Richard L. Cifelli

Dual origin of tribosphenic mammals

Marsupials, placentals and their close therian relatives possess complex (tribosphenic) molars that are capable of versatile occlusal functions. This functional complex is widely thought to be a key to the early diversification and evolutionary success of extant therians and their close relatives (tribosphenidans). Long thought to have arisen on northern continents, tribosphenic mammals have recently been reported from southern landmasses. The great age and advanced morphology of these new mammals has led to the alternative suggestion of a Gondwanan origin for the group. Implicit in both biogeographic hypotheses is the assumption that tribosphenic molars evolved only once in mammalian evolutionary history. Phylogenetic and morphometric analyses including these newly discovered taxa suggest a different interpretation: that mammals with tribosphenic molars are not monophyletic. Tribosphenic molars evolved independently in two ancient (holotherian) mammalian groups with different geographic distributions during the Jurassic/Early Cretaceous: an australosphenidan clade endemic to Gondwanan landmasses, survived by extant monotremes; and a boreosphenidan clade of Laurasian continents, including extant marsupials, placentals and their relatives.

SAUROPOSEIDON PROTELES, A NEW SAUROPOD FROM THE EARLY CRETACEOUS OF OKLAHOMA

Abstract Sauroposeidon proteles, a new brachiosaurid sauropod, is represented by an articulated series of four mid-cervical vertebrae recovered from the Antlers Formation (Aptian–Albian) of southeastern Oklahoma. Most Early Cretaceous North American sauropod material has been referred to Pleurocoelus, a genus which is largely represented by juvenile material and is not well understood. Regardless of the status and affinities of Pleurocoelus, the new taxon is morphologically and proportionally distinct. Among well-known sauropod taxa, Sauroposeidon is most similar to Brachiosaurus; particularly noteworthy are the neural spines, which are set forward on the centra and are not bifurcate, and the extremely elongate cervical ribs. Sauroposeidon and Brachiosaurus also share a derived pattern of pneumatic vertebral ultrastructure and a mid-cervical transition point, at which neural spine morphology changes from very low (anteriorly) to very high (posteriorly). Autapomorphies of Sauroposeidon include posterior placement of the diapophyses, hypertrophied pneumatic fossae in the lateral faces of the neural spines and centra, and an extraordinary degree of vertebral elongation (e.g., C8 = 1.25 m; 25% longer than Brachiosaurus). Additional sauropod material from the Early Cretaceous Cloverly Formation may be referrable to the new Oklahoma sauropod, which appears to be the last of the giant North American sauropods and represents the culmination of brachiosaurid trends towards lengthening and lightening of the neck.

Dinosaur Body Temperatures Determined from Isotopic (13C-18O) Ordering in Fossil Biominerals

Large dinosaurs had body temperatures similar to those of modern mammals and birds. The nature of the physiology and thermal regulation of the nonavian dinosaurs is the subject of debate. Previously, arguments have been made for both endothermic and ectothermic metabolisms on the basis of differing methodologies. We used clumped isotope thermometry to determine body temperatures from the fossilized teeth of large Jurassic sauropods. Our data indicate body temperatures of 36° to 38°C, which are similar to those of most modern mammals. This temperature range is 4° to 7°C lower than predicted by a model that showed scaling of dinosaur body temperature with mass, which could indicate that sauropods had mechanisms to prevent excessively high body temperatures being reached because of their gigantic size.

The Campanian Terlingua local fauna, with a summary of other vertebrates from the Aguja Formation, Trans-Pecos Texas

ABSTRACT The Terlingua local fauna is a rich assemblage of predominantly terrestrial micro vertebrates from the Upper Cretaceous Aguja Formation of Trans-Pecos Texas. Marine invertebrates (which include elements of both Cretaceous Western Interior and Gulf Coast zoogeographic provinces) from conformably underlying strata suggest that the fauna is of late Campanian age, probably correlative with Judithian assemblages of the Western Interior. A Judithian “age” for the fauna is further supported by its mammal and theropod assemblages, and by the faunas of overlying deposits. The previously reported diversity of the Aguja Formation, which we summarize, is significantly enriched by this new fauna. The fauna also fills a major gap in the biogeography of Campanian terrestrial vertebrates. Notable occurrences in the Terlingua local fauna include the therian mammal Gallolestes, previously known only from Baja California, and a hitherto unrecorded type of primitive ‘tribothere.’ At least 4 marsupial and 6 multitube...

Tribosphenic mammal from the North American Early Cretaceous

The main groups of living mammals, marsupials and eutherians, are presumed to have diverged in the Early Cretaceous, but their early history and biogeography are poorly understood. Dental remains have suggested that the eutherians may have originated in Asia, spreading to North America in the Late Cretaceous, where an endemic radiation of marsupials was already well underway. Here I describe a new tribosphenic mammal (a mammal with lower molar heels that are three-cusped and basined) from the Early Cretaceous of North America, based on an unusually complete specimen. The new taxon bears characteristics (molarized last premolar, reduction to three molars) otherwise known only for Eutheria among the tribosphenic mammals. Morphometric analysis and character comparisons show, however, that its molar structure is primitive (and thus phylogenetically uninformative), emphasizing the need for caution in interpretation of isolated teeth. The new mammal is approximately contemporaneous with the oldest known Eutheria from Asia. If it is a eutherian, as is indicated by the available evidence, then this group was far more widely distributed in the Early Cretaceous than previously appreciated. An early presence of Eutheria in North America offers a potential source for the continents Late Cretaceous radiations, which have, in part, proven difficult to relate to contemporary taxa in Asia.

Dentition and Jaw of Kokopellia juddi, a Primitive Marsupial or Near-Marsupial from the Medial Cretaceous of Utah

We add to the knowledge of the dentition and lower jaw of the primitive marsupial or near marsupial, Kokopellia juddi, based on newly collected materials from the medial Cretaceous (Albian–Cenomanian) of central Utah. The dental formula, i4 c1 p3 m4, is primitive for (or with respect to) Marsupialia, as are a number of features of the dentary and dentition: presence of a labial mandibular foramen, ?an inflected angle, ?and a trace of the meckelian groove; lack of “staggering” of the lower incisor series; lack of “twinning” between entoconid and hypoconulid on lower molars; incompletely lingual position of lower molar paraconid; upper molar protocone relatively small and mesiodistally narrow; and conules placed about halfway between the protocone and the paracone–metacone. Other than the stylocone, cusps are lacking from the stylar shelf; we argue that this represents the primitive marsupial condition based on the economy of character change and the stratigraphic record of marsupials in the Cretaceous of North America. Recent discoveries of early marsupials, eutherians, and therians of metatherian–eutherian grade provide data indicating that some derived features of the dentary and dentition (e.g., loss of coronoid, meckelian groove, and labial mandibular foramen; acquisition of strong, “winged” conules, double rank postvallum/prevallid shearing, and stylar cusp D) probably arose independently, in some cases more than once, among the major groups of tribosphenic mammals. In turn, this suggests that a common ancestor for marsupials and placentals was more primitive than has generally been appreciated.

The origin of the dog-like borhyaenoid marsupials of South America

Dog-like marsupials (superfamily Borhyaenoidea) were the largest predacious mammals during the Tertiary period in South America. They are critical to our understanding of marsupial origin, phylogeny and palaeobiogeography because they have been related to various marsupial lineages of several continents: didelphoids (mainly New World, but also Europe, Asia and Africa), pediomyid, stagodontids (North America), dasyuroids (Australia) and deltatheroidans (predominantly Asian). These relationships, based mainly on dental morphology, have been discussed and rejected several times,,,. Here we report the discovery of exceptionally well preserved skulls and skeletons, referrable to the didelphoid Andinodelphys, which shed new light on the phylogenetic and palaeobiogeographic origin of dog-like marsupials. The skulls of Mayulestes (boryhyaenoid), Andinodelphys and Pucadelphys (didelphoids), from the early Palaeocene epoch of Bolivia are the oldest known for American marsupials. Comparison of their basicranial anatomy suggests that dog-like marsupials are closely related to an early didelphimorphian radiation in South America, rather than to Asiatic (deltatheroidan), North American (stagodontid), or Australian (dasyuroid) lineages.

Cretaceous mammals of southern Utah. I. Marsupials from the Kaiparowits Formation (Judithian)

ABSTRACT Marsupial mammals are herein described from the Kaiparowits Formation of southcentral Utah, of probable Judithian (Campanian, Late Cretaceous) age. New materials of Iqualadelphis lactea, previously described from the upper Milk River Formation of Alberta, support the hypothesis that the species is the most primitive known marsupial. A new species, intermediate in morphology between the Aquilan “Alphadon” creber and the Judithian to Lancian “A” lulli, is described, and the three are removed to a new genus, Protalphadon. Two new species closely similar to Judithian “Alphadon” russelli and “A” praesagus, respectively, are described; these, together with the Lancian “A” rhaister share derived morphology not found in other North American Cretaceous peradectids, and are placed in the new genus Turgidodon. New materials of the Judithian Alphadon halleyi and A. sahnii support the distinction of these closely similar species. Knowledge of the diminutive Alphadon attaragos, described on the basis of an iso...

EVOLUTION OF DENTAL REPLACEMENT IN MAMMALS

Abstract We provide a review of dental replacement features in stem clades of mammals and an hypothetical outline for the evolution of replacement frequency, mode, and sequence in early mammalian evolution. The origin of mammals is characterized by a shift from a primitive pattern of multiple, alternating replacements of all postcanines in most cynodonts to a derived pattern of single, sequential replacement of postcanines. The stem mammal Sinoconodon, however, retained some primitive replacement features of cynodonts. The clade of Morganucodon + crown mammals is characterized by the typical mammalian diphyodont replacement in which antemolars are replaced by one generation in anteroposterior sequence, but molars are not replaced. The stem clades of crown mammals including multituberculates and eutriconodonts have an anteroposterior sequential and diphyodont replacement of premolars. By contrast, stem taxa of the trechnotherian clade (Zhangheotherium, Dryolestes, and Slaughteria) are characterized by an alternating (p2 → p4 → p3) and diphyodont replacement, a condition that is shared by basal eutherians. The sequential replacements of premolars in most extant placentals (either anteroposteriorly p2 → p3 → p4 as in ungulates and carnivores, or postero-anteriorly p4 → p3 → p2 as in some insectivores) would represent secondarily derived conditions within eutherians. The single replacement of P3/p3 of metatherians is the most derived for all therian mammals.

Vertebrate paleontology, geology, and geochronology of the Tapera de Lopez and Scarritt Pocket, Chubut Province, Argentina

Fossil land mammals were collected by G. G. Simpson in 1933-1934 at and near the Tapera de L6pez in central Chubut Province, Patagonia, southern Argentina, from rocks now mapped as the Sarmiento Formation. These fossils are assigned to land mammal faunas of Ca- samayoran (Early Eocene), Mustersan (Middle Eocene), and Deseadan (late Early Oligocene through Early Miocene) age. 40K-40Ar age determinations of eight basalt and two tuff units associated with the Deseadan age local fauna at Scarritt Pocket establish a geochronologic framework that calibrates the biostrati- graphic record at this locality. The radioisotope dates obtained at Scarritt Pocket range from 23.4 Ma to about 21.0 Ma, and equate with earliest Miocene time. The Scarritt Pocket local fauna is the youngest dated Deseadan age fauna yet known in South America. Seven other localities have, or were reputed to have, local faunas of Deseadan age associated with dated volcanic units. Six of these localities are in Argentina (Gran Barranca, Cerro Blanco, Valle Hermoso, Pico Truncado, Caiiad6n Hondo, Quebrada Fiera de Malargiie) and one in Bolivia (Estratos Salla in the Salla-Luribay Basin). The stratigraphic relationships of the volcanic units with these local faunas is discussed, and the taxonomic content of each is reassessed. The Deseadan Land Mammal Age is defined by the earliest record of the land mammal genus Pyrotherium, which is from below a basalt dated at 33.6 Ma at Pico Truncado. Other early records of Pyrotherium occur below basalts dated at about 29 Ma at the Gran Barranca and Valle Hermoso, and from a 28.5 Ma level of the Estratos Salla. Thus, the lower boundary for Deseadan time is about 34 Ma. The youngest record of Pyrotherium is in the upper levels of the Estratos Salla dated at about 24 Ma. However, the Scarritt Pocket local fauna, which lacks Pyrotherium, permits placement of the upper boundary for Deseadan time at about 21.0 Ma. Late Deseadan time is surely, and the end of Deseadan time is apparently, marked by the last record of such groups as Proborhyaeninae (Proborhyaena), Rhynchippinae (Rhynchippus), Archaeohyracidae (Archaeohyrax), and the genera Platypittamys (Octodontidae), Scarrittia (Leontiniidae), Propachyrucos and Prohegetotherium (He- getotheriidae), and Argyrohyrax (Interatheriidae), as these taxa are recorded in the Scarritt Pocket local fauna. Thus, Deseadan time extends from about 34.0 Ma to about 21.0 Ma, making it the Land Mammal Age with the longest known duration in South America.