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The Anseriform Relationships of Anatalavis Olson and Parris (Anseranatidae), with a New Species from the Lower Eocene London Clay

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The Anseriform Relationships of Anatalavis Olson and Parris (Anseranatidae), with a New Species from the Lower Eocene London Clay The Anseriform Relationships of Anatalavis Olson and Parris (Anseranatidae), with a New Species from the Lower Eocene London Clay Storrs L. Olson more recent studies of Ericson (1996, 1997) confirmed the lack ABSTRACT of relationship between the Anseriformes and Galliformes, but the ancestry of the Anseriformes was unresolved beyond a All associated partial skeleton, including the skull but lacking legs, from the lower Eocene London Clay of Essex, England, pos- complex of various groups of wading birds, including sesses derived characters of the coracoid and furcula that show it Charadriiformes. Presbyornis, however, was determined to to belong to the Anseranatidae, which previously had no fossil have branched off within the order and constitutes the sister record. Except for its much larger size, the humerus of this speci- group of the Anatidae proper, with the Anhimidae and Anser- men is identical to that of Anatalavis rex (Shufeldt) from the late anatidae being the primitive outliers of the Presbyornithidae/ Cretaceous or early Paleocene of New Jersey. The Eocene speci- men is described as a new species, Anatalavis oxfordi, and the Anatidae clade. genus Anatalavis is transferred from the form-family Graculavidae The giant Paleocene and Eocene groundbirds of the genus to a new subfamily, Anatalavinae, of the Anseranatidae. Anatala- Diatryma, once thought to have been predatory descendants of vis is characterized by a very broad duck-like bill, a proportion- crane-like birds, also may be part of the anseriform radiation ately very short and robust humerus, and an anterior portion of the (Andors, 1988, 1992). The dietary habits of Diatryma, how- pelvis resembling that of ibises and other wading birds more than ever, have been equivocated (Andors, 1992; Witmer and Rose, that of any known anseriform. Other features of its osteology are unique within the order. 1991). Although no fossils of screamers (Anhimidae) had hitherto been reported, a somewhat more primitive genus is now known from the middle Tertiary of Brazil (Alvarenga, this volume), Introduction and I have examined excellent fossils, as yet undescribed, from Waterfowl of the order Anseriformes are among the best the lower Eocene Willwood Formation of Wyoming and from known and most distinctive groups of modern birds. Although the contemporaneous London Clay of England. Thus, of the three major lineages of living Anseriformes, the only one with waterfowl are abundantly represented by Neogene fossils, no early Tertiary (or later) fossil representative is the Anser- much of their early evolutionary history has remained obscure. anatidae, with its sole member being the Magpie Goose, Anser- The most informative fossils until now have been the Pale- anas semipalmata, of Australia. ocene and early Eocene remains of Presbyornis and its close Two bones from the Homerstown Formation of New Jersey, relatives, which were nearly cosmopolitan in the early Paleo- a deposit of debated Late Cretaceous or early Paleocene age gene. Presbyornis was shown to have a duck-like skull on the (Olson, 1994; Hope, this volume), were thought to show some body of a long-legged wading bird and was interpreted as similarities to the Anatidae, but in the absence of associated showing a derivation of the Anseriformes from a charadrii- material they were assigned to the form-family Graculavidae, form-like ancestor (Olson and Feduccia, 1980b) rather than which contains various taxa resembling primitive Charadrii- from the Gafliformes, as had been postulated previously. The formes (Olson and Parris, 1987). A new fossil from the London Clay consisting of much of a skeleton, although lacking legs, Storrs L. Olson, Department of Vertebrate Zoology, National Museum permits positive identification of the New Jersey fossils as not of Natural History, Smithsonian Institution, Washington, D.C. 20560- only belonging to the Anseriformes but belonging to the family 0116, United States. Anseranatidae. Thus, the New Jersey and London Clay fossils 231 232 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY provide the first record of this family, as well as the earliest characteristic "duck-billed" shape that may be modified sec- certain occurrence of the entire order. ondarily for other feeding functions. The following features The present paper is intended to provide a name for the new also are characteristic of the Anseriformes, some being conver- fossil from the London Clay and to place on record its more sa- gent with Galliformes (Olson and Feduccia, 1980b; Ericson, lient osteological features. Full analysis of anatomical details 1996): the configuration of the quadrato-mandibular articula- and phylogenetic significance will have to await the appear- tion; the enlarged, deep, curved, blade-like retroarticular pro- ance of information not presently available for certain extant cess of the mandible; and the enlarged rounded or ovoid "ba- but sequestered fossils, especially those of Eocene Anhimidae, sipterygoid process" on the parasphenoid rostrum, with a Nomenclature for species' binomials and English names of corresponding enlarged facet on the pterygoid. These charac- modern birds herein follows Sibley and Monroe (1990). ters are practically all that exist to demonstrate the anseriform ACKNOWLEDGMENTS.—I must begin by crediting the singu- relationships of both living and fossil screamers (Anhimidae), lar cooperativeness of Andrew Oxford, who collected the spec- in which the bill has either lost or never had the adaptations for imen that is the primary object of the present study and who, at filter-feeding. The holotype of the new species described below my first suggestion, donated it to the Natural History Museum, has a skull with all of the features typical of the order Anseri- London (formerly the British Museum (Natural History)) formes, to which it clearly belongs. (BMNH). In connection with this transaction, I cannot fail to mention that the hospitality shown to me and my family by Mr. and Mrs. Oxford at their domicile at Great Mongeham, Deal, Family ANSERANATIDAE Kent, afforded us some of our most pleasant memories generat- Recognition of the living Australian Magpie Goose, Anser- ed during a pleasant year in England. Mr. Oxford and I each anas semipalmata, as a monotypic family of Anseriformes has owe a debt of gratitude to that veteran collector of fossil birds been supported by several anatomical studies (e.g., Verheyen, of the London Clay, Michael Daniels, who separately intro- 1953; Woolfenden, 1961; Livezey, 1986), and the distinctive- duced us to the incredibly productive mudflats at Wal- ness of this species is confirmed by DNA sequencing as well ton-on-the-Naze, who led me to contact Mr. Oxford about his (Michael Sorensen, University of Michigan, pers. comm., fossil bird, and whose comments on the manuscript inspired me 1996). Two presumably derived characters group the fossil ge- to delete much that was equivocal. The repeated hospitality of nus Anatalavis in the same family as Anseranas. The first is the Mr. and Mrs. Daniels at their home in Holland-on-Sea, Clac- unique V-shaped furcula with a large, deep symphysis. In other ton, Essex, has been of inestimable benefit to me for my members of the order the furcula is an unelaborated U-shaped knowledge of early Eocene birds. structure with a symphysis that is scarcely, if at all, larger than At the BMNH, Angela Milner greatly facilitated the present the rami. I interpret the former condition to be a derived char- study by making arrangements for the incorporation of the specimen into the collections and having it prepared and photo- acter within the Anseriformes. If it is primitive, the only out- graphed. In this connection, there are hardly sufficient words of groups that show any similarity (and this only in a general way) praise for William Lindsay, whose painstaking removal of an are storks (Ciconiidae) and, to an even lesser extent, herons exceedingly fragile and difficult specimen from its envelope of (Ardeidae). The second character is the presence of a distinct, clay and pyrite was undertaken in such a spirit of collaboration large pneumatic foramen in the dorsal surface of the sternal end and sensitivity that his contribution must rank equal to that of of the coracoid. Although a similar condition exists in modern any scientist who studies these bones. Sandra Chapman has re- Anhimidae, which have one of the most pneumatized skeletons peatedly been of assistance during my study of fossil bird col- in any group of birds, this foramen is absent in Eocene Anhim- lections at the BMNH. Robert Prys-Jones and Cyril Walker, of idae, which are evidently completely nonpneumatic (pers. the Bird Group at Tring, were instrumental in lending compar- obs.). Therefore, the condition in modern screamers is obvious- ative modern skeletons. The photographs are by Phil Crabb, ly independently derived and is not an indication of relation- Natural History Museum, London, Photo Unit, except Figure ship with the Anseranatidae. 8c,D, which is by the Smithsonian Photographic Services (SPS). Carl Hansen (SPS) was instrumental in assisting with Subfamily ANSERANATINAE the electronic composition of the figures. INCLUDED GENUS.—Anseranas Lesson. DIAGNOSIS.—Subfamilial characters are intended as diag- Order ANSERIFORMES nostic only within the Anseranatidae. Rostrum strong, deep, The primary adaptation of the order Anseriformes is the and hooked; frontal area with a large bony casque; attachment modification of the bill so that the upper jaw houses an en- for M. depressor mandibulae not greatly developed. Humerus larged tongue, which functions as a double-piston pump used of normal anseriform proportions. Pelvis with anterior portions in filter-feeding (Olson and Feduccia, 1980a), giving rise to a of the ilia narrow. NUMBER 89 233 Subfamily ANATALAVINAE, new subfamily terior margin of nostril to bill tip, 34; maximum width of bill as preserved, 27.5. INCLUDED GENUS.—Anatalavis Olson and Parris, 1987. Skull (measurements taken from dorsal aspect): Width of DIAGNOSIS.—Rostrum very broad and shallow, not hooked; interorbital bridge, 21.5; width of frontals at nasofrontal hinge, frontal area without bony casque; attachment for M.
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