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Norntates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y AMERICAN MUSEUM Norntates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2737, pp. 1-22, figs. 1-1 3 June 29, 1982 Cranial Morphology of the Baenid Turtles EUGENE S. GAFFNEY' ABSTRACT The family Baenidae is a group of extinct cryp- living cryptodires, the Eucryptodira. The arterial todiran turtles known from more than a dozen canals of the skull are preserved in enough spec- well-preserved skulls from the Cretaceous, Paleo- imens to conclude that baenids had an arterial cene, and Eocene. The baenid skull is character- pattern characterized by a large and well-devel- ized by significant generalized features in the skull oped canalis stapedio-temporalis and a small but roofand basicranium that are consistent with their persistent canalis caroticus lateralis. hypothesized relationship as the sister group to the INTRODUCTION The Baenidae is an important group of included diagnoses, synonymies, lists ofspec- Cretaceous to Eocene non-marine turtles that imens, skull and shell figures, and a review have been considered as the sister group to of the literature up to 1972. Descriptions all living cryptodires (Gaffney, 1975). While were lacking and the present work is an effort information on the systematics of this group to fill this lack for skulls. The relationships has been available (Gaffney, 1972), a more of taxa within the Baenidae were also dis- complete study ofthe cranial morphology has cussed in Gaffney (1972), but in 1975 I dealt been lacking. The present paper is intended with the relationships ofbaenids to other tur- to fill this void, particularly in the description tles and the question of the "Amphichely- of basicranial regions. A number of impor- dia," a supposedly ancestral turtle group that tant figures of baenid crania have already was characterized by baenids. Evans and appeared in Gaffney (1972, 1979b) and Ar- Kemp (1975, 1976) described two skulls, chibald and Hutchison (1979). To eliminate Mesochelys and Dorsetochelys, and discussed repetition, I assume that the reader has these relationships of these taxa to baenids and three papers available and is familiar with other turtles. Gaffney (1979a) described the contents of Gaffney (1979b). Glyptops and presented an analysis of rela- tionships involving this genus, Mesochelys, Dorsetochelys, and baenids. The most im- PREVIOUS WORK portant work on taxa within the Baenidae Gaffney (1972) provided the primary basis since 1972 is Archibald and Hutchison's pa- for current systematic work on baenids and per (1979) on Palatobaena which described ' Curator, Department of Vertebrate Paleontology, American Museum of Natural History. Copyright g American Museum of Natural History 1982 ISSN 0003-0082 / Price $1.85 2 AMERICAN MUSEUM NOVITATES NO. 2737 new, well-preserved material of this hereto- MORPHOLOGY fore poorly known form and proposed a new NASAL: Nasals are characteristically re- scheme of its phylogenetic relationships. An tained in baenids and lost in most eucryp- important but as yet unpublished work is todires. In baenids the nasals retain the prim- Archibald (MS), which describes the first itive (for amniotes) posterior contact with the skull-shell associations of Eubaena and Sty- frontals, whereas in those eucryptodires with giochelys and proposes a baenid phylogeny nasals the prefrontals meet medially and pos- based on shell features. Gaffney's cranial terior to the nasals. morphology review (1979b) has whole skull In most baenids (Gaffney, 1979b, fig. 1) drawings of all the baenid genera (but does the nasal contacts the maxilla laterally, the not repeat the important palatal stereopho- frontal posteriorly, and the other nasal me- tographs from Gaffney, 1972), as well as a dially. The anterior edge is free and forms the series of figures showing more detailed, in- dorsal margin ofthe apertura narium externa. ternal features of baenid skulls. Lower jaws Within the baenids, Hayemys is unique in of baenids are dealt with to some extent in having exceptionally large nasals that are lat- Gaffney (1979b) and Archibald and Hutch- erally and posteriorly expanded in compari- ison (1979) but a detailed study is in prepa- son to other baenid nasals. In Baena the na- ration. Also under way is a new phylogenetic sals are either lost or fused to the frontals. analysis and classification of the family. Sutures differentiating nasals are not visible in any ofthe numerous specimens (including ACKNOWLEDGMENTS presumedjuveniles) ofBaena, suggesting loss I am grateful to the curators and collection rather than fusion. The anterior margin of managers of the institutions from which I the apertura narium externa in Baena is have borrowed baenid specimens over the thickened with an anterior facing concavity, years (a complete institutional list is in Gaff- similar to one seen in Palatobaena bairdi. ney, 1972). Dr. David Archibald, Yale Uni- Chisternon is unique among baenids in lack- versity, very kindly allowed me to use infor- ing a nasal-maxilla contact, this being caused mation from his unpublished Ph.D. thesis by an anterior extension of the frontal. and with the aid of Dr. Howard Hutchisop, In Palatobaena the preorbital part of the University of California, Berkeley, made skull is highly modified from the general specimens available to me that I otherwise baenid pattern by extreme shortening antero- would not have been aware of. posteriorly and widening laterally. Nasals are Ms. Lorraine Meeker drew figures 6 and known only in one specimen of Palatobaena 13, Mr. Chester Tarka took the photographs bairdi (CCM 77-11, figured in Gaffney, in figures 3 and 4, Mrs. Jennifer Emry drew 1979b; Archibald and Hutchison, 1979) figures 1 and 2, the American Museum where they are seen to be widely separated Graphic Arts Department did figure 8. 1 very from each other by the extension ofthe fron- much appreciate the assistance of these peo- tals into the apertura narium externa. Al- ple. I took the remaining photographs. though the limits ofthe nasals are not entirely clear in this specimen they do show a dor- ABBREVIATIONS solateral and medial contact with the frontal and ventral contact with the maxilla. The rim AMNH, American Museum of Natural History of the entire apertura narium externa is re- CCM, Carter County Museum cessed in Palatobaena bairdi, whereas in P. MCZ, Museum ofComparative Zoology, Harvard University gaffneyi (Archibald and Hutchison, 1979) PU, Princeton University only the ventral margin is recessed. There are UCMP, University ofCalifornia at Berkeley, Mu- no signs of nasals in this last specimen. seum of Paleontology The ventral morphology of the nasal area USNM, National Museum of Natural History (Gaffney, 1979b, fig. 3) has paired troughs UW, University of Wyoming concave ventrally and separated by a median YPM, Yale Peabody Museum ridge along which the two nasals meet. The 1982 GAFFNEY: BAENID TURTLES 3 E FIG. 1. Disarticulated baenid bones from the Late Cretaceous, Hell Creek Formation, Bug Creek Anthills, McCone County, Montana. A. Ventral view of right frontal (MCZ 355 1), a presumed juvenile, 9.6 mm. in length. B. Ventral view of right frontal (MCZ 3554), 22.7 mm. in length. C. Dorsal view of supraoccipital (MCZ 3558), 12.8 mm. in length. D. Ventral view of squamosal (AMNH 8558), 16.6 mm. in length. E. Lateral view of left parietal (MCZ 3526), presumed juvenile, 21.4 mm. in length, anterior is to left, dashed line indicates foramen nervi trigemini. troughs housed the olfactory bulbs and are primitive chelonian condition, whereas eu- an anterior extension of the sulcus olfacto- cryptodires typically have conspicuously rius. large dorsal prefrontal plates that meet in the PREFRONTAL: The prefrontal in turtles usu- midline. Hayemys, however, does have a ally consists oftwo plates, a horizontal dorsal moderately large dorsal plate that probably one and a vertical ventral one. As empha- approximates the primitive condition, but at sized by Evans and Kemp (1976) and dis- present I interpret this as an autapomorphy cussed by Gaffney (1979a) baenids are gen- for Hayemys. erally characterized (see Gaffney, 1979b, fig. The ventral process of the prefrontal is a 1) by a reduction of the dorsal plate in com- broadly curved plate oriented in a roughly parison with what is hypothesized as the vertical plane, extending anterolaterally, and 4 AMERICAN MUSEUM NOVITATES NO. 2737 is a mere vertical slit in Glyptops, whereas in the Baenidae it is wider but not so wide as in the Testudinidae. In my earlier paper on baenids I figured Trinitichelys as lacking a dorsal exposure of the prefrontal (Gaffney, 1972, fig. 2). Since then I have been convinced by other students ofthis specimen that a small prefrontal lappet is present and have modified the new recon- struction accordingly (Gaffney, 1979b, fig. 164). Hay (1904, 1905, 1908) reported the pres- ence of a lacrimal in Eubaena, Baena, and Chisternon. Hay's "lacrimal" is here identi- fied as the prefrontal in view ofthe following features held in common between the pre- A - B frontal of baenids and living cryptodires: (1) articulates with the vomer ventrome- dially (2) borders the foramen orbito-nasale ventrolaterally (3) meets the palatine between the two above structures (4) has a long suture with the maxilla along the anterior border of the fossa orbi- talis (5) forms the anterior wall of the fossa orbitalis FIG. 2. A. Dorsal view of left maxilla, Pala- The only significant distinction between tobaena bairdi (AMNH 8550). B. Dorsal view of them seems to be the small or absent dorsal left maxilla, Eubaena cephalica (MCZ 3519). C. plate in baenids. Hay's identification of this Medial view of right maxilla (MCZ 3519, re- element as the lacrimal required further iden- versed), Eubaena cephalica. All are Late Creta- tifications to be consistent with that deter- ceous, Hell Creek Formation, Bug Creek Anthills, mination. A reevaluation suggests the follow- McCone County, Montana.
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