Phoca wymani and Other Tertiary Seals (Mammalia: Phocidae) Described from the Eastern Seaboard of North America SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY CLAYTON E. RAY SERIAL PUBLICATIONS OF THE SMITHSONIAN INSTITUTION The emphasis upon publications as a means of diffusing knowledge was expressed by the first Secretary of the Smithsonian Institution. In his formal plan for the Insti- tution, Joseph Henry articulated a program that included the following statement: "It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge." This keynote of basic research has been adhered to over the years in the issuance of thousands of titles in serial publications under the Smithsonian imprint, com- mencing with Smithsonian ContrAutions to Knowledge in 1848 and continuing with the following active series: Smithsonian Annals of Flight Smithsonian Contributions to Anthropology Smithsonian Contributions to Astrophysics Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoology Smithsonian Studies in History and Technology In these series, the Institution publishes original articles and monographs dealing with the research and collections of its several museums and offices and of professional colleagues at other institutions of learning. These papers report newly acquired facts, synoptic interpretations of data, or original theory in specialized fields. These pub- lications are distributed by mailing lists to libraries, laboratories, and other interested institutions and specialists throughout the world. Individual copies may be obtained from the Smithsonian Institution Press as long as stocks are available. S. DILLON RIPLEY Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY • NUMBER 28 Phoca wymani and Other Tertiary Seals (Mammalia: Phocidae) Described from the Eastern Seaboard of North America Clayton E. Ray SMITHSONIAN INSTITUTION PRESS City of Washington 1976 ABSTRACT Ray, Clayton E. Phoca wymani and Other Tertiary Seals (Mammalia: Phocidae) Described from the Eastern Seaboard of North America. Smithsonian Contribu- tions to Paleobiology, number 28, 36 pages, 3 figures, 11 plates, 1976.—Fossil seal remains from Richmond, Virginia, first reported by Wyman in 1850, and named Phoca wymani by Leidy in 1853, have been neglected and unjustifiably regarded as cetacean by most subsequent authors. Recently recognized parts of the holotype and other material, in part recently collected in Richmond, show that the species is a monachine seal, here called Monotherium? wymani (Leidy, 1853a). It is derived from Miocene beds that are definitely older than the York town Forma- tion and probably correlative with the Calvert Formation of Maryland. Thus Monotherium? wymani is probably the oldest known monachine. Other evidence of fossil phocids in eastern North America is reviewed. OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SI PRESS NUMBER 6139. SERIFS COVER DESIGN: The trilobite Phacops rana Green. Library of Congress Cataloging in Publication Data Ray, Clayton Edward Phoca wymani and other Tertiary seals (Mammalia, Phocidae) described from the eastern sea- board of North America. (Smithsonian contributions to paleobiology ; no. 28) Supt. of Docs. No.: SI 1.30:28 1. Phoca wymani. 2. Phocidae, Fossil. 3. Paleontology—Tertiary. 4. Paleontology—North America. I. Title: Phoca wymani and other Tertiary seals ... II. Series: Smithsonian Institution. Smithsonian contributions to paleobiology ; no. 28. QE701.S56 no. 28 [QE882.P5] 560'.8s [569'.74] 75-619303 Contents Page Introduction .... 1 Acknowledgments 1 Historical Review of Tertiary Phocidae in Eastern North America 2 Phoca wymani Leidy 4 Taxonomic History ...... 4 The Holotype 8 Fate of the Specimens 8 Additional Material ... 10 Localities 10 Geology 12 Description 14 Diagnosis .... .... .... 19 Generic Allocation 19 Leptophoca lenis True 20 Addendum 21 Literature Cited .... 22 in Phoca wymani and Other Tertiary Seals (Mammalia: Phocidae) Described from the Eastern Seaboard of North America Clayton E. Ray Introduction The evidence and literature of pre-Pleistocene fossil seals, family Phocidae, in North America have been extremely skimpy and unsatisfactory. Barnes and Mitchell (in press) have now reviewed and clarified the meager evidence for the Pacific coast. With the exception of Leptophoca lenis True, 1906, the published record for the Atlantic seaboard may for the most part be characterized as at best obscure, confused, and without focus. Presented here is a brief historical review of that record, followed by a detailed account of Phoca wymani and other remains of fossil seals from Richmond, Virginia, with notes on, and illustra- tions of, some other relevant specimens. Speci- mens deposited are in the Museum of Comparative Zoology (MCZ) and in the National Museum of Natural History, Smithsonian Institution, the latter under the catalog numbers of the United States National Museum (USNM). ACKNOWLEDGMENTS.—I wish to thank Farish A. Jenkins, Jr., for the loan of the temporal bones of Monotherium? wymani, MCZ 8741, and for per- mission to prepare them and extract the auditory ossicles of the left side. Elaine Anderson and Ralph Lutz provided the essential information Clayton E. Ray, Department of Paleobiology, National Mu- seum of Natural History, Smithsonian Institution, Washing- ton, D.C. 20560. from Jeffries Wyman's catalog that demonstrated conclusively the identity of the specimens. Barbara Lawrence, Antony J. Sutcliffe, Jane Knapp, Alta Copeland, and Robert W. Purdy supplied useful historical data, as did Robert Mayo, Elizabeth Childs, and Mrs. Stuart Gibson, all of the Valen- tine Museum, Richmond. Peter A. McCrery and associates of the Richmond Gem and Mineral So- ciety salvaged fossil vertebrates from the Ballard Street locality in 1968 and donated them to the Smithsonian Institution. Calvin F. Allison was the first to notice and bring to my attention seal re- mains among specimens from Ballard Street. Franklin L. Pearce and Gladwyn B. Sullivan pre- pared all of the phocid fossils from Richmond. James G. Mead granted free access to the modern pinniped collections and permission to extract auditory ossicles. Paul Sartenaer granted access to the collections of the Institut Royal des Sciences Naturelles de Belgique, and permitted my borrow- ing pertinent specimens. Q. Brett Hendey provided excellent casts of many elements of Prionodelphis capensis. Derek Siddons loaned and permitted casting a partial mandible of Leptophoca lenis. Charles A. Repenning reawakened my interest in the problem, made available auditory ossicles of several species of phocids, and read a draft of the manuscript critically. Thomas G. Gibson, Lauck W. Ward, and Blake W. Blackwelder have been very generous in sharing their knowledge of the SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY stratigraphy and correlation of the Chesapeake series. In the Museum and in several rewarding field excursions, L. W. Ward has imparted in a short time an otherwise unobtainable understand- ing of Virginia coastal plain stratigraphy devel- oped by him through years of work, and not yet published. George W. Andrews processed several matrix samples from Richmond and reported up- on the diatoms. Frank C. Whitmore, Jr., provided previously unpublished data on cetaceans from Richmond and elsewhere, and reviewed the manu- script. Victor A. Krantz, assisted by Franklin L. Pearce, made the photographs from which Law- rence B. Isham and Jeffrey Lund prepared the plates and the figures. The work on which this paper is based was supported by the Smithsonian Institution, in part through the Remington Kel- logg Memorial Fund, the Walcott Fund, and the Smithsonian Research Foundation. Historical Review of Tertiary Phocidae in Eastern North America FIGURE 1; PLATE 5: FIGURES 1-6; PLATES 8-11 The first notice of a fossil phocid in the western hemisphere seems to be that by Richard Harlan (in Conrad, 1838:xi, footnote, Dall reprint, 1893: [13]; Harlan, 1842:143), who mentioned "seal" among fossils identified from excavations for marl on the plantation of Lucas Benners, near the north bank of the Neuse River 15-16 miles down- stream from New Bern, North Carolina. Although recent search has failed to identify with certainty any material on which this record was based, new collections from nearby localities and modern bio- stratigraphic interpretations suggest that Harlan's "seal" might well have been derived from beds of late Tertiary age, possibly equivalent to the York- town Formation (Ray, in press). The second report of a fossil phocid of possible Tertiary age in eastern North America is that of Lyell, apparently first recorded in Murchison (1843:551), and subsequently in a series of publi- cations, mostly of similar content by Lyell (1843: 32, 1845:257). The specimen was identified by Owen as a canine tooth, with the crown fractured, of a seal thought to be allied to the modern hooded seal, Cystophora cristata. It was collected by Lyell, apparently from the greensand, at Gay Head, Martha's Vineyard, Massachusetts (Lyell, 1845:257, fig. 6). These beds have been regarded as Miocene and/or Pliocene, possibly correlative with the Yorktown Formation (Dall, 1894:299; Gibson, 1965:980), but are now regarded as older on the basis of the invertebrates (Gibson, personal communication, 1975) and the vertebrates, prin- cipally Squalodon, which is not known to occur in deposits younger than the Calvert Formation in the Atlantic Coastal Plain (Whitmore, personal communication, 1975). Intrusion of the seal tooth from the Pleistocene cannot be discounted. The specimen has been lost sight of and may have been destroyed. The only other pinniped fossil ob- tained by Lyell on Martha's Vineyard, a rolled rostrum of a probably Pleistocene walrus skull (Lyell, 1845:258, pl. 5: fig. 1), was compared, also by Owen, to modern walrus specimens in the Mu- seum of the Royal College of Surgeons, and was deposited there (Flower and Garson, 1884:203), where virtually the entire collection, excepting specimens out on loan, was destroyed during World War II (Sutcliffe, personal communication, 1972). The seal canine may not have been in the Museum of the Royal College of Surgeons as neither Owen (1845) nor Flower and Garson (1884) listed it, but neither was it listed by Lydekker (1885) in the collections of the British Museum (Natural History), nor did I find it there in reviewing the collections of fossil pinnipeds in 1972. Although its affinities can scarcely be determined unless the specimen is found, it seems safe to say that a spe- cial relationship to hooded seal would be difficult to support solely on the basis of a broken canine, particularly if the tooth were as old as Miocene, and not Pleistocene. However, there is no particu- lar reason to doubt that the specimen was indeed the canine of some seal of late Tertiary age, espe- cially in view of the reliability of Owen and Lyell. Confidence in the record is bolstered further by the recent discovery, in the greensand at Gay Head, by Clifford Kaye, U. S. Geological Survey, of a fragmentary, incomplete right humerus, now USNM 214625 (National Museum of Natural His- tory), tentatively assigned to Monotherium aber- ratum, and suggestive of an age greater than that of the Yorktown Formation (Plates 8-11: figures 1, 2). Monotherium was described from the Dies- tian deposits of the Antwerp Basin, Belgium. The major part of a large right humerus, USNM 25874, NUMBER 28 and a few other elements, from the St. Marys For- mation of Maryland, are assigned tentatively to Monotherium affine. Monotherium has not been recognized among the abundant remains of phocids from the Yorktown Formation. Third, from the Miocene of Richmond, Vir- ginia, Wyman (1850a, b, c) reported fossil seal material that included the basis for Phoca wymani named by Leidy in June 1853. This, with recently collected material from Richmond, is the principal subject of the present report, and is discussed at length below. Fourth, at a meeting of the Academy of Natural Sciences of Philadelphia on 2 August 1853 (the proceedings of which, volume six, numbers 9-10, were published prior to 5 September 1853, the date on which the Smithsonian Institution acknowl- edged receipt of this issue, according to Nolan, 1913:xi), Leidy (1853b) reported upon a new species of fossil seal, the complete text of which follows: Mr. Conrad has presented me with an outline drawing (of which the accompanying wood engraving is a copy,) of a tooth, discovered by Mr. Samuel A. Wetherill in the green sand, of the cretaceous series, near Burlington, New Jersey. The specimen was given to Mr. Conrad, who made the drawing indicated, and afterwards loaned it to an acquaint- ance, from whom he has not been able to obtain it again. The figure represents a double-fanged tooth, with a crown divided into five prominent lobes. It is, without doubt, the tooth of a mammal, and resembles very much one of the pos- terior molars of Stenorhynchus serridens, Owen [—Lobodon carcinophagus], an animal of the seal tribe. It may have belonged to a cetacean allied to Basilosaurus, but until further evidence is obtained, I propose to call the species indicated by the tooth Stenorhynchus vetus. Although the incredible Cretaceous age (un- doubtedly antedating the origin of pinnipeds) has been carried into the later literature by some au- thors as a troublesome anomaly (Kellogg, 1922:68, 115; King, 1964:129), Leidy (1865:1-2, ftn; 1869: 416) had long ago expressed the opinion that the tooth was of Miocene age, and had pointed out as well (1865:1) that Pleistocene mammals were known from the same deposits. The geologic age of many fossils from the New Jersey coastal plain is subject to gross misinterpretation (Ray, 1975: 296-298). However, a crabeater seal in the North Atlantic of any geologic age is inexplicable. The drawing (Figure la) does indeed suggest a tooth essentially identical to a lower post-canine of the FIGURE 1.—Lobodon species: a, the holotype of Lobodon vetus, after Leidy (1853b) (X 1), as confirmed by Leidy (1869: 416), though not indicated by Leidy (1853b); b, lower right fourth postcanine of modern Lobodon carcinophagus, USNM 270385, in lingual aspect (X 1). living crabeater, Lobodon carcinophagus (Figure lb), and dissimilar to any tooth of any other seal. The postcanines of some individuals of Phoca vitulina and of Pusa are rather comblike, but are much smaller, lower-crowned, fewer-cusped, and lack the posteriorly directed apical hook of the principal cusp seen in some teeth of modern Lobo- don and apparently in the fossil. Efforts to explain this tooth away, for example as an abraded shark tooth by Murray (1866:124), have not been suc- cessful (Allen, 1880:475^76). As the name has been validly proposed for a seemingly distinctive species, it would seem best, following the lead of Leidy (1869:415), to retain it as a species inquir- enda in the form of Lobodon vetus (Leidy, 1853), of Miocene or later age, pending recovery of the holotype or discovery of new material. One does not work long with fossil marine mam- mals before becoming aware of the chronic con- fusion between remains of cetaceans and seals, from the earliest times at least through the nine- teenth century, if not to the present (see for ex- ample the synonymies of various squalodonts in Kellogg, 1923), particularly pernicious being the penchant of authors until recently to name taxa on the basis of isolated teeth. In some instances it has been possible to resolve the resultant problems, but there remain in limbo too many troublesome taxa, based originally upon nondescript teeth, and subsequently shuttled inconclusively about among various cetacean and pinniped assignments. Among these are Phoca debilis Leidy, 1857 (usually cited as 1856, but the publication not received at the Boston Society of Natural History earlier than 25 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY April 1857, according to Nolan, 1913:xi), and Phoca modesta Leidy, 1869, both named from small isolated teeth of unknown age in the collec- tions of the Academy of Natural Sciences of Phila- delphia, from the Ashley River deposits of South Carolina. Kellogg (1923) listed them as Delphino- don? debilis (p. 13) and Phoca? modesta (p. 26) respectively, and he (1922:127, 128; 1923) and Hay (1902:589) cited the earlier literature. In my opinion both taxa represent small porpoises, neither Delphinodon nor Squalodon, whose affini- ties may or may not be determinable by thorough comparison with other taxa, including modern representatives, but which in any event are not pinnipeds and thus are outside the scope of this paper. The fifth record is that of Leptophoca lenis True, 1906, from the Calvert Formation of Mary- land (and referred material from Virginia), thus far the only eastern North American Tertiary phocid adequately understood in the literature and based upon reasonably satisfactory material, though certainly not "the first authentic remains of American fossil seals" as claimed by True (1905). Considerably more material has accumu- lated since True's description, reinforcing the pho- cine character of the species. This will be reported elsewhere, with the exception of two specimens from Richmond reported herein, one of which, a partial humerus, is illustrated alongside the holo- type (Plates 8-11: figures 3, 4) and a mandibular ramus illustrated for comparison with monachine mandibular fragments from Richmond and North Carolina (Plate 5: figures 1-6). From 1906 onward has been a period of quies- cence for eastern North American Tertiary pinni- peds, completely so in terms of new, published information, and predominantly so in terms of additions to collections, until the past decade, during which collections have grown at an accel- erating rate. New material is known from the Cal- vert, Choptank, and St. Marys Formations, and elsewhere, and most abundantly from the York- town Formation (Ray, in press). Now, on the threshold of significant advancement of knowledge of the subject after more than half a century with- out progress, it seems particularly fortunate to be able to reinstate among the pinnipeds a species that was among the earliest named and most se- curely founded Tertiary phocids, but which has languished, not forgotten, but misused, among the Cetacea virtually from its conception. Phoca wymani Leidy, 1853, had aroused my curiosity years ago, but so long as its holotype was unrec- ognized and presumed missing, and additional material was unknown, there seemed little pros- pect of clarifying its position. These handicaps have now been in part overcome by recovery of at least part of the original material and by discovery of a few additional specimens. Phoca wymani Leidy FIGURES 2, 3; PLATES 1-7 TAXONOMIC HISTORY.—Jeffries Wyman was a native and lifelong resident of Massachusetts, ex- cepting for travel and frequent wintering in the southern United States for reasons of health (see Gray, 1875, for this and most of the biographical information about Wyman). His specific associa- tion with Richmond, Virginia, stemmed largely from his occupying "the chair of anatomy and phy- siology in the medical department of Hampden- Sidney College, established at Richmond, Virginia," from 1843-1847, where he spent the winter and spring of each year in teaching. This was termi- nated in 1847 by his appointment to a position at Harvard, where his affiliation continued until his death on 4 September 1874. It was undoubtedly through acquaintances made while at Hampden- Sidney College (its medical department, since 1853, the Medical College of Virginia, Rucker, 1950:292) that he received the fossils collected by Dr. Martin Burton in Richmond, which in part form the primary subject of the present report. In the Proceedings of the Boston Society of Natural History for 6 February 1850 (Wyman, 1850a:24l), it is recorded that "Prof. Wyman ex- hibited some fossil bones of Seals found in the Miocene deposit beneath the city of Richmond, Va., where they occur in company with the teeth of Sharks and Zeuglodonts [squalodonts?]." Again, in the meeting of 7 August 1850 (Wyman, 1850b: 323), "Prof. Wyman announced that he had re- ceived other fossil remains of seals from the Mio- cene deposit of Virginia, near Richmond .... He had a large portion of a cranium, of a well known genus, but of a species not yet ascertained. As these bones were found at some distance from each NUMBER 28 other, farther discoveries would probably be made." In November of the same year he reported upon vertebrate remains from Richmond, mostly collected by Dr. Martin Burton of Richmond; the complete text on the seal remains (Wyman, 1850c: 229-230) is as follows: Among the most interesting of the relics discovered by Dr. Burton were parts of the cranium of an animal belong- ing to the natural family of Phocidae, a family of which but few remains had been previously detected, and in so far as I have been able to find any record, only in one other locality in the United States. The bones were fragile, and had evidently been crushed previous to exhumation. The pieces in my possession consist of two temporal bones nearly entire, a fragment including a portion of the parietal and occipital bones; and in addition a part of the base of the skull. The reentering angle of the occiput, the well marked depressions corresponding with the cerebral convolutions on the parietal bones, the form of the cranial cavity, the deep fossa above the internal auditory foramen, the vascular canals opening on the occiput, and the inflated tympanic bones, all indicated an affinity to the Phocidae. The size varied but little from that of the common Harp seal, (P. Groenlandica.) The presence of an interparietal crest indi- cating a large development of the temporal muscles, offers a diagnostic sign by which it may be distinguished from P. barbata, P. Groenlandica, P. hispida, P. mitrata, and P. vitulina. From those species of seals which are provided with a crest the fossil presents a well marked difference in having the mastoid process much larger, more rounded, and promi- nent, nearly equalling the tympanic bone in size. The en- trance to the carotid canal is in full view when the base of the skull is turned upwards. The imperfectly divided canal which lodges the Eustachian tube and the tensor tympani muscle is of remarkable dimensions, especially when com- pared with that of P. Groenlandica. The interparietal crest extending from the occiput to the anterior edge of the frontals, is most narrow posteriorly where it is but slightly elevated above the surrounding bones. The fragments of cranium above described were found in the Shockoe creek ravine near the base of Church Hill. In the ravine at the eastern extremity of the city and in the neighborhood of [end p. 229, begin p. 230] the penitentiary, Dr. Burton obtained several other portions of the skeleton of another seal. These consisted of an imperfect cervical vertebra, a lumbar vertebra nearly entire, a fragment of the sacrum, coccygeal vertebra, fragments of ribs and the lower extremity of a fibula. Their generic characters have been satisfactorily made out by comparisons with recent bones. In figure 1, page 232, I have represented the coccygeal vertebra which corresponds in its general characters very accurately with recent bones of P. Groenlandica from the same region of the vertebral column. The small size of the vertebral canal and the imperfect transverse process, the wide spread articulating processes and the blunted spinous process indicate its affinity to the seals. The fragment of a left fibula (figs. 2 and 3) presents at its lower extremity (fig. 3), an oblique regularly concave articulating surface, on its inner face, and on its outer (figs. 2 and 3,) an elevated ridge or crest on either side of which is a groove for the passage of a tendon. Leidy (1853a:8) lists among the Miocene mam- mals of North America: Phoca Wymani, Leidy. Wyman: Am. Journ. Sc. 1850, X. 229. In my opinion this constitutes valid publication of the species under the International Code of Zoo- logical Nomenclature (1961), in that it satisfies Articles 11, 12, and 16, the last of these by "indica- tion" of "a bibliographic reference to a previously published description, definition, or figure." The month and year of publication of Leidy's "The Ancient Fauna of Nebraska . . ." are June 1853 (see Rhees, 1882:8), the latter not 1854 as com- monly cited; the whole volume six of Smithsonian Contributions to Knowledge, of which Leidy (1853a) is part seven, was completed in 1854, as indicated on its title page. In 1857 Leidy (p. 265) published "Notice of remains of two species of Seals" (not two new species!), of which the part relevant here is as follows: 1. Phoca Wymani. Remains of a Seal. Wyman, Am. Jour. Sci. X. 229. Phoca Wymani, Leidy. Anc. Fauna of Nebraska, 8. A tooth, apparently an inferior canine, from the miocene deposite [sic!] of Virginia, recently presented to the Academy by Prof. Tuomey, I suspect to belong to the same species as the remains of a seal from the same deposite, described by Prof. Wyman. It is abundantly clear that Leidy did not regard this as the type description, but as the referral (retracted in 1869) of a specimen to a previously established species. Certainly Prof. Tuomey's tooth from Virginia did not contribute to Leidy's concept of Phoca wymani when he named it in 1853, as the tooth was not donated to the Academy of Natural Sciences of Philadephia until 2 September 1856 (anonymous, 1857:xviii). The proceedings of a meeting of the Academy of Natural Sciences of Philadelphia on 5 November 1867, record (Cope, 1868a: 132; see Nolan, 1913: xiii, for year of publication) that "Squalodon mento Cope was characterized from four molar teeth, which were between two and three time[s] as large as those belonging to the Squalodon wy- SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY manii (Phoca of Leidy) with similar incurved crowns, but much more rugose." Again, more formally, in a paper ordered to be published at a meeting of 31 December 1867, Cope (1868b: 152) stated as follows: Squalodon wymanii m[ihi], Phoca wymanii Leidy. Pro- ceedings Academy N. Sci. 1856, 265. Of this, the smallest species of the genus, three premolar teeth are in the collection, and the type specimen is in the Academy's Museum. . . . These two publications by Cope in 1868 mark the first use of the name Squalodon wymanii (also listed by Cope, 1868b: 144), the first as a nomen nudum, but the second apparently validly pro- posed. In my opinion Cope did not intend to appropriate the species Phoca wymani to the Cetacea, as he did not refer to Leidy's (1853a) type description, but only to the paper in which he (Leidy, 1857) referred the tooth from Virginia, which Cope designated the holotype of Squalodon wymanii, indicating himself as the author of the species. In 1869, Leidy explicitly separated the concepts of the original Phoca wymani and the interloping cetacean. He listed (p. 415) the former as follows: Phoca Wymani. Animal belonging to the Phocidae, Wyman: Am. Jour. Sc. 1850, X, 229, Figs. 1-3. Phoca Wymani, Leidy: Anc. Faun. Neb. 1853, 8. Remains found in a miocene formation at Richmond, Vir- ginia. His account of the latter (pp. 425-426) is in part as follows: Delphinodon Wymani. Phoca Wymani, Leidy: Proc. Ac. Nat. Sc. 1856, 265. Squalodon Wymani, Cope: Proc. Ac. Nat. Sc. 1867, 132, 151, 152. Three teeth, from the miocene formation of Charles County, Maryland, ascribed by Prof. Cope to a species of Squalodon, appear to me, at least in part, to belong to a smaller species of Delphinodon. One of the teeth . . bears a resemblance to that first described of the larger species .... A tooth [ANSP 11225, the "type specimen" of Cope, 1868b, above] . . from the miocene formation of Virginia, orig- inally ascribed by me to the same species as the remains of a Seal described by Prof. Wyman, is very like the one above indicated . . . The remaining two teeth from Charles County, ascribed by Prof. Cope to Squalodon Wymani, are different from the preceding, and it is uncertain whether they belong to the same animal .... Thus Leidy and Cope scrupulously maintained the separate identity of the pinniped and the cetacean. Van Beneden (1877:28) merely men- tioned Phoca wymani as a form described early but not well characterized. Allen (1880:470-473, 480) discussed Phoca wymani at length in a speculative manner, which, by weight of his authority, served largely to con- demn Phoca ivymani to nearly a century of neglect and misinterpretation. He stated first (p. 470) that: some supposed Phocine remains were described by the late Professor Wyman from the Tertiary deposits underlying the city of Richmond, Virginia. They came from two localities, and consisted of quite different materials. The specimens are at present unknown, so that their reexamination is im- possible. A part of these remains were in all probability Squalodont, while others may have been Phocine. Further (p. 470), under the heading "Squalo- dont Remains described as Phocine," he indicated: No less than three species referred originally to "Phoca" are in all probability referable, in part or wholly, to Squalodon, as is more or less explicitly admitted by their original de- scriber. These are Phoca wymani, P. debilis, and P. modesta, of Leidy. On page 471 he quoted Wyman's (1850c:229) description of the skull of Phoca wymani, followed by the assertion: In the description above given there is nothing to prevent the supposition that these cranial fragments are referable to a small species of Squalodont. If, however, they are really Phocine, they represent a type very unlike anything at pres- ent known, either existing or extinct. But other remains are described by Professor Wyman, from the same locality, and in the same paper, which do not seem to admit of such an interpretation. He then quoted Wyman's description of the post- cranial remains, followed (pp. 472-473) by the commentary that: the specimens here described do not appear to have been preserved, or to have been seen by subsequent writers, but Professor Wyman was an osteologist of too well-known pro- ficiency to admit of the supposition that these remains did not present well-marked Phocine affinities. Indeed, his de- scription and rude figures of the fibula above mentioned show clearly that its affinities were rightly interpreted. The vertebra is not so evidently Phocine. Three years later the description of these remains became the basis of Dr. Leidy's "Phoca wymanii," who, in proposing the name, merely cited Wyman's description. In 1856 [1857] he referred to it a tooth. ... In 1867 [1868] Professor Cope referred Phoca wymani, Leidy, to Squalodon. . . . The Squalodon wymani NUMBER 28 of Cope thus, inferentially at least, includes the remains described by Wyman, though direct reference seems to be made only to the tooth referred by Leidy to his Phoca wymani in 1856, and which is that of a squalodont. The Phoca wymani, if not originally a composite species, as was in all probability the case, certainly became so in 1856. In 1869 Dr. Leidy retained, under the name Phoca wymani, the specimens above mentioned as described by Wyman in 1850, separating the tooth referred by him to this species in 1856 under the name Delphinodon wymani. In concluding his account of fossil pinnipeds, Allen (p. 480) stated that: in North America few remains of Pinnipeds have been found, and these, with two exceptions, are all from the Quaternary, and are referable to existing species. The excep- tions are the so-called Phoca wymani, based in part at least upon veritable Phocine remains from the Miocene of Rich- mond, Virginia, and the enigmatical Lobodon vetus .... Judging from the foregoing statements, Allen vacillated radically in his thinking about Phoca wymani; much of what he said is contradictory or wrong. For example: Leidy never "admitted" that Phoca wymani was referable to Squalodon; Allen trusted Wyman's interpretation of the postcranial elements, but cast doubt on that of the skull; the Squalodon wymani of Cope did not, inferentially or otherwise, include the remains described by Wyman; erroneous subsequent (1857) referral of a specimen to a species in no way makes the species composite. Roger listed "Ph. [Phoca] Wymanni," mis- spelled, in 1887 (p. 145), and "Ph. Wymani," correctly in 1896 (p. 74). Toula (1897:49, 53, 55) reviewed the history of Phoca wymani, noting Cope's and Allen's (1880) conclusions in large part correctly, while retaining the species among the fossil seals. Trouessart (1897:388, footnote) indicated under Pinnipedia, "Phoca Wymani, Ph. debilis, Ph. modesta, Leidy (1853-1869) sunt dentes Squalodontium (Cetacea); vide infra: Squalodon Wymani," but when the Cetacea were published (1898:1023) listed it as Delphinodon Wymanii Leidy, 1856. Hay (1902:591) indicated that "the name Del- phinodon leidyi is intended to replace D. wymani Leidy .... The type of D. leidyi is Leidy, J. 1869 A, pl. XXX, fig. 12 [ANSP 11225]." The new name must have been proposed under the erro- neous belief that Phoca wymani and Delphinodon wymanii were specific homonyms, which they were not and are not. In my opinion Delphinbdon leidyi was an unnecessary replacement name and should be relegated to the junior synonymy of Delphinodon wymanii (Cope, 1868b). Hay's biblio- graphic citations under D. leidyi (p. 591) and Phoca wymani (p. 785) do not reflect the distinc- tion between the two—for example, none of the references to teeth (Cope, 1868a, b; Leidy, 1857) has anything to do with Phoca wymani, and no reference to Wyman's 1850 material has anything to do with Delphinodon wymanii. Hay did appar- ently understand the real separation of the two taxa, as evidenced by his listing them separately and in part distinguishing their respective litera- ture, and by listing Phoca wymani again in 1930 (p. 564). Trouessart (1905:762) followed Hay, and made no mention of the species under Phoci- dae (1904:282-288). True (1912:185-186; confirmed by his unpub- lished notes) obviously accepted Allen's (1880) interpretations, implying that Leidy (1856 [ = 1857]) was a validation of Leidy (1854 [ = 1853a]), not the mere referral of a specimen as in fact it was. He also accepted Hay's replacement name, Delphinodon leidyi, for the cetacean, but he did recognize that the seal material of Wyman (1850) was a separate entity. Kellogg (1922:74, 75, 120, 130) perpetuated the unfounded notion that Wyman's original material described as seal was at best composite, and in part cetacean (squalodont). Wilson (1935:126), without reference to Phoca wymani as such, accepted Wyman's (1850c) re- port as a valid record of fossil phocid remains, although erroneously supposing it to be the first for the United States. The taxonomic history of Phoca wymani cannot be concluded without reference to the work of William Palmer, a taxidermist and naturalist in the United States National Museum, and an active collector of Miocene vertebrates along Chesapeake Bay approximately from 1907 until his death in 1921 (Gilmore, 1941:337). His prowess as collector has been adequately noted in published acknowl- edgments (as in True, 1912:166) and in one case by specific epithet (Parietobalaena palmeri Kel- logg, 1924), but his taxonomic insights have gone unremarked. In one of those rare occurrences that occasionally rewards antiquarian research, I found, long after reaching my own conclusions on the literature of Phoca wymani, in perusing notebooks SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY found among the late Remington Kellogg's papers, that William Palmer had covered essentially the same ground more than 50 years earlier, from the viewpoint of his interest in Delphinodon, and had written up his results in a formal manner under the title "Fauna Calvertensis, no. 2. The Fossil Cetacean Genus, Delphinodon." This projected series of publications never materialized, not even the first, which was to have been the description, under almost the same generic name, of the mate- rial ultimately described as Parietobalaena palmeri Kellogg, 1924. In a note written in May 1920 and placed with the holotype tooth of Delphinodon wymani in the collections of the Academy of Nat- ural Sciences of Philadelphia, Palmer recorded the synonymy of that species. The remarkable con- gruence of our conclusions would make it a pleas- ure to enter his name as coauthor of the present notes were he not so many years removed from the option of declining. However, it seems no dis- service to quote the following excerpt from his notes, which will reveal how perceptively he analyzed the main problems and how succinctly he stated them: Delphinodon wymanii Owing largely to careless citations the naming of this species has been remarkably confused. It was described [in the morphological, not taxonomic, sense] but not named by Leidy in 1856 in conjunction with a reference to remains of a seal described though not named by Prof. Wyman in 1850, but named and not described by Prof. Leidy in 1853 as Phoca wymani. It is clear that the porpoise tooth collected by Prof. Tuomey was without a name in 1856. We next find it named by Prof. Cope in 1867 as Squalodon wymanii. On page 132 occurs, "Squalodon wymanii (Phoca of Leidy)." This is a nomen nudum here. On page 144 occurs, "Squalodon wymanii. Cope, miocene." Again a nomen nudum. On page 152 he writes, "Squalodon wymanii m[ihi]. Phoca wymanii Leidy." He mentions the type, compares it with other specimens, and states that it is the smallest of the genus and distinctly uses a new valid name, the first for this type. Leidy had never used wymanii but had incidently discussed this porpoise tooth under the name of Phoca Wymani which of course rightfully belonged to the "seal remains" described by Wyman and named by Leidy in 1853, the first fossil seal named in North America. It is clear that Cope correctly identified this specimen as a cetacean tooth and his language clearly shows that he had written a new name. THE HOLOTYPE.—Leidy (1853a:8) seems clearly to have had in mind the cranial material as the basis for his name Phoca wymani, as he cited the page reference as 229, on which only the cranium is described, not 230, on which the postcranial remains are described, nor the figures, on page 232, in which the latter are in part illustrated. Further, Toula (1897:53) wrote that "Phoca Wymani Leidy ist auf zwei Paukenbeine begrundet; eines der Stticke weist auch noch ein Stuck des Parietale und Occipitale auf. (Wyman: Amer. Journ. 1850, 229.)." If Leidy's intent were not regarded as suffi- ciently definite, then Toula's statement should qualify as subsequent designation of a lectotype, or, if not, then certainly True's (1912:186) state- ment that "in 1902, Dr. O. P. Hay pointed out the fact that the name Phoca wymani really belonged to the seal skull originally described by Wyman in 1850, and could not properly be applied to the teeth from Virginia and Maryland" would so qualify, even if he drew more than was warranted from Hay's bibliographic annotations. In any case, the prerogative of lectotype selection was not avail- able to Kellogg (1922:74) when he asserted, "The form Phoca wymani is here restricted to the fibula and vertebra from the ravine outside of the city limits of Richmond, Virginia." Thus, I feel that a conclusive case exists for the cranial remains as holotype by original indication (Leidy) or as lecto- type by subsequent designation (Toula, Hay, or True), and that Kellogg's later attempt is without force. FATE OF THE SPECIMENS.—Apparently no one has knowingly restudied Wyman's phocid speci- mens since his original reports of 1850. Although Leidy borrowed other specimens from Wyman, in- cluding some of cetaceans from Richmond (see, for example, Leidy, 1869:426, 432, 439; much or all of this material, in part still bearing Wyman's catalog numbers, is now in the collections of the Academy of Natural Sciences of Philadelphia, under the numbers, ANSP 11227, 11257, 11263, and 11268), there is no indication that he ever saw the material on which he based the name Phoca wymani, nor apparently did any other au- thor who wrote about it until more than 100 years later, when I (1958:441) unwittingly reported upon the two temporal bones, cataloged in the collections of the mammal department of the Mu- seum of Comparative Zoology under the number MCZ 8741, labeled "Monachus?, S.C.?, R. W. Gibbes coll. ?." At that time I tentatively accepted NUMBER 28 the questioned data, and suggested that it provided a northerly (Pleistocene?) record for the monk seal. There the matter rested until 1971, when, in connection with comprehensive studies of Tertiary pinnipeds of the Atlantic Coastal Plain, and at the urging of Charles A. Repenning, my interest in Phoca wymani was reactivated. Restudy of the literature suggested that MCZ 8741 might con- ceivably be part of the material. Borrowing and reexamining these temporal bones showed that they did in fact differ somewhat from those of modern Monachus, and showed as well the pres- ence of the faint but legible old numbers 825 on both specimens. At my request, Elaine Anderson, with the aid of Ralph Lutz, then of the Boston Museum of Science (successor to the Boston Soci- ety of Natural History) found in the library of that institution the Catalog of Jeffries Wyman's Collection of Comparative Anatomy in Boylston Hall (at Harvard University). The entries for fossil seal remains in the catalog are as follows: 824. A mass of clay presenting a cast of the interior of the cranium of a seal, from the Tertiary at Richmond, Va. [illegible word]. 825. Right and left temporal bones, with a portion of the parietal bones fitting the preceding specimen. From Dr. Martin Burton of Richmond. 826. Lumbar vertebrae of a fossil seal found at Richmond. From Dr. Martin Burton. 827. Fragment of the sacrum of a seal. Fossil. From Dr. Martin Burton. 828. Cervical vertebra of a seal. Fossil. From Dr. Martin Burton. 829. Dorsal vertebra of a seal. Dr. Burton. 830. Caudal vertebra of a seal. Dr. Burton. 831. Lumbar vertebra of a seal. Dr. Burton. 833. Ribs of a seal from Dr. Burton. 834. Metatarsal bone of a seal from Dr. Burton. 844. Extremity of the fibula of a seal, from Richmond. Presented by Dr. Burton. [This last entry lined out and replaced by an unrelated specimen.] During recent curation of the collections of fossil marine mammals in the National Museum of Natural History there were found two casts of the distal part of a single left fibula of a seal, one with the number 929 and the other 930, from the old catalog of vertebrates, now in the Division of Mammals, NMNH, entered in 1852 as follows: Fossil seal cast; Richmond, Va.; bone of hand; Dr. J. Wyman. These are noted as item 36 in the "List of Additions to the Museum of the Smithsonian Institution during the year 1851" (Baird, 1852:62). They are now cataloged in the collections of the Department of Paleobiology under the single num- ber USNM 214650. Possibly representing the same bone may be the cast "of the inferior extremity of the radius of a fossil mammal obtained near Richmond, Va. From Dr. J. Wyman." listed among "Donations to Museum" at the stated meeting of 5 January 1847 of the Academy of Natural Sciences of Philadelphia (Anonymous, 1847:141). The cast has not been found in recent examinations of the collections of the Academy. Thus far then, of the seal remains in Wyman's catalog, only the temporal bones, no. 825, now MCZ 8741, and the casts of the distal part of the fibula, no. 844 (stricken), now USNM 214650, have been relocated. We know by Wyman's own assertion (Gray, 1875:105) that, during the sum- mer of 1874, "he had gone through his own museum of comparative anatomy, which had some- what suffered in consequence of the alterations in Boylston Hall, and had put the whole into perfect order," just prior to his death on 4 Sep- tember 1874, and that "The collections . . . forming a part of the late Prof. Wyman's Anatomical Mu- seum in Boylston Hall, have been deposited in the Museum [of Comparative Zoology] by the corpo- ration of the College [Harvard]" in 1875 (Agassiz, 1876:8). The entry for the temporal bones in the catalog of the mammal department of the MCZ is in the handwriting of G. M. Allen (Barbara Lawrence, prrsonal communication, 1975), and was made probably in 1907-1909, during his first, part-time employment at MCZ, when his chief activity was curation of the mammalian osteologi- cal collections, including fossils, which he com- pleted by mid-1909 (Henshaw, 1908:7; 1909:8), obviously after the deterioration of the "perfect order" of Wyman's collection of comparative anatomy, else he would undoubtedly have salvaged Phoca wymani. G. M. Allen was Wyman's younger cousin and said to resemble him in personal man- ner, including meticulous scholarship (Barbour, et al., 1943:299, 303, 304). Not so readily under- stood is the role of J. A. Allen, who surely was in a better position to have saved Phoca wymani for posterity than anyone could ever be again. On arriving at the MCZ in 1862 he was "to attend the course by Jeffries Wyman on comparative anatomy" 10 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY (Allen, 1916:8), and he mentions Wyman as "a frequent caller" in Agassiz's laboratory where he worked (p. 9). Wyman and Allen overlapped at Harvard from 1862 until Wyman's death in 1874. Both were active and held various responsible positions in the Boston Society of Natural History during the period (Gray, 1875:106; Allen, 1916: 44). Allen was curator of birds and mammals in- cluding fossils) both in the MCZ and the Boston Society at Wyman's death and at the time of trans- fer of his collections to the MCZ. Allen had published major papers on pinnipeds prior to 1874 and began work for his "History of North American Pinnipeds" no more than three years thereafter. All of these factors notwithstanding, he stated in 1880 that "the specimens are at pres- ent unknown, so that their reexamination is impossible" (p. 470) and "the [postcranial] specimens here described do not appear to have been preserved, or to have been seen by subse- quent writers" (p. 472). It is difficult to imagine how he could have failed to see and preserve the specimens. ADDITIONAL MATERIAL.—On 18 January 1968, vertebrate remains were salvaged from excavations in downtown Richmond (Figure 2, locality 5) by Peter A. McCrery and other members of the Rich- mond Gem and MineralrSociety, including Rich- ard May, William Packard, and Donald Woolford, and were donated to the Smithsonian Institution by the Society. Some months later in the vertebrate paleontological laboratory of the Smithsonian Institution, Peter McCrery, working as a volunteer, prepared out the most promising specimen that turned out to be the major part of the skull of the small porpoise, Kentriodon pernix, now USNM 171077. Nothing further was done with the collec- tion until the fall of 1973 when Calvin Allison, another volunteer in the Smithsonian laboratory, discovered the distal part of a phocine humerus, now USNM 187409, loose in a box of miscella- neous material from the site. This led to close examination of all remaining material, which yielded a large cetacean lumbar vertebra, a few fragments of fishes, and most importantly, from a single block of matrix, several parts, poorly pre- served, probably from a single individual of a monachine seal, all now USNM 187410, including a very incomplete right mandibular ramus, a left ulna lacking the distal epiphysis, a left tibia and most of the shaft of a left fibula, each lacking the distal epiphysis, a left ectocuneiform, and several fragments of ribs, all carefully salvaged and skill- fully restored by F. L. Pearce. A right lower canine tooth from the same block seems not to belong with this material. LOCALITIES.—Various sources for Phoca wymani have been cited in error, for example, "Carolina merid." by Trouessart (1898:1023) and "Mary- land" by Trouessart (1905:591) and by Hay (1930:564). In 1958 (p. 441) I erroneously re- corded the source of the temporal bones as "South Carolina?" on the basis of queried entries in the MCZ catalog—to be ignored now that the true identify of the specimens has been recognized. Wyman (1850c: 229) recorded the locality for the partial cranium as "Shockoe creek ravine near the base of Church Hill." A plan of Richmond made by Charles S. Morgan in 1848, showing the principal topography of the area by hachures, places the base of Church Hill on the Shockoe Creek (northwest) side approximately at Twenty- first Street, and the southwest side, paralleling the James River, near Franklin Street, essentially similar to the present situation. Broad Street, paralleling Franklin Street, and trending approxi- mately S 53.5° E, traverses the center of Church Hill as labeled on the map of 1848. The inter- section of Broad Street and Twenty-first Street is central to the Shockoe Creek side of the base of Church Hill. Thus the type locality of Phoca wy- mani may be fixed as "Shockoe Creek ravine near the base of Church Hill," at or near the inter- section of Broad Street and Twenty-first Street (Figure 2, locality 1). According to Wyman (1850c:229-230), "In the ravine at the eastern extremity of the city and in the neighborhood of the penitentiary, Dr. Burton obtained several other portions of the skeleton of another seal." These included all the postcranial remains listed in Wyman's catalog (above), of which only the casts of the fibula in the USNM have been relocated at this time. His reference to "portions of the skeleton of another seal" suggests a single individual, but reference to "the ravine at the eastern extremity of the city" and "the neigh- borhood of the penitentiary" entails two widely separated localities. The ravine in question can only be that of Bloody Run, bordering Chimbor- azo Hill on the west (Figure 2, locality 2), as the NUMBER 28 11 1000 0 i—i i—< r=r |vw CONTOUR INTERVAL 10 FEET FIGURE 2.—Map of a part of the city of Richmond, Virginia, based on the Richmond Quad- rangle, 7.5 minute series (topographic) map, 1968, of the U. S. Geological Survey. (Segments of the city limits as of 1848 according to Morgan are indicated by a heavy broken line. The line A-A' indicates the line of section along the Church Hill Tunnel, shown in Figure 3. The line of section beyond the ends of the tunnel (broken) is imprecisely known. Localities for fossil pinnipeds and other vertebrates are as follow: 1, the type locality of Phoca wymani; 2, the ravine (of Bloody Run) at the (1848) eastern extremity of the city; 3, the neighborhood of the penitentiary; 4, downtown expressway excavations, Byrd Street between Third and Fourth streets; 5, Ballard Street, now Fourteenth Street, between Grace and Broad streets.) easternmost boundary of the city at that time fol- lowed the middle of Bloody Run ravine (Morgan, 1848). The penitentiary in question could not have been the Libby Prison of Civil War infamy, which stood on the south corner of Twenty-first and Cary Streets, but which was in 1850 a ship chandlery and tobacco warehouse, having been commandeered as a prison only after the first battle of Bull Run (21 July 1861), and dismantled and moved to Chicago shortly after the War. Instead the "penitentiary" in question undoubtedly was the state penitentiary which then (Morgan, 1848) as now occupied an irregular area bounded ap- proximately by Belvidere, Byrd (now the down- town expressway), Second, and Spring Streets, (Figure 2, locality 3), and at that time directly on the western boundary of the city, approximately 1.5 miles slightly north of west of the type locality. In February 1975, vertebrate remains, representing Squalodon, were again encountered in this area, less than 0.25 mile east of the penitentiary, at 113 feet above sea level in excavations for the down- 12 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY town expressway (Whitmore, personal communi- cation), immediately north of Byrd Street between Third and Fourth Streets (Figure 2, locality 4). Thus, Wyman's "several other portions of the skel- eton of another seal" probably came in part from Bloody Run ravine (locality 2) and in part from near the penitentiary (Locality 3), at the then eastern and western limits of the city. The material salvaged in 1968 by the Richmond Gem and Mineral Society was obtained from a road cut made in widening the former Ballard Street, now Fourteenth Street, from two to four lanes, in the block between Broad and Grace streets, on the west side of Shockoe Creek ravine, directly opposite, and approximately 0.5 mile northwest of, the type locality, on the east side of Shockoe Creek ravine (Figure 2, locality 5). A por- tion of this roadcut remains exposed along the west side of Fourteenth Street. GEOLOGY.—Hay (1930:564) listed Phoca wymani as "upper Miocene (Calvert); Maryland," and Clark and Miller (1912:167) listed Phoca wymani from the Calvert Formation of Richmond, but the presence of other Neogene beds in Richmond was not understood at that time. The exact horizon from which any of the fossil pinnipeds of Rich- mond was derived is unknown, but only Paleocene, Miocene, and Quaternary deposits are present. However, as no phocid is known otherwise before the Miocene, that may be taken as the lower limit stratigraphically in Richmond, and the Paleocene beds of Richmond, including those known in and near the valley of Shockoe Creek (Aquia Forma- tion of Darton, 1911:16-17, pl. 1; Nanjemoy For- mation of Daniels and Onuschak, 1974:21, pl. 3; definitely Aquia Formation, and largely if not entirely Paleocene, Hazel, 1969:C64, and Ward and Blackwelder, personal communication, 1975), may be eliminated from serious consideration. There is no evidence of Oligocene deposits in Richmond. The latest Tertiary or Quaternary sands and gravels capping Church Hill and other uplands and the Quaternary alluvium flooring the lower end of Shockoe Creek Valley, the James River Val- ley, and other lowlands, are virtually nonfossili- ferous and largely nonmarine (Daniels and Onus- chak, 1974:28-36, pl. 3; Lafayette Formation and Columbia Group of Darton, 1911:27-31, pl. 1), and are thus exceedingly improbable sources for the fossil phocid material. Here and there in the older literature the indi- cation "Yorktown" age or epoch occurs in con- nection with the Miocene of Virginia (Cope, 1868a: 131; 1868b: 138, 150) and in a few instances explicitly with Phoca wymani (for example, Dana, 1863:521; Guiscardi, 1871:8). The "Yorktown Epoch" was a regional time-term proposed and used by Dana but not widely adopted, essentially coextensive with the Miocene as then understood, and applied to the time of deposition of rocks of the Chesapeake group (Calvert, Choptank, St. Marys, and Yorktown formations). Most of the evolution of the stratigraphic nomenclature of the Chesapeake group in Virginia was noted by Mans- field (1948:3-4); the Yorktown Formation of modern usage was named formally by Clark and Miller only in 1906 (p. 19) and is now regarded in whole or in large part as Pliocene in age. The westernmost known occurrence of the Yorktown Formation in the Richmond area is at Quinton, New Kent County, approximately 18 miles due east of Richmond (Ward and Blackwelder, per- sonal communication, 1975). Thus the Yorktown Formation is not a possible source of Phoca wymani. The probable source beds for fossil pinnipeds in Richmond are limited to the strata of Miocene age, younger than the time of deposition of the Aquia Formation and older than that of the York- town Formation. They have been assigned tradi- tionally to the Calvert Formation (Darton, 1911: 18) or Calvert and questioned St. Marys Forma- tions (Daniels and Onuschak, 1974:21). These Miocene beds overlying the largely diatomaceous layers correlated with the Calvert Formation of Maryland are well represented in sections in and near Richmond, are under regional biostrati- graphic study at present, and are assignable neither to the Calvert nor to the St. Marys Formation (Ward and Blackwelder, personal communication, 1975). The beds in Richmond universally assigned to the Calvert Formation include a conspicuous dia- tomaceous layer, visible at distance on both flanks of Shockoe Creek ravine because of its light color on weathered outcrops and its sparse vegetation. This bed was made known by W. B. Rogers in 1841, and has been written about extensively since (see Roberts, 1942, for citations to the many pub- lications and reprintings). It has been correlated NUMBER 28 13 FIGURE 3.—Vertical section from northwest to southeast along the line A-A', extended, of Figure 2 along the course of the Church Hill Tunnel of the Chesapeake and Ohio Railway. (Modified from Coryell, 1876, pl. 5, and Darton, 1911, fig. 1. The letters AD indicate the sources of Coryell's samples analyzed for diatoms, of which only C was rich in diatoms. The stratum labeled "Yellow Clay" corresponds to Darton's Lafayette Formation; the "Diatomatic Sand" to the Calvert Formation, and possibly in part younger Miocene beds; the "Green Sand" to the Aquia and (below) the Patuxent Formations.) with the Fairhaven member of the Calvert Forma- tion in Maryland, on the basis of the abundant diatoms. Among the areas noted especially for the diato- maceous bed are the slopes of Church Hill. Rogers (1841; reprinted 1881:59; 1884:452) refers espe- cially to the area "at the foot of the abrupt bare bank which has been cut into north of Main street, on Church hill, and thence south nearly to the next street—Indeed, along the slope of Church hill, as on the other side of the valley, this stratum may be found in all the ravines and cuttings, at the proper level." He refers also to the overlying sands and clays with plant remains and molds of scallops and other Miocene shells. His most exten- sive account of the diatomaceous deposits of Shoc- koe Creek Valley is that of 1859. Along with Rogers' publications, perhaps the most useful in connection with Phoca wymani is that of Coryell (1876; reprinted 1881), particu- larly his geologic profile (his pl. 5; here modified in Figure 3) through Church Hill from northwest to southeast along the course of the Church Hill Tunnel of the Chesapeake and Ohio Railway. This tunnel, which was begun in February 1872, opened in 1874, and largely disused from 1901 to 1925, collapsed in part during repairs on 2 October 1925, burying a work train and several workmen, after which it was mostly filled with sand and the ends sealed with concrete in 1926 (cf. Teal and Armit- age, 1950:775, and Blackford, 1973, Bower, 1975, and Heite, 1964). Its portals are still preserved, located immediately north of the intersection of Nineteenth and Marshall streets on the northwest (visible at surface), and near the head of Bloody Run ravine on the southeast (roofed, by a concrete extension of the tunnel some 241 feet in length), and indicated on the map (Figure 2) by the points A and A'. Coryell's section was reproduced with modifications by Darton (1911: fig. 1), who assigned all beds between his Aquia and Lafayette Formations to the Calvert Formation. Coryell (1876:231-232) described the section in Church Hill very well, in part as follows: 14 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Upon this granite bed is a sandy clay some fifty feet thick, of various colors and composition, then a ferruginous sand [Aquia Formation of Darton], so compact in places as to resist the pick and crowbar, and which, when struck, pro- duces sparks of fire, and is classed with the rocks. Upon this stratum is the blue deposit [Calvert Formation] through which the tunnel was made. The thickness of the stratum in the tunnel was found to be eighty feet; and the color; when first exposed, a dark blue, but on long exposure be- coming nearly white. Upon this is a yellow clay [Lafayette Formation of Darton], very distinct in color, and filled with water-worn silicious boulders, nearly the size of hens' eggs. This stratum is thirty feet in thickness, and upon it are various strata of clays which make the soil of the upland country, and under present cultivation do not indicate much fertility. In excavating the tunnel, large quantities of bones and teeth were found, not interspersed through the material, but generally in pockets. . . . Mr. Peticolas, recognizing the importance and value of the true position and thickness of the strata from a known base-line, very kindly devoted his time to establish these facts for this paper, and, with the knowledge and assistance of Major Channing M. Bolton, he was furnished with material from well-established points in the tunnel and the railroad excavations, marked on the profile A, B, C, D. To the eye, there is a uniform stratum eighty-five feet in thickness, blue in color when first exposed, and becoming nearly white after long exposure; this was generally denomi- nated the diatomaceous stratum. Material from the bottom of the tunnel and lower portion of the blue stratum (A and B), was nearly barren, and the remains of an inferior type; at the point C diatoms were found in great abundance, and at D, near the top, the clay was quite barren. Most important is the probable 80-85 feet of Calvert Formation, extending on the east side of Shockoe Creek ravine (Figure 2, locality 1), from an altitude of approximately 50 feet to 135 feet, which is to say from below the base of Church Hill well up its slopes. The cranium of Phoca wy- mani inevitably came from the lower part of this stratum (through which the tunnel was later ex- cavated, yielding numerous vertebrate remains) if it was collected in place, as seems probable in view of the improbability of such a delicate object surviving a tumble from high up the slope. Matrix recovered from the auditory bullae was insuffi- cient for analysis, but rediscovery and study of the associated endocranial cast would almost certainly yield significant results. Darton (1911:24) described the extensive devel- opment of the diatomaceous Calvert Formation on the southeast side of Church Hill, on the nose of the slope bordering Libby Hill Terrace (Figure 2, locality 2) and extending along the west side of Bloody Run ravine to the old tunnel entrance. These beds constitute a likely source for the seal remains found by Dr. Burton "in the ravine at the eastern extremity of the city" (Wyman, 1850c: 229). Darton (1911:25), in discussing the western margin of the Calvert Formation in Richmond, noted several exposures "in the neighborhood of the penitentiary" (Figure 2, locality 3). These include the "east side of Hollywood Cemetery," "railroad cuts on the north bank of the river at the foot of Pine and Laurel streets," "Bank near the foot of First Street," "formerly exposed in the steep slopes of the park just opposite [Gambles or Gimbles Hill Park]," and "penetrated by the railroad tunnel under Byrd Street between Third and Fourth streets." This last locality (Figure 2, locality 4) is exactly that of the present roadcut for the downtown expressway, from which Squalo- don remains were recovered recently. The cut bank exposed still at the Ballard (Four- teenth) Street locality (Figure 1, locality 5) con- sists of up to 12 feet of diatomaceous, Calvert?, clay, overlain by a six-inch bed of bone and other rubble, grading upward into a bed of sandy clay up to 10 feet in thickness. Matrix associated with the phocid remains (described below) from this locality indicates that the partial humerus of Lep- tophoca came from the lower clay, with abundant well preserved diatoms; and the monachine re- mains, from the upper, sandy clay with sparse, generally fragmentary diatoms. The diatoms in the upper bed may be reworked and not strati- graphically significant, those in the lower are sug- gestive of zone 15 or 16 of the uppermost Calvert or lowermost Choptank Formation (Andrews, pers. comm., 1975). The upper bed is probably equivalent to the horizon from which the Squalo- don material was recovered at locality 4 to the west (Ward, pers. comm., 1975), and therefore is probably assignable to the Calvert Formation. Thus, all specimens of fossil seals from Rich- mond probably came from the Calvert Formation, rather than from the overlying Miocene beds under study by Ward and Blackwelder. DESCRIPTION.—Wyman's identification and de- scription (1850c, quoted in full above) of his fossil seal remains have been taken unjustifiably lightly, especially by his associate, J. A. Allen. Besides being a professional comparative anatomist, Wy- NUMBER 28 15 man had long and special acquaintance with phocids, as revealed by a superficial review of the record. For example, his first recorded communi- cation to the Boston Society of Natural History, on 20 January 1841, was on a skull of the crabeater seal (Wyman, 1844; Gray, 1875:107), and in a letter to Spencer Fullerton Baird of 28 March 1851, in the archives of the Smithsonian Institu- tion, he stated, "I have a plenty of crania of the Harp Seal which I brought from Labrador [in 1849], but obtained but two crania of the Hooded Seal and one of the Ph. vitulina." Further, his description of the Richmond material was highly pertinent in terms of phocid cranial anatomy, particularly in view of his limited comparative material, including no fossil phocids and no Monachus. There could be little or no justifica- tion for the suggestion that he had misidentified cetacean cranial material (Allen, 1880:471; Kel- logg, 1922:75) particularly as he recorded a ceta- cean petrosal at the same time (1850c:231). Of cranial features noted by Wyman and no longer available on the remnants of the holotype, most are characteristic of phocids generally, or are not susceptible to evaluation without the material, as "the reentering angle of the occiput [lambdoi- dal crest], the well marked depressions correspond- ing with the cerebral convolutions on the parietal bones [presumably in internal aspect, as the con- volutions are not clearly reflected externally in phocids, as they are in many mustelids], the form of the cranial cavity, . . .the vascular canals open- ing on the occiput. . . ." However, "the interparie- tal crest extending from the occiput to the anterior edge of the frontals . . . most narrow [lowest?] posteriorly where it is but slightly elevated above the surrounding bones" is a feature that clearly separates the fossil from all modern phocines ex- cept adult Halichoerus and some old individuals (males only?) of Phoca vitulina, and aligns it with the fossil monachines in which the skull roof is known (Monotherium gaudini, Pliophoca etrusca, and Prionodelphis capensis) and with the living monachines, Monachus, Hydrurga, and Leptony- chotes. My observations on modern phocids do not coincide completely with those of King (1972:98) who states, "In phocids other than . . . [Pusa, His- triophoca and Pagophilus'] the temporal ridges meet to form a sagittal crest . . . ." It seems safe to assert that, in addition to Pusa, Histriophoca, and Pagophilus, among living phocines, Erignathus and probably Cystophora never form a sagittal crest. In the USNM collections there are two skulls of very old male Cystophora (USNM 38233 and 188846) in which high parasagittal lips, separated by a narrow trough, are developed on the frontals; it is not inconceivable that exceptional individuals of Cystophora could be found in which these lips unite to form a sagittal crest, but I have seen none. Among modern monachines, some individ- uals of Mirounga develop a very short, low sagittal crest on the parietals only; I have seen no ap- proach to formation of a sagittal crest in Lobodon, but the small USNM collection includes no very old individuals; Ommatophoca characteristically not only does not develop a sagittal crest but has only a weak, unfused sagittal contact between left and right frontal and (anteriorly) parietal bones, with a strong tendency to retain in adulthood a fontanelle, especially between the unfused fron- tals (cf. King, 1969:27-28). Other cranial features of Phoca wymani remain- ing available are limited to the temporal bones (and auditory ossicles, discussed below), of which examples are at hand for all the living species of phocids. It is immediately apparent through com- parison with these that Phoca wymani is a mona- chine seal. Temporal bones are known for very few fossil monachines, including an incomplete left temporal from the Scaldisian of the Antwerp Basin, referred to Callophoca obscura; some half dozen well preserved temporals and many frag- ments from the Yorktown Formation of North Carolina, referred to Callophoca (Ray, in press); the incomplete left temporal in the type skull of Pliophoca etrusca from the late Pliocene of Italy, inadequately described and illustrated for present purposes, but apparently essentially as in modern Monachus (Tavani, 1941:100, fig. 1; pl. 14(1): figs. 2a, 2d, 3); and one well-preserved right tem- poral, and some fragments, of Prionodelphis ca- pensis from the Pliocene of South Africa (Hendey and Repenning, 1972:80-81, pl. 6: fig. B). Naturally, as is intuitively clear to anyone who has looked critically at pinniped skulls, and as well demonstrated for the oceanic species of Phoca by Burns and Fay (1970:389), any conclusions based on small numbers of skulls, to say nothing of iso- lated temporal bones, must be regarded as highly tentative in view of the great variation. Compari- 16 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY son of limited numbers of temporal bones, espe- cially limited for ones with the intracranial surface exposed to view, among the living monachine seals shows that they are highly diagnostic at the generic level, indicating that their characteristics do have some taxonomic, and possibly phylogenetic, utility. The temporal bones of the southern monachine genera, including Mirounga, are highly diverse, with each exhibiting unique modifications. For example, in Mirounga the petrosal apex is greatly swollen much as in all phocines (except Erigna- thus), and the auditory process (ectotympanic) forms a crescentic ossification ventral to the exter- nal acoustic meatus and posterior to and conform- ing to the retroarticular process of the glenoid fossa. In Ommatophoca the auditory process of the bulla is greatly thickened, long, and prominent, there is a massive mastoid-like swelling of the squamosal region dorsal to the external acoustic meatus and the mastoid region, and the petrosal is much reduced, especially the apex (absolutely smaller than in any other phocid). In Lobodon the bulla is globose, and the auditory process short and inconspicuous in relation to the large bulla. In Leptonychotes the petrosal is remarkably broad, including the apex, which is widely rounded and flat. In Hydrurga the petrosal apex is somewhat swollen, the bulla is triangular in outline in ven- tral aspect, and its walls are enormously thickened (as in Ommatophoca, King, 1969:11). No claim to originality is implied in these observations of what are in any case only some of the most obvious characters in the temporals of southern mona- chines, as most, if not all, have been noted before, for example, by Turner (1887:65) and King (1969:29) for Ommatophoca, and by Hendey and Repenning (1972:81, pl. 7: fig. C) for Leptony- chotes. However, it seems not to have been em- phasized that this observed diversity is in harmony with the strongly divergent adaptive radiation re- flected in the overall biology and morphology of southern monachines. The petrosal apices illus- trate the point forcefully, including as they do the most reduced (Ommatophoca), the broadest and flattest (Leptonychotes), and one of the most swollen (Mirounga), among the phocids. In most features, including overall proportions, the temporals of Phoca wymani are generally most similar to those of Monachus among living seals, but bear an equally or even more striking resem- blance in virtually every feature to that of Priono- delphis capensis and in many respects to referred temporals of Callophoca. In my opinion these similarities reflect an expectable community of fea- tures and lack of extreme modification indicative of real relationship among the fossils, Phoca wy- mani and Prionodelphis capensis, and the living Monachus, which I regard as persistently primitive. Because of the complexity of the temporal bone, many features are subtle and difficult to describe, but many of the similarities and differences are obvious on inspection (Plates 1 and 2). Salient features of the temporal bones of Phoca wymani are noted below, and compared and contrasted with those of other seals. As revealed in ventral aspect (Plate 1), the bulla is rather small, little inflated, triangular in outline, and bulking little more than the large mastoid (as noted by Wyman, 1850c:229), which is shaped as in monachines generally, rather than in phocines, but is unusually swollen, perhaps ex- ceeding somewhat those of Prionodelphis capensis and Callophoca obscura (referred), with which it compares best. King (1966:387, fig. 1) indicated that in northern phocids (including Monachus) the posterior extremity of the petrosal is visible externally behind the bulla when the skull is viewed in ventral aspect, whereas in southern phocids the bulla essentially conceals the petrosal. Hendey and Repenning (1972:81) stated, "The latter condition is very evident in P. capensis, strongly suggesting an affinity with the Antarctic seals [as opposed to Monachus]." Phoca wymani is very similar to Prionodelphis capensis in this character, and neither is very different from Cal- lophoca (referred) or Monachus. It seems to me that the features of this complex region do not lend themselves readily to reduction to "key" char- acters, and that weighing the spatial relations among its components indicates that the distinc- tions among Monachus, Prionodelphis capensis, Callophoca obscura (referred), and Phoca wymani are meager, and that those between Monachus on the one hand, and Prionodelphis capensis and the living southern monachines on the other, are neither so clearcut nor significant as might be sup- posed. The southern monachines do tend to have little or none of the petrosal exposed adjacent to the posterior lacerate foramen behind the bulla, and instead generally have the exoccipital and the NUMBER 28 17 bulla in close approximation lateral to the fora- men. However, lateral to that approximation the condition is variable and uncertain, depending in part on the generally unclear boundary between petrosal and mastoid. Additionally, in at least some individuals of Ommatophoca and Leptonychotes (with which Prionodelphis capensis has been espe- cially compared) the petrosal is exposed immedi- ately adjacent to the posterior lacerate foramen. In any event, based on the totality of its charac- ters, the temporal bone of Prionodelphis capensis appears to be aligned best with the northern (espe- cially fossil) monachines, not the southern. Also in ventral aspect, the posterior carotid fora- men opens in full view (as noted by Wyman, 1850c:229) in contrast to most phocines, in which it lies partially concealed on the medial wall of an inflated bulla (cf. King, 1972:98-99), except in Erignathus in which it is clearly exposed and pos- teriorly situated. The foramen in Phoca wymani also opens far anterior to the posterior extremity of the bulla, in contrast to Monachus and Priono- delphis capensis, Callophoca (referred), and most other monachines, in which it is situated more posteriorly, in Monachus approximating the pos- teromedial corner of the triangular bulla. Not visible in strictly ventral aspect, but opening anteriorly near the anterior apex of the bulla, is the auditory canal, exceptionally large as noted by Wyman (1850c: 229) in contrast to that of Phoca groenlandica. This canal is quite large in Phoca wymani, as it is in Monachus and in most mona- chines, and is generally quite small in most pho- cines. Its variable and irregular shape makes quan- tification of the differences unfeasible. King (1966:387) has indicated that in northern phocids, including Monachus, the mastoid region is visible when the skull is viewed in dorsal aspect, whereas it is not visible dorsally in southern pho- cids. As with the ventral exposure of the petrosal, the situation with the mastoid region seems less clearcut than one might wish and not entirely susceptible to treatment as a key character, at least as defined. For example, although the mastoid is visible dorsally in Monachus, the entire configura- tion of the region is much more like that of Phoca wymani, Prionodelphis capensis, and Callophoca obscura (referred), and other monachines, than of any phocine. In exception to King's generalization also, the mastoid region of Ommatophoca is broadly visible dorsally. Also in dorsal or intracranial aspect, the floccu- lar or cerebellar fossa is extremely large in Phoca wymani (noted by Wyman, 1850c:229) as it is in Prionodelphis capensis and in phocids generally, although considerably constricted in Monachus, Ommatophoca, and Hydrurga. The anterior part of the petrosal in Phoca wymani is flat (not swol- len dorsally) and the apex is broadly rounded, much as in Prionodelphis capensis (Plate 2), although perhaps not quite as broad. In all pho- cines except Erignathus, the petrosal is greatly swollen anteriorly into a more (Cystophora cri- stata and Phoca groenlandica for example) or less (Phoca vitulina and Phoca hispida for example) globular mass. Among monachines, the petrosal is swollen anteriorly, greatly in Mirounga, and less so, but considerably, in Hydrurga. In Erignathus the petrosal is rounded anteriorly and only slightly swollen dorsoventrally, less so than in any other phocine or in Mirounga or Hydrurga. Among those monachines having more or less flat petro- sals, Leptonychotes represents the extreme in broad and rounded apex, Monachus the extreme in narrow and pointed apex, and Ommatophoca the extreme (among all phocids) in reduced apex. Clearly the various features of the temporal bone by which phocines and monachines and northern and southern monachines have been asserted or implied to be separable are far from absolute. However, it is equally clear that the temporal bones of Phoca wymani, Prionodelphis capensis, Callophoca (referred), and Monachus are more similar to one another than are any of them to those of any other phocid, and that of these, Phoca wymani and Prionodelphis capensis seemingly are closest to one another. With the permission of Dr. Farish A. Jenkins, Jr., the left auditory bulla of MCZ 8741 was opened along previously existing fractures for re- moval of matrix and search for auditory ossicles. The incus and malleus were recovered in good condition, except for the absence of the distal end of the manubrium of the malleus. Each element is similar to its counterpart in modern Monachus and in referred specimens of Callophoca from North Carolina. Studies in progress of phocid auditory ossicles are not yet sufficiently advanced to warrant secure generalizations, but prelimi- nary observations on small numbers of sets of 18 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY ossicles of most genera and subgenera of modern phocids suggest that the ossicles will offer reliable features for distinguishing phocines from mona- chines (see also King, 1966:387, for example), and some genera from one another. In phocine seals examined thus far the maximum linear dimension of the incus is less than 80 percent of that of the malleus, whereas it is more than 90 percent in the monachines. The neck of the malleus is relatively long in phocines, short in monachines. The head of the malleus is relatively large in comparison to the body of the incus in phocines, whereas in monachines the body of the incus seems to be ex- panded, and dwarfs the bulk of the head of the malleus. In all of these features the incus and mal- leus of Phoca wymani correspond to those of monachines, and in addition show further special similarities to Monachus and Callophoca (refer- red) in all details of morphology (Plates 3 and 4) and most notably in the two simple incudomalleo- lar articulations, contrasting strongly with the multifaceted or complexly curved articular sur- faces in most phocines and some monachines, and with the single confluent saddle-shaped articula- tion in Ommatophoca (King, 1969:12; confirmed on additional specimens). The distal end of the left fibula from Wyman's original collection, represented by casts, USNM 214650 (Plate 6: figure 2; Plate 7: figure 1), affords no obviously diagnostic features. It represents a stoutly developed fibula with two strongly marked tendinal grooves on its lateral face, defining a crest or process between them. It is curious that these topographic features, widespread in both phocines and monachines, are conspicuously little developed in Monachus, Mirounga, and specimens from North Carolina referred to Callophoca, in all of which the lateral face of the distal end of the fibula is one of low relief. The fibula does afford the one small point of morphological overlap between Wyman's original material and the Ballard Street sample of 1968. The fibula is represented in the latter collection by most of the left diaphysis, lacking the proximal part (Plate 6: figure 1; Plate 7: figure 2), of an individual (USNM 187410) in which the distal epiphysis had yet to fuse, and was not recovered. About all that can be determined is that the two pieces are compatible in size and morphology as far as revealed by the meager comparisons possi- ble, and are far too large for Leptophoca^ lenis, as are all the elements, presumably of a single indi- vidual, cataloged under USNM 187410, excepting the right lower canine, which may represent Leptophoca lenis. The left tibia of USNM 187410 (Plate 6: figure 1; Plate 7: figure 2) is essentially complete except for the unfused and missing distal epiphysis. The proximal epiphysis is present and tightly fused to the diaphysis. The proximal end of the fibula is present and tightly fused to the tibia, as it is in all known phocids except Monachus schauinslandi and possibly a tibial fragment from the Scaldisian of Belgium referred to Gryphoca similis (Van Beneden, 1877, pl. 13: figs. 19 and 20; Ray, in press). The tibia is short and heavily built, with the shaft relatively straight, as in most monachines (except Leptonychotes), and in contrast to pho- cines in which the tibia is long and slender, and generally curved or spiraled in appearance. The length of the tibia, minus the distal epiphysis, is approximately 3.5 times or less the breadth of the_ proximal end in monachines (except Leptony- chotes) and 4 times or more the breadth in pho- cines. The pretibial and posttibial fossae are shal- low in USNM 187410, with a thick body of bone between as in monachines generally (including Leptonychotes), and in contrast to phocines in which these fossae are deeply excavated, develop- ing in some instances overhanging lips on their medial borders, and in some instances reducing the bony wall between the fossae to an extremely thin lamina. In sharp contrast to most monachines including the Richmond specimen, the tibia of Prionodelphis capensis resembles that of phocines in development of fossae and extreme thinning of the intervening bone. The proximal articular facets in the fossil are of low relief, as in monachines, and in contrast to phocines in which the condyles are curved and their lips elevated adjacent to the intercondyloid area. The tibia from Richmond is very similar in character to those of Monachus, Mirounga, and Callophoca (referred), and in size to those of Monachus. The left ulna of USNM 187410 (Plate 5: figures 8, 10) is essentially complete except for the ab- sence of the distal epiphysis. The bone is relatively short and broad (craniocaudally) as in most monachines and in contrast to phocines (and NUMBER 28 19 Leptonychotes). An unusual feature of uncertain significance is seen in the medially sloping, cranial part of the proximal surface of the olecranon proc- ess (Plate 5: figure 10). In all other phocid skele- tons examined, this surface is transversely oriented (if approximately planar), or rounded or irregu- lar, except in a single fragment from Belgium re- ferred to Monotherium affine (Van Beneden, 1877, pl. 16: fig. 10), in which the surface resembles that of the specimen from Richmond, although not clearly reflected in Van Beneden's illustration. Considerable intraspecific variation may be ex- pected in this surface of strong muscular insertion. The left ectocuneiform of USNM 187410 is severely eroded and has not been compared in detail. The incomplete right mandibular ramus is heav- ily constructed and preserves the alveoli of two large double-rooted postcanine teeth (Plate 5: fig- ures 1, 4). Its robust character, evidence of large double-rooted cheek teeth, and details of configu- ration set it apart from known phocines, and sug- gest alliance with monachines, among which it can be matched almost exactly by specimens from North Carolina referred to Callophoca obscura. This similarity is taken to imply general, not precise, relationship, as the mandibles of Mona- chus, Callophoca, Pristiphoca, Prionodelphis ca- pensis, and other fossil monachines are rather similar, and this fragment does not include the anterior and posterior parts in which characters thought to be diagnostic are found. DIAGNOSIS.—A fully satisfactory specific diagnosis of Phoca wymani is not yet feasible, owing to the meager material of it and other fossil seals. Still, the holotype is better, and the additional material more confidently referred, than is the case with most fossil phocids. Thus, as so often in the system- atics of vertebrate fossils, science may be better served not by adhering to the strictest taxonomic procedures, but by conserving the taxon with con- fidence that further discoveries will vindicate its retention. In the meantime, Phoca wymani may be diag- nosed as a middle Miocene monachine, generally Monachus-like in size and morphology of the skull, mandible, ulna, tibia, and fibula; with long, well- developed sagittal crest; temporal bone similar to that of Monachus but with unreduced floccular fossa, rounded petrosal apex, carotid canal open- ing more anteriorly, and petrosal less exposed pos- teriorly in ventral aspect; temporal bone similar to that of Prionodelphis capensis in almost all fea- tures, except more anterior opening of carotid canal; ulna with proximal surface of olecranon sloping medially, as in Monotherium; fibula with pronounced lateral crest at distal end.- GENERIC ALLOCATION.—Clearly Wyman's seal cannot remain in the genus Phoca, which should not be made to include, even in its broadest pale- ontological usage, a species that is definitively monachine. Even the tacitly accepted practice of using the genus to accommodate a variety of mostly poorly understood fossil phocines (as Phoca vin- dobonensis, P. couffoni, P. moori, and others) has fostered incorrect conclusions, for example, that modernized Phoca extends well back into Miocene time (Davies, 1958:486; King, 1972:96). With regard to Wyman's generic assignment to Phoca, it should be noted that the genus was very broadly construed at the time, explicitly so by Wy- man (1850c); he mentioned, as species of Phoca, the bearded, harp, ringed, hooded, and common seals, each accorded its own genus in most subse- quent literature, and even by the most inclusive concepts of the present, arrayed among three tribes and three genera, one of which includes four subgenera (Burns and Fay, 1970:390). Also, it should be noted that there was almost certainly not a single specimen of Monachus in North America at the time (Allen, 1887). There is no entirely satisfactory option avail- able, in view of the limited material of Phoca wymani and the poor knowledge of other Tertiary phocids, for most of which the temporal bone is unknown. The possibility must be considered that Wy- man's seal could represent Leptophoca. This would be the most economical hypothesis, in that it would account for the apparent absence of monachines in the Calvert Formation of Maryland and northern Virginia, and for the presence of the leptophocine humerus (and canine) in Richmond. It is phylogenetically conceivable in that mona- chines and phocines must surely have diverged from a common ancestor, and surely not long be- fore the time of deposition of the Calvert Forma- tion. This would call for a seal combining a small, generally phocine skeleton (Leptophoca lenis) with a monachine temporal region (Phoca wy- 20 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY mani). This hypothesis is rejected here because other remains from Richmond, including Wyman's fibular fragment and the new specimens (except the partial humerus), are typically monachine, and as far as comparable parts are available, con- trast in size and subfamilial character with re- mains assigned with some assurance to Leptophoca lenis. Thus it does seem that there are two very distinct seals in the Calvert Formation, indicative of well-marked divergence between phocines and monachines by that time, and there is need for a generic assignment for Wyman's seal that will serve best to reflect its relationships until it and other Miocene seals are sufficiently better known to determine whether a separate genus is warranted. Following past practices, one might utilize the genus Monachus for poorly known fossil mona- chines as Phoca has been used, but this would be similarly misleading, as even the conservative Monachus does not extend back into the Miocene, and as there are recognizable morphological dif- ferences in the parts preserved. Of course G. M. Allen (curatorially) and I (1958:441) inadvert- ently assigned the remnants of the holotype to Monachus. Of fossil monachines, the temporal bone is known only for Pliophoca etrusca (incomplete), Callophoca obscura (referred), C. ambigua (re- ferred), and Prionodelphis capensis. The element has not been fully described in Pliophoca etrusca, but in all other respects the species is almost identi- cal to Monachus monachus. The temporal bone of Phoca wymani is extremely close to those of Prio- nodelphis capensis and Callophoca (referred) in most respects. However, I refrain from assigning the species to Prionodelphis on the following grounds: 1. The genus Prionodelphis and the assignment of the South African species to it are so tenu- ously based at present as to offer little prospect of stability. 2. The tibias referred to Phoca wymani and to Prionodelphis capensis are very dissimilar. 3. The two species are widely separated geolog- ically (probably early middle Miocene vs. middle or late Pliocene) and geographically (the two liv- ing species of Mirounga could however be cited as a modern analogy). Phoca wymani could with reasonable justifica- tion be placed in the genus Callophoca for the present, but I am reluctant to do so because: 1. The temporal bones of Callophoca affording best comparison to Phoca wymani are referred speci- mens from North Carolina, with only a highly incomplete referred specimen from Belgium. 2. The postcranial elements referred to Phoca wymani are in part, notably the ulna and fibula, not closely similar to their referred counterparts in Callophoca, and the temporal bones differ in some respects as well. 3. Phoca wymani is geologically older than Cal- lophoca (probably early middle Miocene vs. early Pliocene). The striking similarity of the temporal bone to those of Prionodelphis capensis and Callophoca cannot be disregarded. The probable derivation of the two from Monotherium suggests that the latter ought to be considered for reception of the Richmond seal, an attractive possibility because: 1. Monotherium is the geologically oldest well- founded fossil monachine genus available, and is in fact the only one unquestionably of Miocene age. 2. The referred ulna from Richmond shares an apparently unique feature with one from Belgium assigned to Monotherium. 3. Monotherium is the monachine known from the most nearly contemporaneous, though younger, deposits, the Diestian of Belgium and (referred specimens from) the St. Marys Formation and Gay Head greensand of eastern United States; these occurrences are geographically reasonable in terms of fossil and modern phocid distribution around the North Atlantic. Thus, Phoca wymani is here reassigned tenta- tively to Monotherium, and should now be written Monotherium? wymani (Leidy, 1853). As the tem- poral bone of Monotherium unfortunately is un- known, this generic assignment must be regarded as no more than a temporary device, based on scanty immediate evidence and on a weakly docu- mented general concept of phocid evolution. Nevertheless, in light of the present alternatives, this placement affords the most plausible reflection of probable affinities, pending discovery and study of additional material. Leptophoca lenis True PLATES 8-11: FIGURES 3, 4 The right lower canine found in the block with NUMBER 28 21 the skeletal elements under USNM 187410 seems to be too small for a monachine of the size repre- sented by the other elements, and is on the other hand comparable in size and character to canines referred to Leptophoca lenis, to which species this specimen is tentatively assigned. The right humerus, USNM 187409 (Plates 8-11; figure 3), lacking the proximal end, from the Bal- lard Street locality, but not directly associated with the above specimen, compares satisfactorily with the holotype of Leptophoca lenis, to which it is assigned. In any event, this specimen clearly represents a small primitive phocine, close to if not conspecific with L. lenis. Addendum After this paper had gone to press, an electro- print copy of the Catalogue of Collection of Com- parative Anatomy in Boylston Hall [at Harvard University] Belonging to Jeffries Wyman" was re- ceived from the Boston Museum of Science and has been placed in the Remington Kellogg Library of Marine Mammalogy of the Smithsonian Institu- tion Libraries. Availability of the facsimile of the complete catalog revealed two additional entries for remains of fossil seals beyond those listed above under "Fate of the Specimens," as follow: 473. Cast of the lower extremity of the fibula of a seal—the original, No. [blank] was found at Richmond, Va. [Entire entry lined out and replaced by an unrelated specimen, as was number 844, above]. 909. Ossicula of the ear of a Fossil Seal. Richmond. See 824- 825. The specimens listed under number 909 must have been the auditory ossicles of the right side of MCZ 8741, which Wyman, with his usual care, undoubtedly noticed and preserved in cleaning the right temporal bone, the bulla of which was in- complete, widely open, and essentially empty of matrix when it came to me (cf. Plate 1: Figure 2). Unfortunately these first-discovered fossil phocid auditory ossicles apparently were lost after Wy- man's death, and have yet to be relocated. Furthermore, review of the papers of Jeffries Wyman housed in the Francis A. Countway Li- brary of Medicine, Harvard Medical Library, Boston, Massachusetts, revealed correspondence from Dr. Martin Burton of Richmond, Virginia, to Wyman. Relevant quotations from these letters follow. Letter of 10 April 1850: .... You will also receive a drawing, showing the upper and under view of a skull that I found not long since in the blue earth. It was about to crumble, and I got Mr. Peticolas to make a drawing of it. . . . An opportunity has just unexpectedly offered of sending you these things, and allows me no time to be more explicit. in haste yr very sincere friend Richmond-Wednesday Night-April 10th Burton I am afraid you will be able to make nothing of the drawing. Letter of 11 May 1850 (obviously, from state- ments in the complete letter, in reply to a letter from Wyman, responding to his of 10 April): .... I have sent you by Mr. Blake the tympanic bones of the head, the drawing of which you have, with the lump of dirt that filled the cavity of the skull. The dirt is a perfect cast of the brain, representing its size and shape, and even its convolutions. I should be pleased to know the conclusion at which you may arrive (if any) as to the head. . . . Your very sincere friend M. Burton Richmond May 11th 1850 Letter of 28 May 1850, quoted here in its en- tirety: Dr. Wyman Dear Sir The blue earth containing fossils is (as you know) com- posed almost exclusively of the exuvia of infusoria, and lies at the bottom of the hills and valleys in and around Rich- mond. In whatever direction wells or pits have been sunk to a sufficient depth, this earth has been reached, and ap- pears (from such observations as I have made) to have the form of the surface of the country—rising in the center of the hills, sloping on the sides, and appearing upon or near the surface in the valleys. Where gullies have been formed on the slopes of the hills by the action of torrents, or a small stream flows through the valleys between the hills, this earth has been exposed and worn to variable depths. Fossils have been found in this earth in so many places, as to render it probable that it contains them wherever found—many more in some places than in others. Besides the ravine near the Penitentiary, I got many fos- sils from a well sunk on the western slope of the hill west of the Penitentiary—many from a well sunk on the top of the hill east of the Penitentiary—some from the foot of Church hill—and some from the gullies north of the Medi- cal College. I have never found any except in the blue earth. It is gratifying to me to have been the means of affording you the pleasure (I know you have experienced) in determin- 22 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY ing the nature of the different fossils I have given you, and of contributing to the advancement of your much cherished science. The seal skull was found in the blue earth at the foot of Church hill—the vertebrae of the porpoise, also of the seal, were found in the ravine near the Penitentiary— the two localities being at least a mile apart. I thank you for your kindness in offering to give me the credit of the discoveries in the communication you are pre- paring for Silliman's Journal, and in return, can only promise that, I shall not be less diligent or anxious to serve you than heretofore. It gives me pleasure to collect that you may interpret. Your very sincere friend M. Burton Richmond May 28th Unfortunately, search for the drawings of the seal skull (in the Countway Library and in the Boston Museum of Science thus far) has not yet been suc- cessful, but other possible repositories are being investigated. The last of Dr. Burton's letters quoted above confirms the conclusion that the seal skull was found in place in the Calvert Formation, i.e., "in the blue earth at the foot of Church hill." The same letter resolves the dilemma discussed under "Localities" regarding "portions of the skele- ton of another seal" coming ostensibly from two widely separated localities, one "the ravine at the eastern extremity of the city," the other "in the neighborhood of the penitentiary" (Wyman, 1850c: 229-230). Obviously "eastern" was a simple lapsus on Wyman's part for "western." Thus "locality 2" (Figure 2) is incorrectly placed in Bloody Run ravine at the (then) eastern extremity of the city and should apply to "the ravine near the peniten- tiary," probably the deep ravine south of the peni- tentiary and west of Gambles or Gimbles Hill, near localities 3 and 4, and near the (then) western extremity of the city (Figure 2). For locating information about Jeffries Wyman, it is a pleasure to acknowledge the assistance of Charles P. Lyman and David Gunner of the War- ren Anatomical Museum, Richard J. Wolfe of the Francis A. Countway Library of Medicine, and Barbara Wiseman of the Boston Museum of Science. Mr. Wolfe also granted permission to pub- lish the quotations from Dr. Burton's letters to Jeffries Wyman. Literature Cited Agassiz, Alexander 1876. Report of the Curator to the Museum Committee. Annual Report of the Museum of Comparative Zoology at Harvard College, 1875(A):5-14. Allen, J. A. 1880. History of North American Pinnipeds. Volume 12 in U. S. Geological and Geographical Survey of the Territories Miscellaneous Publications, xvi + 785 pages, 60 figures. 1887. The West Indian Seal. Science, 9(206):35. 1916. Autobiographical Notes and u Bibliography of the Scientific Publications of Joel Asaph Allen, xii + 215 pages, frontispiece. New York: American Mu- seum of Natural History. Anonymous 1847. Donations to Museum. Academy of Natural Sciences of Philadelphia Proceedings, 3(7): 141. 1857. Donations to Museum in September and October, 1856. Academy of Natural Sciences of Philadelphia Proceedings, 8(6):xviii-xx. Baird, S. F. 1852. Report of the Assistant Secretary in Charge of the Museum, etc. Smithsonian Institution Annual Re- port, 6:39-65. Barbour, T., B. Lawrence, W. E. Schevill, S. L. Washburn, and M. B. Cobb 1943. Glover Morrill Allen, 1879-1942. Journal of Mam- malogy, 24(3):297-304, plate. Barnes, L. G., and E. Mitchell In press. Late Cenozoic Northeast Pacific Phocidae. Con- seil International pour I'Exploration de la Mer, Rapports et Proces-verbaux des Reunions, 167:34- 42, 6 figures. Blackford, F. R. 1973. The Train That Will Never Move. The Virginian- Pilot (Norfolk, Portsmouth, Virginia Beach, and Chesapeake, Virginia), 108(299:16 September):C3, 2 figures. Bower, Suzanne 1975. Collapse on Church Hill. Richmond Mercury, 3(26: 5 March):3, 10-11, 5 figures. Burns, J. J., and F. H. Fay 1970. Comparative Morphology of the Skull of the Rib- bon Seal, Histriophoca fasciata, with Remarks on Systematics of Phocidae. Journal of Zoology (Lon- don), 161 (3): 363-394, 3 figures, 4 plates. Clark, W. B., and B. L. Miller 1906. The Geology of the Virginia Coastal Plain. Chapter I (pages 11-24) in Part I in Heinrich Ries, The Clay Deposits of the Virginia Coastal Plain. Geo- logical Survey of Virginia Geological Series Bulle- tin, 2: 184 pages, 15 plates, 10 figures. 1912. Physiography and Geology of the Coastal Plain Province of Virginia. Virginia Geological Survey Bulletin, 4: 274 pages, 19 plates. NUMBER 28 23 Cope, E. D. 1868a. [Four Extinct Species of Mammalia, Which Were Discovered by Jas. T. Thomas, in the Miocene Deposits of the Yorktown Epoch in Charles Co., Maryland.] Academy of Natural Sciences of Phila- delphia Proceedings 19(4):131-132. 1868b. An Addition to the Vertebrate Fauna of the Mio- cene Period, with a Synopsis of the Extinct Cetacea of the United States. Academy of Natural Sciences of Philadelphia Proceedings, 19(4): 138-156. Coryell, Martin 1876. Diatomaceous Sands of Richmond, Virginia. Trans- actions of the American Institute of Mining Engi- neers, 4:230-232, 1 plate. [Reprinted, 1881, in The Virginias, A Mining, Industrial, and Scientific Journal, 2(l):6-7, 1 figure.] Dall, W. H. 1894. Notes on the Miocene and Pliocene of Gay Head, Martha's Vineyard, Mass., and on the "Land Phos- phate" of the Ashley River District, South Carolina. American Journal of Science, series 3, 48(286):296- 301. Dana, J. D. 1863. Manual of Geology, xvi + 798 pages, 984 figures, frontispiece, folded map. Philadelphia: Theodore Bliss. Daniels, P. A., Jr., and Emil Onuschak, Jr. 1974. Geology of the Studley, Yellow Tavern, Richmond, and Seven Pines Quadrangles, Virginia. Virginia Division of Mineral Resources Report of Investiga- tions 38: 75 pages, 13 plates, 25 figures, 5 tables. Darton, N. H. 1911. Economic Geology of Richmond, Virginia, and Vicinity. United States Geological Survey Bulletin, 483: 48 pages, 1 figure, 10 plates. Davies, J. L. 1958. The Pinnipedia: An Essay in Zoogeography. Geo- graphical Review, 48(4):474-493, 10 figures. Flower, W. H., and J. G. Garson 1884. Class Mammalia, Other Than Man. Part II in Catalogue of the Specimens Illustrating the Osteol- ogy and Dentition of Vertebrated Animals, Recent and Extinct, Contained in the Museum of the Royal College of Surgeons of England, xliii + 779 pages. London: J. Sc. A. Churchill. Gibson, T. G. 1965. Eocene and Miocene Rocks off the Northeastern Coast of the United States. Deep-Sea Research, 12(6): 975-981, 2 figures. Gilmore, C. W. 1941. A History of the Division of Vertebrate Paleon- tology in the United States National Museum. Proceedings of the United States National Museum, 90(3109):305-377, 5 plates. Gray, Asa 1875. Memorial of the Late Prof. Jeffries Wyman. Bos- ton Society of Natural History Proceedings, 17:96- 124. Guiscardi, Guglielmo 1871. Sopra un Teschio Fossile di Foca. [Memoir ex- tracted from] Atti delta R. Accademia delle Scienze Fisiche e Matematiche, Napoli, 5(6): 1-8, 2 plates. Harlan, R. 1838. [Footnote.] Page xi in T. A. Conrad, Fossils of the Medial Tertiary of the United States, xvi + 89 pages, 49 plates. Philadelphia: Judah Dobson. [Re- published, with an introduction by Dall, as Special Publication of the Wagner Free Institute of Science, Philadelphia, 1893. xviii + 136 pages.] 1842. Notice of Two New Fossil Mammals from Bruns- wick Canal, Georgia; with Observations on Some of the Fossil Quadrupeds of the United States. American Journal of Science and Arts, 53(1): 141- 144, 1 plate. Hay, O. P. 1902. Bibliography and Catalogue of the Fossil Verte- brata of North America. United States Geological Survey Bulletin, 179: 868 pages. 1930. Second Bibliography and Catalogue of the Fossil Vertebrata of North America. Carnegie Institution of Washington Publication, 390, volume II: xiv + 1074 pages. Hazel, J. E. 1969. Faunal Evidence for an Unconformity between the Paleocene Brightseat and Aquia Formations (Mary- land and Virginia). United States Geological Survey Professional Paper, 650-C:C58-C65, 5 figures, 1 table. Heite, E. F. 1964. The Tunnels of Richmond. Virginia Cavalcade (winter), 14(3):42-47, 18 figures. Hendey, Q. B., and C. A. Repenning 1972. A Pliocene Phocid from South Africa. South African Museum Annals, 59(4):71-98, 17 plates, 2 figures, 7 tables. Henshaw, Samuel 1908. Report. Pages 3-8 in Annual Report of the Curator of the Museum of Comparative Zoology at Harvard College to the President and Fellows of Harvard College for 1907-1908. 43 pages, 1 plate. 1909. Report. Pages 3-9 in Annual Report of the Curator of the Museum of Comparative Zoology at Harvard College to the President and Fellows of Harvard College for 1908-1909. 49 pages, 3 plates. Kellogg, Remington 1922. Pinnipeds from Miocene and Pleistocene Deposits of California. University of California Publications, Bulletin of the Department of Geological Sciences, 13(4):23-132, 19 figures, folding table. 1923. Description of Two Squalodonts Recently Discov- ered in the Calvert Cliffs, Maryland; and Notes on the Shark-toothed Cetaceans. Proceedings of the United States National Museum, 62(16): 69 pages, 20 plates. 1922. 24 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY King, J. E. 1964. Seals of the World. 154 pages, 17 figures, 14 plates, frontispiece, 30 maps. London: British Museum (Natural History). 1966. Relationships of the Hooded and Elephant Seals (Genera Cystophora and Mirounga). Journal of Zoology (London), 148(4):385-398, 7 figures. 1969. Some Aspects of the Anatomy of the Ross Seal, Ommatophoca rossi (Pinnipedia:Phocidae). Volume 63 in British Antarctic Survey Scientific Reports. 54 pages, 10 plates, 39 figures, 14 tables. 1972. Observations on Phocid Skulls. Pages 81-115, 25 figures in volume 1 in R. J. Harrison, editor, Func- tional Anatomy of Marine Mammals, xvii + 451 pages. London and New York: Academic Press. Leidy, Joseph 1853a. The Ancient Fauna of Nebraska. Smithsonian Con- tributions to Knowledge, 6(7): 126 pages, 24 plates, 3 figures, frontispiece. 1853b. [Description of Stenorhynchus vetus.] Academy of Natural Sciences of Philadelphia Proceedings, 6(10): 377, 1 figure. 1857. Notice of Remains of Two Species of Seals. Acad- emy of Natural Sciences of Philadelphia Proceed- ings, 8(6):265. 1865. Cretaceous Reptiles of the United States. Smith- sonian Contributions to Knowledge, 14(6): 142 pages, 20 plates, 35 figures. 1869. The Extinct Mammalian Fauna of Dakota and Nebraska, Including an Account of Some Allied Forms from Other Localities, Together with a Synopsis of the Mammalian Remains of North America. Academy of Natural Sciences of Phila- delphia Journal, second series, 7:i-vii + 8-472, 30 plates, folding map. Lydekker, Richard 1885. Catalogue of the Fossil Mammalia in the British Museum (Natural History), Part I: Primates, Chi- roptera, Insectivora, Carnivora, and Rodentia. xxx + 268 pages, 33 figures. London: British Museum (Natural History). Lyell, Charles 1843. On the Tertiary Strata of the Island of Martha's Vineyard in Massachusetts. Geological Society of London Proceedings, 4, part 1(92): 31-33. [Re- printed, 1843, London, Edinburgh, and Dublin Philosophical Magazine and Journal of Science, 33 (new series, 23): 187-189. Also, 1844, American Jour- nal of Science and Arts, 46:318-320.] 1845. Travels in North America; with Geological Obser- vations on the United States, Canada, and Nova Scotia. Volume I, xiii + 316 pages, 4 plates, 6 fig- ures. London: John Murray. [Also New York: Wiley and Putnam, with different pagination.] Mansfield, W. C. 1948. Stratigraphy of the Miocene of Virginia and the Miocene and Pliocene of North Carolina. Pages 1-19, in Julia Gardner, Mollusca from the Miocene and Lower Pliocene of Virginia and North Caro- lina, Part I: Pelecypoda. United States Geological Survey Professional Paper 199-A: 178 pages, 23 plates, 4 figures, 2 tables. Morgan, C. S. 1848. Plan of Richmond (Henrico County), Manchester and Spring Hill, Virginia. (Scale 550 feet to one inch (1:6600), 70 X 96 mm.) [Located in Library of Congress, Geographic and Map Division.] Murchison, R. I. 1843. Anniversary Address on the Progress of Geology [delivered February 17, 1843, to the Geological Society of London]. London, Edinburgh, and Dublin Philosophical Magazine and Journal of Science, third series, 22(148, supplement):511-567. Murray, Andrew 1866. The Geographical Distribution of Mammals, xvi + 420 pages, frontispiece, diagram, 101 maps. London: Day and Son. Nolan, E. J., editor 1913. An Index to the Scientific Contents of the Journal and Proceedings of the Academy of Natural Sci- ences of Philadelphia, 1812-1912. xiv + 1419 pages. Academy of Natural Sciences of Philadelphia. [Owen, Richard] 1845. Descriptive and Illustrated Catalogue of the Fossil Organic Remains of Mammalia and Aves Contained in the Museum of the Royal College of Surgeons of England, vii + 391 pages, 10 plates. London: Richard and John E. Taylor. [Published anony- mously.] Ray, C. E. 1958. Additions to the Pleistocene Mammalian Fauna from Melbourne, Florida. Museum of Comparative Zoology Bulletin, 119(7):419-450, 5 figures, 3 tables. 1975. The Relationships of Hemicaulodon effodiens Cope 1869 (Mammalia:Odobenidae). Biological Society of Washington Proceedings, 88(26):281-304, 6 plates. In press. Seals and Walruses (Mammalia: Pinnipedia) of the Yorktown Formation of Virginia and North Carolina. Smithsonian Contributions to Paleobiol- ogy. Rhees, W. J. 1882. Catalogue of Publications of the Smithsonian Insti- tution, (1846-1882), with an Alphabetical Index of Articles. Smithsonian Miscellaneous Collections, 27(4): xiv + 328 pages. Roberts, J. K. 1942. Annotated Geological Bibliography of Virginia, xi + 726 pages. Charlottesville, Virginia: Alderman Library. [University of Virginia Bibliographical Series Number Two.] Roger, Otto 1887. Verzeichniss der Bisher Bekannten Fossilen Sauge- thiere. (Zusammengestellt von Dr. Otto Roger.) Bericht des Naturwissenschaftlichen Vereins fiir Schwaben und Neuburg (a.V.) (Augsburg), 29:1- 162. NUMBER 28 25 1896. Verzeichniss der Bisher Bekannten Fossilen Saiigc- thiere. (Neu zusammengestellt von Dr. Otto Roger, etc.) Bericht des Naturwissenschaftlichen Vereins fiir Schwaben und Neuburg (a.V.) (Augsburg), 32: 1-272. Rogers, W. B. 1841. Infusorial Stratum and Associated Tertiary Beds in the Vicinity of Richmond. Section IV in Report of the Progress of the Geological Survey of the State of Virginia for the Year 1840. 132 pages. Richmond, Virginia. [Reprinted, 1881, in The Vir- ginias, A Mining, Industrial, and Scientific Journal, 2(4):58-59; also, 1884, in Geology of the Virginias, xv + 832 pages, 15 plates, geologic map, New York: D. Appleton.] 1859. [Infusorial Earth from the Tertiary Strata of Vir- ginia and Maryland.] Boston Society of Natural History Proceedings, 7(4):59-64. Rucker, M. P. 1950. Medical Sciences. Pages 277-332 in The James River Basin: Past, Present and Future, xxxi + 843 pages. Richmond: Virginia Academy of Science. Tavani, G. 1941. Revisione dei Resti del Pinnipede Conservato nel Museo di Geologia di Pisa. Palaeontographia Italica, Memorie di Paleontologia, 40(new series 10): 97-113, 2 plates, 23 figures. Teal, J. E., and L. E. Armitage 1950. Chesapeake and Ohio Railway Company. Pages 715-803 in The James River Basin: Past, Present and Future, xxxi + 843 pages. Richmond: Virginia Academy of Science. Toula, Franz 1897. Phoca vindobonensis n. sp. von Nussdorf in Wien. Beitrage zur Paldontologie und Geologie Osterreich- Ungarns und des Orients, Mittheilungen des Pald- ontologischen Institutes der Universitat Wien, 11 (2): 47-70, 3 plates. Trouessart, E.-L. 1897. Carnivora, Pinnipedia, Rodentia I. Fascicle II (pages 219-452) in Catalogus Mammalium tarn viventium quam fossilium. Nova editio. Berlin: R. Friedlander & Sohn. 1898. Sirenia, Cetacea, Edentata, Marsupialia, Allotheria, Monotremata. Fascicle V (pages 999-1264) in Catalogus Mammalium tarn viventium quam fos- silium. Nova editio. Berlin: R. Friedlander & Sohn. 1904. Primates, Prosimiae, Chiroptera, Insectivora, Carni- vora, Pinnipedia. Fascicle I (iv + 288 pages) in Catalogus Mammalium tarn viventium quam fos- silium. Quinquennale supplementum, anno 1904. Berlin: R. Freidlander & Sohn. 1905. Cetacea, Edentata, Marsupialia, Allotheria, Mono- tremata. Fascicle IV (pages vii + 753-929) in Catalogus Mammalium tarn viventium quam fos- silium. Quinquennale supplementum (1899-1904). Index alphabeticus. Berlin: R. Friedlander & Sohn. True, F. W. 1905. The First Discovery of Fossil Seals in America. Science, new series, 22(572): 794. 1906. Description of a New Genus and Species of Fossil Seal from the Miocene of Maryland. Proceedings of the Untied States National Museum, 30(1475): 835-840, 2 plates. 1912. Description of a New Fossil Porpoise of the Genus Delphinodon from the Miocene Formation of Maryland. Academy of Natural Sciences of Phila- delphia Journal, second series, 15:165-194, 10 plates. Turner, William 1887. Report on the Seals Collected during the Voyage of H.M.S. Challenger in the Years 1873-76. Part 68 in volume 26 in Zoology in Report on the Scientific Results of the Voyage of H.M.S. Challenger. 138 pages, 10 plates, 2 figures, 14 tables. Van Beneden, P.-J. 1877. Description des Ossements Fossiles des Environs d'Anvers, premiere partie: Pinnipedes ou Amphi- theriens. Musee Royal d'Histoire Naturelle de Bel- gique Annates, 1: 88 pages, 18 plates, 17 figures, 9 maps. Wilson, L. E. 1935. Miocene Marine Mammals from the Bakersfield Region, California. Peabody Museum of Natural History Bulletin, 4: 143 pages, 23 figures. Wyman, Jeffries 1844. [On the Cranium of a Seal.] Boston Society of Nat- ural History Proceedings, 1:2-3. [Verbal report at meeting of 20 January 1841, printed no later than 1843; whole volume published 1844.] 1850a. [Some Fossil Bones of Seals.] Boston Society of Natural History Proceedings, 3:241-242. 1850b. [Other Fossil Remains of Seals.] Boston Society of Natural History Proceedings, 3:323. 1850c. Notice of Remains of Vertebrated Animals Found at Richmond, Virginia. American Journal of Science and Arts, second series, 10(29):228-235, 1 plate. 26 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY PLATE 1.—Temporal bones of some phocid seals in ventral aspect: 1, Prionodelphis capensis, cast of South African Museum L15652; 2, 3, Monotherium? wymani, holotype (part), MCZ 8741; 4, Phoca groenlandica, USNM 188766; 5, Monachus schauinslandi, USNM 243854; 6, Callophoca obscura, referred, Yorktown Formation of North Carolina, USNM 187581. (Right side except figure 3, all XI.) NUMBER 28 27 V PLATE 3.—Stereophotographs of left mallei of some phocid seals, oriented with head toward top of page, articulating facets equally exposed to view, neck directed toward bottom of page, and neck and manubrium parallel to plane of page: 1, Monotherium? wymani, holotype (part), MCZ 8741; 2, Callophoca obscura, referred, Yorktown Formation of North Carolina, USNM 205516; 3, Monachus schauinslandi, USXM 213841; 4. Cystophora cristala, I'SNM 188963. (All approxi- mately X 4.) ***> 5* f WfWfw - % M ;gw PLATE 4.—Stereophotographs of left incudes of some phocid seals, oriented with body toward top of page, articulating facets equally exposed to view, long process directed toward bottom of page, and long process parallel to plane of page: 1, Monotherium? wymani, holotype (part), MCZ 8741; 2, Callophoca obscura, referred, Yorktown Formation of North Carolina, USNM 205516; 3, Monachus schauinslandi, USNM 243841; 4, Cystophora cristata. I'SNM 188963. (All approximately X 4.) - 6 %~ /S/770 PLATE 5.—Right mandibular rami and left ulnae of some Monachine seals (X 0.75): 1, 4, Monotherium? wymani, incomplete right mandibular ramus, in occlusal and lingual aspects, USNM 187410 (part) (arrow in figure 4 indicates posteriormost alveolar border); 2, 5, Leptophoca lenis, incomplete right mandibular ramus, in occlusal and lingual aspects, private collection of Derek Siddons, from beach at Stratford Hall Plantation, Virginia, Calvert? Formation (arrow in figure 5 indicates posteriormost alveolar border); 3, 6, Callophoca obscura, referred, Yorktown Formation of North Carolina, incomplete right mandibidar ramus with fifth postcanine, in occlusal and lingual aspects, USNM 181770; 7, 9, Monachus monachus, left ulna, in medial and cranial aspects, USNM 219059; 8, 10, Monotherium? wymani, left ulna, lacking distal epiphysis, in medial and cranial aspects, USNM 187410 (part). NUMBER 28 31 * * *IC ^^^ytxjg^ O 0*- 01 PLATE 6.—Left tibiae and fibulae of some monachine seals in cranial aspect (X 0.75): 1, Monotherium? wymani, associated tibia and fibula, lacking distal epiphyses, and the fibula lack- ing part of shaft, USNM 187410 (part); 2, Monotherium? wymani, cast of distal fragment of fibula, USNM 214650; 3, Monachus monachus, tibia and fibula (fused proximally) USNM 219059. 32 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY 1 ' V PLATE 7.—Left tibiae and fibulae of some monachine seals in caudal aspect (X 0.75): 1, Monotherium? wymani, cast of distal fragment of fibula, USNM 214650; 2, Monotherium? wymani, associated tibia and fibula, lacking distal epiphyses, and the fibula lacking part of shaft, USNM 187410 (part); ;5, Monachus monachus, tibia and fibula (fused proximally), USNM 219059. NUMBER 28 33 "° r: c-f -5 « e . be ~ . M T3 _, S -3 rt £ « 2 ?i <» © "^ •*= 5; rt o ■2 S,g .§ ^ § Q eo" P ' ■■•&%.-:$L '''"- •.v» u § z .y - pi 3 .- G"1 ^ "5b u: <_• ^ in ,-; *J CN rt ,C _52 Z "rt .. Z S rt 5 O rt> n op ffi Ml rt2 ^ 4J CA a E x S O O o c „, * -S o bo-2 c c B 2 £ -C I-I TJ ^ c .. ■^ ^ cr, O ■rt Gi t/j .^ NUMBER 28 35 CO *u Z "2 5 *J ^j K* ;*-"j?.">- i# cu a 13 2 2 r °-~ rt bo-2 a s o 1/5 O 2 _, C c "3 rt « 3 . O 1-1 "O "5 c h o a a .3 —• —. s «a ■»< - .; U y ■- I rt * _■ o G _ ->*i oO ;2 < Sg P 5 ■3|s Q M D 36 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY SN.S s rt > u T3 3 O 6 Z a -3 u o rt 5" >s '5b o « rt u O CJ T3 *j CO GJ J- o 13 S =3 Z a P •3 ■s rt o "» -3 3 wa "rt ■S.S ° cy c3 a s o » 3 U (LI a a = o bo £ rt1 - a 51 — ' *- CN 5 73 <=■> rt J2 S to W '•g CV *s 3 ■« Z g C CO ■C « D CO — rt ^J CV 3 a 3- if bo . 3 CJ ^J 3 -O 1 „- t A. £*'£ -• £ « sc - *,2 G 5 *i *> ^ .§^ § Q eo S ft U.S. GOVERNMENT PRINTING OFFICE: 1 976— 211 -S59 /e Publication in Smithsonian Contributions to Paleobiology Manuscripts for serial publications are accepted by the Smithsonian Institution Press, sub- ject to substantive review, only through departments of the various Smithsonian museums. Non-Smithsonian authors should address inquiries to the appropriate department. If submission is invited, the following format requirements of the Press will govern the preparation of copy. Copy must be typewritten, double-spaced, on one side of standard white bond paper, with \l/i" top and left margin, submitted in ribbon copy with a carbon or duplicate, and accom- panied by the original artwork. Duplicate copies of all material, including illustrations, should be retained by the author. There may be several paragraphs to a page, but each page should begin with a new paragraph. Number consecutively all pages, including title page, abstract, text, literature cited, legends, and tables. The minimum length is 30 pages, including typescript and illustrations. The title should be complete and clear for easy indexing by abstracting services. Taxonomic titles will carry a final line indicating the higher categories to which the taxon is referable: "(Ammonoidea: Goniatitidae)." Include an abstract as an introductory part of the text. Identify the author on the first page of text with an unnumbered footnote that includes his professional mailing address. A table of contents is optional. An index, if required, may be supplied by the author when he returns page proof. Two headings are used: (1) text heads (boldface in print) for major sections and chapters and (2) paragraph sideheads (caps and small caps in print) for subdivisions. Further headings may be worked out with the editor. In taxonomic keys, number only the first item of each couplet; if there is only one couplet, omit the number. For easy reference, number also the taxa and their corresponding headings throughout the text; do not incorporate page references in the key. In synonymy, use the short form (taxon, author, date:page) with a full reference at the end of the paper under "Literature Cited." Begin each taxon at the left margin with sub- sequent lines indented about three spaces. Within an entry, use a period-dash (.—) to separate each reference. Enclose with square brackets any annotation in, or at the end of, the entry. For references within the text, use the author-date system: "(Jones, 1910)" and "Jones (1910)." If the reference is expanded, abbreviate the data: "Jones (1910:122, pl. 20: fig. 1)." Simple tabulations in the text (e.g., columns of data) may carry headings or not, but they should not contain rules. Formal tables must be submitted as pages separate from the text, and each table, no matter how large, should be pasted up as a single sheet of copy. Use the metric system instead of, or in addition to, the English system. Illustrations (line drawings, maps, photographs, shaded drawings) can be intermixed throughout the printed text. They will be termed Figures and should be numbered con- secutively; however, if a group of figures is treated as a single figure, the components should be indicated by lowercase italic letters on the illustration, in the legend, and in text references: "Figure 9b." If illustrations (usually tone photographs) are printed separately from the text as full pages on a different stock of paper, they will be termed Plates, and individual components should be lettered (Plate 9b) but may be numbered (Plate 9: figure 2). Never combine the numbering system of text illustrations with that of plate illustrations. Submit all legends on pages separate from the text and not attached to the artwork. An instruction booklet for the preparation of illustrations is available from the Press on request. In the bibliography (usually called "Literature Cited"), spell out book, journal, and article titles, using initial caps with all words except minor terms such as "and, of, the." For capitalization of titles in foreign languages, follow the national practice of each language. Underscore (for italics) book and journal titles. Use the colon-parentheses system for volume, number, and page citations: "10(2) :5-9." Spell out such words as "figures," "plates," "pages." For free copies of his own paper, a Smithsonian author should indicate his requirements on "Form 36" (submitted to the Press with the manuscript). A non-Smithsonian author will receive 50 free copies; order forms for quantities above this amount with instructions for payment will be supplied when page proof is forwarded.