-39; text figs. Z, LL.1887. Hyla copii Boulenger, Ann. and Mag. Nat. Hist., ser. 5, vol. 20, no. 115,p. 53, July (El Paso, Tex.).
?
Boulenger, 1888, Ann. and Mag. Nat. Hist.,ser. 6, vol. 1, no. 3, p. 189, Mar.
?
Gxjnther, 1901, Biologia Centrali-Americana, Reptilia and Batrachia, p. 266, pi. 70, fig. C, June.1888. Hyla coper [Cope], Amer, Nat., vol. 22, no. 253, p. 80, Jan. [errore typogr.].1899. Hyliola digueti Mocqtjard, Bull. Soc. Philom. Paris, ser. 9, vol. 1, no.4, p. 165, pi. 1, fig. 4 (territory of Tepic, Mexico).1905. Hyla copei Gadow, Proc. Zool. Soc. London, vol. 2, p. 194.Type locality.?Northern Sonora, Mexico.Range.?From northern boundary of Sonora and Chihuahuasouthward to Michoacan (Tupataro) and to Guerrero (Chilpancingo),to the Federal District (Tacubaya), and to Vera Cruz (Presidio nearMotzorongo, about 20 miles south of Cordoba). Of the three Mexi-can hylas that occur within the limits of Southwestern United States,this rough-skinned species is the most widely distributed, occurringfrom southern Utah to northern Lower Cahfornia.Remarks.?The cotypes of Hyla affinis may have been collected byJ. H. Clark within the present boundaries of Arizona, although theyare listed by Baird as coming from northern Sonora. A brief itin-erary of Clark, while attached to Colonel Graham's party of theUnited States and Mexican Boundary Survey, will be found in theremarks under Bufo pundatus (p. 62). The original U.S.N.M.number assigned to the cotypes of H. affinis is 3261, and somewhatlater this lot was reentered under 11410. There were originally fourspecimens, but the type bottle now has one additional young specimen.In 1866 Cope published the results of his dissections on theabdominal integuments of Amphibia and described their union withthe superficial fascia of the muscles by an areolar or fibrous network.These closely approximated areolae, which "secrete an adhesive fluidas aid in maintaining the pecuhar positions assumed," are formed bya thickening of the abdominal skin, and they indicate the extent ofabdominal attachment. Cope lists the hylids that he studied accord-ing to the extent of the dermal attachment, and in his second grouparenicolor is proposed as a substitute name for H. affinis, which ispreoccupied by a prior use of the same specific name by Spix.Boulenger was misled by errors in the published description ofHyla affinis and concluded that a specimen of this tree frog from ElPaso, Tex., represented a different species, which he named Hyla copii.Boulenger later insisted that "iJ. arenicolor Cope is a mere name,proposed to replace that of affinis, Baird, preoccupied," and that his66785?32 11
158 BULLETIN 160, UNITED STATES NATIONAL MUSEUMspecimen differed from the original description of Hyla affinis in threeimportant points. According to Baird, in H. affinis the tympanumis two-thirds the size of the eye, the web of hand extends only to thethird joint of the second finger, and there is no vermiculation onanterior and posterior faces of hind legs. On the other hand, H. copii,is described by Boiiienger as having a "tympanum hardly one-thirdthe size of the eye, fingers without web, and hinder side of thighsmottled or vermiculated with brown."The measurements of the four cotypes of H. affinis (U.S.N.M. No.11410) are, respectively, as follows: Head-and-body length, 41.7, 40,37.2, and 31.8 mm.; transverse diameter of tympanum, 3, 2.8, 2.6,and 2.1 mm.; transverse diameter of eye, 5, 4.9, 4.5, and 4 mm.;anterior edge of eye to nostril, 4.1, 3.8, 3.9, and 3.2 mm. On all fourof these cotypes, the transverse diameter of the tympanum is slightlymore than one-half that of the eye. Not one of them possesses anyvestige of a web between the fingers at the base. The presence orabsence of a brownish vermiculation on the posterior surface of thethigh of specimens preserved in spirits is of slight diagnostic value inthis species, though as a general rule this area is flesh or orange col-ored in life and normally is without any brown spotting or marbling.In Mexican specimens, at least, the relative size of the tympanum tothat of the eye seems to be quite variable. The transverse diameterof the tympanum is generally about one-half that of the eye, occa-sionally as much as three-fourths, and rarely only one-third.The five cotypes of Hyliola digueti (M.N.H.P. Nos. 492m, 492n,parchment labels Nos. 98-257, 98-258, 01-343, 01-344, 01-345) werecollected by Leon Diguet in the territory of Tepic [ = Nayarit], Mexico.They comprise four adults and one young individual, and all arereferable to Hyla arenicolor. The head-and-body lengths of the fouradults are, respectively, 41.4, 33, 33.7, and 29.5 mm.; the hind limbbeing carried forward along the body, the tibio-tarsal joint reaches tobetween the eye and tip of snout in three individuals, and slightlybeyond tip of snout in one; tarsal fold sharp-edged; large inner andsmall outer metatarsal tubercles; no web between fingers at base;transverse diameter of tympanum slightly more than half that ofeye; vomerine teeth in two small groups between the choanae; skinof upper parts coarsely granular.Relatively little individual morphological variation, except as notedin the preceding remarks, was observed in the series examined, althoughconsiderable color variation was seen.Specimens examined.?Ninety-four, as follows:
MEXICAN TAILLESS AMPHIBIANSHyla arenicolor 159
Museum CatalogueNo. Num-ber ofspeci-mens Locality collected Date collected By whom collected orfrom whom received
U.S.N.M..M.C.ZU.S.N.M-M.C.ZU.S.N.M..M.C.Z....U.S.N.M.-Do....M.C.Z. ...F.M.N.H.U.S.N.M..Do.-.-C.Q.E.M..U.S.N.M..M.H.N.P..U.S.N.M..M.C.Z....Do....Do....Do....A.M.N.H.Do....Do....Do....U.S.N.M-.A.M.N.H.U.S.N.M-.A.M.N.H-Do...Do...Do...C.G.E.M.U.S.N.M-Do...Do...Do...M.C.Z....U.S.N.M.M.C.Z....
620767372114102386470487649-51470508322967713764743447435646824492m,492n4791712165
1663
O.A.SA.M.N.H-U.S.N.M-.N.M.W...M.C.Z...
.
U.S.N.M-.Do.-..
166510784
12490-911248912114-15189891255833547-4812492-96
12486-?812483-8412485
989799161135926156-679028322958367-6850191325471183-851877-11-1976321025614598
Sonora:NogalesSaa Jose de GuaymasNorthern SonoraNo definite localityChihuahua:Near BallezaBarranca del Cobre, SierraTarahumare.Near BatopUas.Between Mexico and Chi-huahua.MoralesSanzDuRANGO: ElSaltoSinaloa:PlomosasNo definite locality.Zacatecas: PlateadoNayarit:No definite locality.San Luis Potosi:AhualulcoAlvarez (Kilo 58) near SanLms Potosi, altitude 8,000feet.Mountains of Mezquitic, 22miles north of San LuisPotosi.Near San Luis PotosiSan Luis PotosiJalisco:Aquilar Mine, Hostotipa-quillo.AtemajacCerro del ColCerro Pelon, Rio Blanco,north of Zapopan.ChapalaWest and above Hostotipa-quillo.Rio Blanco near Guadalajara.On trail from Tequesquiteto Hostotipaquilla (Sierrade Nayarit).South of Tlaquepaque.TonalaBetween Tonala and Tla-quepaque.No definite locality.Guanajuato:No definite localitydo -dodo -Hidalgo: MiguelMexico:Valleys of Mexico and Tol-uca.San Juan TeotihuacanFederal District:Mexico City -.Santa Fe (IJ^ miles west)Near Tacubaya, Santa Fe...Morelos: CuernavacaMichoacan:Camp near ApatringanTupataro? State: No definite locality..
June, 191918511903Sept. 23, 18981921Oct. 11,1898
1923Juiy'20,"i898July 17,1897Mar., 1920Sept. 2,1897
Aug. 28, 1892/June 23, 1926\Sept. 30, 19261879
18791925Nov. 7,1919Sept. 23, 1919Sept. 18, 1919Sept. 12, 1919Feb. 24, 1892Nov. 2, 1919Sept. 30, 1903Sept. 25, 1919
Sept. 14, 1919Oct. 6, 1919Sept. 17, 1919July 15,191918771877
19131884-851922July 8, 1925May 18, 19191921
Francis J. Dyer.W. W. Brown.J. H. Clark.Nelson and Goldman*Hans Gadow.E. A. Goldman.John Potts.W. W. Brown.S. E. Meek.Nelson and Goldman.Do.J. M. Gallegos.Nelson and GoldmaaL6on Diguet.E. W. Nelson.|w. W. Brown.Edward Palmer.Do.W. W. Brown.Paul D. R. Ruthling.Do.Do.Do.P. L. Jouy.Paul D. R. Ruthling.J. N. Rose and J. H.Painter.Paul D. R. Ruthling.Do.Do.Do.C. Curota Terron.Alfred Dug6s.Do.Do.Do.W. M. Mann.C. T. HoegeE. R. Dunn.Joseph R. Slevin.Paul D. R. Ruthling.A. L. Herrara.
19211879 Hans Gadow.Alfred Dugfis.Do.
1 Cotypes of Hyla affinis. 2 Cotypes of Hyliola digueti.
160 BULLETIN 160, UNITED STATES NATIONAL MUSEUMHYLA BAUDINn Dum^rU and BibronFigures 1, c; 19, e; 20, c1841. Hijla haudinii DumI^ril and Bibron, Erp6tologie g6nerale, vol. 8, pp. 564,565.1854. Hyla vanvUetii Baird, Proc. Acad. Nat. Sci. Philadelphia, vol. 7, p. 61,Apr. (Brownsville, Tex.).?Baird, 1859, Report on the United States andMexican Boundary Survey, vol. 2, ReptUes, p. 29, pi. 38, figs. 1-3.1862. H[yla] muricolor Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 14, p. 359,Sept. (Mirador, Vera Cruz).1882. Hyla baudini Brocchi, Mission scientifique au Mexique et dans I'AmeriqueCentrale, recherches zoologiques, pt. 3, sect. 2, pp. 29-31, pi. 14, figs. 4,4a-b.?GtJNTHER, 1901, Biologia Centrali-Americana, Reptilia and Batrachia,p. 270, pi. 71, figs, a-d, Sept.1906. Smilisca haudinii Dickerson, The frog book, p. 151, pi. 57, figs. 178-180.Type locality.?Mexico.Range.?From Bexar County, Tex., south through Vera Cruz toYucatan and Central America; and on the west coast from Sinaloa(Mazatlan) southward through JaUsco, Guerrero, Oaxaca, and Chiapasto Guatemala.Remarks.?The type of Hyla haudinii (M.H.N.P. No. 401, parch-ment label No. 4798), according to the catalogue, came from Mexico.It exhibits the typical features of this species, as is shown by thefollowing notes: Head-and-body length, 54.8 mm.; transverse diam-eter of tympanum, 3.6 mm.; transverse diameter of eye, 6 mm.; thehind limb being carried forward along the body, the tibio-tarsal jointreaches to anterior margin of eye; a long sharp-edged tarsal fold; alarge inner but no outer metatarsal tubercle; toes webbed as usual;large apical disks; fingers webbed at base; tip of first finger reachesto apical disk of second; two external vocal vesicles; vomerine teethin two transverse rows between and at about level of middle ofchoanae; throat, abdomen, and under surface of thighs coarselyareolate or granular; black crossbars on hind limbs; an interorbitalblack band with lateral streaks that connect posteriorly with irregularblack markings on upperparts.Although the type of Hyla vanvlietii (presumably U.S.N.M. No.3239) is now lost, the description and figures published by Bairdindicate that he had in hand a specimen of tliis species. Accordingto the entry in the catalogue, this specimen was collected at the mouthof the Rio Grande, Tex., by J. H. Clark, while he was employed bythe United States and Mexican Boundary Survey under the super-vision of Maj. W. H. Emory. The type locahty is given as Browns-ville, Tex., in the original description.The type of Hyla muricolor (U.S.N.M. No. 25097) is a large indi-vidual, which can be matched in size, color, and structure with speci-mens of haudinii taken elsewhere in Mexico. It was collected byDr. Carlos Sartorius in the hills west of Mirador, Vera Cruz. Thefollowing notes were made on this type: Head-and-body length, 72.5
MEXICAN TAILLESS AMPHIBIANS 161mm. ; transverse diameter of tympanum, 5.5 mm. ; transverse diameterof eye, 8.3 mm.; anterior edge of eye to nostril, 6.2 mm.; the hindlimb being carried forward along the body, the tibio-tarsal jointreaches to center of eye; a sharp-edged tarsal fold from inner meta-tarsal tubercle to tibio-tarsal joint; large inner and vestigial outermetatarsal tubercle; vomerine teeth in two transverse series nearlevel of middle of choanae; upperparts \vith dark confluent spots andinterorbital black band; fore and hind limbs with dark cross bands;skin on crown of head split longitudinally to expose fronto-parietalregion; body wall on left side damaged, exposing egg mass.Ruthven ^^ says of this species:There was much variation in color in the specimens of H. baudini which weexamined in life. Occasional individuals were a dull orange *9 (about 153d)above, but in by far the greater majority of the ground color of the back was darkolive green (230), dark olive (180), dark brown (130 or 155) or dark orange (105).The head was a shade lighter than the ground color in most specimens, and thepale spot varied from a pale dull greenish tint (203d) to a very pale yellow (221)or bright green (336). The belly was nearly always a light orange tint (116, 121,136), and the light spots on the sides, light orange yellow (191). The dark mark-ings vary greatly in distinctness. In the paler individuals they are usuallyindistinct or even entirely wanting, except for faint traces. In those which havea darker ground color they are usually distinct, particularly the black stripe fromthe tympanic region to the shoulder. They also vary in arrangement, for theymay either be connected into a more or less irregular dorsal stripe or stripes,beginning with the cross band between the eyes, or they may be in the form ofirregularly extended spots.We found H. baudini common at Cuatotolapam. Most of the specimens weretaken during night rains on the banana trees at San Juan. At these times theywere very noisj'. During the day we found them secreted under boards, in thebases of such large leaved plants as the "elephant ears," bananas, etc. Theywere observed breeding in a pond near La Laja Creek on July 17.Dr. E. W. Nelson states in his field catalogue that in wet weatheron June 10, 1895, three of these tree frogs were found on the groundat an altitude between 900 and 1,200 feet in the mountains of SantoDomingo, Oaxaca. On July 19, 1897, he found two of them hidingunder brick tiles on the roof of a veranda, where they uttered hoarsecroakings during the evening and at nightfall.As an illustration of the elusiveness of tree frogs. Dr. Hans Gadow ^?contrasted an occurrence of countless numbers of Hyla haudiniispawning in a wet meadow near Presidio south of Cordoba in VeraCruz with his fruitless search for this species on his subsequent travelsin Mexico. This experience is described as follows:Whilst rambling along the edge of the forest we became conscious of a noise,at first resembling the mutter of a distant sawmill; but on our reaching the other
" Ruthven, A. G., The amphibians and reptiles collected by the University of Michigran-Walker Expe-dition in southern Vera Cruz, Mexico. Zool. Jahrb. (Syst. Abt.), vol. 32, pt. 4, pp. 310, 311, 1912.M The numbers refer to the "Code des Couleurs," by Klincksiek and Vallete.60 Oadow, H., Through southern Mexico: Being an account of the travels of a naturalist, London, pp.75,76, 1908.
162 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
side of a cluster of trees this sound grew into a roar, like that of steam escapingfrom many engines, mingled with the sharp and piercing scream of saws. Itcame from a meadow containing a shallow pool of rainwater. In the wet grass,on its stalks, and on the ground, hopped about hundreds of large green tree-frogs;nearer the pool they were to be seen in thousands, and in the water itself there weretens of thousands. Hopping, jumping, crawling, shding, getting hold of eachother, or sitting still. Most of them were in amplexus, and these couples werequiet, but the solitary males sat on their haunches and barked solemnly, withtheir resounding vocal bags protruding. Every now and then one was makingfor a mate, and often there were three or four hanging on to each other and rollingover. The din was so great that it was with difficulty that we caught the remarksthat we shouted, although we were standing onlj' a few feet apart. Each sweepof a butterfly net caught at least half-a-dozen frogs.Now the grassy pool, where the frogs were closest, was about 30 yards square(900 square yards), rather more than the area of a tennis lawn, and each squareyard held from 50 to 100 frogs?many square yards certainly held several hundredseach. At the very lowest computation this gives 45,000 frogs; and there was,besides, an outer ring of some five hundred square yards where frogs were fairlynumerous, say from 5 to 10 to the square yard, mostly spent females, but thesefew thousands we may leave out of the reckoning, to understate rather than over-estimate the number. Supposing there were onh^ 20,000 females, each spawningfrom 5,000 to 10,000 eggs?say only 5,000?the total would amount to just 100,000,000 eggs. The spawn literally covered both ground and water thickly. Butthe greatest surprise awaited us on the following morning, when we went to photo-graph the scene. There was not a single frog left; the water had all evaporated,and the whole place was glazed over with dried-up spawn! The prospectivechance of millions of little frogs was gone, their expectant parents having beendeceived in calculating their day of incarnation. That was on the 4th of July,several weeks after the beginning of the rather fitful rainy season.Specimens examined.?Eighty-four, as follows
:
Hyla haudinii
Museam
A.M.N.H...U.S.N.M....Do.Do....C.O.E.M.N.M.W...U.S.N.M..Do.DoM.O.ZU.S.N.MC.A.SDoU.S.N.MF.M.N.H..-.U.S.N.MM.C.ZU.S.N.M.N.M.W.
.
Do...
CatalogueNo.
MEXICAN TAILLESS AMPHIBIANS 163Hyla baudinii?Continued
Museum
164 BULLETIN 160, UNITED STATES NATIONAL MUSEUMdark bluish plumbeous and purplish, respectively, indicates that thegeneral color of the upperparts in both species "in life is some shadeof green. There are, however, some differences in the general bodybuild of these tree frogs that are very difficult to describe.The structual features that appear to distinguish them are as follows:In Hyla taeniopus, the vomerine teeth are in small clusters betweenthe choanae ; transverse diameter of tympanum about one-half thatof the eye; a tarsal fold extends from inner metatarsal tubercle totibio-tarsal joint; tip of first finger reaches to apical disk of second;fingers one-fourth webbed; upper surface of hind Umb with darkcrossbars. The type of H. bistinda differs from the cotypes of H.taeniopus in having the vomerine teeth in two slightly oblique rows atlevel of posterior margins of choanae; transverse diameter of tym-panum one-third that of the eye; tarsal fold indistinct; first fingerconsiderably shorter than second; fingers wdth vestigial web at base;upper surface of hind Umb without any trace of dark crossbars.Specimens examined.?One, the type.HYLA EXIAHA Baird1854. Hyla eximia Baird, Proc. Acad. Nat. Sci. Philadelphia, vol. 7, p. 61, Apr.
?
Baird, 1859, Report on the United States and Mexican Boundary Survey, vol.2, Reptiles, p. 29, pi. 38, figs. 8-10.
?
Brocchi, 1882, Mission scientifique auMexique et dans rAm<5rique Centrale, recherches zoologiques, pt. 3, sect. 2,pp. 32, 33, pi. 1, fig. 2; pi. 13, figs. 4, 4a.1858. Hyla euphorbiacea GtiNXHER, Catalogue of the Batrachia Salientia in thecollection of the British Museum, p. 109, pi. 10, fig. C (Cordoba and Cordil-leras, Mexico).1901. Hyla nana Gunther, Biologia Centrali-Americana, Reptilia and Batrachia,pp. 263, 264, pi. 73, fig. A, June (Cuernavaca, State of Morelos, Mexico).[Nee Hyla nana Boulenger, 1889, Ann. Mus. Civ. di Storia Nat. Genova,ser. 2, vol. 7, p. 249, pi. 2, fig. 2 (Colonia Resistencia, South Chaco, Argen-tine Republic).]1902. H[yla] smithii Boulenger, Zool. Rec, vol. 38, Rept. and Batr., p. 33.Type locality.?City of Mexico, Federal District, Mexico.Range.?From Durango southward through Guanajuato, Hidalgo,and Puebla to Vera Cruz (Cordoba), and on the west coast south-ward through Nayarit to Michoacan and Oaxaca.Remarks.?The two cotypes of Hyla eximia (U.S.N.M. No. 3248)were collected by Maj. William Rich, in the vicinity of the City ofMexico, Federal District. Both of these cotypes are young individ-uals and measure 19.2 and 19.5 mm. in length, respectively. Theoriginal coloration of these specimens has disappeared, and the bodiesare soft and nearly transparent. The bodily proportions and struc-tural features agree with young individuals of the tree frog currentlyrecognized as Hyla eximia. The original drawing of the adult treefrog figured by Baird in 1859 does not have any notation as to thecatalogue number or locality. This specimen has not been recognized
MEXICAN TAILLESS AMPHIBL^NS 165among the specimens of this species now deposited in the UnitedStates National Museum.Giinther's Hyla euphorbiacea was based on five specimens, whichare clearly identical with H. eximia. The following notes were madeon these cotvpes:B.M. No. 1930. 4. 10. 2: Collected by Auguste Salle at Cordoba inVera Cruz and figured by Gtinther in 1858; head-and-body length,30.2 mm.; the hind limb being carried forward along the body, thetibio-tarsal joint reaches to posterior margin of eye; normal colorpattern. B.M. No. 57. 10. 28. 51: From the Cordilleras of Mexicoand purchased from fi. Parzudaki; head-and-body length, 33.8 mm.;the hind limb being carried forward along the body, the tibio-tarsaljoint reaches to center of eye; normal color pattern. B.M. No.58. 11. 22. 5: From the Cordilleras of Mexico and purchased from?. Parzudaki; label states that it was found "dried up" in 1906.B.M. Nos. 1858. 6. 15. 4-8: From Hugh Cuming's Mexican collection;head-and-body length, respectively, 39.8 and 40.3 mm.; the hind Hmbbeing carried forward along the body, the tibio-tarsal joint reaches toposterior margin of eye in one, and almost to center of eye in theother; a black canthai stripe edged above ^^'ith white; and a post-ocular lateral dark stripe edged above with white, which tends todisappear on posterior half of side of body.The cotypes (B.M. Nos. 1901. 12. 19. 76-82) of Hyla nana Giinther(pTeoccupied = Hyla smithii Boulenger) are likewdse referred to thisspecies. Dr. E. R. Dunn carefully checked a series of specimens ofH. eximia with the cotypes of H. nana and concluded that this specieswas not valid. Converseh', H. W. Parker, who had previously exam-ined the cot^^pes, wrote that H. smithii seemed to differ from bothH. eximia and H. gracilipes. The seven cotj^pes of H. nana [ = smithii]were collected by H. H. Smith at Cuernavaca in Morelos. My ownobservations on one of these cotypes are as follows: Head-and-bodylength, 25.8 mm.; transverse diameter of tympanum, 1.2 mm.;transverse diameter of eye, 2.5 mm.; anterior edge of eye to nostril,2.6 mm.; the hind limb being carried forward along the body, thetibio-tarsal joint reaches to anterior margin of eye; no tarsal fold; anelongate inner, but no outer metatarsal tubercle; vomerine teeth quitesmall and in two clusters between the choanae; side of head from tipof snout to eye and from margin of upper lip to canthus rostralisdark brownish black, but edged above with a narrow light line; ablack streak between eye and tympanum; behind tympanum thisdark lateral band tapers posteriorly to a narrow ribbon and disappearson fianks; upperparts j-ellowish or olive-green; abdomen and undersurface of thighs yellowish; skin on abdomen and under surface ofthighs granular. The development of the vomerine teeth and thesize of these cot3"pes indicate that none of them are fully adult. Thelargest individual lacks a tarsal fold and has a dark loreal region.
166 BULLETIN 160, UNITED STATES NATIONAL MUSEUMThis combination has not been exactly matched in the series of H.eximia available for study. The tarsal fold on immature individualsof eximia is occasionally indistinct and when viewed under a micro-scope can be detected only by utilizing the light to the best advantage.A distinct tarsal fold was observed in a representative series ofeximia from Vera Cruz, Guanajuato, Nayarit, and Michoacan, and inat least 13 of these 35 specimens this tarsal fold was distinctly sharp-edged. The tarsal fold extends in a somewhat curved line from theinner metatarsal tubercle nearly to the tibio-tarsal joint.Individual variation in a tree frog as widely distributed as H.eximia occasions no surprise. In this species the dark markings onthe upperparts generally run more to longitudinal dark streaks thanto distinct dorsal spots. A specimen taken at Ciudad in Durango(U.S.N.M. No. 14083), however, has an unusual color pattern, con-sisting of closely spaced dark spots and streaks that extend from tipof snout to vent. Another specimen taken at Oaxaca in Oaxaca(U.S.N.M. No. 47908) has a large inner and a minute outer meta-tarsal tubercle. In at least one specimen, labeled as coming fromGuanajuato (U.S.N.M. No. 9898), the tibio-tarsal joint reaches onlyto the tympanum.It is often possible to point out pecularities that seem to be diag-nostic characters in a limited series, but such distinctions frequentlydisappear when a large number is available. While the extremevariants of H. eximia may seem quite distinct, and this variabilitymay account in part for several of the synonyms, there are so manyintermediate types of color pattern that differences of this sort canhardly be considered of diagnostic importance. The average indi-vidual of eximia has a black spot on each upper eyelid; paired longi-tudinal dark streaks on back; dark spots in sacral region; a lateraldark streak, edged above with white, from nostril to eye, and fromposterior edge of upper eyelid through tympanum and along sides toabout level of middle of sacrum; and a white or light-colored stripealong upper lip below eye from end of snout to axilla. On some of thespecimens, the color pattern of the upperparts consists of scattereddark spots, without any tendency toward uninterrupted streaks.Many specimens have a pair of longitudinal sacral streaks. A fewspecimens were noted that have uniformly colored upperparts withoutany trace of dorsal dark spots. A peculiarly marked specimen fromNayarit (U.S.N.M. No. 65890) had longitudinal dark streaks and alsonumerous fine black specks on upper surface of fore and hind limbs,and a lesser number on sides and back. A few individuals have com-plete transverse dark bars on upper surfaces of hind limbs, whileothers have dark spots, but no complete crossbars. At least 10specimens having a light-edged dark streak along outer surface oftibia were noted. The hinder surface of the thigh occasionallyexhibits a distinct color pattern consisting of a darker area more or
MEXICAN TAILLESS AMPHIBIANS 167less marbled with light and often with a light upper edge. Otherindividual peculiarities less marked than those listed above might becited, but they indicate the variation that may be expected.Specimens examined.?One hundred and ninety-three, as follows:Hyla eximia
Museum
U.S.N.MF.M.N.HM.C.Z.Do.
M.C.Z.Do.B.MM.C.Z...U.S.N.M.Do...
F.M.N.H...DoU.S.N.M...-M.C.ZDo.B.M...
CatalogueNo.
U.S.N.MDoA.M.N.H---Do.Do.Do.Do.Do.Do.Do-F.M.N.H....DoU.S.N.MDoDoDoDo
F.M.N.H....A.M.N.H....DoDoU.S.N.M...A.M.N.H.-Do
A.M.N.H....C.O.E.M....A.M.N.H...U.S.N.MM.N.W ,O.A.SDoA.M.N.HU.S.N.M..
140831512
16611003746830-3465889-9012068-8112082120671206212063-6612049-5812059-6112083-90
961962987598981135712682102373906-119029-31
1930.4.10.2708731289-9932396-9815221326612042-4813257-6432481325512005-41
136432296-972264 A-B22651901. 1219. 76-8213862313863113641910-61610761241392747908
Num-ber ofspeci-mens Locality collectedDURANGO:Ciudad, altitude 8,100 feet...CoyotesSan Luis Potosi:Mountains of AlvarezEbanoNayarit:Santa Teresa, altitude 6,800feet.TepicJalisco:Aqua Delgada reservoir, 4miles north of Guadalajara.Jamay..Laguna de MagdalenaOcotlanSouth of Tlaquepaquedo.Tonala .-.Grassy fields between Tonalaand Tlaquepaque.Guanajuato:AcambaroCelaya .-Guanajuatodododo -SilaoHidalgo:GuerreroHacienda de Velasco, north-east of Real del Monte.Vera Cruz:CordobaOrizaba .-.Vicinity of OrizabaPotrero near CordobaSuchilFederal District:Chapultepee ParkNorth of Guadalupe.-do...City of MexicoSan JuanicoOne mile west of Tlalpam(rain pools).Mexico:ChalcoLermaValley of Mexico and TolucaSan Juan Teotihuacan, Val-ley of Mexico.do -Morelos: CuernavacaPuebla:Los Reyes, near Santa Cata-lina.PueblaSanta CatalinaMichoacan: TupataroGuerrero: TacubayaOaxaca:Cosolapado. --MixtequilloOaxaca..
Date collected
Apr. 3, 1885
1879May 10,1922Aug. 8, 1897June 22, 1923Aug. 10,1919Aug. 21,1919Sept. 29, 1919Aug. 16,1919Sept. 10, 1919Sept. 14, 1919Oct. 6, 1919Sept. 17, 1919May 27,1901May 28,190118771877Jan. 30,18801881187919131913
1920/Dec. ? , 1904-IJan. ?.1905May 3, 1919July 8, 1919May 5, 1919May 7,1919July 13,1919/Apr. 30,1901-IMay 1, 19011884-8518901890
July 28,1920Sept. ?, 1919July 31,19201881
July 14,1925doMay 19,1920June 20, 1894
By whom collected orfrom whom received
Alphonso Forrer.E. Heller and C. M.Barber.Edward Palmer.W. W. Brown.Nelson and Goldman.W. M. Mann.Paul D. R. Ruthling.Do.Do.Do.Do.Do.Do.Do.
S. E. Meek.Do.Alfred Dug6s.Do.Do.Do.Do.W. M. Mann.Do.Auguste Sal]6.W. S. Blatchley.Francis Sumichrast.Do.E. Heller and C. M.Barber.Paul D. R. Ruthling.Do.Do.William Rich.Paul D. R. Ruthling.Do.
?S. E. Meek.Do.C. T. Hoege.G. O. Rogers.Do.H. H. Smith.
Paul D. R. Ruthling.H. Ruano.Paul D. R. Ruthling.Alfred Dup6s.Hans Gadow.Joseph R. Slevin.Do.Paul D. R. Ruthling.Nelson and Goldman.
1 Cotypes of Hyla euphorbiacea. 2 Cotypes of Hyla eximia. ' Cotypes of Hyla nana.
168 BULLETIN 160, UNITED STATES NATIONAL MUSEUMHyla eximia?Continued
Museum
MEXICAN TAILLESS AMPHIBIANS 169somewhat oblique or transverse rows between the choanae, andseparated medially by a short interval; tongue large, broad, andfeebly notched behind; snout broad, rounded; canthus rostralisdistinct; loreal region sloping more or less obliquely; distance betweenanterior edge of eye and narial opening greater than, equal to, orsomewhat less than, transverse diameter of eye; interorbital spacesomewhat wider than upper eyelid; pupil oval, broader than high;iris dark yellow, spotted with bronze, and without cross line; greatestdiameter of tympanum equal to two-thirds to three-fourths the trans-verse diameter of eye; fingers united at base by dermal fringe orvestigial web; first finger shorter than second; adhesive disks offingers somewhat larger than those of toes and about one-half aslarge as tympanum; toes two-thirds webbed; tarsal fold weakly orstrongly developed; inner metatarsal tubercle distinct, egg-shaped,and about as long as first joint of first toe; outer metatarsal tuberclesmall and conical, or flat, or absent; subarticular tubercles distinct;the hind limb being carried forward along the body, the tibio-tarsaljoint reaches to or slightly beyond tip of snout; the heels touch oroverlap when the hind limbs are brought together at right angles tolong axis of body; a discoidal dermal fold across chest; a dermal foldextends above and downward behind tympanum to axilla; upperpartsdark green, grass green, olive-green, or brownish in life (dirty blue inalcohol) with paired oblong white-edged dark spots on hinder back;sides of snout green; upper lip with dark gray stripe, edged above,or above and below with white; loreal stripe blacldsh, edged abovewith white, bronze, or light brown; from eye to tympanum and behindtympanum a dark lateral streak of regular or irregular wid th extendsbackward to about halfway length of body, where it either disappearsor is broken up on hinder half of flank into a series of spots; green offlanks not so sharply contrasted with whitish underparts; inguinalregion either whitish or udth bluish tinge; anterior surface of thighwhitish or bluish; hinder surface of thigh always without darkermarkings, with the flesh color showing through the bluish tinge:outer edge of tibia and tarsus with lengthwise light-edged dark streaks;cross bands on limbs indistinct, or incomplete (small spots), or welldeveloped; a dark spot on inner side of upper arm; underparts whitish,without darker markings; throat of female white, of males eitheruniformly light gray or else rather dark greenish gray, and occasionallyspotted; region of vent rather dark or intense gray; head-and-bodylength of males, 35 to 38 mm., of females, 37.8 to 50 mm.; greatestdiameter of tympanum of males, 2.2 to 2.5 mm., of females, 2.1 to2.7 mm.; transverse diameter of eye of males, 3.2 to 3.7 mm., offemales, 3.4 to 3.8 mm.; w^dth of head of males, 12.6 to 13 mm., offemales, 13 to 15.5 mm.
170 BULLETIN 160, UNITED STATES NATIONAL MUSEUMSpecimens examined.?Thirteen, as follows
:
Hyla gracilipes
Museum
MEXICAN TAILLESS AMPHIBIANS 171tion has since been confirmed by Dr. E. R. Dunn, who has reexaminedthe cotypes of H. godmani in the British Museum of Natural History.Dunn reports that he was unable to distinguish Giinther's species,and my own subsequent comparisons resulted in the same conclusion.One of the cotypes of H. godmani (B.M. No. 1901. 12. 19. 96) wascollected at Misantla in Vera Cruz in June, 1888, and was presentedby F. D. Godman. It is described in my notes as follows: Head-and-body length, 36.5 mm.; transverse diameter of tympanum, 1.7mm.; transverse diameter of eye, 4 mm.; anterior edge of eye tonostril, 3.3 mm.; width of head at level of posterior angles of jaws,13 mm.; the hind limb being carried forward along the body, thetibio-tarsal joint reaches to end of snout; toes four-fifths webbed, thelast phalanx of fourth toe free; a long tarsal fold from inner meta-tarsal tubercle to tibio-tarsal joint; tip of first finger barely reachesto apical disk of second; fingers one-third webbed; abdomen andunder surface of thighs coarsely areolate; vomerine teeth in slightlyoblique, posteriorly converging rows, between the choanae.The remaining eight cotypes of H. godmani (B.M. Nos. 1901. 12.19. 88-95) came from Jalapa in Vera Cruz, and were likewise presentedby F. D. Godman. This series ranges in head-and-body length from25 mm. to 38.3 mm. In alcohol they are quite variable in color, thefollowing varieties being noted: Uniform gray; gray with black specks;purplish gray with white specks; purplish gray with lighter blotches;reddish purple with darker confluent marks; brownish with blackspecks; and purplish brown. In case of the largest individual, thehind limb being carried forward along the body, the tibio-tarsal jointreaches to between eye and tip of snout; and on the next to largestindividual it reaches only to anterior margin of eye. There is a sharp-edged tarsal fold on both of these individuals that extends from innermetatarsal tubercle to tibio-tarsal joint. The vomerine teeth are intransverse rows between the choanae.The coloration of this tree frog seems to be quite variable. Theupperparts may be unicolored, with white supralabial and lateralline, and with whitish underparts, as in U.S.N.M. Nos. 57740 and16568-76; or the upperparts may be finely speckled, as in U.S.N.M.No. 71090. Dr. E. W. Nelson found two of these tree frogs thatwere paired in a small stream in the woods near Mirador on February11, 1894. They were pale olive-brown above and whitish below.On July 9, 1926, a party from the University of Michigan took oneof these tree frogs in a second growth of forest jungle near brookcascades in the vicinity of Necaxa, Puebla.
172 BULLETIN 160, UNITED STATES NATIONAL MUSEUMSpecimens examined.?Eighty-two, as follows:Hyla miotympanum
Museum
U.S.N.M.B.M -U.S.N.M.Do.--B.MM.C.Z...U.S.N.M.?Do...Do...Do...Do...Do...Do...Do...
CatalogueNo.
3826&-681901. 12.19. 88-954696647228-291901. 12.19.967038-4216568-7616577-8216583-9471090-10971150-527110932379-4013243U.M.U.M.... 5M.N.B 6657U.S.N.M ' 30302-04U.S.N.M ! 57740
Num-ber ofspeci-mens Locality collectedVera Cruz:Cordoba..JalapaJicoMirador..Misantla.OrizabadodododoOrizaba, alpine regiondoPotrero(?) MoRELOs: 40 leagues southof City of Mexico.Puebla:NecaxaNo definite localityOaxaca: TehuantepecChiapas
Date collected By whom collected orfrom whom received
Apr. 28, 1908-June 29, 1893Feb. 11, 1894June ?, 1888
July 9, 1926-
-
June 5, 1905.
Frederick Knab.F. D. Qodman.E. W. Nelson.Nelson and Goldman.F. D. Godman.W. S. Blatchley.Francis Sumiclirast.Do.Do.Do.Do.Francis Sumichrast.John Potts.H. B. Baker.Herr Berkenbusch.Francis Sumichrast.J. Hurter.Cotypes of Hyla godmani.
' Cotypes of Hyla microtis. HYLA PHLEBODES Stejneger1906. Hyla phlebodes Stejneger, Proc. U. S. Nat. Mus., vol. 30, no. 1471, pp. 817,818, June 4.
?
Barbour and Cole, 1906, Bull. Mus. Comp. Zool., vol. 50,no. 5, p. 154, Nov.Type locality.?San Carlos, Costa Rica.Range.?From Costa Rica northward through Guatemala tosouthern Yucatan, Mexico.Remarks.?As with most of the tree frogs occurring in Mexico, verylittle is known in regard to the habits of this little delicate-skinnedspecies. Joseph R. Slevin, of the California Academy of Sciences,observed numbers of these tree frogs clinging to reeds in a Guatemalaslough and captured a large series with the aid of an insect net. Theupperparts (in alcohol) of the type (U.S.N.M. No. 29970) are darkvinaceous-drab with a network of dark purplish reticulations. Themeasurements of this specimen are as follows: Head-and-body length,28.5 mm.; transverse diameter of tympanum, 1.4 mm.; transversediameter of eye, 3.5 mm.; anterior edge of eye to nostril, 2.3 mm.Two specimens (M. C. Z. No. 2463) from Chichen Itza in Yucatanare referred to this species, and they differ from the type in havingmore conspicuous darker markings. The color pattern is unsym-metrical and consists of a network of irregularly edged dorsal darkstreaks in addition to scattered minute dark specks; interorbital darkband irregular. A curved dark streak with anterior and posteriorprojection crosses back near fore end of sacrum, and behind this areone or more dark blotches.
MEXICAN TAILLESS AMPHIBIANS 173Specimens examined.?Tv^o (M.C.Z. No. 2463), collected at ChichenItza, Yucatan, in 1904, by L. J. Cole.HYLA PUCATA Brocchi1877. Hylaplicata Brocchi, Bull. Soc. Philom. Paris, ser. 7, vol. 1, no. 3, p. 126.Brocchi, 1882, Mission scientifique au Mexique et dans I'Amerique Cen-trale, recherches zoologiques, pt. 3, sect. 2, pp. 35, 36, pi. 12, fig. 1.Type locality.?Mexico.Range.?Unknown.Remarks.?This little tree frog is described as having the vomerineteeth situated at the level of the anterior edges of the choanae.This peculiarity alone distinguishes this species from all other knownMexican hylas. The type (M.H.N.P. No. 380a, parchment labelNo. 6317), according to the label was collected somewhere in Mexicoby Firmin Bocourt, and is described in my notes as follows: Head-and-body length, 38.7 mm.; transverse diameter of tympanum, 2.4mm.; transverse diameter of eye, 4.3 mm.; the hind limb being car-ried forward along the body, the tibio-tarsal joint reaches to betweenthe eye and tip of snout; a tarsal fold; toes webbed to base of penulti-mate phalanx; inner metatarsal tubercle distinct, outer one missing orelse very minute; no supernumerary tubercles on plantar surface offoot; fingers free, no web at base; first finger shorter than second;vomerine teeth in tvro transverse rows near level of anterior mar-gins of choanae; coloration of upperparts apparently gj-eenish; under-parts yellowish; a narrow light yellow stripe along canthus rostralisfrom nostril to eye, and from posterior edge of upper eyelid backwardabove tympanum, and then obliquely downward to middle of side;this stripe seems to be the upper margin of a broad dark band thatextends obliquely downward behind the tympanum; throat, abdomen,and under surface of thighs coarsely granulated.Specimens examined.?One, the type.HYLA STAUFFERI Cope1865. Hyla staufferi Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 17, p. 195, Oct.Type locality.?Orizaba, State of Vera Cruz, Mexico.Range.?Known to occur onl}^ at three localities in western VeraCruz?Orizaba, Potrero, and Motzorongo.Remarks.?This puzzling little tree frog seems to dift'er from Hylaeximia in several respects. It has no tarsal fold, the loreal region ismore oblique, and the interval between eye and nostril is noticeablygreater than the transverse diameter of the eye. The original colorpattern of the type (U.S.N.M. No. 15317) has disappeared. Accord-ing to Cope, the upperparts were originally dark olive, with inter-orbital dark bar, a longitudinal black bar over each scapula, and anindistinct dark stripe along the coccyx. An approach to this arrange-66785?32 12
174 BULLETIN 160, UNITED STATES NATIONAL MUSEUMment of the darker markings is found in some lightly marked indi-viduals of Hyla eximia. On the other hand, the head of H. staufferiseems to have a different outline than that of eximia. The type ofstaufferi is approximately the same size as one of the cotypes of H.Tiana [ = smithii], and all these specimens lack a tarsal fold. Thecotypes of H. smithii, however, have a dark lateral band with whiteupper margin and thus agree with eximia.Three specimens, one young and two supposed adults (U.S.N.M,Nos. 32396-98), collected by Francis Sumichrast at Potrero nearCordoba in Vera Cruz, are referred to this species. The followingobservations are based on the two adults : Head-and-body length, 27.3and 30.3 mm.; transverse diameter of tympanum, 1.2 and 1.2 mm.;transverse diameter of eye, 2.6 and 3.1 mm.; anterior edge of eye tonostril, 2.9 and 3.5 mm.; width of head at level of posterior angles ofjaws, 9.2 and 9.6 mm.; no tarsal fold; tip of first finger does not orbarely reaches to base of second ; the hind Hmb being carried forwardalong the body, the tibio-tarsal joint reaches to center or to anteriormargin of eye; a dermal fold across chest; an interorbital dark bar;a curved dark streak over vertebral margin of scapula; a lateral darkstreak from posterior edge of upper eyelid, backward above tympanumand along sides to about halfway length of body; dark spots orsplotches in sacral region; upper surface of fore and hind limbs withdark crossbars.The measurements of the type of Hyla staufferi are as follows:Head-and-body length, 25.5 mm.; transverse diameter of tympanum,1.3 mm. ; transverse diameter of eye, 2.4 mm. ; anterior edge of eye tonostril, 3 mm.; width of head at level of posterior angles of jaws, 8.5mm. The exact status of staufferi can not be settled by comparisonwith the available series of eximia, and in absence of conclusive evi-dence that it is merely a variant of the latter, this small tree frog istentatively recognized as a distinct species.A specimen (B. M. No. 1903. 9. 30. 286) from Motzorongo in VeraCruz is considered by Dunn to be another example of H. staufferi.Specimens examined.?Four, as follows
:
Hyla staufferi
Museum
MEXICAN TAILLESS AMPHIBIANS 175HYLA TAEMOPUS GUnther1901. Hyla taenio'pus Gunther, Biologia Central!-Americana, Reptilia andBatrachia, pp. 269, 270, pi. 72, fig. f, Sept.Type locality.?Jalapa, State of Vera Cruz, Mexico.Range.?Not known to occur elsewhere than at the type locahty.Remarks.?The status and relationships of this species are quitepuzzling. The two cotypes (B. M. Nos. 1901. 12. 19. 86-87) werecollected by a Mexican Indian, Mateo Trujillo, who was employed asa collector by F. D. Godman. The development of the vomerineteeth indicates that they may possibly be immature individuals. Thegeneral shape of the body, the extent of the web between fingers andtoes, as well as the coloration of the upperparts and sides, are super-ficially like a young Phyllomedusa. The shape of the pupil of the eye,however, is uncertain. After studying these cotypes, H. W. Parkeris inclined to regard taeniopus as being specifically distinct from bavdi-nii. One of the cotypes differs from a juvenile Hyla baudinii in havinga tympanum that is slightly less than one-half the transverse diameterof the eye, and a more depressed and more prominent snout.The following comments on the cotypes of this species were fur-nished by Dr. E. R. Dunn:I found it difficult to make up my mind about Hyla taeniopus. I believe thatthey are not H. baudinii, but it is diflScult to decide. The cotypes are both youngand differ from each other, but are obviously the same species. They lack thepost-tympanic black mark and show more black on the sides. The lateral mar-bling is very striking, more so on one than on the other. The size of the tympanumis not the same on both cotypes, and one has a longer snout than the other.These cotypes are described in my notes as follows: Larger indi-vidual: Head-and-body length, 30 mm.; transverse diameter of tym-panum, 1.7 mm. ; transverse diameter of eye, 3.3 mm. ; anterior edge ofeye to nostril, 3.1 mm. ; width of head at level of posterior angles of jaws,10.7 mm.; the hind limb being carried forward along the body, thetibio-tarsal joint reaches to center of eye; fingers one-fourth webbedat base; tip of first finger reaches to apical disk of second; toes two-thirds webbed; large inner and small outer metatarsal tubercle;a tarsal fold from inner metatarsal tubercle to tibio-tarsal joint;vomerine teeth in small clusters between the choanae; tongue broad,circular; abdomen and under surface of thighs coarsely granular;upperparts (in alcohol) rather uniform purplish; sides creamy whitewith anastomosing blackish-brown spots or l^lotches, the white areasbeing slightly larger than the dark-brown areas; three indistinctdark crossbars on upper surface of thigh and tibia, and two on tarsus;throat and chest with brown specks. The coloration of the smallercotype is the same as the preceding, except that the dark spots onsides do not anastomose, and there are similar dark crossbars onhind limbs. The head-and-body length of this cotype is 24.8 mm.The vomerine teeth of both have barely penetrated the gums.
176 BULLETIN 160, UNITED STATES NATIONAL MUSEUMSpecimens examined.?Two cotypes (B.M. Nos. 1901. 12. 19.86-87), collected at Jalapa, Vera Cruz, by Mateo Trujillo.HYLA VENULOSA (Lanrenli)Figures 19, c; 20, d1768. Rana venulosa Laurenti, Synopsin reptilium, p. 31.1858. Hyla Uchenosa Gunther, Catalogue of the Batrachia Salientia in thecollection of the British Museum, p. 102, pi. 8, fig. c (Vera Cruz and Cordoba,Vera Cruz, Mexico).
?
Brocchi, 1882, Mission scientifique au Mexique etdans rAm6rique Centrale, recherches zoologiques, pt. 3, sect. 2, p. 33, pi. 14,fig. 2.1877. Hyla spilomma Cope, Proc. Amer. Philos. Soc, vol. 17, no. 100, p. 86,July 20 (Cosamaloapam, State of Vera Cruz, Mexico).1882. Hyla nigropunctata Boulenger, Catalogue of the Batrachia Salientia s.Ecaudata in the collection of the British Museuna, 2d edit., p. 366 (Jalapa,State of Vera Cruz, Mexico).Type locality.?Indiis [incorrect = ? South America],Range.-?-From Cordoba in Vera Cruz westward to the Presidioin Sinaloa, and southward through Oaxaca and Tabasco to Centraland South America.Remarks.?Laurenti based his Rana venulcsa on the tree frogfigured by Seba " and states that its habitat is in "Indiis," which isobviously incorrect if subsequent authors are right in allocating thisname to the large tree frog that frequents South and Central America.According to Boulenger (op. cit., p. 366) the cotypes ofGunther 's Hyla Uchenosa were b-e, half-grown and young speci-mens from Vera Cruz;/, half-grown specimen from Cordoba (Sall6collection) ; and g, half-grown specimen from Mexico. These cotypesare described in my notes as follows
:
Four cotypes (B.M. Nos. 54. 11. 13. 14-17) from the collectionmade by Hugh Cuming and labeled as coming from Vera Cruz. Twohalf-grown and two young individuals comprise this series. Head-and-body length, respectively, 36.9, 35.6, 25, and 23.3 mm.; lateralwhite stripes from posterior edge of upper eyelid to thigh quiteprominent; black pustules on back; pustules on top of head the samecolor as rest of upperparts; brownish-black pustulose specks onexposed surfaces of fore and hind limbs ; vomerine teeth in transverserows at level of posterior margins of clioanae; fingers with vestigeof web at base; the hind limb being carried forward along the body,the tibio-tarsal joint reaches to center of eye; no tarsal fold; skin onabdomen and under surface of thighs coarsely granular.One cotype (B.M. No. 56. 3. 17. 24) from Cordoba, Vera Cruz,collected by Auguste Salle. Head-and-body length, 44.7 mm. ; fingers
61 Seba, A., Locupletissimi rerum naturalimn thesauri accurata descriptio, et iconibus artiflciossimisexpressio, per universam physices historiam, Amsterdam, vol. 1, p. 120, pi. 76, fig. 1, 1734.
MEXICAN TAILLESS AMPHIBIANS 177
with vestigial web at base; vomerine teeth in transverse rows betweenchoanae; other characters similar to preceding diagnosis.Two cotypes (B.M. No. 57. 10. 28. 49) from Mexico, a half-grownand a young individual, purchased from ?. Parzudald. Head-and-body length, 47.5 mm. ; the hind limb being carried forward along thebody, the tibio-tarsal joint reaches to between eye and tip of snout;vomerine teeth in transverse rows at level of posterior margins ofchoanae ; coloration faded (in alcohol) ; skin similar to other cotypes.Boulenger based his Hyla nigropunctata in part on these cotypesof H. lichenosa, which he states differ from H. venulosa in having thetympanum one-third the diameter of the eye, and in part on threeadditional specimens. One of the cotypes of this species (B.M.Nos. 81. 10. 31. 20) was collected at Jalapa in Vera Cruz by C. T.Hoege, a collector employed by F. D. Godman. The following noteswere made on this specimen: Head-and-body length, 62.8 mm.;transverse diameter of tympanum, 3.6 mm.; transverse diameterof eye, 5 mm.; anterior edge of eye to nostril, 5.8 mm.; the hind limbbeing carried forward along the body, the tibio-tarsal joint reachesto anterior margin of eye; throat, chest, and abdomen, and undersurface of thighs coarsely areolate; vomerine teeth in transverserows at level of posterior margins of choanae.The two remaining cotypes (B.M. No. 59. 9. 20. 2), a half-grownand a young individual, have a general coloration (in alcohol) thatis very similar to Giinther's Vera Cruz specimens. The larger indi-vidual has a head-and-body length of 42 mm. ; the hind limb beingcarried forward along the body, the tibio-tarsal joint reaches toanterior margin of eye; and vomerine teeth are in transverse rows atlevel of posterior margins of the choanae.Giinther ?^ has already remarked that "this species shows a some-what perplexmg amount of variation in the shape of the vomerineseries, size of the tympanum, development of cuticular glands, andcoloration." Topotype specimens agree with the original descriptionof lichenosa and are sufficiently like Hyla venulosa to be consideredthe same species. Giinther placed Boulenger's H. nigropunctata inthe synonymy of //. venulosa.The type of Cope's Hyla spilomma, which was collected by FrancisSumichrast and is now lost, seems to have been a young individualwith fingers united at base by a vestigial web. Another young speci-men collected at Santa Teresa m Nayarit (U.S.N.M. No. 46829) hasa small tympanum, and the vomerine teeth are abnormal in that theyare located between the choanae.A living specimen collected at Presidio in Sinaloa by AlphonsoForrer is described by Boulenger ?^ as follows
:
M Gunther, A. C. L. Q., Biologia Centrali-Americana, ReptUia and Batrachia, p. 273, Sept., 1901.
?3 Boulenger, O. A., Description of a new genus and species of frogs of the family Uylidae. Ann. and Mag.Nat. Hist., ser. 5, vol. 10, no. 58, pp. 327, 328, Oct., 1882.
178 BULLETIN 160, UNITED STATES NATIONAL MUSEUMUpper surfaces light brown, with large chestnut-brown spots; these occupyentirely the hinder part of the back and the hind limbs; upon the latter the lighterground-color appears in the form of transverse lines; on the flanks the spots arebordered with pure white; lower surfaces of a rather dirty white, the throat withbrown vermiculations. The eye is very beautiful, the iris being golden with blackreticulations, and a vertical and a horizontal black bar, forming a cross. Thevocal bladders are black; they can not be retracted, as in the Ranae with externalvocal sacs; and when empty they hang on each shoulder like a cutaneous lobe.As is well known, the bones of this frog are of a beautiful "vert de gris"; this coloris seen on the vomerine groups and on the borders of the lower jaw when themouth is open.The attitudes and movements of H. venulosa are much the same as those of itsEuropean congener; but it is entirely noctural, remaining concealed the whole ofthe day. It is by no means shy, but, when handled, exudes a great quantity ofpoisonous fluid, more so than any Batrachian I have liad before. Besides, thisfluid, of a milky appearance, coagulates instantaneously, sticking to the fingersin a very disagreeable manner; it has a strong odor, resembling that of peaches,and afifects very disagreeably the mucous membrane of the nostrils, causing astrong itching.Specimens forwarded to museums are rarely accompanied by fieldobservations, and it is therefore of interest that Frederick Knabnoted that a specimen taken at Cordoba was found in tillandsia.A description of the gliding position assumed by the frog when it isfalling is given by Cott ^*
:
A preliminary experiment was made by launching one of the frogs from off theveranda of the house, whence it had a faU of about 25 feet on to the hard floorof the courtyard below. The little creature spread its arms and legs out widelyas it fell, and landed the "right way up"; it appeared to be quite unhurt.The animal was next flung high upwards, from the ground, and I was surprisedand delighted to find that it invariably managed, by means of a violent wriggle,to establish itself in a definite balanced position, so that it always fell belly down-wards, with its limbs stretched in a constant and characteristic attitude. Manytimes it was thrown in such a manner that it spun over and over on leaving thehand, but it never once failed to get into what I shall call its "gliding position"before reaching the culminating point of the trajectory, and this position, onceassumed, was invariably maintained without a visible movement until the impactof landing.At this early stage in the experiments, it was noticed that the frog appeared tofall much more slowly than might have been expected, and it was surprising tofind that the impact of hitting a hard stone floor after a fall of some 35 or 40 feetdid not in any way hurt the creature. * * *On the 22d of October I set out in the direction of Souza, a suburb of Para,accompanied by a small Barbadian boy, named Geraldo, and taking with me somespecimens of H. venulosa. At Souza there is a high water tower, and up thisGeraldo climbed with instructions to release the frog from the top, on a givensignal, while I remained below to pick up the pieces. The tower is not less than140 feet high, and it is situated in an open space where the ground is hard andcovered at this time of the year by a scanty growth of grass.I stood near the foot of the tower, expecting the frog to fall nearly, if not quite,vertically. But on a wave from my handkerchief, I saw the little creature leap
?* Cott, H. B., Observations on the life habits of some batrachians and reptiles from the Lower Amazon.Proc. Zool. Soc. London, 1926, pt. 4, pp. 1164, 1165, Dec, 1926.
MEXICAN TAILLESS AMPHIBIANS 179into the air, immediately spread out its limbs, and sail slowly down in fine style,passing far over my head and landing at a spot 90 feet behind me. It was quiteuninjured. (There was a slight breeze blowing at the time in the same directiontoward which the frog glided, but not enough, I fancy, to account for more thana few yards of lateral movement.)The experiment was repeated, and it was found that the frog took 7% secondsto reach the ground. On this occasion I was at the top of the tower, and was ableto observe that after it had dropped the first 12 feet up [sic] or so, there was,apparently, no further acceleration in the speed of descent; for at this point astate of equilibrium is reached in which the upward pressure of the air on theventral surface of the frog is equal to the animal's weight.Specimens examined.?Nineteen, as follows
:
Hyla venulosa
Museum
M.C.Z.-.-U.S.N.M..DoB.MDoA.M.N.H.M.C.ZB.MU.S.N.M..Do.-.A.M.N.H.Do.-.-B.MDo....
CatalogueNo.
8375382643830356.3.17.2481.10.31.206300976154. 11. 13.14-174682946917134491384157.10.28.4959. 9. 20. 2
Num-ber ofspeci-mens Locality collectedVera Crtjz:Cerro del QalloCordoba;""]do"-i"iii"i'iiii""i^ii!JalapaOne mile south of NewCemetery.PanucoNo definite locality1 Nayarit: Santa Teresa.1 Oaxaca: Tuxtepec
I ?State:1 t No definite locality...1 I do... -.-2 do2 do
Date collected
1921Mar. 21, 1908Mar. 25, 1908June 16,1919Apr. 10,1923Aug. 9, 1897Apr. 11,1894
By whom collected orfrom whom received
E. R. Dunn.Frederick Knab.Do.Auguste Sallf.C. T. Hoege.Paul D. R. Ruthling.W. W. Brown.Hugh Cuming.Nelson and Goldman.Do.PaulD.R.Ruthling.
. Do.E. Parzudaki.
> Cotypes of Hyla lichenosa. ' Cotypes of Hyla nigTopundaia.Genus HYLELLA Reinhardt and Liitken1862. Hylella Reinhardt and Ltjtken, Vidensk. Meddel. Nat. Forening, 1861,Kj0benhavn, pt. 1, p. 199. [Included species: Hylella tenera Reinhardt andLtJTKEN, idem, p. 200; and Hylella pundatissima Reinhardt and Ltjtken,idem, p. 200, pi. 4, fig. 5 (from vicinity of Lagoa Santa, Brazil).]1879. Exerodonta Brocchi, Bull. Soc. Philom. Paris, ser. 7, vol. 3, p. 20. [Geno-type, Exerodonta sumichrasti Brocchi, idem, p. 20 (Santa Efigenia, Mexico) .]Two species of Hylella are known to occur in Mexico, and theydiffer from all other Mexican Hyhdae in the absence of vomerineteeth. These diminutive tree frogs must escape the notice of collec-tors, since they are rarely found in collections made in that country.Very few specimens of either of these species have thus far foundtheir way into American museums. The eastern form, Hylella pida,is the most distinctively marked of the two Mexican species. Thepresence or absence of vomerine teeth in the young of some species ofHyla is sometimes difficult to determine without dissection, and theyoccasionally may be confused mth adult Hylella.
180 BULLETIN 160, UNITED STATES NATIONAL MUSEUMKEY TO MEXICAN SPECIES OF HYLELLA1. No tarsal fold; a narrow white stripe bordered below by darkerstreak commences at tip of snout and extends backward oncanthus rostralis, across eyelid, and above tympanum,becoming fainter as it approaches the groin; upper surfaceslight purplish, unicolored, or with scattered, round, darkpurple light-edged spots of the size of the tympanum; undersurfaces whitish; limbs mottled, witli concealed portionspigmentless; skin of upperparts smooth; underparts, withexception of gular region, granulated; tympanum slightly morethan one-third diameter of eye; no vomerine teeth; tonguenearly circular, scarcely notched behind; canthus rostralisangular; loreal region subvertical; fingers with vestigialbasal web; toes three-fourths webbed, the membrane extend-ing to disks of the third and fifth toes and leaving free thelast phalanx of the fourth; adhesive disks smaller than or aslarge as the tympanum; the hind limb being carried forwardalong the body, the tibio-tarsal joint reaches between the eyeand the end of the snout; head-and-body length, 24 mm picta (p. 180)A tarsal fold present; no lateral white stripe; upper surfaces ashcolored; under surfaces yellowish or light orange; no markingson sides or on concealed surfaces of limbs, or on superiorsurfaces of limbs; skin of upperparts smooth; underparts,including gular region, granulated; tympanum one-third toone-half diameter of eye; no vomerine teeth; tongue circular,scarcely notched behind; canthus rostralis distinct, obtuse;loreal region subvertical; fingers one-third webbed; toes three-fourths webbed, the membrane reaching the disks of thethird and fifth toes and leaving free the last phalanx of thefourth; adhesive disks as large as the tympanum; the hindlimb being carried forward along the body, the tibio-tarsaljoint reaches the eye; head-and-body length, 28-32 mm-sumichrasti (p. 181)HYLELLA PICTA Ganther1901. Hylella picta Gunther, Biologia Centralia-Americana, Reptilia and Ba-trachia, pp. 286, 287, pi. 73, fig. c, Sept.Type locality.?Jalapa, State of Vera Cruz, Mexico.Range.?From Jalapa in Vera Cruz southward to Frontera inTabasco; not known to occur on the Pacific drainage area.Re7narks.?The type of this species (B.M. No. 1901. 12. 19. 100)was collected by Mateo Trujillo at Jalapa in Vera Cruz. It is de-scribed in my notes as follows: Head-and-body length, 24.1 mm.;transverse diameter of tympanum, 1 mm.; transverse diameter ofeye, 2.3 mm.; anterior edge of eye to nostril, 2.2 mm.; the hind limbbeing carried forward along the body, the tibio-tarsal joint reaches tothe anterior margin of the eye; no tarsal fold; large inner but no outermetatarsal tubercle; fingers with vestige of web at base; no vomerineteeth; canthus rostrahs rounded; loreal region subvertical; upper-parts (in alcohol) hght purplish red, with somewhat widely scatteredblack dots.
MEXICAN TAILLESS AMPHIBIANS 181A notation on the label accompanying the small series in the UnitedStates National Museum indicates that Doctor Townsend found thistree frog on vegetation in the woods near Frontera in Tabasco.Specimens examined.?Nine, as follows:Hylella picta
Museum
182 BULLETIN 160, UNITED STATES NATIONAL MUSEUMbase; skin on throat, abdomen, and under surface of thighs coarselygranulated.It has been reported that the eggs of this tree frog are deposited inthe axils of the leaves of Spanish beard (Tillandsia) , where theyundergo their whole metamorphosis.Specimens examined.?Six, as follows:Hylella sumichrasti
Museum
MEXICAN TAILLESS AMPHIBIANS 183
vocal vesicle large; some vinaceous spots behind angle ofmouth; the fore limb being carried forward along the head,the wrist reaches beyond the tip of the snout; fingers moder-ately elongated?in order of increasing lengths: 1, 2, 4, and 3;the hind limb being carried forward along the body, the tibio-tarsal joint reaches to front of scapula or between shoulder andeye; hind limb stout; toes with blunt tips and distinct subartic-ular tubercles; two very prominent, oval, compressed metatarsaltubercles, the inner sublongitudinal and protuberant, theouter subtransverse; skin perfectly smooth or irregularlyroughened by tiny asperities; a dermal fold extends acrosshead behind the eyes; body length, snout to vent, 28 to 40 mm.Hypopachus variolosus (p. 184)Toes free, without vestigial web at base; fingers free, blunt orslightly dilated at tips; terminal phalanges simple; precora-coids absent; coracoids united by a simple cartilage; no omo-sternum; sternum cartilaginous; underparts yellowish or brown-ish, stippled or marbled with brown Gastrophryne (p. 186). _22. A prominent outer mietatarsal tubercle and a somewhat largeroval inner metatarsal tubercle; snout acuminate, moderatelyprojecting, and not twice as long as the diameter of the eye;the hind limb being carried forward along the body, the tibio-tarsal joint reaches the shoulder or slightly beyond; toes quitefree, with blunt tips and distinct subarticular tubercles; skinsmooth or with small tubercles on back; a more or less dis-tinct dermal fold across the head behind the eyes; upperpartsolive or brownish; generally a broad light band along eachside of the body, and a narrow vertebral line; limbs lighter incolor; thigh and leg each with a broad dark cross band; awhite line from angle of mouth to insertion of fore limb; malewith subgular sac; throat deep black; body length, snout tovent, 26 to 28 mm Gastrophryne usta (p. 187)No outer metatarsal tubercle; inner metatarsal tubercle verysmall, indistinct; snout pointed, projecting, and twice as longas the diameter of the eye; the hind limb being carried for-ward along the body, the tibio-tarsal joint reaches in front ofthe shoulder; toes quite free, with blunt tips and distinct sub-articular tubercles; skin perfectly smooth; a dermal foldacross the head behind the eyes; upperparts rose colored orlight brown; back with an irregular broad dark band; sidesof head and body dark brown; hip with a large dark-brownround spot; thigh, leg, and tarsus each with a dark-edgedcross band; belly marbled with brown; throat brown, mar-bled with white; body length, snout to vent, 32 mmGastrophryne elegans (p. 187)Genus HYPOPACHUS Keferstein1867. Hypo-pachus Keferstein, Nachr. konigl. Ges. Wiss. Univ. Gottingen,no. 18, p. 351, July 24. [Genotj^pe, Hypopachus seebachii Kefekstein,idem, p. 352 (Costa 'R,icsL)?Engystoma variolosum Cope, 1866, Proc. Acad.Nat. Sci. Philadelphia, vol. 18, p. 131, May (Arriba, Costa Rica).]Three species of Hypopachus?cuneus, inguinalis, and variolosus?are now recognized. The Texan species, H. cuneus, has been taken
184 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
at Brownsville, Tex., and future collecting may show that it occursalso south of the Rio Grande in the State of Tamauhpas, Thisspecies is not Usted in the key since there are no Mexican records.H. variolosus seems to be restricted in Mexico to the Pacific coast,where it has been found as far north as southern Sinaloa. A quitedifferently marked and much brighter-colored species, H. inguinalis,is known to occur as far north as the Lake Peten district of northernGuatemala. This species may possibly occur also in the adjoiningstates of Mexico. The habits and life histories of these narrow-mouthed toads are not on record. All three have loose leatheryskins, and they may hide, as does Gastrophryne, under old logs andother debris. HYPOPACHUS VARIOLOSUS (Cope)1866. Engystoma variolosum Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 18,p. 131, May.1867. Hypopachus seebachii Keferstein, Nachr. konigl. Ges. Wiss. Univ. Gottin-gen, no. 18, p. 352, July 24 (Costa Rica).1883. Hypopachus oxyrrhinus Boulenqer, Ann. and Mag. Nat. Hist. ser. 5, vol.11, no. 65, p. 344, May (Presidio near Mazatlan, Sinaloa).Type locality.?Arriba, Costa Rica.Range.?From near Mazatlan in Sinaloa southward through Jaliscoto Costa Rica.Remarks.?The type specimen of Cope's Engystoma variolosum(U.S.N.M. No. 6486) was collected at Arriba in 1863 by Charles N.Riotte, who was at that time the United States minister to CostaRica. It is fairly well preserved, although the original color patternhas been somewhat obscured by the preservative and the light, andthe left hind leg is missing below the knee. The measurements ofthe type are as follows: Head-and-body length, 35.5 mm.; transversediameter of eye, 3.2 mm.; anterior edge of eye to nostril, 2.4 mm.;width of head at level of posterior angles of jaws, 9.8 mm.Hypopachus seebachii, which was collected by Professor von Seebachduring his travels in Costa Rica in 1865, was recognized as a newgenus and species by Peters. Two years later. Cope ?^ pointed outthat H. seebachii was a synonym of his H. variolosus. Boulengerthought that specimens collected by Alphonso Forrer at Presidionear Mazatlan in Sinaloa represented a new species, which he namedHypopachus oxyrrhinus. The following notes relate to these twocotypes: Smaller individual (B.M. No. 1882. 12. 5. 8): Head-and-body length, 27.5 mm.; transverse diameter of eye, 3.3 mm.; ante-rior edge of eye to nostrU, 1.9 mm.; the hind limb being carried for-ward along the body, the tibio-tarsal joint reaches to barely beyond
" Cope, E. D., Seventh contribution to the herpetology of Tropical America. Proc. Amer. Philos.Soc, vol. 11, no. 82, p. 167, July 16, 1869.
MEXICAN TAILLESS AMPHIBIANS 185the axilla; side of head below canthus rostralis and shoulder dark;dark cross bands on hind limbs, and when femur, tibia, and tarsusare folded back at right angles to long axis of body, the ends of thesedark crossbars meet and simulate a continuous band. The othercotype (B.M. No. 1883. 4. 5. 3) does not differ in any essential detailfrom the preceding specimen and has a body length, snout to vent,of 29.8 mm.The key prepared by Cope ^^ in 1889 to distinguish oxyrrhinus fromvariolosus apparently was incorrectly drawn up from memory, for theheel of oxyrrhinus does not reach to end of the muzzle, and the sidesof variolosus are blackish brown anteriorly, as stated in the originaldescription. In describing oxyrrhinus, Boulenger observes that theheel or tibio-tarsal joint reaches to between the shoulder and the eye.The distinctions made by Cope do not exist. Giinther ^^ correctlyplaced H. seebachii and H. oxyrrhinus in the synonymy of //. variolosus.A juvenile specmien (M.N.B. No. 7585) collected by AdolpheBoucard in Mexico is described in my notes as follows: Head-and-body length, 20 mm.; the hind limb being carried forward along thebody, the tibio-tarsal joint reaches to the eye; very large subarticulartubercles on fingers and toes; inner metatarsal tubercle quite large,the outer one smaller, but larger than the subarticular tubercles;skin of underparts minutely areolate; a quite narrow dark interor-bital stripe; remainder of upperparts (in alcohol) grayish; a blackstripe along canthus rostralis from tip of snout to eye, and from eyeto above tympanum and to behind the latter on the sides; upperanterior surface of thighs pink; dark cross bands on hind limbs indis-tinct; underparts dark, with numerous closely spaced minute lightspots.Specimens examined.?Seven, as follows:Hypopachus variolosus
Museum
186 BULLETIN 160, UNITED STATES NATIONAL MUSEUMGenus GASTROPHRYNE ?8 Fitzinger1828. Microps Wagler, Isis von Oken, vol. 21, p. 744. [Genotype, Micropsunicolor Wagler, idem, p. 744= [7?ana] ovalis Schneider, 1799, Historiaeamphibiorum, naturalis et literariae, fasc. 1, p. 131; Microps Wagler pre-occupied by Microps Megerle, 1823, Coleoptera.]1838. Stenocephalus Tschudi, M6m. Soc. Sci. Nat. de Neuchatel, vol. 2, pp. 49,86. [Genotype, Microps unicolor Wagler, 1828, Isis von Oken, vol. 21,p. 744= [iJana] ovalis Schneider, 1799, Historiae amphibiorum naturalis etliterariae, fasc. 1, p. 131; Stenocephalus Tschudi preoccupied by Stenocepha-lus Latreille, 1825, Hemiptera.]1841. Engystoma Dum^ril and Bibron, Erpetologie g^nerale, vol. 8, pp. 738,740. [Genotype, [Rand\ ovalis Schneider, 1799, Historiae amphibiorum,naturalis et literariae, fasc. 1, p. 131.] [Nee Engystoma Fitzinger, 1826=Breviceps Merrem, 1820.]1843. Gastrophryne Fitzinger, Systema reptilium, fasc. 1, p. 33. [Genotype,Engystoma rugosum Dumeril and Bibron, 1841, Erpetologie gen^rale, vol. 8,p. 744 (des parties miSridionales de I'Am^rique du Nord) = Engystomacarolinense Holbrook, 1836, North American herpetology, ed. 1, vol. 1, p. 83,pi. 11 (Charleston, S. C.).]Tliirteen species of tlie genus Gastrophryne were enumerated byDoctor Stejneger in 1910, the northernmost being the Uttle G. caroli-nensis of Southeastern United States and the southernmost Boulen-ger's G. leucosticta from Santa Catharina, Brazil. The 12 Americanspecies form a natural assemblage with a rather widespread range, andhence the occurrence of Boulenger's G. borneensis in Borneo has longproved a puzzling obstacle to zoogeographers. This record has oftenbeen cited as one of the remarkable instances of discontinuous distri-bution in the Amphibia. A restudy of the type of G. borneensis in theBritish Museum of Natural History has been made by Doctor Noble,^^who finds that the internal anatomy of the Bomean species differsfrom that of the American forms in a number of important details.He has shown that although G. borneensis resembles the species ofGastrophryne externalh'^, the structural features indicate that it is notclosely related to the American species, and be therefore proposes anew generic name, Gastrophrynoides, for the species that occurs inBorneo.There are in American institutions relatively few specimens of nar-row-mouthed toads from Mexican localities, and lack of adequatematerial, especially from critical areas, has made it rather difficult todraw any definite conclusions as to the number of species of this genusthat may be expected to occur within the boundaries of the MexicanRepublic. It is possible that Strecker's G. areolata and Girard's G.texensis may occur in northern Mexico, particularly south of and along
?' For use of the name Qastrophryne in place of Engystoma, see Stejneger, L., Proc. Biol. Soc. Washington,vol. 23, pp. 165-167, Dec. 29. 1910.?? Noble, Q. K., An analysis of the remarkable cases of distribution among the Amphibia, with descrip-tions ofnew genera. Amer. Mus. Nov., no. 212, pp. 20-23, 1926.
MEXICAN TAILLESS AMPHIBIANS 187the Rio Grande. No intensive herpetological collecting has beencarried on in recent years in the States of Tamanlipas and Chihuahua.Very little has been published in regard to the habits and hfe his-tories of these httle narrow-mouthed toads. The Texan species areknown to hibernate in small groups under rotten logs and under pilesof rubbish. During the breeding season, it frequents pools of stag-nant water and may be observed floating on the surface, if oneapproaches quietly. Doctor Wright has published observations onthe eggs ^? and the tadpoles ^^ of two species found within the Umits ofthe United States. GASTROPHRYNE ELEGANS (Boulenger)1882. Engystoma elegans Boulenger, Catalogue of the Batrachia Salientia s.Ecaudata in the collection of the British Museum, 2d edit., p. 162.
?
GtJNTHER, 1900, Biologia Centrali-Americana, Reptilia and Batrachia, p. 210,pi. 62, fig. D, Feb.1910. Gastrophryne elegans Stejneger, Proc. Biol. Soc. Washington, vol. 23,p. 166, Dec. 29.Type locality.? Cordoba, State of Vera Cruz, Mexico.Range.?Not known to occur elsewhere than at the type locality.Remarks.?An adult female narrow-mouthed toad collected byAuguste Salle at Cordoba in Vera Cruz and previously identified byGiinther ^^ as Engystoma carolinense was found by Boulenger to lackthe outer metatarsal tubercle. This single character is the basis forthe recognition of this species. The type, so far as known, remainsunique. Not until a series of specimens from the type locality andvicinity is available for study, will it be possible to determine whetherthe outer metatarsal tubercle is normally absent in narrow-mouthedtoads of that region.Specimens examined.?None.GASTROPHRYNE USTA (Cope)1866. Engystoma ustum Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 18, p. 131,May.
?
Brocchi, 1882, Mission scientifique au Mexique et dans I'Am^riqueCentrale, recherches zoologiques, pt. 3, sect. 2, p. 94, pi. 10, fig. 1.1869. Engystoma mexicanum Peters, Monatsber. k. preuss. Akad. Wiss. Berlin,p. 881, Dec. (State of Puebla, probably Matamoros).1903. Eupemphix gadovii Boulenger, Ann. and Mag. Nat. Hist., ser. 7, vol. 12,p. 552, Nov. (San Mateo del Mar, Tehuantepec)
.
1910. Gastrophryne usta Stejneger, Proc. Biol. Soc. Washington, vol. 23, p. 166,Dec. 29.Type locality.?Guadalaxara [ = Guadalajara], State of Jalisco,Mexico.
'" Wright, A. H., and Wright, A. A., A key to the eggs of the Salientia east of the Mississippi River.Amer. Nat., vol. 58, no. 657, p. 378, 1924." Wright, A. H., Synopsis and description of North American tadpoles. Proc. U. S. Nat. Mus., vol. 74,no. 2756, pp. 2, 3, 13-16; pi. 6, fig. 3; pi. 6, fig. 1, June 4, 1929." Giinther, A. C. L. O., Catalogue of the Batrachia Salientia in the collection of the British Museum,pp. 51, 52, 1858.
188 BULLETIN 160, UNITED STATES NATIONAL MUSEUMRange.?From southern Sinaloa southward through San LuisPotosi and Jahsco to Tehuantepec in Oaxaca and southern VeraCruz (Minatitlan).Remarks.?The type of Cope's Engystoma ustum (U.S.N.M. No.24965) is partially macerated, the fleshy portions are largely dis-integrated, and the skeleton is more or less disarticulated. J. J.Major, the collector of the type, in 1861 sent a collection of reptilesto the Museum, and this type was presimiably included in the ship-ment, since no other accessions are credited to him.The three cotypes of Eupemphix gadorii (B. M. Nos. 1903. 9. 30.259-261) were obtained by Dr. and Mrs. Hans Gadow at San IMateodel Mar in Oaxaca. H. W. Parker, of the British Museum of NaturalHistory, has recently examined the cotypes and reports that they arenarrow-mouthed toads belonging to the genus Gastrophryne and thatalthough very young they seem to be referable to the species usta.According to my own observations, all these cotypes are very youngindividuals, the head-and-body length of the largest being 12.4 mm.;one has a Ught vertebral stripe; an inner metatarsal tubercle present;a light silvery streak extends along canthus rostrahs from nostril toeye and is continued backward from eye along side to near level ofanterior end of sacrum as a very irregular wavy band.In the introduction to the report on the herpetological specimenscollected by Berkenbusch in Puebla, Peters states that they weretaken in the vicinity of Matamoros and at other localities. Nospecific type locaUty is given, except that of Puebla in the title of thepaper, for his supposedly new species, Engystoma mexicanum. Cope ^^seems to have been the first to recognize that Peters's species wasidentical with usta, but nevertheless the last-mentioned species wasmistakenly allocated to the genus Systoma.Specimens examined.?Twenty, as follows:Gastrophryne usta
Museum
MEXICAN TAILLESS AMPHIBIANS 189Family RANIDAE Bonaparte1831. Ranidae Bonaparte, Giorn. Arcad. Sci. Lett, ed Arti, Roma, vol. 49,pp. 65, 75.The members of the Ranidae have a wide distribution in temperateand tropical regions, but are not foimd in Australia, Tasmania, NewZealand, eastern Polynesia, or Hawaii. Frogs are generally thoughtof as being largely aquatic, but there are some with large digital disksthat are arboreal in habit. Others, like the North American woodfrog {Rana sylvatica), prefer a land life and are generally found on theground in the woods. As ^vith the true tree frogs, there are a fewspecies that burrow in the ground. AU the Mexican species of thegenus Rana are more or less aquatic. One Mexican species, R. monte-zumae, is commonly sold for food in the markets of Mexico City.The Ranidae have maxillary but no mandibular t?eth, a tirmi-sternal type of shoulder girdle, cylindrical or slightly expanded sacraldiapophj-ses, and a biconvex sacral vertebra with double condyle forthe coccyx. The tongue of the ranids differs from the type found inother families of tailless amphibians in that it is provided vrith. twolateral posterior horns. In rare instances ranas have been foundwith a nearly trilobed tongue. Frogs of the genus Oxyglossus havea single median horn on the posterior free end of the tongue.Genus RANA Linnaeus1758. Rana Lixxaeus, Systema naturae, ed. 10, vol. 1, p. 210. [Genotype,[Rana] temporaria Lixxaetts, p. 212 (Europe); based upon Linnaeus, 1746,Fauna Svecica, p. 94.]1814. Ranaria Rafixesque, Specchio Sci., Palermo, vol. 2, fasc. 7. [Substitutename for Ra7ia Linnaeus.]1843. Pelophylax Fitzinger, Systema reptLlium, fasc. 1, p. 31. [Genotj-pe,Rana esculenta Linnaeus, 1758, Systema naturae, ed. 10, vol. 1, p. 212.]1859. Ranula Peters, Monatsber. k. preuss. Akad. Wiss. Berlin, p. 402, May.[Genotype, Ranula gollmerii Peters, idem, p. 402 (Caracas. Venezuela).]1867. Pohlia Steindachner, Reise der Oesterreichischen Fregatt^ Novara, Zool-ogischer theil, vol. 1, Amphibien, p. 15. [Genotype, Pohlia palmipes Stein-dachner, idem, p. 15, pi. 1, figs. &-S? Rana palmipes Spix, 1824, Animalianova, sive species novae testudinum et ranarum quas in itinere per Bra-siliam, etc., p. 5, pi. 5, fig. 1 (Amazon River, Brazil).]1868. Trypheropsis Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 20, p. 117.[Genotype, Trypheropsis chrysoprasinus Cove? Ranula chrysoprasina Cope,1866, Proc. Acad. Nat. Sci. Philadelphia, vol. 18, p. 129 (Arriba, Costa Rica).]1891. Crotaphitis Schulze, Jahresber. Naturw. Ver. Magdeburg, 1890, p. 176.[Genotype, Rana temporaria Linnaeus.]1891. Baliopygus Schulze, Jahresber. Naturw. Ver, Magdeburg, 1890, p. 177.[Genotype, [Rana] esculenta Linnaeus, 1758, Systema naturae, ed. 10, vol. 1,p. 212; based upon " Ranae viridis aquaticae" Roesel von Rosenhop, 1758,Historia naturalis ranarum nostralium, p. 53, pi. 13 (Europe).]66785?32 13
190 BULLETIN 160, UNITED STATES NATIONAL MUSEUMWith the exception of the generic names that are based on Linnaeanspecies from Europe, the synonyms of the genus Rana here enumeratedare restricted to those based on New World species.There are at least six distinct species of the genus Rana on conti-nental Mexico. The largest of these is the well-known North Ameri-can bullfrog {R. catesbeiana) , which is here for the first time recorded inMexico. The wide-ranging leopard frog {R. pipiens) seems to be themost abundant rana in Mexico. One South American species, R.palmipes, reaches its northern limit in Vera Cruz. A Mexican relative,R. tarahumarae, of the Pacific coaSt yellow-legged frog (R. boylii) hasbeen found in the high sierras of western Chihuahua. R. montezumaeundoubtedly possesses the most peculiar color pattern of all Mexicanranas, and notwithstanding its characteristic appearance nothing hasbeen published in regard to its life history, and all that is known of itshabitat is that it frequents the lakes in the Valley of Mexico. Therarest of all Mexican frogs in collections is one which Boulenger namedRana pustulosa.All known species of the genus Rana, except one, deposit their eggsin the water, where they develop into tadpoles. The single exceptionof this general habit is a frog (R. opisthodon) that occurs on theSolomon Islands, and this species lays its eggs on land in moist crevicesof rocks close to water. These eggs develop directly into frogs with-out any aquatic tadpole stage. The regular larval stage and subse-quent metamorphosis are hurried through within the egg. Tho buccalapparatus of the larvae of these ranas is quite characteristic for eachspecies, and many of them can be recognized in life by their colorpattern. The mouth disk is devoid of horny teeth in some species.In others there are more than the normal number of rows. The tad-pole of R. catesbeiana may have as many as three rows of horny teethin the upper and in the lower jaws.KEY TO MEXICAN SPECIES OF RANA1 . No trace of dorso-lateral glandular dermal folds 2Dorso-lateral glandular dermal folds present 3
*?. Tympanum distinct, with or without small asperities, its trans-verse diameter two-fifths to one-half that of eye; upperparts,including top of head, brown or olive, marked with irregularblotches, or small black spots, or numerous dots; limbs withirregular dark crossbars; abdomen white; throat and chesteither white or mottled with brown; limbs mottled with grayishbrown; skin of upperparts either smooth or with smallpustules; a feeble curved glandular fold from eye to shoulder,but no dorso-lateral glandular fold; the hind limb being carriedforward along the body, the tibio-tarsal joint reaches to ornearly to end of snout; tarsal fold distinct or indistinct; distancefrom snout to vent 1% to 2 times length of tibia; tips of toesswollen into small disks; a broad web between the toes; outermetatarsals separated nearly to base; inner metatarsal tubercle
MEXICAN TAILLESS AMPHIBIANS 191
elliptic, feebly prominent, one-third or more the length of innertoe; no outer metatarsal tubercle; subarticular tubercles largeand prominent; tips of fingers feebly swollen; first finger longerthan second; males with thick blackish pad on inner side offirst finger; no external vocal vesicles; vomerine teeth in twosmall groups within and behind level of choanae; head broaderthan long and much depressed; snout rounded, as long astransverse diameter of eye, and barely projecting beyondmouth; canthus rostralis indistinct; loreal region very oblique,slightly concave; nostril equidistant from end of snout and fromeye; interorbital width equal to or a little less than that ofupper eyelid; head-and-body length of adults, 45 to 78 mm.tarahumarae (p. 214)Tympanum very distinct, its transverse diameter nearly equalto that of eye in females and greater than the latter in males;upperparts brown or olive-brown, rarely green, uniform orspotted, or marbled with dark brown; limbs with or withoutdark cross bands; hinder side of thighs often marbled withblack and yellow; chin yellow; rest of underparts white, some-times marbled with brown, or spotted with white and yellow;skin of upperparts fairly smooth and leathery, with scatteredwarts; a strong glandular fold from eye to shoulder, but nodorso-lateral fold; sides often with pustulose warts; the hindlimb being carried forward along the body, the tibio-tarsaljoint reaches to eye or tympanum; no tarsal fold; distance fromsnout to vent 2 to 2}^ times length of tibia; tips of toes slightlyswollen; toes fully webbed, with exception of distal joint offourth; outer metatarsals separated to base; inner metatarsaltubercle elliptic, feebly prominent, one-third or more thelength of inner toe; noouter metatarsal tubercle; subarticulartubercles small; tips of fingers pointed or obtuse; first fingeras long as or a little longer than second; males with retract-ing pad on inner side of first finger; internal vocal vesiclespresent; vomerine teeth in small clusters or short oblique rowsbetween or rarely behind level of choanae; head broader thanlong and much depressed; snout rounded, as long as or alittle longer than transverse diameter of eye, and clearlyprojecting beyond mouth; canthus rostralis indistinct; lorealregion very oblique, slightly concave; nostril equidistant fromend of snout and from eye, or a little nearer the former; inter-orbital width varies from one-half to nearly once that of uppereyelid; head-and-body length of adults, 100 to 200 mm.catesbeiana (p. 195)Ground color of upperparts as well as the dark spots stippled ordotted with closely approximated minute white spots or specks;underside of thighs occasionally reddish, but likewise coveredwith small white dots; upperparts purplish brown or olive-brown, uniform or speckled, vermiculated, or mottled withlighter and darker, with or without large rounded or elongatedocellated dark brown spots, which may be grass green in life andare often light edged or encircled with ring of white dots;tympanum occasionally stippled with white specks; limbs withor without dark cross bands; underparts grayish white, ash,dirty yellow, or brownish, conspicuously spotted, blotched, orvermiculated with spots and sinuations of white; anterior, pos-terior, and inferior surfaces of thighs marked with numerous
192 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
white dots and blotches; skin of back between dorso-lateralglandular dermal folds smooth or granular, and with smallwarts; a narrow or moderately broad, feebly prominent, dorso-lateral glandular dermal fold from above tympanum to sacralregion or a little beyond, sometimes interrupted, and occasion-ally absent; sides with flattened warts; underparts generallysmooth; abdomen and thighs sometimes granular; a broadglandular fold from below tympanum to above arm; uppersurface of tibia with or without longitudinal glandular ridges;fingers long and slender, pointed, and sometimes bordered bynarrow dermal fold; first finger usually as long as second, occa-sionally slightly longer or slightly shorter; the hind limb beingcarried forward along the body, the tibio-tarsal joint reachesto tympanum or eye or bej^ond; distance from snout to vent2 to 2% times length of tibia; toes rather long, pointed, webbedto tips or with last joint of fourth free; outer metatarsals sep-arated nearly to base; subarticular tubercles small, feeblyprominent; tarsal fold feeble, but often a distinct ridge; innermetatarsal tubercle elliptic or subtriangular and prominent,one-fourth to one-third the length of inner toe; no outermetatarsal tubercle; width of head at level of posterior anglesof jaws greater than distance between end of snout and poste-rior margin of tympanum; snout rounded, scarcely projectingbeyond mouth, as long as or a little longer than transversediameter of eye; canthus rostralis indistinct; loreal region veryoblique, scarcely concave; nostril equidistant from eye andfrom end of snout or a little nearer the former; interorbitalwidth less than that of upper eyelid; tympanum moderately orvery distinct, two-thirds to once the diameter of the eye; vomer-ine teeth in small transverse or oblique groups between or barelybehind posterior borders of choanae; male with a large blackishexternal vocal vesicle on each side, forming loose folds frombehind angle of mouth to shoulder; head-and-body length ofadults, 100 to 145 mm montezumae (p. 197)Ground color of upperparts and underparts never stippled orspotted with white dots 44. Skin of back between dorso-lateral glandular dermal folds smoothor granular, with more or less conspicuous elongated wartsor interrupted glandular ridges; a prominent, narrow ormoderately broad glandular dorso-lateral dermal fold, extend-ing backward from upper eyelid at most to level of acetabulum,but generally interrupted or else ending at about level ofmiddle of sacrum; a glandular fold from below eye to vocalvesicle or to above arm; longitudinal glandular ridges bearingsmall conical tubercles sometimes present on upper surface oftibia; hinder surface of thigh rugose or granular; sides ofbody with low papillae; lower parts smooth, abdomen excep-tionally feebly granular; upperparts brown, gray, olive, orbright green, with numerous rounded or elongated dark brownor black spots strongly contrasted with lighter ground colorand with or without light margins; upperparts rarely withoutdark spots; a dark central streak, a dark blotch on upper eye-lid, and a rounded dark spot on top of snout sometimes pres-ent; tympanum often reddish or bronzed, occasionally withlight spot in center; dorso-lateral glandular dermal folds
MEXICAN TAILLESS AMPHIBIANS 193golden, bronzed, or grayish; sides of body paler than backand variously spotted or marbled with black; upper surfacesof limbs with large dark spots, or with continuous or inter-rupted dark cross bands; hinder surface of thighs yellow orgrayish white, vermiculated, marbled, or spotted with black,or else black with reddish-yellow spots; underparts white oryellow; throat and chest often spotted, marbled, or mottledwith gray or brown; longitudinal dark streak or dark mar-bling may or may not be present on anterior surface of thigh;fingers obtusely pointed, sometimes bordered by feeble lateralfringe; first finger equal to or longer than second; the hind limbbeing carried forward along the body, the tibio-tarsal jointreaches to eye or to end of snout, or a little beyond, and usuallyto between eye and end of snout; distance from snout to ventiVs to 2 times length of tibia; toes obtusely pointed, two-thirdsto nearly entirely webbed, usually two phalanges of fourth free,occasionally one or three; outer metatarsals separated nearlyto base; subarticular tubercles rather small, moderately prom-inent; tarsal fold distinct, feeble, or absent; inner metatarsaltubercle narrow, more or less prominent, or compressed, one-third to one-half the length of inner toe; outer metatarsaltubercle absent, or small and indistinct; width of head at levelof posterior angles of jaws generally less than distance betweenend of snout and posterior margin of tympanum ; snout roundedor obtusely or acutely pointed, projecting more or less beyondthe mouth, and as long as or longer than the transverse diam-eter of eye; canthus rostralis distinct or obtuse; loreal regionoblique, feebly concave; nostril equidistant from eye andfrom end of snout, or a little nearer the former; interorbitalwidth one-third to once that of upper eyelid; tympanum verydistinct, smooth surfaced, and one-half to three-fourths diam-eter of eye; vomerine teeth in short transverse or posteriorlyconverging oblique groups usually between the choanae (fig.24), occasionally on a level with their posterior borders, orextending a little beyond their level, and rarely on a level withtheir anterior borders; male with vocal vesicles internal, ormore or less developed externally, but not retractile, forminga flabby pouch behind angle of mouth and above arm; head-and-body length of adults, 65 to 110 mm pipiens (p. 203)Skin of back between dorso-lateral glandular dermal folds with-out elongated warts or glandular ridges 5Tips of fingers swollen; vomerine teeth in oblique posteriorlyconverging rows, the apex of which projects behind level ofposterior borders of choanae; head broader than long, ratherstrongly depressed; snout rounded, scarcely projecting beyondthe mouth, as long as transverse diameter of eye; canthus ros-tralis distinct; loreal region moderately oblique and deeplyconcave; nostril a little nearer to end of snout than to eye;tympanum very distinct, its transverse diameter less thanhalf that of eye; first finger longer than second; the hind limbbeing carried forward along the body, the tibio-tarsal jointreaches to between eye and end of snout; distance from snoutto vent 1% times length of tibia; toes with tips swolleninto small disks, webbed to tips; outer metatarsals separatednearly to base; subarticular tubercles rather large and promi-
194 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
nent; no tarsal fold or ridge; inner metatarsal tubercle oval, two-fifths length of inner toe; no outer metatarsal tubercle; skin ofback between dorso-lateral glandular dermal folds covered withsmall pustules; minute sharp pointed asperities as well aslongitudinal rows of minute tubercles on upper surface ofthigh, tibia, and tarsus; a strong glandular fold from postero-external angle of upper eyelid to shoulder, and its upper branchis continued backward as a moderately prominent narrowdorso-lateral fold to about level of middle of sacrum; under-parts smooth; upperparts (in alcohol) olive, with indistinctdark spots; sides marbled with dark and light; hinder side ofthighs blackish, marbled with gray; throat and chest withgrayish suffusion; abdomen whitish; head-and-body lengthof t3'pe, 111 mm pustulosa (p. 213)Tips of fingers obtuse or rather pointed; vomerine teeth in smalloblique groups or short transverse series between the choanae,or on a level with the posterior borders of the latter, equidis-tant from each other and from choanae, or nearer to each other;width of head at level of posterior angles of jaws less thandistance between end of snout and posterior margin of tym-panum; snout rounded or obtusely pointed, projecting beyondthe mouth, as long as or longer than transverse diameter of eye;canthus rostralis distinct; loreal region moderately oblique,concave; nostril equidistant from eye and from end of snout,or a little nearer the latter; interorbital width greater than,equal to, or less than that of upper eyelid; tympanum verydistinct, one-half to five-sixths diameter of eye; fingers obtuseor rather pointed, with more or less distinct lateral dermalridge; first finger longer than second; male during breedingseason with large velvety brown horny pad on inner side offirst finger; internal vocal vesicles present; terminal phalangesfeebly expanded distally; the hind limb being carried forwardalong the body the tibio-tarsal joint reaches to eye, to end ofsnout, or to between these two points; distance from snout tovent 1% to 2 times length of tibia; tips of toes swollen into verysmall disks; toes webbed to tips or with two distal joints offourth free; outer metatarsals separated nearly to base; subar-ticular tubercles rather small, fairly prominent; no tarsal fold,sometimes an indistinct ridge; inner metatarsal tubercle elliptic,fairly prominent, one-fourth to one-half the length of inner toe;no outer metatarsal tubercle; skin of back between dorso-lateral glandular dermal folds either smooth, or with minutehispid tubercles, or granular with small pearllike tubercles; astrong glandular fold from upper eyelid to shoulder, and itsupper branch is continued backward as a narrow or moderatelybroad dorso-lateral fold to about level of acetabulum or notquite so far; a more or less distinct glandular fold, often inter-rupted posteriorly, from below eye to above arm; longitudinalor sinuous glandular ridges bearing numerous small hispidtubercles on upper surface of thigh, tibia, and tarsus; under-parts smooth; upperparts green, olive, gray, or brown, some-times green in front and brown behind, uniform or with smallblackish spots and specks; dark canthal, supratympanic, andoblique post-tympanic streaks; loreal and temporal regionsoften dark brown, with light streak below eye and tympanum to
MEXICAN TAILLESS AMPHIBIANS 195
rictus oris; tympanum occasionally reddish brown; a shortoblique dark bar extends downward upon front of forelimb;hind limbs with or without more or less regular dark crossbands; hinder surface of thighs often vermiculated with blackand yellow, or marbled with dark brown and yellow; undersurface of thighs mottled or marbled with dark brown; throatand chest sometimes entirely brown or heavily marked withblotches and fine specks of same color; abdomen generallyuniform white, often yellow and mottled or spotted with somedarker color; sides yellowish, vermiculated or mottled withdark brown or black; head-and-body length of adults, 95 to125 mm palmipes (p. 200)RANA CATESBEIANA ShawCommon BullfrogFigures 21, 22, and 231802. Rana catesbeiana Shaw, General zo-ology, vol. 3, pt. 1, p. 106, pi. 33.?Storer, 1925, Univ. California Publ.Zool., vol. 27, p. 276, pi. 18, figs.53-56.?Wright, 1929, Proc. U. S.Nat. Mus., vol. 74, no. 2756, pp. 8,37, pi. 8, figs. 7-9.1889. Rana catesbiana Cope, U. S. Nat.Mus. Bull. 34, p. 424, fig. 108.?DicKERSON, 1906, The frog book, p.227, pis. 87, 89-96.Type locality.?South Carolina.Range.?Probably restricted inMexico to the coastal lowlands ofTamaiilipas and northern Vera Cruz,and to the tributaries of the RioGrande in the northern MexicanStates.Remarks.?According to Stejneger and Barbour, ^* "Shaw evidentlyknew of the Bull Frog, principally from Catesby, but he speaks ofit only as being North American and apparently saw no actualspecimens himself." It seems desirable to select a more definitetype locality than North America for this species. The assumptionthat Shaw based his description upon the account of this species byCatesby appears well founded. According to his brief itinerary,Mark Catesby arrived in Virginia, where he had relatives, on April 23,1712, and remained there for seven years. He had not as yet givenany thought to the publication of a work on natural history and hadevidently made no drawings, although during this sojourn he didcollect a few plants for a friend in England. On his next visit toNorth America, Catesby landed on May 23, 1722, at Charleston,
Figure 21.?Diagrammatic ventral view ofshoulder girdle of Rana catesbeiana; flrmisternalin structure. The inner extremities of theclavicle and coracoid are attached to theepicoracoid cartilage, and the opposite halvesof the girdle are incapable of movementCI., clavicle; Co., coracoid; E., episternum;Mes., sternum or mesosternum; O., omoster-num; Sc, scapula; .Y., xiphisternura
" Stejneger, L., and Barbour, T., A check list of North American amphibians and reptiles, p. 35, 1923.Cambridge, Mass.
196 BULLETIN 160, UNITED STATES NATIONAL MUSEUMPmx,
Max
Q.J.-1
Figure 22.?Diagrammatic dorsal view of skull of Rana catesbeiana: C,condyle; Ex. oc, exoccipital; Fr., frontoparietal; Max., maxillary; Na.,nasal; Pmx., premaxillary; Pr., prootic; Pt., pterygoid; Q. J., quadra-tojugal; S., sphenethmoid; Sg., squamosalPmx.
Figure 23 .?Diagrammatic ventral view of skull of Rana catesbeiana: Ex. oc,exoccipital; Fr., frontoparietal; Max., maxillary; Pa., parasphenoid; Pal.,palatine; Pmx., premaxillary; Pr., prootic; Pt., pterygoid; Q., quadrate car-tilage; Q. J., quadratojugal; S., sphenethmoid; Sg., squamosal; Vo., vomer
MEXICAN TAILLESS AMPHIBIANS 197
S. C. For about one year he searched for animals and plants alongthe seacoast, and then journeyed up the Savannah River to FortMoore, where he continued to paint and describe the fauna and flora.After nearly three years in Carolina, Georgia, and Florida, he departedfor the Bahama Islands, and later in 1726 returned to England.Although Catesby in his "Natural History of Carolina, Florida, andthe Bahama Islands," 1731, frequently includes observations thatdate back to the time of his visit to relatives in Virginia, the descrip-tions and drawings were based upon animals actually observed onthe second visit. The bullfrog that served as a basis for the drawingwas no doubt captured somewhere in South Carolina, and the in-clusion of observations on the habits of this frog in Virginia merelyindicates that Catesby 's knowledge of the habits of this species wasnot limited to his field work in South Carolina. Therefore SouthCarolina is here designated as the type locality for Rana catesbeiana.Specimens examined.?Six, as follows:Rana catesbeiana
Museum
198 BULLETIN 160, UNITED STATES NATIONAL MUSEUMNo. 3344, this number was originally assigned to "12 + " frogs. Anotation made some years later states that 15 of these specimenshad been reentered under Nos. 39383-97. Fourteen paratypes(Nos. 39383-96), in addition to the electotype, are now in the pos-session of the museum. One paratype (M.C.Z. No. 2600) wasevidently sent to the Museum of Comparative Zoology as an exchange.Of these 14 paratypes, nine have a head-and-body length that isnot greater than 65 mm. The largest (U.S.N.M. No. 39383) hasa head-and-body length of 95 mm.Inasmuch as some diagnostic importance has been assigned to therelative lengths of the toes, the average ratios for this series of 14specimens have been computed as follows: Lengths of first, second,and third toes, measured from inner metatarsal tubercle to tip ofcorresponding toe, are equivalent, respectively, to 28.33, 51.18, and74.22 per cent of the fourth toe. In the largest paratype (No. 39383)the distance between the inner metatarsal tubercle and the tip of thefourth toe is 46.5 mm., and the corresponding measurement for thesmallest (No. 39389) is 30.2 mm. The following measurements weretaken from the electotype and the largest paratype: Head-and-bodylength, 77 and 95 mm.; transverse diameter of tympanum, 6 and 7.4mm.; transverse diameter of eye, 7.5 and 8 mm.; and anterior edgeof eye to nostril, 5.1 and 5.9 mm.The three paratypes of Rana adtrita Troschel were collected byBaron J. W. von Miiller somewhere in Mexico. As described byTroschel, these have the same peculiar and characteristic color patternthat so readily distinguishes R. montezvmae from other Mexican frogs.The underside of the thigh is described as being reddish, but markedwith numerous white spots. Cope has mentioned the presence of pinkspots on the under surface of the thighs of a specimen he examined.Cope failed to designate a type specimen for his supposedly distinctsubspecies concolor and merely stated that it was collected by Dr.Alfred Duges at Guanajuato. The original description is limited tothe following words: "Similar to the typical form, but of a uniformfawn-brown color without the spots of the typical form." The sub-species concolor was either overlooked or ignored by Cope two yearslater, since it does not appear in the published synonymy of Ranamontezumae, although at the end of the accompan3ang description itis stated that an unspotted brown variety had been found nearGuanajuato by Dug^s. Some of the specimens from Guanajuato areconsiderably larger than those from Mexico City. The head-and-body length of the largest specimen (U.S.N.M. No. 12684) is 125 mm.
;
transverse diameter of tympanum, 9.6 mm.; transverse diameter ofeye, 11.5 mm.; and anterior edge of eye to nostril, 7 mm. Most ofthese specimens are so faded that the distinctive color pattern hasbeen obliterated, and a few of them are bleached almost white. Dis-
MEXICAN TAILLESS AMPHIBIANS 199tinct broad dark crossbars can be seen on several when the skin is wet.The skin on the back between the dorso-lateral glandular folds,unlike those from the City of Mexico, is distinctly warty. In viewof the unsatisfactory condition of these specimens it is obviously diffi-cult to determine their exact relationships, and they are tentativelyreferred to this species. Besides these larger individuals, there is inthe collection one lot of four immature frogs (U.S.N.M. No. 10247)that are unquestionably R. montezumae. All are somewhat faded,and on two and possiblj' on a third this bleaching process has revealeddistinct traces of darker spots on the back between the dorso-lateralglandular folds. The darker spots are not visible on the fourthindividual and this (No. 81101) has been selected as the electotypefor Rana montezumae concolor Cope.Specimens examined.?One hundred and six, as follows:Rana montezumae
Museum
200 BULLETIN 160, UNITED STATES NATIONAL MUSEUMRANA PALMIPES Spix1824. Rana palmipes Spix, Animalia nova, sive species novae testudinum etranarum quas in itinere per Brasiliam, etc., p. 5, pi. 5, fig. 1.
?
Boulenger,1882, Catalogue of the Batrachia Salientia s. Ecaudata in the collection ofthe British Museum, 2d edit., p. 48.
?
Gunther, 1900, Biologia Centrali-Americana, Reptilia and Batrachia, p. 202, Feb.
?
Noble, 1918, Bull. Amer.Mus. Nat. Hist., vol. 38, art. 10, p. 316, pi. 14.?Boulenger, 1920, Proc.Amer. Acad. Arts and Sci., vol. 55, no. 9, p. 473, Aug.
?
Miranda-Ribeiro,1926, Archiv. Mus. Nac. Rio de Janeiro, vol. 27, p. 165, fig. 92.?Lutz, 1927,Mem. Inst. Oswaldo-Cruz, vol. 20, fasc. 1, pp. 46, 62, pi. 9.1859. Rana affinis Peters, Monatsber. k. preuss. Akad. Wiss. Berlin, p. 403,May (Caracas, Venezuela).
?
Cope, 1866, Proc. Acad. Nat. Sci. Philadel-phia, vol. 18, p. 130.
?
Boulenger, 1919, Ann. and Mag. Nat. Hist., ser. 9,vol. 3, no. 16, p. 416.1859. Ranula gollmerii Peters, Monatsber. k. preuss. Akad. Wiss. Berlin, p.402, May (Caracas, Venezuela).1863. Rana clamata var. guianensis Peters, Monatsber. k. preuss. Akad. Wiss.Berlin, p. 412.1867. Pohlia palmipes Steindachner, Reise der Oesterreichischen FregatteNovara, Zoologischer theil, vol. 1, Amphibien, p. 15, pi. 1, figs. 5-8.1874. Ranula brevipalmata Cope, 1874, Proc. Acad. Nat. Sci. Philadelphia, p. 131(Nauta, Peru). [Nee Rana brevipalmata Peters, 1871, Monatsber. k.preuss. Akad. Wiss. Berlin, p. 646.]1874. Ranula nigrilatus Cope, Proc. Acad. Nat. Sci. Philadelphia, p. 131 (Nauta,Peru).1877. Rana vaillanti Brocchi, Bull. Soc. Philom. Paris, ser. 7, vol. 1, no. 4, p. 176(Mullins River, near Belize, British Honduras).
?
Brocchi, 1882, Missionscientifique au Mexique et dans l'Am6rique Centrale, recherches zoologiques,pt. 3, sect. 2, p. 11, pi. 2, figs. 1-lc.1882. Rana copii Boulenger, Catalogue of the Batrachia Salientia s. Ecaudatain the collection of the British Museum, 2d edit., p. 49. [Substitute namefor Rana brevipalmata Cope, preoccupied.]1900. Rana bonaccana Gunther, Biologia Centrali-Americana, Reptilia andBatrachia, p. 201, pi. 60, fig. B, Feb. (Bonacca Island, off coast of BritishHonduras).1900. Rana melanosoma Gunther, Biologia Centrali-Americana, Reptilia andBatrachia, p. 203, pi. 63, fig. B, Feb. (Duenas, Guatemala, and HaciendaRosa de Jericho; altitude, 3,250 feet, Nicaragua).1913. Rana brevipalmata rhoadsi Fowler, Proc. Acad. Nat. Sci. Philadelphia,p. 166, pi. 9 (Bucay, Province of Guayas, Ecuador).Type locality.?Amazon River, Brazil.Range.?From Vera Cruz (Jalapa) and Morelos (Cuernavaca) south-ward through Oaxaca and Chiapas to the Mexican border, and thencesouth through Central America and South America to northern Peru(Nauta, Maranon River), western Brazil (Matto Grosso or Villa Bella),and to Pernambuco on the Atlantic coast of Brazil.Remarks.?Boulenger's monograph on the American frogs of thegenus Rana has been followed in compiling the synonymy of Ranapalmipes because sufficient material from critical areas in Central andSouth America is not now available in American institutions to permita review of the forms that are obviously closely related or identical
MEXICAN TAILLESS AMPHIBIANS 201
with R. palmipes. Central American and Mexican specimens nowreferred to this species vsivy considerably in the extent of the webbetween the toes, the texture of the skin between the dorso-lateralglandular folds, the relative transverse diameter of the tympanum ascompared wdth that of the eye, the shape of the snout, and the generalcoloration. Until the limits of variation with respect to characterssuch as those just mentioned can be determined by aa examination ofadequate series from one or more localities in South and CentralAmerica, respectively, it is impossible to arrive at any definiteconclusion.This frog is especially interesting because it is the only representa-tive of the genus Rana in South America. Since its discovery bySpix in stagnant water along the Amazon River in Brazil, its range hasbeen extended to western Brazil, to the Pacific coastal region of Ecua-dor, and north as far as central Mexico. In spite of its rather exten-sive range, comparatively little is known in regard to its habits andstructural variation. With one exception, all the specific names here-inafter enumerated are based on young or immature individuals.Banula gollmerii, according to Boulenger, "was described from arecently transformed young, from Caracas, measuring 50 mm. fromsnout to vent." A somewhat larger frog from Caracas, which mea-sured 63 mm. from snout to vent, became the type of Peters's Ranaaffinis. The describer was inclined to regard the last-mentionedspecies as a local variety of the European Rana temporaria. Peterssome years later allocated these two species to Rana palmipes.The types of Ranula brevipalmata and Ranula nigrilatus were col-lected by Prof. James Orton at Nauta on the Maranon River innortheastern Peru. Both of these specific names were based onyoung frogs, and they have head-and-body lengths of 49 and 48 mm.,respectively. The toes of hrevipalmata are not fully webbed, sincethe membrane reaches to the bases of the distal joints on all thetoes except the fourth, on which it extends to the penultimate joint.The tympanum is described as being as large as the eye. The toes ofnigrilatus are fully webbed, and the tympanum is two-thirds as largeas the eye.The type of Rana vaillanti was collected by Firmin Bocourt on thebanks of MulHns River, near Belize in British Honduras. It is alarge adult, with a head-and-body length of 125 mm. The toes arefully webbed, the vomerine teeth are in oblique groups at the level ofthe posterior borders of the choanae, and the coloration of the under-parts and limbs is similar to other Central American specimens ofRana palmipes.Giinther seems to have overlooked the small terminal disks of thetoes when he described Rana bonaccana. Bonacca Island lies off thecoast of British Honduras and not off Yucatan as stated by Giinther.It is the eastern one of a group known as the Bay Islands. No char-
202 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
acters are given in the description of this species that can not bematched in specimens from Central America. Giinther based hisRana melanosoma on smooth-skinned specimens from Duenas inGuatemala and from Hacienda Rosa de Jericho in Nicaragua. Thespecimens are immature and the largest individual has a head-and-body length of 65 mm. The toes are webbed to the distal end of thepenultimate joint, or nearly so.The type of Rana brevipalmata rhoadsi (A.N.S.P. No. 18051) camefrom Bucay, Province of Guayas, in western Ecuador. Fowler recog-nized the resemblance in coloration between his supposedly new formand Cope's R. brevipalmata, but held that a new name was warrantedbecause the diameter of the tympanum was only two-thirds that ofthe eye and the snout was shorter. The type is a young frog with ahead-and-body length of 43 mm.The habitat of this species in Vera Cruz is described by Ruthvenas follows : ^^This species was observed in some numbers along La Laja Creek and at LakeCatemaco, but specimens were very diiBcult to secure. It was found only wherethe shores were densely covered with vegetation, usually where there were busheshanging low over the water. In such places the individuals stayed well out fromshore, upon sticks or low branches, and upon the slightest alarm leaped into thewater, where they were very difficult to see even when partially above the surface.The following observations on the general coloration of this frog inlife are based on specimens collected by C. R. Halter in Nicaragua:^*The variation in color of our specimens is limited to the fading out of the groundtone and to the developing of spots on the posterior part of the back. In lifethe color above was grass-green from the tip of the snout to the tympanum andolive from the tympanum to the ends of the appendages. In certain lights abronze-like shimmer was present on the head. The posterior sides of the legswere mottled with dark brown and white. The lower parts of the legs were lessdistinctly mottled with brown. The throat was pearly white; the belly andventral surfaces of the appendages were washed with lemon yellow. The pos-terior appendages were heavily mottled with brown below.Specimens examined.?Fourteen as follows:Rana palmipes
Museum
M.C.Z..U.S.N.MDo..Do..Do..Do...Do...Do..
CatalogueNo.
8433479147n5-16201654711930329-3130485-8846762
Num-ber ofspeci-mens Locality collectedVera Ceuz:A 1varadoJalapa _TlacotalpamMORELOs: CuernavacaOaxaca:Mountains near Santo Do-mingo, altitude 1,600 feet.City of TehuantepecdoChiapas: Mountains nearTonala.
Date collected
May 18,1894Aug. ?.1892June 16,1895
Aug. 15,1895
By whom collected orfrom whom received
E. R. Dunn.R. Monies d'Oca.Nelson and Goldman.P. L. Jouy.Nelson and Goldman.Francis Sumichrast.Do.Nelson and Goldman.
" Ruthven, A. O., Zool. Jahrb. (Syst. Abt.), vol. 32, pt. 4, p. 306, 1912.
'? Noble, Q. K., The amphibians collected by the American Museum expedition to Nicaragua in 1916
.
Riill Amftr lVTii<! Nat VfWt. vr>l SS art in n 317 1Q1?
MEXICAN TAILLESS AMPHIBIANS 203RANA PIPIENS SchreberCommon Leopard FrogFigures 1, a, and 241782. Rana pipiens Schreber, Der Naturforscher, Halle, vol. 18, p. 185, pi. 4.
?
DicKERSON, 1906, The frog book, p. 171, pis. 43-50. col. pis. 11, 12.?Wright,1914, Carnegie Inst. Washington Publ. No. 197, p. 52, pi. 15, Sept. 8.?Storer, 1925. Univ. California Publ. Zool., vol. 27, p. 266, pi. 17, fig. 52.1803. Rana halecina Daudin, Histoire naturelle des rainettes, des grenouilleset des crapauds, p. 41.
?
-Holbrook, 1836, North American herpetology, vol. 1,p. 89, pi. 13.?HoLBROOK (1842), 1844, North American herpetology, vol. 4,p. 91, pi. 22.
?
Boulenger, 1882, Catalogue of the Batrachia Salientia s.Ecaudata in the collection of the British Museum, 2d edit., p. 41.
?
Brocchi,1882, Mission scientifique au Mexique et dans I'Amerique Centrale, recher-ches zoologiques, pt. 3, sect. 2, p. 10.
?
Cope, 1886, Proc. Amer. Philos.Soc, vol. 23, no. 124, p. 517, Oct. 1.?GtJNTHER, 1900, Biologia Centrali-Americana, Reptilia and Batrachia, p. 198, Feb. 1.
?
Boulexger, 1920, Proc.Amer. Acad. Arts and Sci., vol. 55, no. 9, p. 433, Aug.1826. Rana uiricnlarius Harlan, Amer. Journ. Sci. and Arts, vol. 10, p. 60(Pennsylvania and New Jersey)
.
1856. Rana oxyrhynchus Hallowell, Proc. Acad. Nat. Sci. Philadelphia, vol. 8,p. 142, Aug. (Florida; nee A. Smith).1858. Rana lecontii Gijnther, Catalogue of the Batrachia Salientia in the col-lection of the British Museum, p. 15 inec Baird and Girard, 1853).Botjlenger, 1882, Catalogue of the Batrachia Salientia s. Ecaudata in thecollection of the British Museum, 2d edit., p. 42.1859. Rana herlandieri Baird, Report on the United States and Mexican Bound-ary Survey, vol. 2, Reptiles, p. 27, pi. 36, figs. 7-10 (Brownsville, Tex.).1877. Rana macroglossa Brocchi, Bull. Soc. Philom. Paris, ser. 7, vol. 1, no. 4,p. 177 (Plateau of Guatemala).?Brocchi, 1882, Mission scientifique auMexique et dans I'Amerique Centrale, recherches zoologiques, pt. 3, sect. 2,p. 12, pl. 3, figs. 1-lc.1877. Rana maculata Brocchi, BuU. Soc. Philom. Paris, ser. 7, vol. 1, no. 4, p. 178(Totonicapan, Guatemala).
?
Brocchi, 1882, Mission scientifique au Mexiqueet dans I'Amerique Centrale, recherches zoologiques, pt. 3, sect. 2, p. 13, pl.3, figs. 2-2c.1882. Rana nigricans Brocchi, Mission scientifique au Mexique et dans I'Amer-ique Centrale, recherches zoologiques, pt. 3, sect. 2, p. 15, pl. 4, fig. 3 (neeHallowell, 1854).1882. Rana lecontei Brocchi, Mission scientifique au Mexique et dans I'AmeriqueCentrale, recherches zoologiques, pt. 3, sect. 2, p. 14, pl. 4, figs. 1-lc. {neeBaird and Girard, 1853).1883. Rana forreri Botjlenger, Ann. and Mag. Nat. Hist., ser. 5, vol. 11, no. 65,p. 343, May (Presidio, Sinaloa).?GiJNTHER, 1900, Biologia Centrali-Amer-icana, Reptilia and Batrachia, p. 199, pl. 60, fig. A, Feb.1886. R. [ana] h. [alecina] auslricola Cope, Proc. Amer. Philos. Soc, vol. 23, no.124, p. 517, Oct. 1.?Cope, 1887, U. S. Nat. Mus. Bull. 32, p. 19.1886. Rana h. [alecina] sphenocephala Cope, Proc. Amer. Philos. Soc, vol. 23,no. 124, p. 517, Oct. (Substitute name for Rana oxyrhynchus Hallowell.)1889. R. [a7ia] v. [irescens] auslricola Cope, U. S. Nat. Mus. Bull. 34, pp. 398, 399.1889. Rana virescens sphenocephala Cope, U. S. Nat. Mus. Bull. 34, p. 399, fig. 99.1889. Rana virescens virescens Cope, U. S. Nat. Mus. Bull. 34, p. 401, fig. 100.1889. Rana virescns brachycephala Cope, U. S. Nat. Mus. Bull. 34, p. 403, fig. 101.(Yellowstone River, Mont,).
204 BULLETIN 160, UNITED STATES NATIONAL MUSEUM1899. Rana trilobata Mocquard, Bull. Soc. Philom. Paris, ser. 9, vol. 1, no. 4,p. 158, pi. 1, fig. 1 (Jalisco, Mexico).1900. Rana omiltemana Gunther, Biologia Centrali-Americana, Reptilia andBatrachia, p. 200, pi. 61, fig. A, Feb. (Omilteme, Guerrero, Mexico).1922. Rana burnsi Weed, Proc. Biol. Soc. Washington, vol. 35, p. 108, Oct. 17(New London, Kandiyohi County, Minn.).1922. Rana kandiyohi Weed, Proc. Biol. Soc. Washington, vol. 35, p. 109, Oct. 17(New London, Kandiyohi County, Minn.).Type locality.?Raccoon, Gloucester County, N. J.Range.?From Nova Scotia, southern New Brunswick (St. CroixRiver), Ontario (Ottawa River), Keewatin (Norway House), north-em Alberta (Fort Smith on Slave River), the Kootenay district ofsoutheastern British Col-umbia, Snake River ofwestern Idaho and downthe Columbia River asfar as Fort Dalles, Oreg.,northeastern California(Alturas, Modoc County),Pyramid Lake in Nevada,and the Lake Tahoe re-gion (Placer and EldoradoCounties) of eastern Cali-fornia, south to the lowerColorado River Valley inArizona and California,Florida, the Gulf coast,Mexico, Guatemala, andNicaragua.Remarks .?Boulenger "has preferred to use Ranahalecina for the Americanleopard frogs, citing Lin-naeus^* as the authorityfor this name, but is careful to state that the name appears in thesynonymy of Rana ocellata. In explaining the derivation of the namehalecina, Boulenger remarks that the " Sillhoppetosser " of Kalm ^'has been latinized by Linnaeus. Kalm states that the Swedes inAmerica call these frogs "sillhoppetosser," or herring hoppers, inallusion to the fact that they make their appearance early in thespring at the beginning of the herring season. Rana halecina isclearly employed by Linnaeus as a vernacular descriptive term in
" Boulenger, Q. A., A monograph of the American frogs of the genus Rana. Proc. Amer. Acad. Artsand Sci., vol. 55, no. 9, p. 433, Aug., 1920; Burt, C. E., Proc. Biol. Soc. Washington, vol. 44, p. 13, Feb.21, 1931.
'? Linnaeus, C, System* naturae, ed. 12, vol. 1, p. 356, 1766.
'? Kalm, P., En Resa til Norra America, vol. 3, p. 45, 1761; Forster, Kalm's travels into North AmericSfLondon, vol. 1, pp. 379, 380, 1772.
Figure 24.?Opened mouth of Rana pipiens, showing positionof vomerine teeth; c, Choana; e., orifice of eustachian tube; t.,tongue, with bilobed posterior extremity and with attachmentat anterior end
MEXICAN TAILLESS AMPHIBIANS 205Latin form, in allusion to the comment on the habits of this frog bythe original author. There are numerous similar usages of Latindescriptive terms in this edition of Linnaeus. They have no statusunder any code of nomenclature.The specific name Rana pipiens seems to have been published firstby Schreber, and he likewise based the description of this species onthe " Sillhoppetosser " {op. cit., vol. 3, p. 46), which Kalm observedon March 11 at Raccoon, Gloucester County, N. J.Harlan failed to state whether the original description of Ranautricularius was based on preserved specimens, and inasmuch as hewas at that time somewhat interested in the life histories of frogs, itis quite likely that this diagnosis was based on captive individuals.It is stated, however, in a footnote that "a specimen preserved inspirits in the cabinet of the Philadelphia Acad, of Nat. Sc. is erron-eously labelled R. halecina." The presence of an external greenishvocal vesicle extending from the lower jaw to above the shoulderjoint is a modification that permits a greater distension of this ap-paratus, and for such males Harlan proposed the name Rana utricu-larius, and those that show no such modification of the outer skinwere allocated to R. halecina.In 1856, Edward Hallowell gave the name Rana oxyrhynchus to amale frog collected by Mr. Ashmead "in a sulphur spring, near theSt. John's River, about three hundred miles from Key West," Fla.Inasmuch as this name was preoccupied. Cope in 1886 proposedRana h.[alecina] sphenocephala as a substitute name. The type ofthis subspecies is presumably in the collection of the Academy ofNatural Sciences of Philadelphia. The diagnostic characters of thesubspecies sphenocephala were defined by Cope in 1889 as follows:Head entering length of head and body two and a half or less than three times;males with external vesicles; muzzle more acuminate; no cross bars on tibia;spots smaller.Hallowell, however, describes the type of R. oxyrhynchus as beingentirely destitute of vocal vesicles, head narrow and acute, and hindlimbs with white-margined transverse black bars.Baird gives "southern Texas generally" as the habitat of Ranaberlandieri. The 11 cotypes of this species (U.S.N.M. No. 3293)were collected by Capt. Stewart Van Vliet, at Brownsville, Tex.Two of them (M.C.Z. No. 155) were sent to the Museum of Com-parative Zoology at Cambridge, Mass., and there now remain in thecollection of the United States National Museum 1 adult, 2 young,and 6 tadpoles. The adult specimen in this lot w^as figured byBaird on Plate 36, Figures 7-10, of the report on the reptiles of theUnited States and Mexican Boundary Survey.66786?32 14
206 BULLETIN 160, UNITED STATES NATIONAL MUSEUMAccording to the characters employed by Cope in 1889 for dis-tinguishing the subspecies of virescens [ = 'pi/piens], Rana berlandieriis distinguished as follows: Head narrow; muzzle acuminate; inter-orbital space two-thirds the width of upper eyelid; short glandulesand Unear series of glandulose tuberosities on back between dorso-lateral glandular dermal folds; dark spots on upperparts irregular inoutline and narrowly edged with hght yellow; hind limbs with light-edged dark crossbars; longitudinal dark streak on anterior face of thigh;large external vocal vesicles; the hind Umb being carried forwardalong the body, the tibio-tarsal joint reaches the end of the snout;body length, snout to vent, 79 mm.The three cotypes (M.H.N.P. No. 169a, parchment label No. 632)of Rana macroglossa Brocchi are labeled as coming from the plateauof Guatemala and were received from the ''Sociedad Economica deGuatemala." Among other characters assumed to be of diagnosticvalue, Brocchi called attention to the rather wdde tongue with promi-nent posterior horns, the vomerine teeth in two small rounded groupsbetween the choanae, the smooth skin, and a first finger longer thanthe second. These cotypes are described in my notes as follows:Head-and-body length, respectively, 64, 66.5, and 46.5 mm.; the hindlimb being carried forw^ard along the body, the tibio-tarsal jointreaches to between the eye and end of snout; terminal phalanges claw-like; transverse diameter of tympanum one-half to three-fourths that ofeye; muzzle rounded; interorbital space as wdde as upper eyelid; skinof upperparts fairly smooth, without prominent elongate glanduleson back between dorso-lateral glandular dermal folds; dark spots onupperparts indistinctly outlined and somewhat effaced; throat spottedwith brown or gray; sides marbled, spotted with black on brownishor grayish background; interrupted transverse dark bands on uppersurfaces of hind limbs. Two of these cotypes are clearly referable toRana pipiens. The third, to which the parchment label is attached,has the under surface of the thighs brown and stippled with Ughtspecks. This specimen is questionably referred to this species.Paul Brocchi gave the name Rana maculata to three frogs (M.H.N.P.parchment label No. 6412) collected by Firmin Bocourt, a memberof the staff of the "Mission Scientifique au Mexique et dans I'AmeriqueCentrale" at "Totonicapam (Mexique)" [ = Totonicapan, Guate-mala]. In the description of this species, Brocchi mentioned the smalloval tongue with posterior horns of moderate size, nearly subequalfirst and second fingers, unusually swollen first finger, slightly devel-oped subarticular tubercles, and vomerine teeth in two obhque row^s,with apex of open chevron thus formed on a level with choanae.These cotypes are characterized as follows: Head-and-body length,respectively, 61, 58.5, and 54.5 mm.; the hind limb being carried for-ward along the body, the tibio-tarsal joint reaches beyond end of
MEXICAN TAILLESS AMPHIBIANS 207
snout; transverse diameter of tympanum about two-thirds that ofeye; muzzle acuminate; interorbital space as wide as upper eyelid;well marked dorso-lateral glandular dermal folds; black spots onupperparts small and closely approximated; throat marbled withdark gray; sides and posterior surfaces of thighs marbled with darkbrown or black on a grayish white background; continuous narrowtransverse dark bands on upper surfaces of hind limbs.The type of Rana nigricans Brocchi seems to be lost, and no entrywith corresponding data was found in the catalogues of the MuseumNational d'Histoire Naturelle in Paris.A female frog collected by Alphonso Forrer at Presidio [de Mazatlan]in Sinaloa, which was subsequently acquired by the British Museumof Natural History, became the type of Ranaforreri Boulenger (B.M.No. 1882. 12. 5. 7). This specimen is described in my notes as follows:Head-and-body length, 71.6 mm.; transverse diameter of tympanum,6.5 mm.; transverse diameter of eye, 8.1 mm.; anterior edge of eyeto nostril, 5.5 mm.; the hind limb being carried forward along thebody, the tibio-tarsal joint reaches nearly to end of snout; a distincttarsal fold; a large elongate internal metatarsal tubercle; muzzlebroad and rounded; interorbital space narrow, not quite half thewidth of upper eyelid; numerous large light-edged black spots; largeprominent elongate glandules on back between dorso-lateral glandulardermal folds; limbs with black crossbars; under surface of thighsareolated. This is the specimen figured by Giinther on Plate 60,Figure A, Biologia Centrali-Americana.Cope, in 1886, established the subspecific name austricola by remark-ing that the "Mexican specimens referred to in these citations [i. e.,the synonymy for "Rana halecina Daudin (Kalm)"] constitute asubspecies, which I call R. h. austricola." When we examine these11 citations to the literature we find that mention of Mexican speci-mens is made in only three. Giinther and Boulenger both refer sixMexican specimens in the British Museum collection to Rana lecontii,while Brocchi figures a frog from Vera Cruz, which is referred toR. lecontei on the captions for the plate and in the text. Inasmuchas it is generally the practice to select as the lectotype either a speci-men or a published illustration actually seen by the original describerbefore the description is published, I definitely choose the specimenfigured by Brocchi^" on Plate 4, Figure 1, of his "Etude des Batraciensde I'Amerique Centrale" as the type of this subspecies. The illus-tration published by Brocchi, if correctly drawn, shows that thespecimen, upon which it is based, differs from the diagnosis for thesubspecies austricola in having the muzzle obtuse at the end and the
* Brocchi, P., Mission scientiflque au Mexique et dans rAmfirique Centrale, recherches zoologiquesParis, pt. 3, sect. 2, p. 14, pi. 4, fig. 1, 1882,
208 BULLETIN 160, UNITED STATES NATIONAL MUSEUM
crossbars on the tibia interrupted. The diagnosis published by Copein 1889 reads as follows:Head entering length of head and body two and a half to nearly three times;no external vocal vesicles; muzzle more or less acuminate; spots less distinct;tibia generally crossbarred; no longitudinal band on front of femur.In 1886, Cope ^^ states that the "Rana halecina, Kalm, subspeciesberlandieri" in his *^ "Check-List of North American Batrachia andReptiha " is not the Rana berlandieri of Baird. Again in 1889, Cope **remarks that the "last named subspecies [i. e., Rana virescens hrachy-cephala Cope] is the one I called R. v. berlanderi fsic], but it turns outthat the typical specimens of that species belong to the subspeciesvirescens." Taken at its face value, this statement is rather contra-dictory, for Cope listed the cotype of R. berlandieri Baird (U.S.N.M.No. 3293) among the specimens referred to his subspecies R. virescensbrachycephala. What Cope seemingly intended to say was that "thetypical specimens" of the Rana halecina berlandieri of his check listactually belong to the subspecies virescens.The subspecific name brachycephala appears for the first time in1886 in Cope's "Synonymic List of North American species of Bujoand Rana" where its habitat is given as the "Central and Sonoranregions." No description accompanies the name, and the synonymyincludes two citations, one of which is the "2?. h. berlandieri Cope" ofhis "Check List" pubhshed in 1875 and the other, "Rana halecinaBoulenger, Cat. Batr. Brit. Mus., ed. ii, p. 41, nee Kalmii." Untilthe publication of Cope's "Batrachia of North America," the sub-specific name brachycephala was strictly a nomen nudvm, inasmuchas the characters that distinguished this subspecies from supposedlytypical halecina [ = virescens Cope, 1889] were not indicated. Cope'sdiagnosis of the subspecies brachycephala published in 1889 is seem-ingly based upon one specimen, and this inference is drawn from hisintroductory remark that "I select as typical a specimen from theYellowstone River (No. 3363)." There are, however, 15 specimenscatalogued under that number, although Cope mentions only 5 inhis list of specimens examined, and all of them were collected by Dr.F. V. Hayden in 1857 along the Yellowstone River in Montana.This lot consists of 2 adults, 1 young, and 12 tadpoles. Inasmuch asit is now impossible to recognize the actual individual upon which Copebased his description, all these specimens have been designated asthe cotypes of the subspecies brachycephala. The following notesapply to the largest cotype: Head-and-body length, 86; transverse
" Cope, E. D., Synonymic list of North American species of Bufo and Rana, with descriptions of somenew species of Batrachia, from specimens in the National Museum. Proc. Amer. Philos. Soc, vol. 23,no. 124, p. 517, Oct. 1, 1886." Cope, E. D., Check-list of North American Batrachia and Reptilia. U. S. Nat. Mus. Bull. 1,p. 32, 1875." Cope, E. D., U. S. Nat. Mus. Bull. 34, p. 398, 1889.
MEXICAN TAILLESS AMPHIBIANS 209diameter of tympanum, 6; transverse diameter of eye, 8; anterioredge of eye to nostril, 5.4 mm.; the hind limb being carried forwardalong the body, the tibio-tarsal joint reaches to anterior margin ofeye. Cope in 1889 published the following short diagnosis of thesubspecies brachycephala
:
Head shorter and more obtuse, entering the length [of head and body] threeand a half times; males without or with rudimental external vocal vesicles;dorsal spots larger, widely yellow bordered; tibial crossbands complete; nolongitudinal band on the front of the thigh.Mocquard's Rana trilobata (M.H.N.P. No. 169a, parchment labelNo. 97-189) is based on a single 3^oung individual collected by LeonDiguet at Guadalajara in Jalisco. The specific name alludes to thetrilobed form of the posterior end of the tongue. Boulenger^* hasexamined the type in the Paris Museum and states that "there is ashort furrow between the horns of the tongue, such as I have observedin some R. escvlenta." This specimen may be briefly characterizedas follows: Head-and-body length, 33.5 mm.; the hind limb beingcarried forward along the body, the tibio-tarsal joint reaches to aboutthe center of the eye; a dorso-lateral glandular dermal fold present;posterior end of tongue has three projections as described by Mocquard
;
muzzle rounded; no linear glandular folds on back between dorso-lateral glandular dermal folds; dark spots and markings on upperpartsindistinct; posterior limbs indistinctly crossbarred ; tympanum slightlymore than half the diameter of the eye.Five adult males (B.M. Nos. 1895. 7. 15. 31-35), which werecollected at Omilteme in Guerrero by H. H. Smith, a collector em-ployed b}'^ Godman and Salvin, become the cotypes of Giinther'sRana omiltemana. The largest one of these cotj^pes has a head-and-body length of 60.5 mm. Giinther's published description of thisspecies may be summarized as follows: Snout narrow, acuminate;interorbital space narrow, a little more than half the width of theupper eyelid; rather small rounded or subquadrangular dark spotswith narrow light edges on back; small glandules on back betweennarrow dorso-lateral glandular dermal folds; hind limbs with lightedged dark crossbars; external vocal vesicles small; hind limb, withouttarsus, as long as two-thirds length of body.Weed has published descriptions of tw^o well-marked color variantsof the leopard frog. These specimens were selected from a tankcontaining several thousand frogs. Color characters alone wereemployed to distinguish Rana hurnsi (F.M.N.H. No. 3065) andRana kandiyohi (F.M.N.H. No. 3066) from Rana pipiens. Theupperparts of Rana hurnsi are described as being more or less immacu-late, with some shade of green or brown, while Rana kandiyohi hasblack spots that apparently fuse vnth. a mottled ground color.M Boulenger, Q. A., Proc. Amer. Acad. Arts and Sci., vol. 55, no. 9, p. 435, footnote, Aug., 1920.
210 BULLETIN 160, UNITED STATES NATIONAL MUSEUMThe various forms that have been enumerated are here interpretedas individual variations. This conclusion has been reached after acritical examination and tabulation of many specimens. No con-stant geographic peculiarities were encountered among the specimensstudied. Unique variations in color pattern were found in almostevery local series studied, and as none of these variations are exactlyalike, it seems necessary to disregard such peculiarities and groupall the leopard frogs of North and Central America under one name.The ground color of the upperparts varies greatly and is rarelyuniform green or brown. There are two prominent dorso-lateralglandular dermal folds, which are either bronze or yellow, sometimesa very light tint of yellow, and which extend almost parallel to eachother from the upper eyelid to the sacral region. These folds oftenbecome silvery white after long immersion in alcohol, and betweenthem are brown, olive-green, or nearly black spots, which are gener-ally irregular in shape and arrangement. In many instances thearrangement of the darker marldngs on one side is totally differentfrom that on the opposite side.A tabulation of the Mexican series of leopard frogs demonstratedthe existence of an endless variety of patterns. One of the variantshas numerous, closely aggregated, dark spots. On another the darkdorsal spots are unusually elongated, and several spots apparentlyare fused. Other specimens have small widely separated dark spots.Fourteen frogs have the dorsal dark spots indistinctly outlined, andin a few instances the spots were nearly effaced. A few individualswere noted that have no apparent dark marldngs between the dorso-lateral glandular dermal folds. Large dark dorsal spots with distinctyellow borders were observed on 45 specimens. At least 50 frogswere counted that have the dorsal dark spots distinctly outlined, buton these the light margins are either very narrow or lacking. Wideor narrow transverse dark bars on the hind limbs, which may becontinuous or interrupted, occur more frequently than dark oval spots.The dark transverse bars were continuous on 70 individuals and in-terrupted on 35. A dark marbling on the anterior surface of thethigh is quite noticeable on some specimens and in eight instances adark band was formed. Many of the frogs, however, lack this darkermarbling. Most of the specimens have the throat and chest uni-formly light colored, though there were a number of frogs amongthose examined that have the throat and chest variegated or marbledwith some darker color.The Mexican specimens generally have an acuminate snout and arather narrow head. Other specimens taken at the same localitiesas those with narrow heads, however, have a much wider head andan obtuse snout. One series of 13 specimens from Durango have theskin of the upperparts unusually rough and pustulose. This condi-
MEXICAN TAILLESS AMPHIBIANS 211tion exists also on immature individuals and has been observed onspecimens with a body length as short as 35 mm. More than 20specimens were found that have distinct longitudinal rows of pus-tules on the upper surface of the tibia. On the majority of the adultfrogs the skin of the upperparts is granular, with more or less promi-nent longitudinal glandular folds on back between the dorso-lateralfolds of the skin. Young frogs and occasional adults have the skinof the upperparts nearly smooth. A dark spot is generally presenton each upper eyelid, and in 40 instances a dark spot was noted onthe upper surface of the snout.The absence of a well-defined streak that extends from end of snoutto end of glandular fold behind angle of the mouth on each side of thehead above the dark or mottled upper lip is stated by Boulenger inhis monograph on the American frogs of the genus Rana to be geo-graphically distinctive for specimens from Arizona, Texas, Mexico,and Central America. This light streak is absent or only distinctfrom below the eye posteriorly in many of the Mexican specimens.Nevertheless, there are leopard frogs in the United States NationalMuseum from Tamaulipas, Vera Cruz, Guanajuato, Jalisco, Guerrero,and Tabasco that have this light streak as well defined as any indi-vidual from Eastern United States or Canada. The general color-ation of live frogs of this species taken at Cuatotolapam in Vera Cruzis described as follows by Ruthven:?^Ground color of upper parts dark brownish olive, occasionally speckled withbright green; stripe on canthus rostralis and dorsal spots black or blackish, thelatter margined with greenish; stripe along upper lip light greenish yellow, fre-quently much broken up; dorso-lateral glandular fold light greenish yellow toorange brown; belly white.The length of the hind limb has very little diagnostic value. In aseries of 50 individuals, 10 were found in which the tibio-tarsal jointreached to the middle or anterior margin of the eye, when the hindlimb was carried forward along the side of the body, 15 in which itreached to between the eye and the end of the snout, 17 to the end ofsnout, and 8 beyond the end of the snout.Noble ^^ has published the following comments on a small series offrogs that are referred to Rana austricola and that were collected byC. R. Halter in Nicaragua:The variation in color is not limited to an intensif.ying or fading of the tones.There is also a slight variation in pattern. The spots on the back may be moreor less confluent. In the smallest and the largest specimens in the collection thespots have been extended to form an unbroken but irregular streak on each sideof the back. Each pair of spots may be confluent in a transverse direction, butit is more common for the spots in each longitudinal row to run together. ThereM Ruthven, A. G., Zool. Jahrb. (Syst. Abt.), vol. 32, pt. 4, pp. 305, 306, 1912.w Noble, O. K., The amphibians collected by the American Museum Expedition to Nicaragua in 1916.Bull. Amer. Mus. Nat. Hist., vol. 38, art. 10, p. 316, 1918.
212 BULLETIN 160, UNITED STATES NATIONAL MUSEUMdoes not seem to be any progressive development of pattern during the growthof an individual, for specimens of the same size may have very different colorpatterns.Specimens examined.?Two-hundred and sixty-seven, as follows
:
Rana pipiens
Museum
U.S.N.MDoDo
DoDoDoDoDoDoDoM.C.ZDoU.S.N.MDoDoDoDoDoDoM.C.ZDoDoDoU.S.N.MDoDoDoDoDoA.M.N.HB.MA.M.N.HU.S.N.MDoDoM.H.N.PU.S.N.MDoDoDoDoDoDo
* Cotypes of Rana berlandieri.* Type of Rana forreri.
' Type of Rana irilobata.
CatalogueNo.
MEXICAN TAILLESS AMPHIBIANSRana pipiens?Continued 213
214 BULLETIN 160, UNITED STATES NATIONAL MUSEUMin having the tips of the fingers distinctly swollen. The followingnotes were made on the type: Head-and-body length, 110.8 mm.;transverse diameter of tympanum, 5.8 mm.; transverse diameter ofeye, 12.2 mm.; anterior edge of eye to nostril, 7.8 mm.; the hind limbbeing carried forward along the body, the tibio-tarsal joint reachesto between eye and end of snout; a prominent glandular fold frompostero-external angle of upper eyelid to shoulder, and its upperbranch is continued backward as narrow dorso-lateral dermal foldto about level of middle of sacrum; no tarsal fold; skin of upperpartscovered with small pustules; minute sharp pointed asperities as wellas longitudinal rows of minute tubercles on upper surface of tibia;vomerine teeth in oblique posteriorly converging rows, the apex ofwhich projects behind level of posterior borders of choanae; upper-parts (in alcohol) olive, ^^^th indistinct dark spots; sides marbledwith dark and light; hinder side of thighs blackish, marbled withgray; throat and chest with grayish suffusion; abdomen whitish.Specimens examined.?One, the type.RANA TARAHUMARAE Boulenger1917. Rana tarahumarae Boulenger, Ann. and Mag. Nat. Hist., ser. 8, vol. 20,no. 120, pp. 416, 417.
?
Boulenger, 1919, idem, ser. 9, vol. 3, no. 16,p. 411.
?
Boulenger, 1920, Proc. Amer. Acad. Arts and Sci., vol. 55, no. 9,p. 468, Aug.Type locality.?loquiro and Barranca del Cobre, Sierra Tara-humare, State of Chihuahua, Mexico.Range.?Not known to occur elsewhere than in the Sierra Tara-humare in western Chihuahua.Remarks.?Giinther based his description of this species uponsix cotypes, four of which came from loquiro [ = ? Yoquiva] andtwo from Barranca del Cobre. The coloration of the upperpartsalone will distinguish this species from any other Mexican Rana.The characters possessed by this frog indicate that it is an isolatedrace of the boylii group. Boidenger correctly summarized therelationships of this frog in the following words: "This species isvery closely allied to R. boylii, differing in the larger eye, the moreoblique loreal region, the more distinct tympanum, the shorter tibia,and the absence of vocal sacs." Although the distributional limitsof this frog are unknown, it is clear that its range is isolated fromthat of boylii by several well-defined barriers, of which the arid regionof southwestern Arizona and the Mohave Desert of California arethe most effective.These six cotypes of Rana tarahumarae are described in my notesas follows:Four cotypes: B.M. Nos. 1911.12.12.36-39; loquiro [ = ? Yo-quival. Sierra Tarahumare; Neilly, collector. Largest individual:Head-and-body length, 78 mm.; transverse diameter of tympanum,
MEXICAN TAILLESS AMPHIBIANS 2153.5 mm.; transverse diameter of eye, 8.4 mm.; anterior edge of eyeto nostrU, 6.2 mm.; the hind limb being carried forward along thebody, the tibio-tarsal joint reaches to end of snout; vomerine teethwithin and behind level of posterior margins of the choanae; a largeelongate internal metatarsal tubercle; no dorso-lateral glandulardermal fold; a feeble, curved glandular fold from eye to shoulder;upperparts, including top of head, grayish (in alcohol) with numeroussmall black spots or dots; skin on back pustulose on one cotype;throat and limbs heavily mottled with brownish; abdomen white.Two cotypes: B.M. Nos. 1914. 1. 28. 148-149; Barranca del Cobre,Sierra Tarahumare, altitude 3,000 feet; Dr. Hans Gadow, collector.Largest individual: Head-and-body length, 61.3 mm.; transversediameter of tympanum, 3.7 mm.; transverse diameter of eye, 7.7mm.; anterior edge of eye to nostril, 4.8 mm.; the hind limb beingcarried forward along the body, the tibio-tarsal joint reaches to barelybeyond the nostril; a distinct tarsal fold; elongate internal metatarsaltubercle, but no outer one; vomerine teeth in two small clustersmostly behind, but within level of choanae; no dorso-lateral glandulardermal fold; upperparts olive brown, marked with black spots withlight centers; black spots on upperparts tend toward a linear arrange-ment; black crossbars on upper surfaces of thigh, tibia, and tarsus,and outer surface of foot; dark crossbars on forearm.Boulenger erred when he stated that this species lacked a tarsalfold. This fold is quite distinct on one of the larger cotypes, but isless sharply defined on some of the smaller ones.Specimens examined.?Six, as follows:Rana tarahumarae
Museum

INDEX
(Principal references'Jare given in boldface figures)abbotti, Eleutherodactylus, 91.Acrodytes, 148.acutirostris, Biifo, 27.Adenomus, 28.badioflavus, 28.adtrita, Rana, 197, 198.Aelurophryne, 18.affinis, Hyla, 5, 156, 157.Rana, 200, 201.Agalychnis, 140.helenae, 145.moreletii, 146.agua, Bufo, 28, 53, 55.albilabris, Cystignathus, 83, 84.Leptodactylus, 82 (key), 83, 85.albomarginata, Hyla, 149.alfredi, Eleutherodactylus, 8, 14, 95(kev), 99.Hylodes, 99.alvariiis, Bufo, 5, 29, 30, 31 (key), 35.americanus, Bufo, 27.Amphibia, 16.amulae, Tomodactylus, 13, 120, 131.Anaxyrus, 28.melancholicus, 28.anomalus, Bufo, 13, 29, 44, 46.Drorooplectrus, 29.Ansonia, 29.penangensis, 29.arborea, Hvla, 168.Rana, '147, 148.arboreus, Calamita, 147.arenicolor, Hyla, 142, 149, 153 (key),156.areolata, Gastrophryne, 186.argillaceus, Bufo, 7, 58.asper, Bufo, 28.aspera, Hylomantis, 149.augusti, Eleutherodactylus, 8, 91, 94(kev), 100.Hylodes, 14, 100.Auletris, 148.aurata, Hvla, 148.austricola^ Rana, 203, 207, 211.badioflavus, Adenomus, 28.Baliopygus, 189.Batrachyla, 90.leptopus, 90.longipes, 90, 107.baudinii, Hyla, 9, 11, 18, 130, 142, 149,155 (key), 160, 175.Smilisca, 160.beatae, Eleutherodactylus, 14, 93 (kev)
,
104.Hylodes, 104.
berkenbuschii, Hylodes, 11, 116.berlandieri, Rana, 203, 205, 206, 208.bicolor, Hyla, 139.Phyllomedusa, 140.Rana, 139.biporcatus, Strabomantis, 90.bistincta, Hyla, 7, 150, 151 (key), 163.Boana, 147.boans, Hvla, 147, 148.Rana, 147, 148.bocourti, Hyla, 168.Bombinator, 24.bonaccana, Rana, 200, 201.Borborocoetes mexicanus, 13, 108, 109,110, 112.borneensis. Gastrophryne, 186.boylii, Rana, 190, 214.brachvcephala, Rana, 36, 203, 208, 209.Brevicipitidae, 17 (key), 182 (key),brevipalmata, Rana, 200.Ranula, 200, 201.Bufo, 27, 31 (key), 75.acutirostris, 27.agua, 28, 53, 55.alvarius, 5, 29, 30, 31 (key), 35.americanus, 27.anomalus, 13, 29, 44, 46.argiUaceus, 7, 58.asper, 28.bufo, 28.calamita, 29.canaliferus, 8, 34 (key), 40, 59.carens, 28.coccifer, 35 (key), 41.cognatus, 29, 32 (key), 41.compactilis, 11, 28, 29, 30, 31, 33(key), 44, 45.coniferus, 29.cristatus, 11, 29, 33 (key), 48.cruentatus, 28.debilis, 5, 6, 30, 31 (key), 50.dipternus, 41, 44.d'orbignyi, 29.dorsalis, 72, 73.frontosus, 72, 73.granulosus, 27, 28, 68, 70.guttatus, 28.haematiticus, 30.horribilis, 11, 53, 55.insidior, 4, 50, 52.intermedins, 63, 64, 65.kelaartii, 28.lateralis, 14, 58, 69.lentiginosus frontosus, 72.lentiginosus woodhousei, 72.leschenaultii, 28. 217
218 BULLETIN 160, UNITED STATES NATIONAL MUSEUMBufo levifrons, 44, 46, 47.margaritifer, 28.marinus, 30, 32 (key), 53.marmoratus, 148.marmoreus, 11, 34 (key), 58.mexicanus, 12, 44, 46, 47.monksiae, 8, 63, 66.naricus, 27.nasutus, 27.nebulifer, 68, 70.occidentalis, 63, 65..occipitalis, 14, 48, 49.peltocephalus, 28, 55.proboscideus, 27, 28.punctatus, 5, 29, 31, 33 (key), 60,157.(Microphryne) pustulosus, 78.quercicus, 30.semilineatus, 27.simus, 30, 34 (key), 63.speciosus, 5, 44, 45.stentor, 77.sternosignatus, 13, 68, 70.trachypus, 68, 60.typhonius, 28.valliceps, 11, 29, 35 (key), 49, 68,80.valliceps microtis, 68, 70.vulgaris, 27, 28.woodhousii, 29, 32 (key), 73.bufo, Bufo, 28.Rana, 27.Bufonidae, 17 (key), 27, 77.burnsi, Rana, 204, 209.Calamita, 147.arboreus, 147.calamita, Bufo, 29.Epidalea, 29.calcitrans, Eleutherodactylus, 13, 108,109, 110.Hylodes, 108, 110.caliginosus, Leptodactylus, 88, 89.callidryas, Hyla, 140.canaliferus, Bufo, 8, 34 (key), 40, 59.carens, Bufo, 28.carolinense, Engystoma, 186, 187.carolinensis, Gas'trophryne, 186.catesbeiana, Rana, 190, 191 (key), 195.Cauphias, 76 (key), 118.crassum, 12, 118.guatemalensis, 119.Centrotelma, 149.Ceratophrys, 24.Chaunus, 28.marmoratus, 28.Chilophryne, 28.conifera, 29.chrysoprasina, Ranula, 189.chrysoprasinus, Trypheropsis, 189.Cinclidium, 149.granulatum, 149.Cincloscopus, 149.coccifer, Bufo, 35 (key), 41.coerulescens, Ollotis, 29.
cognatus, Bufo, 29, 32 (key), 41.compactilis, Bufo, 11, 28, 29, 30, 31, 3$(key), 44, 45.concoior, Rana, 8, 197, 198.conifera, Chilophryne, 29.coniferus, Bufo, 29.conspicillatus, Hylodes, 99.copei, Hyla, 157.coper, Hyla, 157.Cophomantis, 149.punctillata, 149.copii, Hvla, 157.Raiia, 200.cornutue, Limnophys, 90.coronata, Gastrotheca, 133.couchii, Scaphiopus, 6, 20.Cranopsis, 29.fastidiosus, 29.crassa, Hyla, 119.crassum, Cauphias, 12, 118.Craugastor, 90.Crepidius, 29.epioticus, 29.crepitans, Hyla, 149.Crinia, 77.cristatus, Bufo, 11, 29, 33 (key), 48.Crotaphitis, 189.cruentatus, Bufo, 28.cuneus, Hypopachus, 183.Cystignathidae, 77.cystignathoides, Phyllobates, 127.Syrrhophus, 7, 126 (key), 127, 129.CystignathuR, 81.albilabris, 83, 84.fragilis, 83, 85.gracilis, 83, 85.labialis, 8, 83, 84.labyrinthicus, 81.melanonotus, 88.microtis, 8, 88.pachypus, 81.perlaevis, 8, 88, 89.dacnicolor, Phyllomedusa, 7, 140, 142(key), 143.daulinia, Smilisca, 149.debihs, Bufo, 6, 6, 30, 31 (key), 60.Dendrohj^as, 148.Dendropsophus, 148.Diaglena, 130, 132 (key), 137.spatulata, 13, 137.digueti, Hyliola, 12, 157, 158.diplotropis, Leptophis, 136.dipternus, Bufo, 41, 44.Docidophryne, 28.d'orbignyi, Bufo, 28.dorsalis, Bufo, 72, 73.Rhinophrynus, 8, 11, 26.Dromoplectrus, 29.anomalus, 29.Dryomelictes, 148.Dryophytes, 148.dugesii, Scaphiopus, 8, 12, 22.dunnii, Eleutherodactylus, 10, 91,^97(key), 105.
INDEX 219
elegans, Engystoma, 187.Gastrophryne, 13, 183 (key), 187.Eleutherodactylus, 75, 76 (key), 90,92 (in key), 123.abbotti, 91.alfredi, 8, 14, 95 (key), 99.augusti, 8, 91, 94 (key), 100.beatae, 14, 93 (key), 104.calcitrans, 13, 108, 109, 110.dunnii, 10, 91, 97 (key), 105.flavescens, 91.inoptatus, 92.laticeps, 11, 91, 93 (key), 106.latrans, 91, 101, 102.longipes, 6, 91, 95 (key), 107.martinicensis, 90.melanostictus, 91.mexicanus, 75, 91, 92, 99 (key),108.plicatus, 13, 112.rhodopis, 7, 91, 92, 98 (key), 106,111, 113.rugulosus, 8, 91, 92, 96 (key), 116.sallaei, 13, 112.venustus, 13, 91, 97 (key), 117.Elosia, 90.empusa, Peltaphryne, 29.Otaspis, 29.Engystoma, 186.carolinense, 186, 187.elegans, 187.mexicanum, 11, 187, 188.rugosuDQ, 186.ustum, 187, 188.variolosum, 183, 184.Engystomops, 17, 75, 76 (key), 77.petersi, 77.petersii, 77.pustulosus, 77, 78.stentor, 77.Entomoglossus, 81.pustulatus, 81.Enydrobius, 91.Epedaphus, 149.Epidalea, 29.calamita, 29.epioticus, Crepidius, 29.Epirhexis, 90.longipes, 107.esculenta, Rana, 189, 209.Eupemphix, 77, 78, 79.gadovii, 14, 187, 188.nana, 77.nattereri, 77.pustulosa, 78, 79.Euhyas, 90.euphorbiacea, Hyla, 13, 164, 165.Eurhina, 28.Exerodonta, 179.sumichrasti, 12, 179, 181.eximia, Hyla, 5, 130, 149, 150, 153(key), 164, 168, 173, 174.fastidiosus, Cranopsis, 29.fitzingeri, Hylodes, 90.flavescens, Eleutherodactylus, 91.fodiens, Pternohyla, 13, 135.
forreri, Rana, 13, 203, 207.fragilis, Cystignathus, 83, 85.Leptodactylus, 12, 83.frontalis, Hyla, 148.frontosus, Bufo, 72, 73.fuscomaculata, Paludicola, 77, 79.gabbi, Hyla, 142.gadovii, Eupemphix, 14, 187, 188.galdi, Pristimantis, 90.Gastrophryne, 182, 183 (key), 186.areolata, 186.borneensis, 186.caroHnensis, 186.elegans, 13, 183 (key), 187.leucosticta, 186.texensis, 186.usta, 6, 183 (key), 187.Gastrophrynoides, 186.Gastrotheca, 130, 131 (key), 133.coronata, 133.marsupiata, 133.ovifera, 133.gigas, Rana, 53, 54.Gnatliophysa, 81.godmani, Hvla, 13, 170, 171.goUmerii, Ranula, 189, 200, 201.graciUpes, Hvla, 6, 150, 154 (key), 165,168.gracilis, Cystignathus, 83, 85.granulatum, CincHdium, 149.granulosus, Bufo, 27, 28, 68, 70.gratiosa, Hyla, 149.guatemalensis, Cauphias, 119.Plectrohyla, 118.guianensis, Rana, 200.gtintheri, Leij'la, 90.guttatus, Bufo, 28.guttilatus, Malachylodes, 123.Syrrhophus, 8, 125 (key), 127, 129.haematiticus, Bufo, 30.halecina, Rana, 203, 204, 205, 207, 208.hallowellii, Hylodes, 90.hanr ondii, Scaphiopus, 18, 19, 22, 23.hebes, Scytopis, 149.helenae, Agalychnis, 145.Phyllomedusa, 8, 142 (key), 145.Hesperomys sonoriensis, 5.holbrookii, Scaphiopus, 19.holochlora, Hyla, 146.horribilis, Bufo, 11, 53, 55.Hyas, 148.Hyla, 130, 132 (key), 147, 150 (key),179.affinis, 5, 156, 157.albomarginata, 149.arborea, 168.arenicolor, 142, 149, 153 (key), 156.aurata, 148.baudinii, 9, 11, 18, 130, 142, 14?,155 (kev), 160, 175.bicolor, 139.bistincta, 7, 150, 151 (kev), 163.boans, 147, 148.bocourti, 168.
220 BULLETIN 16 0, UNITED STATES NATIONAL MUSEUMHyla callidryas, 140.copei, 157.coper, 157.copii, 157.crassa, 119.(Hylomedusa) crepitans, 149.euphorbiacea, 13, 164, 165.eximia, 5, 130, 149, 150, 153 (key),164, 168, 173, 174.frontalis, 148.gabbi, 142.godmani, 13, 170, 171.gracilipes, 6, 150, 154 (key), 165,16S.gratiosa, 149.holochlora, 146.(Centrotelma) infulata, 149.lactea, 148.lafrentzi, 168.lichenosa, 13, 176.marmorata, 148.marsupiata, 133.microtis, 11, 170.miotympamim, 6, 130, 142, 149,156 (key), 163, 170.moreletii, 146.muricolor, 6, 160.nana, 13, 164, 165, 174.nasus, 90.nigropunctata, 13, 176, 177.palmata, 147, 148.phlebodes, 149, 155 (key), 173.plicata, 12, 150, 152 (key), 173.ranoides, 90.regilla, 149, 168.rudis, 12, 136.septentrionalis, 148.smithii, 13, 164, 165, 174.spilomma, 7, 151 (key), 163, 176,177.staufFeri, 7, 150, 152 (key), 173,174.taeniopus, 13, 150, 156 (key), 163,164, 175.vanvlietii, 160.venulosa, 130, 142, 148, 149, 151(key), 155 (key), 176.versicolor, 148.viridis, 147.xerophyllum, 148.zonata, 149.Hylaria, 147.Hylella, 130, 133 (key), 179, 180 (key).picta, 13, 179, 180.platycephala, 8, 181.punctatissima, 179.sumichrasti, 8, 180 (key), 181.tenera, 179.Hylidae, 17 (key), 139, 131 (key).Hyliola, 149.
. digueti, 12, 157, 158.regilla, 149.Hylodes, 90, 114, 115, 119.alfredi, 99.augusti, 14, 100.beatae, 104.berkenbuschii, 11, 116.calcitrans, 108, 110.
Hylodes conspiciUatus, 99.fitzingeri, 90.hallowellii, 90.laticeps, 106.lineatus, 90, 108.martinicensis, 90, 115.plicatus, 112, 114.rhodopis, 112, 115.ricordii, 90.sallaei, 112, 113, 114.venustus, 117.Hylodidae, 119.Hylomantis, 149.aspera, 149.Hylomedusa, 149.Hypodictvon, 123, 129.Hypopachus, 182, 183.cuneus, 183.inguinalis, 183, 184.oxvrrhinus, 13, 184, 185.seebachii, 183, 184.variolosus, 183, (key), 184.Hypsiboas, 148.Hypsipsophus, 148.Incilius, 29.infulata, Hyla, 149.inguinalis, Hypopachus, 183, 184.inoptatus, Eleutherodactylus, 92.insidior, Bufo, 4, 50, 53.intermedins, Bufo, 63, 64, 65.kandiyohi, Rana, 204, 309.kelaartii, Bufo, 28.labialis, Cystignathus, 8, 83, 84.Leptodactylus, 85.labyrinthica, Rana, 81.labvrinthicus, Cystignathus, 81.lactea, Hyla, 148.lacteus, Sphaenorhynchus, 148.lafrentzi, Hyla, 168.lateralis, Bufo, 14, 58, 59.Schmismaderma, 28.laticeps, Eleutherodactylus, 11, 91,93,(kev), 106.Hylodes, 106.latrans, Eleutherodactylus, 91, 101, 102.Lithodytes, 102, 103, 104.lecontei, Rana, 203, 307.lecontii, Rana, 203, 307.Leiuperus, 121.mexicanus, 12, 108.Leiyla, 90.guntheri, 90.leprus, Syrrhophus, 8, 125 (key), 138.Leptodactyhdae, 17 (key), 74, 76 (key),77.Leptodactylus, 75, 76 (key), 81,82 (key).albilabris, 82 (key), 83, 85.caliginosus, 88, 89.fragihs, 12, 83.labialis, 85.melanonotus, 83 (key), 88, 127,128.pentadactylus, 87.typhonia, 81.
INDEX 221Leptophis diplotropis, 136.Leptophr3'ne, 28.leptopus, Batrachyla, 90.leschenaultii, Bufo, 28.leucosticta, Gastrophryne, 186.levifrons, Bufo, 44, 46, 47.lichenosa, Hyla, 13, 176.Limnophys, 90.cornutus, 90.napaeus, 90.lineata, Rana, 90.lineatus, Hylodes, 90, 108.Liohyla, 90.ranoides, 90.Lithodytes, 90, 114.latrans, 102, 103, 104.ranoides, 90.rhodopis, 112, 113.Liuperus nitidus, 11, 120, 121.Liyla, 90.rugulosa, 116.Lobipes, 148.longipes, Batrachyla, 90, 107.Eleutherodactylus, 6, 91, 95 (key),107.Epirhexis, 107.Lophopus, 148.marmoratus, 148.macrodon, Rana, 87.macroglossa, Rana, 203, 306.maculata, Rana, 203, 306.Malachylodes, 123.guttilatus, 123, 137.margaritifer, Bufo, 28.margaritifera, Bufo, 28.marina, Rana, 28, 53, 54.marinus, Bufo, 30, 32 (key), 53.marmorata, Hyla, 148.marmoratus, Bufo, 148.Chaunus, 28.Lophopus, 148.Trachycephalus, 148.marmoreus, Bufo, 11, 34 (key), 58.marnockii, Syrrhophus, 123.marsupiata, Gastrotheca, 133.Hyla, 133.martinicensis, Eleutherodactylus, 90.Hylodes, 90, 115.Megalophrys, 18.melancholicus, Anaxyrus, 28.melanonotus, Cystignathus, 88.Leptodactylus, 83 (key), 88, 127,128.melanosoma, Rana, 200, 303.melanostictus, Eleutherodactylus, 91.mexicana, Pleurodema, 108.mexicanum, Engystoma, 11, 187, 188.mexicanus, Borborocoetes, 13, 108, 109,110, 112.Bufo, 12, 44, 46, 47.Eleutherodactylus, 75, 91, 92, 99(key), 108.Leiuperus, 12, 108.Microphryne, 77.pustulosa, 78.Microps, 186.
Microps, unicolor, 186.microtis, Bufo, 68, 70.Cystignathus, 8, 88.Hyla, 11, 170.miotvmpanum, Hvla, 6, 130 ,142, 149,156 (key), 163, 170.modesta, Rana, 87.monksiae, Bufo, 8, 63, 66.montezumae, Rana, 5, 189, 190, 192(key), 197.moreletii, Agalychnis, 146.Hyla, 146.'Phyllomedusa, 140, 142, 143 (key)146.multiplicatus, Scaphiopus, 6, 19, 33.muricolor, Hyla, 6, 160.mystaceus, Syrrhopus, 10, 104.
nana, Eupemphix, 77.Hyla, 13, 164, 165, 174.Nannophryne, 29.variegata, 29.napaeus, Limnophys, 90.naricus, Bufo, 27.nasus, Hyla, 90.nasutus, Bufo, 27.nattereri, Eupemphix, 77.nebuhfer, Bufo, 68, 70.Nesomantis, 18.nigricans, Rana, 203, 307.nigrilatus, Ranula, 200. 301.nigromaculatus, Trachycephalus, 148.nigropunctata, Hyla, 13, 176, 177.nitidus, Liuperus, 11, 120, 131.Paludicola, 120.Tomadactylus, 120.Notodelphys, 133.ovifera, 133.Nototrema, 133.oviferum, 133, 134.
ocellata, Rana, 204.occidentalis, Bufo, 63, 65.occipitalis, Bufo, 14, 48, 49.Ollotis, 29.coerulescens, 29.omiltemana, Rana, 13, 204, 309.omiltemanus, Syrrhaphus, 13, 108, 109,110.Opisthodelphys, 133.ovifera, 133.opisthodon, Rana, 190.Osilophus, 28.typhonius, 28.Osteocephalus, 149.taurinus, 149.Osteopilus, 148.Otaspis, 29.empusa, 29.Otolophus, 28.ovalis, Rana, 186.ovifera, Gastrotheca, 133.Notodelphys, 133.Opisthodelphys, 133.oviferum, Nototrema, 133, 134.Oxyglossus, 189.66785?32- -15
222 BULLETIN" 16 0, UNITED STATES NATIONAL MUSEUMOxyrhynchus, 27.oxyrhynchus, Rana, 203, 205.oxyrrhinus, Hypopachus, 13, 184, ISL,pachypus, Cystignathus, 81.Rana, 81.palmata, Hyla, 147, 148.palmipes, Pohlia, 189, 200.Rana, 189, 190, 195 (kev), 300,213.Paludicola', 77, 79, 121.fuscomaculata, 77, 79.[Liuperus] nitidus, 120.pustulosa, 77, 78.signifera, 78.Pelobatidae, 17 (key), 18.Pelophylax, 189.Peltaphryne empusa, 29.peltocephalus, Bufo, 28, 55.Peltophryne, 28.penangensis, Ansonia, 29.pentadactyla, Rana, 81.pentadactylus, Leptodactylus, 87.Peralaimos, 77.Stentor, 77.perlaevis, Cystignathus, 8, 88, 89.perlata, Phyllomedusa, 140.petasatus, Pharyngodon, 138.Triprion, 8, 138.petersi, Engystomops, 77.petersii, Engystomops, 77.Pharyngodon, 138.petasatus, 138.phlebodes, Hyla, 149, 155 (key), 173.Phryne, 28.Phrynohyas, 149.Phrynoidis, 28.Phrynomorphus, 28.Phyllobates cystignathoides, 127.ridens, 123.verruculatus, 11, 129.Phyllobius, 149.bicolor, 140.PhyUomedusa, 130, 132 (kev), 139, 142(key), 175.dacnicolor, 7, 140, 142 (key), 143.helenae, 9, 142 (key), 145.moreletii, 140, 142, 143 (key), 146.perlata, 140.picta, Hvlella, 13, 179, 180 (key), 180.pipiens, Rana, 18, 190, 193 (kev), 203.platycephala, Hylella, 8, 181.Plectrohyla guatemalensis, 118.Pleurodema, 77, 78.mexicana, 108.pUcata, Hyla, 12, 150, 152 (key), 173.plicatus, Eleutherodactylus, 13, 112.Hylodes, 112, 114.Pohha, 189.palmipes, 189, 200.Pristimantis, 90.galdi, 90.proboscideus, Bufo, 27, 28.Pseudophryne, 77.Pternohyla, 130, 131 (key), 135.fodiens, 13, 135.punctatissima, Hylella, 179.
punctatus, Bufo, 5, 29, 31, 33 (key),60, 157.punctillata, Cophomantis, 149.pustulatus, Entomoglossus, 81.pustulosa, Eupemphix, 78, 79.Microphryne, 78.Paludicola, 77, 78.Rana, 14, 190, 194 (key), 213.pustulosus, Bufo, 78.Engystomops, 77, 78.quercicus, Bufo, 30.Rana, 150, 189, 190 (kev).adtrita, 197, 198.afRnis, 200, 201.arborea, 147, 148.berlandieri, 203, 205, 206, 208.bicolor, 139.boans, 147, 148.bonaccana, 200, 301.boyhi, 190, 214.brevipalmata, 200.brevipalmata rhoadsi, 200, 202.bufo, 27.burnsi, 204, 209.catesbeiana, 190, 191 (key), 195.clamata guianensis, 200.copii, 200.esculenta, 189, 209.forreri, 13, 203, 207.gigas, 53, 54.halecina, 203, 204, 205, 207, 208.halecina austricola, 203, 207, 211.halecina berlandieri, 208.halecina sphenocephala, 203, 205.kandiyohi, 204, 209.labyrinthica, 81.lecontei, 203, 207.lecontii, 203, 207.lineata, 90.macrodon, 87.macroglossa, 203, 206.maculata, 203, 206.melanosoma, 200, 202.marina, 28, 53, 54.modesta, 87.montezumae, 5, 189, 190, 192(key), 197.montezumae concolor, 8, 197, 198.nigricans, 203, 207.ocellata, 204.omiltemana, 13, 204, 209.opisthodon, 190.ovalis, 186.oxyrhynchus, 203, 205.pachypus, 81.palmipes, 189, 190, 195 (kev), 200,213.pentadactvla, 81.pipiens, 18, 190, 193 (key), 203.pustulosa, 14, 190, 194 (key), 213.sylvatica, 189.tarahumarae, 190, 191 (key), 214.temporaria, 189, 201.trilobata, 12, 204, 209.typhonia, 28, 81.
INDEX 223Rana utricularius, 203, 305.vaillanti, 200, 301.venulosa, 148, 176.virescens austricola, 203.virescens berlandieri, 208.virescens brachycephala, 36, 203,208.virescens sphenocephala, 203.virescens virescens, 203, 206, 208.Ranaria, 189.Ranidae, 17 (key), 189.Ranula, 189.brevipalmata, 200, 301.chrysoprasina, 189.gollmerii, 189, 200, 301.nigrilatus, 200, 301.ranoides, Hyla, 90.Liohyla, 90.Lithodytes, 90.rectifrenis, Scaphiopus, 6, 30.regilla, Hyla, 149, 168.Hyliola, 149.Rhacophorus, 147.Rhaebo, 28.Rhinophrynidae, 17, 34.Rhinophrynus, 17, 34.dorsalis, 8, 11, 36.rostratus, 12, 36.rhoadsi, Rana, 200, 303.rhodopis, Eleutherodactylus, 7, 91, 92,98 (key), 106, 111, 113.Hylodes, 112, 115.Lithodytes, 112, 113.Rhynophrynus dorsalis, 36.ricordii, Hylodes, 90.ridens, Phyllobates, 123.rostratus, Rhinophrynus, 12, 36.rudis, Hyla, 12, 136.rugosum, Engystoma, 186.rugulosa, Liyla, 116.rugulosus, Eleutherodactylus, 8, 91, 92,96 (key), 116.Salientia, 16, 17 (key).saUaei, Eleutherodactvlus, 13.Hylodes, 112, 113, 114.Scaphiopodidae, 18.Scaphiopus, 18, 19 (key), 79.couchii, 6, 30 (key).dugesii, 8, 12, 33.hammondii, 18, 19, 22.hammondii multiplicatus, 19 (key),33.holbrookii, 19.multiplicatus, 6, 19 (key), 33.rectifrenis, 6, 30.Schmismaderma, 28.lateralis, 28.Scinax, 148.Scytopis, 149.hebes, 149.seebachii, Hypopachus, 183, 184.semilineatus, Bufo, 27.septentrionalis, Hyla, 148.signifera, Paludicola, 78.simus, Bufo, 30, 34 (.key), 63.
Snailisca, 149.baudinii, 160.daulinia, 149.smithii, Hyla, 13, 164, 165, 174.sonoriensis, Hesperomys, 5.Sooglossus, 18.spatulata, Diaglena, 13, 137.spatulatus, Triprion, 137.Spea, 19.speciosus, Bufo, 5, 44, 45.Sphaenorhynchus, 148.lacteus, 148.sphenocephala, Rana, 203, 305.spilomma, Hyla, 7, 151 (key), 163, 176,177.staufferi, Hyla, 7, 150, 152 (key), 173.174.Stenocephalus, 186.stentor, Bufo, 77.Engystomops, 77.Peralaimos, 77.sternosignatus, Bufo, 13, 68, 70.Strabomantis, 90.biporcatus, 90.sumichrasti, Exerodonta, 12, 179, 181.Hylella, 8, 180 (key) 181.sylvatica, Rana, 189.Syrrhaphus, 123.omiltemanus, 13, 108, 109, 110.Syrrhophus, 75, 77 (key), 133, 124 (key).cystignathoides, 7, 126 (key), 137,129.guttilatus, 8, 125 (key), 137, 129.leprus, 8, 125 (key), 138.marnockii, 123.mystaceus, 10, 104.omiltemanus, 13.verrucipes, 127 (key), 138.verruculatus, 126 (key), 139.Systoma, 188.taeniopus, Hyla, 13, 150, 156 (key),163, 164, 175.tarahumarae, Rana, 190, 191 (key),314.taurinus, Osteocephalus, 149.temporaria, Rana, 189, 201.tenera, Hylella, 179.texensis, Gastrophryne, 186.Tomodactvlus, 75, 77 (kev), 130.amulae, 13, 120, 131.nitidus, 130.Trachycephalus, 148.marmoratus, 148.nigromaculatus, 148.trachvpus, Bufo, 68, 69.trilobata, Rana, 12, 204, 309.Triprion, 130, 132 (key), 138.petasatus, 8, 138.spatulatus, 137.Trypheropsis, 189.chrj'soprasinus, 189.typhonia, Leptodactvlus, 81.Rana, 28, 81.typhonius, Bufo, 28.Osilophus, 28.
224 BULKETIN 16 0, UNITED STATES NATIONAL MUSEUM
unicolor, Microps, 186.usta, Gastrophryne, 6, 183 (key), 187.ustum, Engystoma, 187, 188.utricularius, Rana, 203, 305.
vaillanti, Rana, 200 201.vaUiceps, Bufo, 11, 29, 35 (key), 49,68, 80.vanvlietii, Hyla, 160.variegata, Nannophryne, 29.variolosum, Engystoma, 183, 184.variolosus, Hypopachus, 183 (key),184.venulosa, Hyla, 130, 142, 148, 149, 151(key), 155 (key), 176.Rana, 148, 176.
venustus, Eleutherodactylus, 13, 91,97 (key), 117.Hylodes, 117.verrucipes, Syrrhophus, 127 (key), 138.verruculatus, Phyllobates, 11, 129.Syrrhophus, 126 (key), 139.versicolor, Hyla, 148.virescens, Rana, 203, 206, 208.viridis, Hyla, 147.vulgaris, Bufo, 27, 28.woodhousii, Bufo, 29, 32 (key), 73.xerophyllum, Hyla, 148.
zonata, Hyla, 149.o



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