Anurans from the Early Tertiary of Patagonia
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ANURANS FROM THE EARLY TERTIARY OF PATAGONIA BOBB SCHAEEFER BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY VOLUME 93: ARTICLE 2 NEW YORK: 1949 ANURANS FROM THE EARLY TERTIARY OF PATAGONIA ANURANS FROM THE EARLY TERTIARY OF PATAGONIA BOBB SCHAEFFER Assistant Curator of Fossil Vertebrates Department of Geology and Paleontology PUBLICATIONS OF THE SCARRITT-PATAGONIAN EXPEDITION, NUMBER 37 BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY VOLUME 93 : ARTICLE 2 NEW YORK: 1949 BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Volume 93, article 2, pages 41-68, text figures 1-7, plates 16-19, table 1 Issued March 28, 1949 Price: $0.50 a copy CONTENTS INTRODUCTION . 47 SYSTEMATIC DESCRIPTIONS .... ... ... .. 49 Family Leptodactylidae . 49 Eophractus, new genus . 49 Eophractus casamayorensis, new species .49 Calyptocephalella Strand . .50 Calyptocephalella canqueli, new species .50 Eupsophus Fitzinger . 55 Eupsophus sp. 55 Neoprocoela, new genus . 57 Neoprocoela edentatus, new species.57 DISCUSSION ..... ...... 59 Anuran Differentiation ...... ..... .... ... .. 59 Anuran Dispersal ..... 64 Trends in Anuran Evolution .. 65 REFERENCES ..... 67 45 Page___ is blank. INTRODUCTION TEE: SOUH AMEixcAN CONTINENT has thus From this incomplete but representative list far yielded a small number of fragmentary it is evident that relatively few characters fossil anurans. Teracophrys (nomen nudum) are actually available in working out the was listed byAmeghino (1901) from theUpper affinities of a fossil anuran. The nature of Oligocene of Patagonia. The specimens, ap- the vertebrate column can often be deter- parently referable to the Leptodactylidae, mined, as well as a few skull characters, un- cannot be located in the Ameghino Collection fortunately rarely including the structure of of the Museo Argentino de Ciencias Natu- the prevomer. By using the criteria men- rales (Miss Noemi Cattoi, personal communi- tioned by Noble (1930), it may be possible cation). Ceratophrys prisca has been de- to postulate an arciferal or firmisternal con- scribed by Ameghino (1899) and Rovereto dition for the shoulder girdle. The terminal (1914) from the Upper Pliocene of Monte phalanges may or may not be present. Thus Hermoso, Argentina, and C. ensenadensis by the opportunity for exact identification is Rusconi (1932) from the Pleistocene (Ense- dependent upon the nature and completeness nadan) near Buenos Aires. of preservation and on the possibility of mak- Both Noble (1928) and Dunn (1931) have ing such identification without recourse to emphasized the importance of paleontologi- structures usually not fossilized. cal discovery in determining the approximate Parker (1940) has pointed out the difficul- times and routes of anuran dispersal. They ties associated with the interpretation of fos- have also pointed out the danger of assuming sil anuran material, and it is obvious that a that the failure to discover fossil anurans conservative approach in the matter of tax- belonging to particular groups is proof that onomic relationship is desirable unless a posi- such groups never existed in a given area. tive identification can be made, as in the case Dunn wisely states (ibid., p. 118), ". great of the Calyptocephalelka described in this weight should be given to positive evidence, paper. It might be pointed out that detailed and very little to negative evidence." It is of information on the osteology of the modern course true that in the absence of fossil data anurans is widely scattered and far from certain reasonable conclusions can be drawn complete. Herpetologists could render pale- on the basis of the distributional pattern of ontology (and possibly themselves) a great the present day faunas. There is, for instance, service by making comprehensive studies good reason for believing that the Discoglos- available. A single fossil discovery may be of sidae never entered North America and that the greatest importance in working out the the Pelobatidae did not invadeSouthAmerica. ramifications of anuran evolution and in de- The pelobatids were present in NorthAmerica termining routes of dispersal, providing a by the Pliocene (Taylor, 1937) but apparently reasonably accurate identification can be have migrated no farther south than Mexico. made. This is too often not possible and, as a The main taxonomic subdivisions of the survey of the literature indicates, has led to Anura now generally recognized are based many differences of opinion among competent on the five different types of vertebral col- students of the subject. umn occurring within the order (Nicholls, The present study is based on a collection 1916; Noble, 1922, 1931). The smaller cate- of lower Eocene and Middle and Upper gories are usually defined on suites of char- Oligocene anurans obtained by Dr. George acters involving the skull (particularly the Gaylord Simpson in 1930 and 1934 while on prevomer, the presence or absence of teeth, the First and Second Scarritt Patagonian and the middle ear), the shoulder girdle, the Expeditions. The specimens consist of nature of the terminal phalanges, the tongue, crushed and mostly dissociated skeletons the thigh musculature, the parotid gland, the embedded in very fine lacustrine or fluviatile presence or absence of Bidder's organ, and sediments. certain external features and proportions. The writer is greatly obligated to Dr. E. 47 48 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 93 R. Dunn for a number of helpful suggestions supplied important information from his and for reading the manuscript. He is like- field books and from preliminary notes made wise Indebted to Mr. K. P. Schmidt, Mr. on this collection. The drawings were pre- Arthur Loveridge, Mr. C. M. Bogert, and pared by Dorothea Kay Barlow and the Dr. J. A. Oliver for the loan of Recent speci- photographs were taken by Elwood Logan. mens for comparative purposes. Dr. Simpson SYSTE,MAT,I'IC DESCRIPTIONS FAMILY ?LEPTODACTYLIDAE lescrilbes the "Bird Clay" in his field notes BOPBRACTUS, NEW i as a "hardl, mlassive, bright green clay with intercalatedl )andss of whiite tuff." This la- GECNOTYPE;: EIophrahtus cas5amayorensis, new custrine deposit is exposel at the south end species. of Caffad16n 1londo near Paso Niemann, (;JNERIC DIA(GNOSIS: Ani anuirtan of leIpto- Trerritory of Chuhut, Arge¢ntina (see locating dactylidl or possibly hlylidi affinities. Ouiter diagram, Schaeffer, 1947). The anuran frag- surface of known (lermial hones of sktll(cov- ments were found in the white tuff, ere(l with (lee) rotinledi (lepressions which SPECIFIC DIAGNOSIS: Same as for genus. are surround( 'd by naIrrow, sharp, connecting DEscRIPToN: These fragments, consisting ridges. Portion of nasal bicone between orbit of the associated elements representing the an(I nares widle; nasals in 'ontItact alonig nidl- type andl in adldition a second partial maxil- dorsal line. Anterior atn(l msedlian border of lary andi a single vertebra (A.M.N.H. No. I'JO. 1. Eaphrttdus casumayorensis, new genus and species. A..NI.IN . No. .3165, type specitnens. A. )orsal view of isolated le(ft. nsilKal in artivulation with Issoxiatednmaxillary illustrated be- low. 1Probsahl}e shape., of fronrit part of skull indicated. B. I.ateral view of inconipilete left mtiaxillary. X3/2. orbit elevtated. M.axillary deep, with teeth; 3164), represent the oldest known anuran ascen(ling l)roe3s >of m. xillary robust, indli- remains from South America. Tlhe few really cating at lea-st partital roofing of the temnporal diagnostic characters observable indicate area, an(d separated from thiickene(d(ldeig- affinity with either the Leptodactylidae or erous portion by shallow gr(x)v. Single the Hiylilae. Of the six anuran families repre- known vertebrapr(xa(pIreous, with cylindrical sented in South America having an entirely centrum. or partly procoelous vertebral column, the Eophractus tenolency towards a secondlary deposition of castamayerensit, niew species dlermal bone over the temporal area occurs Typjr: A.M.N.HI. No. 3165, associated left only in the two just mentioned. The ascendling nasal and left maxillary. process on the Eophractus maxillary is broa(l HORIZON AND LoCALITY: "B3ird (lay, andl high andi may possibly have an articular Casamayoran StagRe, Lower Loerne. Simpson surface for the squanlosal, but this is not 49 so BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 93 certain. In any case, at least partial roofing Calyptocephdeltak STRAND, 1926, Arch. Natur- of the temporal area is clearly indicated. The gesch., div. A, vol. 92, no. 8, p. 55 (new name size and depth of the maxillary and the form for CalyptocephalusDumeril and Bibron, 1841, of the ascending process are very similar to preoccupied). the condition in Calyptocephalelka, the only GENOTYPE: Calyptocephalus gayi Dum6ril Recent South American leptodactylid with a and Bibron. roofed temporal region. The Australian Cyclorana has a similar broad articulation Calyptocephalella canqueli, new species between the maxillary and squamosal (Par- TYPE: A.M.N.H. No. 3429, partial skele- ker, 1930). Among the Neotropical Hemi- ton consisting of crushed skull, right half of phractinae there are various degrees of roof- pectoral girdle, right fore limb, and first three ing, ranging from Amphignathodon with none vertebrae, also isolated vertebrae and limb to Hemiphractus in which the temporal area fragments belonging to same individual. is completely covered. HoRizoN AND TYPE LOCALITY: Sarmientan The nature of the dermal bone sculpturing Group, Deseadan Stage, Lower Oligocene. is highly variable in