Bulletin~ of the American Museum of Natural History Volume 87: Article 1 New York: 1946 - X X |! |

GEORGE GAYLORD SIMPSON BULLETIN~ OF THE AMERICAN MUSEUM OF NATURAL HISTORY VOLUME 87: ARTICLE 1 NEW YORK: 1946 - X X |! | - -s s- - - - - -- -- --| c - - - - - - - - - - - - - - - - - -- FOSSIL PENGUINS FOSSIL PENGUINS GEORGE GAYLORD SIMPSON Curator of Fossil Mammals and Birds PUBLICATIONS OF THE SCARRITT EXPEDITIONS, NUMBER 33 BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY VOLUME 87: ARTICLE 1 NEW YORK: 1946 BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Volume 87, article 1, pages 1-100, text figures 1-33, tables 1-9 Issued August 8, 1946 CONTENTS INTRODUCTION . 7 A SKELETON OF Paraptenodytes antarcticus. 9 CONSPECTUS OF TERTIARY PENGUINS . 23 Patagonia. 24 Deseado Formation. 24 Patagonian Formation . 25 Seymour Island . 35 New Zealand. 39 Australia. 42 COMPARATIVE OSTEOLOGY OF MIOCENE PENGUINS . 43 Skull . 43 Vertebrae 44 Scapula. 45 Coracoid. 46 Sternum. 49 Humerus. 49 Radius and Ulna. 53 Metacarpus. 55 Phalanges . 56 The Wing as a Whole. 56 Femur. 59 Tibiotarsus. 60 Tarsometatarsus 61 NOTES ON VARIATION. 65 TAXONOMY AND PHYLOGENY OF THE SPHENISCIDAE . 68 DISTRIBUTION OF MIOCENE PENGUINS. 71 SIZE OF THE FOSSIL PENGUINS 74 THE ORIGIN OF PENGUINS. 77 Status of the Problem. 77 The Fossil Evidence . 78 Conclusions from the Fossil Evidence. 83 A General Theory of Penguin Evolution. 84 A Note on Archaeopteryx and Archaeornis 92 ADDENDUM . 96 BIBLIOGRAPHY 97 5 INTRODUCTION FEW ANIMALS have excited greater popular basis for comparison, synthesis, and gener- and scientific interest than penguins. Their alization, in spite of the fact that relatively quaintly human appearance, their peculiar few distinguishable species are represented. habits, the ability of some of them to thrive Several years elapsed before the collection in the bitter Antarctic cold, their association could be fully prepared, and it was then with remote coasts and islands, and their hoped that some specialist on penguins would radical differences from all other birds have be able to undertake the description of these long made them favorite subjects of layman materials, but such an opportunity did not and student alike. They appear regularly in arise. With the reorganization of paleontol- cartoons, advertisements, moving pictures, ogy in the American Museum in November, popular books, and magazine articles, and 1944, curatorial responsibility for fossil birds with about equal regularity, if less frequently, was returned to the Department of Geology in the technical literature, where their habits and Paleontology and was assigned specifi- and structure have been described in great, cally to me within the department. In partial but still incomplete, detail and where their fulfillment of this responsibility and in con- origin and history have been and still are bit- tinuation of our program on the Tertiary terly disputed. faunas of South America, the present study Fossil penguins were made known as long has been prepared in order to make available ago as 1859 and by no less an authority than the essential facts about these important fos- T. H. Huxley. Since then, some hundreds of sils. specimens have been collected in several In the course of identification and descrip- widely distant parts of the world, and many tion of our collection, it was necessary to re- of these have been exhaustively described, view literature on other known fossil pen- but knowledge has remained very incomplete guins. It quickly became obvious that a gen- and in part quite confused. The known speci- eral taxonomic revision could not be made mens are all incomplete. No fragment of the under existing circumstances, but it has skull has hitherto been described and in few seemed worth while to present notes on tax- instances have two or more surely associated onomy and to review available knowledge of bones of one individual been found. From the all fossil penguins and not alone those in our scattered literature it has been most difficult, collection. Although it is not only, or even on some points impossible, to gather a clear mainly, dependent on study of the extinct and correct idea of the variety and characters forms, the vexatious problem of penguin of the known fossil penguins as a whole. A origins is also clearly pertinent and some complete and satisfying review cannot now comments on it have been appended. be made, but a contribution toward this end Comparative osteological material of re- is presented. cent penguins has been provided by the De- The present study grew out of the discov- partment of Birds of this Museum, with the ery in 1933 by the Second Scarritt Patago- cooperation of Dr. Murphy and Mr. O'Brien. nian Expedition of a fossil penguin skeleton, The osteological collection unfortunately far from complete but nevertheless much the lacks the important genera Eudyptula and best that has hitherto been found. It includes Megadyptes, but it is rich in most of the spe- for the first time the greater part of a skull, cies of all other living genera. Previous stud- and it also comprises more different parts of ies of penguin osteology have, of course, also the skeleton of one individual than had ever been utilized throughout, even when not ex- been found in association. In addition to this plicitly cited on a given point. The avail- unique skeleton, the expedition also col- ability of these works, especially Gervais lected more than a hundred other specimens and Alix (1877), Pycraft (1898), Shufeldt of fossil penguins, constituting an admirable (1901), Virchow (1931), and Watson (1883), 7 8 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 87 makes it unnecessary to review the osteology In resuming work on Patagonian fossils, I of the Spheniscidae as a whole and possible wish again to express gratitude to Mr. H. S. to concentrate on some of the more striking Scarritt, the principal sponsor of our collect- and pertinent comparative differences within ing in that region. The specimens involved in the group.' this study were collected by Coleman S. Wil- liams, Justino Hernandez, and me, and were 1 Except for the skull, even the most detailed osteo- prepared mostly by Albert Thomson. The il- logical studies of recent penguins have,-however, paid too little attention to their variation and to their lustrations have been prepared by John C. taxonomic differentiation. Even Watson's careful de- Germann. scription is markedly deficient in detecting and stating differences between the various species and genera. A SKELETON OF PARAPTENODYTES ANTARCTICUS OCCURRENCE pointed out in the description below, the THE SPECIMEN HERE TO BE DESCRIBED iS homologous bones of our specimen differ A.M.N.H. No. 3338. It was found by Justino from the type in various minor details. By Hernandez, on the Second Scarritt Patago- standards that have been applied to fossil nian Expedition, in November, 1933. The penguins, especially by Ameghino, these dif- locality is on the south side of the Rio ferences would justify specific or even generic Chubut, about 3 leagues above (west of) separation, but I am convinced that this is Trelew and directly opposite the settlement not correct and that the specific reference is of Gaiman, in Chubut Territory, Argentina. well founded. Equally great differences occur Part of the specimen had weathered out, but within a single species among modern pen- parts were in situ and were removed in a guins. The general agreement in size and block of matrix. structure is close, and no other Patagonian The stratigraphy of this region has been species in the long list established by Ame- discussed elsewhere (Simpson, 1935). The ghino is liable to confusion with this. The penguin skeleton was found in the lower part type is from Santa Cruz Territory, from the of "3" of Section 1 of the paper cited (fig. 1 Middle Patagonian at the mouth of the Rio and pp. 6-7). This stratigraphic unit con- Santa Cruz, according to Ameghino, and our sists of about 35 meters of thick-bedded, specimen is from the Lower Patagonian of pale, yellowish tuff. It represents the first the Chubut Valley, but all the evidence is depositional phase in this region of the that the penguin faunas are identical in Patagonian marine incursion, of disputed age these two regions and at these different levels, but now considered Early Miocene by a which differ little if at all in age. Moreno and strong consensus. It immediately overlies the Mercerat (1891) also referred to this species terrestrial Trelew beds, of late Colhue-Huapi three isolated bones, ulna, femur, and tibio- age (probably latest Oligocene, perhaps tarsus, from Chubut-exact locality not earliest Miocene), with an erosional but not given but doubtless near Trelew or Gaiman angular uncomformity apparently represent- and from the same beds as our specimen. ing only a slight hiatus in time. The species is relatively but not extremely Besides this skeleton, numerous isolated rare. The type, our skeleton, and the three penguin bones were found in the same bed, isolated bones mentioned by Moreno and along with remains of sharks, rays, whales, Mercerat are the only surely identified re- and sparse Ostrea hatcheri. Our party found mains reported in the literature or known to no land mammals at this level, but these were me. Ameghino (1895) also mentioned a meta- probably correctly reported by Frenguelli carpus and the anterior part of a mandible, (1926), although there is some lingering of unstated origin, but, as he later (1905) re- doubt about this because of his failure to dis- marked about the mandible, the reference is tinguish the terrestrial Trelew from the mar- not well established. As noted below, the ine Patagonian at this locality. In any case, humerus referred by Ameghino (1905) to there is little doubt that the deposit was Perispheniscus wimani probably belongs to formed in shallow water and is littoral or Paraptenodytes antarcticus.

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