Malcolm C. McKenna

A Theropod Dinosaur Embryo and the Affinities of the Flaming Cliffs Dinosaur Eggs

An embryonic skeleton of a nonavian theropod dinosaur was found preserved in an egg from Upper Cretaceous rocks in the Gobi Desert of Mongolia. Cranial features identify the embryo as a member of Oviraptoridae. Two embryo-sized skulls of dromaeosaurids, similar to that of Velociraptor, were also recovered in the nest. The eggshell microstructure is similar to that of ratite birds and is of a type common in the Djadokhta Formation at the Flaming Cliffs (Bayn Dzak). Discovery of a nest of such eggs at the Flaming Cliffs in 1923, beneath the Oviraptor philoceratops holotype, suggests that this dinosaur may have been a brooding adult.

Eocene paleolatitude, climate, and mammals of Ellesmere island

Abstract During the early and middle Eocene Ellesmere Island was situated at almost the same latitude as at present. Marine paleotemperatures locally reached 15°C. Mean paleo-obliquity of Earths spin axis was substantial and there is no physical model for any other value than the present mean value. In agreement with geochemical and additional biological data, the Ellesmere mammalian fauna indicates a warm climate in the Arctic about 50 m.y. ago. How this can be compatible with significant periods of winter darkness remains a problem, but decreased solar energy production after the middle Eocene is an uninvestigated possibility. The Donn and Shaw (1977) model of Eocene paleoclimate is in conflict with several lines of evidence presented here.

Epipubic bones in eutherian mammals from the Late Cretaceous of Mongolia

An important transformation in the evolution of mammals was the loss of the epipubic bones. These are elements projecting anteriorly from the pelvic girdle into the abdominal region in a variety of Mesozoic mammals, related tritylodonts, marsupials and monotremes but not in living eutherian (placental) mammals,,. Here we describe a new eutherian from the Late Cretaceous period of Mongolia, and report the first record of epipubic bones in two distinct eutherian lineages. The presence of epipubic bones and other primitive features suggests that these groups occupy a basal position in the Eutheria. It has been argued that the epipubic bones support the pouch in living mammals,,, but epipubic bones have since been related to locomotion and suspension of the litter mass of several attached, lactating offspring. The loss of the epipubic bones in eutherians can be related to the evolution of prolonged gestation, which would not require prolonged external attachment of altricial young. Thus the occurrence of epipubic bones in two Cretaceous eutherians suggests that the dramatic modifications connected with typical placental reproduction,, may have been later events in the evolution of the Eutheria.

Earliest Eutherian Ear Region: A Petrosal Referred to Prokennalestes from the Early Cretaceous of Mongolia

Abstract A right petrosal from the ?Aptian or Albian Khoobur locality is referred on the basis of size and morphology to Prokennalestes trofimovi, the earliest eutherian previously known only from dentigerous elements. The petrosal shows a mosaic of primitive and derived features, bearing on the purported therian and eutherian morphotypes. Among the primitive features shared with the Early Cretaceous prototribosphenidan Vincelestes and other more basal taxa that are modified in later eutherians and metatherians are the pattern of basicranial arterial and venous circulation, including a prootic canal and an intrapetrosal inferior petrosal sinus; a vertical paroccipital process; and a fenestra semilunaris, an incomplete wall between the cavum epiptericum and cavum supracochleare. Among the derived features shared with therians is a cochlea coiled through a minimum of 360°, with Prokennalestes extending the range of the oldest occurrence of such a coiled cochlea by at least 10 million years. Shared with Late Cretaceous eutherians is a shallow internal acoustic meatus with a thin prefacial commissure. The petrosal referred to Prokennalestes is intermediate in having a reduced anterior lamina and lateral flange, both of which are well developed in Vincelestes and essentially lacking in later eutherians and metatherians. Features previously held to be part of the therian and eutherian morphotypes, such as the absence of the anterior lamina and lateral flange, may have been lost independently in metatherians and in post-Prokennalestes eutherians.

New evidence for late mesozoic-early Cenozoic evolution of the Chilean Andes in the upper Tinguiririca valley (35 °S), central Chile

Abstract New geologic, paleontologic and isotopic geochronometric results from the Termas del Flaco region in the upper Tinguiririca River valley in central Chile demand considerable revision of the accepted geotectonic history of the Andean Main Range in this region. A diverse, transitional Eocene-Oligocene aged, land-mammal fauna was recovered from several sites in volcaniclastic sediments of the Coya-Machali (=Abanico) Formation. Major results of our study include: 1) The 1000 + m thick studied deposits, previously attributed to the Cretaceous Colimapu Formation, belong to the Coya-Machali (=Abanico) Formation. Radioisotopic data from levels immediately above (31.5 Ma) and below (37.S Ma) the fossiliferous horizon indicate a latest Eocene to early Oligocene age for the basal part of the formation and the fauna contained in it. 2) The fossiliferous unit rests with slight angular offset on different Mesozoic units: “Brownish-red Clastic Unit” (BRCU) and Banos del Flaco Formation; in a limited area it also overlies a white tuff dated at 104 Ma. 3) The contacts just discussed (none of which is attributable to faulting), demonstrate the existence of two, or possibly three, unconformities in the region. 4) Sedimentological criteria argue against reference of the BRCU to the Colimapu Formation, and imply correlation of the former unit to basal levels with in the late Cretaceous Neuquen Group of western Argentina. 5) The Coya-Machali Formation, previously viewed as representing the western volcanic equivalent of Riogrdndico Supercycle deposits of western Argentino, is likely coeval to much younger units in that region such as the Agua de la Piedra Formation. 6) Paleomagnetic results from the fossil producing horizon indicate about 20 ° of post-early Oligocene, counterclockwise rotation. 7) Fossil mammals from the Coya-Machali Formation near Termas del Flaco represent a distinct biochronologic interval not heretofore clearly recognized from elsewhere on the continent. This new fauna helps fill the long recognized post-?middle Eocene, pre-late Oligocene faunal hiatus between the Mustersan and Deseadan South American Land Mammal Ages (SALMA). In addition, it records the earliest known presence of rodents in South America and otherwise differs strongly from the enigmatic Divisaderan SALMA.

MORPHOLOGY AND RELATIONSHIPS OF APTERNODUS AND OTHER EXTINCT, ZALAMBDODONT, PLACENTAL MAMMALS

Abstract We describe and illustrate new, middle Cenozoic fossils of dentally zalambdodont, North American placentals, including six relatively complete crania of Apternodus and two of Oligoryctes, as well as many partial skulls, mandibles, and teeth of these and other taxa. Several of the new Apternodus specimens are also associated with postcrania. We recognize seven species of Apternodus, three of which are new, formally propose the combination Oligoryctes altitalonidus, and recognize two other genera of small, North American, anatomically zalambdodont placentals, Parapternodus and Koniaryctes. We regard two other taxa previously associated with North American fossil zalambdodonts, one Bridgerian and the other Tiffanian, as valid but do not name them in this paper. In addition, we argue that dental zalambdodonty entails a primary occlusal relationship between the paracone and the ectoflexid, and the reduction or absence of the metacone and talonid basin. A phylogenetic analysis of cranial, dental, and postcranial characters of 30 fossil and Recent taxa leads us to conclude that (1) the Apternodontidae as defined in previous literature is not monophyletic and should be restricted to seven species of Apternodus, (2) the genus Oligoryctes contains at least two species and has a considerably longer geologic record than Apternodus, (3) neither Micropternodus nor currently known Paleocene taxa are closely related to Apternodus or Oligoryctes, and (4) a case can be made for a close relationship among modern soricids, Parapternodus, Koniaryctes, Oligoryctes, and Apternodus to the exclusion of other insectivoran-grade taxa. With the use of ordered, multistate character transformations, Solenodon comprises the sister taxon to a soricid-fossil zalambdodont clade.

Paleogene Pseudoglyptodont Xenarthrans from Central Chile and Argentine Patagonia

Abstract Herein we describe a new, large-bodied species of Pseudoglyptodon, a close sloth ally, from volcaniclastic deposits of the Abanico ( =  Coya-Machalí) Formation of the central Chilean Andean Main Range. This species, P. chilensis, is a rare element of the Tinguiririca Fauna, on which the recently formalized Tinguirirican South American Land Mammal “Age” is founded, being known from just two specimens. The holotype of P. chilensis, a partial skull and largely complete mandibles (preserving seemingly complete upper and lower dentitions), is by far the best-preserved specimen referable to Pseudoglyptodon known. As such, this material permits a more refined phylogenetic placement of this enigmatic xenarthran than has been possible previously, with Pseudoglyptodon representing the proximal outgroup to the clade including the most recent common ancestor of Choelepus and Bradypus, plus all its descendants (i.e., crown clade sloths). A fragmentary specimen from Argentina is removed from Glyptatelus and referred to Pseudoglyptodon. Although this specimen is distinct from P. chilensis and other previously recognized species of Pseudoglyptodon, it offers too meager a basis for formally establishing a new name. Finally, phylogenetic definitions of the names Phyllophaga and Tardigrada are proposed. Historically these terms have been used largely interchangeably, but here we advocate linking the latter to the crown clade.

Amphicticeps and Amphicynodon (Arctoidea, Carnivora) from Hsanda Gol Formation, Central Mongolia and Phylogeny of Basal Arctoids with Comments on Zoogeography

Abstract Amphicticeps shackelfordi and Amphicynodon teilhardi are two small carnivorans from the early Oligocene Hsanda Gol Formation of central Mongolia, and as basal arctoids (infraorder Arctoidea) in Asia, feature unique combinations of morphologies that offer insights into early diversification and zoogeography of the arctoids. Lack of adequate study of Amphicticeps and incomplete knowledge about Amphicynodon, however, prevented them from being figured in the discussions of arctoid relationships. New associated dental and cranial materials collected during recent expeditions in the 1990s substantially enrich our knowledge of the two genera and their stratigraphic positions, and serve as an impetus for a study of their phylogenetic relationships in the broad perspective of basal Arctoidea. Hsanda Gol arctoids are represented by six small- to medium-sized species: Amphicticeps shackelfordi Matthew and Granger 1924, A. dorog, n.sp., A. makhchinus, n.sp., Amphicynodon teilhardi Matthew and Granger 1924,? Cephalogale sp., and Pyctis inamatus Babbitt, 1999. The three species of Amphicticeps apparently form an endemic clade confined to central Asia, whose zoogeographic origin is currently unknown. Amphicynodon has a much higher diversity in Europe than in Asia, and phylogenetically the Asian A. teilhardi seems to be nested within the European congeneric species, indicating an eastward dispersal for this group, linking the European “Grande Coupure” and the Asian “Mongolian Reconstruction” events. To avoid excessive homoplasies in crown groups, we attempted a phylogenetic reconstruction based mostly on stem arctoids. Twenty genera of primitive arctoids occupying basal positions of nearly all major clades are selected for the analysis. The resulting tree, based on 39 characters, approximates the initial divergence of the arctoids. The traditionally dichotomous Arctoidea, formed by sister clades Ursida and Mustelida, is recovered in our analysis. Mustelida is also largely dichotomous with mustelid-like forms on one side and procyonid-like forms on the other. Despite its rather hypercarnivorous dentition, Amphicticeps is found on the Ursida side of the arctoids, although support for such a topology is relatively weak. Amphicynodon is a stem taxon of the Ursida and is a sister to an ursid–pinniped clade.

Gobiconodonts from the Early Cretaceous of Oshih (Ashile), Mongolia

Abstract We describe here the first discovered mammalian remains from the Mongolian Early Cretaceous locality Oshih (Ashile). Four fragmentary, tooth-bearing specimens, probably corresponding to three individuals, have been recovered. All the fossils can be assigned to the family Gobiconodontidae (Chow and Rich, 1984). The specimens include three lower jaw fragments and one upper jaw fragment, and represent at least two different taxa. Gobiconodon hopsoni n. sp., is described and diagnosed here. This new species is larger than G. ostromi (Early Cretaceous Cloverly Formation, USA); thus, it is the largest triconodont and one of the largest Mesozoic mammals known. Gobiconodon sp., found also at Oshih, is slightly larger than G. borissiaki from the Early Cretaceous of Khoobur, Mongolia, but smaller than G. ostromi The specimens of this second species are poorly preserved and provide insufficient data for a diagnosis. The status of the different species of Gobiconodon and the new gobiconodontid Hangjinia is reviewed. In gobiconodontids and Triconodontidae, the maxillae appear to make a significant contribution to the orbital rim, a condition unusual among basal mammals, in which the lacrimal and jugal are the main components. Other triconodonts such as Jeholodens likely an “amphilestid”, appear to show the primitive mammalian condition for this feature. We present a brief consideration of triconodont relationships and discuss alternative placements of Gobiconodon among Mammaliaformes.

Land-Mammal Ages, Faunal Heterochrony, and Temporal Resolution in Cenozoic Terrestrial Sequences

Traditionally, land-mammal biochronology has subdivided the Cenozoic into discernible time intervals with an average duration of about