Joseph E. Peterson

Influence of Microbial Biofilms on the Preservation of Primary Soft Tissue in Fossil and Extant Archosaurs

Background Mineralized and permineralized bone is the most common form of fossilization in the vertebrate record. Preservation of gross soft tissues is extremely rare, but recent studies have suggested that primary soft tissues and biomolecules are more commonly preserved within preserved bones than had been presumed. Some of these claims have been challenged, with presentation of evidence suggesting that some of the structures are microbial artifacts, not primary soft tissues. The identification of biomolecules in fossil vertebrate extracts from a specimen of Brachylophosaurus canadensis has shown the interpretation of preserved organic remains as microbial biofilm to be highly unlikely. These discussions also propose a variety of potential mechanisms that would permit the preservation of soft-tissues in vertebrate fossils over geologic time. Methodology/Principal Findings This study experimentally examines the role of microbial biofilms in soft-tissue preservation in vertebrate fossils by quantitatively establishing the growth and morphology of biofilms on extant archosaur bone. These results are microscopically and morphologically compared with soft-tissue extracts from vertebrate fossils from the Hell Creek Formation of southeastern Montana (Latest Maastrichtian) in order to investigate the potential role of microbial biofilms on the preservation of fossil bone and bound organic matter in a variety of taphonomic settings. Based on these analyses, we highlight a mechanism whereby this bound organic matter may be preserved. Conclusions/Significance Results of the study indicate that the crystallization of microbial biofilms on decomposing organic matter within vertebrate bone in early taphonomic stages may contribute to the preservation of primary soft tissues deeper in the bone structure.

FACE BITING ON A JUVENILE TYRANNOSAURID AND BEHAVIORAL IMPLICATIONS

Abstract The juvenile tyrannosaurid specimen BMR P2002.4.1 possesses a series of four partially healed, oblong lesions along the left maxilla and nasal bones. The morphology of the lesions and their positioning and orientation are compatible with the jaws of the specimen, suggesting that the lesions may have been the result of a bite from an attacker of similar size and species as the bite victim. Bone remodeling of the lesions indicates partial healing and demonstrates that the injury was not fatal. The injury appears to have affected the growth of the skull, causing a slight warping of the left maxilla and a slight leftward curvature of the nasal. Previous examples of intraspecific behavior in theropods have noted various injuries on skulls; the majority of documented injuries are in adults. The presence and nature of these wounds suggests that juvenile tyrannosaurid behavior included intraspecific aggression among similar-sized individuals, as has been observed in modern crocodilians. Although it is not possible to unequivocally infer behavior from a fossil specimen, as a juvenile animal it is unlikely that the behavior directly reflects sexual competition or conflict, although it may represent learning behavior for later conflict as competing adults or territoriality over food or spatial resources.

An azhdarchid pterosaur cervical vertebra from the Hell Creek Formation (Maastrichtian) of southeastern Montana

An azhdarchid pterosaur cervical vertebra from the Hell Creek Formation (Maastrichtian) of southeastern Montana Michael D. Henderson & Joseph E. Peterson To cite this article: Michael D. Henderson & Joseph E. Peterson (2006) An azhdarchid pterosaur cervical vertebra from the Hell Creek Formation (Maastrichtian) of southeastern Montana, Journal of Vertebrate Paleontology, 26:1, 192-195 To link to this article: https://doi.org/10.1671/0272-4634(2006)26[192:AAPCVF]2.0.CO;2

Cranial pathologies in a specimen of Pachycephalosaurus.

Background A frontoparietal dome of a large pachycephalosaurid collected from the Upper Cretaceous Hell Creek Formation in 2001 is identified as Pachycephalosaurus wyomingensis. The specimen features two large oval depressions on the dorsal surface, accompanied by numerous circular pits on the margin and inner surface of the larger depressions. Methodology/Principal Findings In order to identify the origin of these structures, computed tomography (CT) data and morphological characteristics of the specimen are analyzed and compared with similar osteological structures in fossil and extant archosaurs caused by taphonomic processes, non-pathologic bone resorption, and traumatic infection/inflammatory origins. The results of these analyses suggest that the structures are pathologic lesions likely resulting from a traumatic injury and followed by secondary infection at the site. Conclusions/Significance The presence of lesions on a frontoparietal dome, and the exclusivity of their distribution along the dorsal dome surface, offers further insight into frontoparietal dome function and supports previously hypothesized agonistic behavior in pachycephalosaurids.

Distributions of Cranial Pathologies Provide Evidence for Head-Butting in Dome-Headed Dinosaurs (Pachycephalosauridae)

Pachycephalosaurids are small, herbivorous dinosaurs with domed skulls formed by massive thickening of the cranial roof. The function of the dome has been a focus of debate: the dome has variously been interpreted as the product of sexual selection, as an adaptation for species recognition, or as a weapon employed in intraspecific combat, where it was used in butting matches as in extant ungulates. This last hypothesis is supported by the recent identification of cranial pathologies in pachycephalosaurids, which appear to represent infections resulting from trauma. However, the frequency and distribution of pathologies have not been studied in a systematic fashion. Here, we show that pachycephalosaurids are characterized by a remarkably high incidence of cranial injury, where 22% of specimens have lesions on the dome. Frequency of injury shows no significant difference between different genera, but flat-headed morphs (here interpreted as juveniles or females) lack lesions. Mapping of injuries onto a digitial pachycephalosaurid skull shows that although lesions are distributed across the dome, they cluster near the apex, which is consistent with the hypothesis that the dome functioned for intraspecific butting matches.

METHODS OF MICROVERTEBRATE SAMPLING AND THEIR INFLUENCES ON TAPHONOMIC INTERPRETATIONS

Abstract Localized accumulations of small vertebrate fossils are relatively common in the Hell Creek Formation (latest Maastrichtian–Paleocene) and have traditionally been collected using nonstandardized field methods. The lack of established protocols has resulted in biased datasets and assemblage interpretations. Two commonly used collection methods were compared from two different microvertebrate localities in the Upper Cretaceous part of the Hell Creek Formation of southeastern Montana in order to identify their influence on taphonomic interpretations. The first of these methods was surface collection, in which weathered specimens were picked from the surface of the rock and in talus piles. This method commonly involves the selection and collection of larger and more easily identifiable specimens. The second method used in this study was quarrying and disaggregation, which requires excavation, collection, and total screen rinsing of a large volume of rock; this method was utilized in order to recover all fossils present. Both methods were applied at a sandstone-dominated and a mudstone-dominated site to assess an absolute concentration of fossils for each technique. Results of a quantitative analysis of these two common collection methods demonstrate how each one influences fossil recovery according to shape and size by skewing the characteristics of collected fossil assemblages and the taphonomic interpretations of microvertebrate accumulations. Although the study sites reflect only two possible types of microvertebrate localities, the results are applicable to many microvertebrate studies.

HYDRODYNAMIC BEHAVIORS OF PACHYCEPHALOSAURID DOMES IN CONTROLLED FLUVIAL SETTINGS: A CASE STUDY IN EXPERIMENTAL DINOSAUR TAPHONOMY

ABSTRACT The hydrodynamic behaviors of isolated dinosaur bones have been largely overlooked in the paleontological literature. Investigations into the hydrodynamic properties of dinosaur remains with unique taphonomic signatures, such as pachycephalosaurid frontoparietal domes, have the potential to aid in the interpretation of preservation for skeletal elements for which modern analogues are not available. For this study, a series of transport experiments were conducted to assess the entrainment velocities and settling orientations of a collection of pachycephalosaurid specimens. Casts of four pachycephalosaurid frontoparietal domes and skulls were composed of a urethane resin with a comparable average density to compact and cancellous bone, and placed in a flume with manual velocity control. Data were recorded for competent velocity, transport distance, and settling orientations upon resting and burial of specimens for 35 trials per cast. Though specimens vary considerably in mass, the results suggest specimen shape has a greater influence on transport and hydrodynamic behavior than size; significantly lower velocities are required to transport complete skulls than isolated domes. Resting and burial orientations of specimens vary significantly for domes and complete skulls. The highly variable transport velocities and settling orientations of pachycephalosaurid crania offer insight into pachycephalosaurid taphonomy and illustrate the importance of future taphonomic studies on large fossil vertebrate remains.

Fluvial transport potential of shed and root-bearing dinosaur teeth from the late Jurassic Morrison Formation

Shed dinosaur teeth are commonly collected microvertebrate remains that have been used for interpretations of dinosaur feeding behaviors, paleoecology, and population studies. However, such interpretations may be biased by taphonomic processes such as fluvial sorting influenced by tooth shape: shed teeth, removed from the skull during life, and teeth possessing roots, removed from the skull after death. As such, teeth may behave differently in fluvial systems due to their differences in shape. In order to determine the influence of fluvial processes on the preservation and distribution of shed and root-bearing dinosaur teeth, the hydrodynamic behaviors of high-density urethane resin casts of shed and root-bearing Allosaurus and Camarasaurus teeth were experimentally tested for relative transport distances at increasing flow velocities in an artificial fluviatile environment. Results show that tooth cast specimens exhibited comparable patterns of transport at lower velocities, though the shed Camarasaurus teeth transported considerably farther in medium to higher flow velocities. Two-Way ANOVA tests indicate significant differences in the mean transport distances of tooth casts oriented perpendicular to flow (p < 0.05) with varying tooth morphologies and flow velocities. The differences exhibited in the transportability of shed and root-bearing teeth has important implications for taphonomic reconstructions, as well as future studies on dinosaur population dynamics, paleoecology, and feeding behaviors.

New data towards the development of a comprehensive taphonomic framework for the Late Jurassic Cleveland-Lloyd Dinosaur Quarry, Central Utah

The Cleveland-Lloyd Dinosaur Quarry (CLDQ) is the densest deposit of Jurassic theropod dinosaurs discovered to date. Unlike typical Jurassic bone deposits, it is dominated by the presence of Allosaurus fragilis. Since excavation began in the 1920s, numerous hypotheses have been put forward to explain the taphonomy of CLDQ, including a predator trap, a drought assemblage, and a poison spring. In an effort to reconcile the various interpretations of the quarry and reach a consensus on the depositional history of CLDQ, new data is required to develop a robust taphonomic framework congruent with all available data. Here we present two new data sets that aid in the development of such a robust taphonomic framework for CLDQ. First, x-ray fluorescence of CLDQ sediments indicate elevated barite and sulfide minerals relative to other sediments from the Morrison Formation in the region, suggesting an ephemeral environment dominated by periods of hypereutrophic conditions during bone accumulation. Second, the degree of abrasion and hydraulic equivalency of small bone fragments dispersed throughout the matrix were analyzed from CLDQ. Results of these analyses suggest that bone fragments are autochthonous or parautochthonous and are derived from bones deposited in the assemblage rather than transported. The variability in abrasion exhibited by the fragments is most parsimoniously explained by local periodic re-working and re-deposition during seasonal fluctuations throughout the duration of the quarry assemblage. Collectively, these data support previous interpretations that the CLDQ represents an attritional assemblage in a poorly-drained overbank deposit where vertebrate remains were introduced post-mortem to an ephemeral pond during flood conditions. Furthermore, while the elevated heavy metals detected at the Cleveland-Lloyd Dinosaur Quarry are not likely the primary driver for the accumulation of carcasses, they are likely the result of multiple sources; some metals may be derived from post-depositional and diagenetic processes, and others are potentially produced from an abundance of decomposing vertebrate carcasses. These new data help to support the inferred depositional environment of the quarry as an ephemeral pond, and represent a significant step in understanding the taphonomy of the bonebed and Late Jurassic paleoecology in this region.

Role of Sediment Size and Biostratinomy on the Development of Biofilms in Recent Avian Vertebrate Remains

Microscopic soft tissues have been identified in fossil vertebrate remains collected from various lithologies. However, the diagenetic mechanisms to preserve such tissues have remained elusive. While previous studies have described infiltration of biofilms in Haversian and Volkmann’s canals, biostratinomic alteration (e.g., trampling), and iron derived from hemoglobin as playing roles in the preservation processes, the influence of sediment texture has not previously been investigated. This study uses a Kolmogorov Smirnov Goodness-of-Fit test to explore the influence of biostratinomic variability and burial media against the infiltration of biofilms in bone samples. Controlled columns of sediment with bone samples were used to simulate burial and subsequent groundwater flow. Sediments used in this study include clay-, silt-, and sand-sized particles modeled after various fluvial facies commonly associated with fossil vertebrates. Extant limb bone samples obtained from Gallus gallus domesticus (Domestic Chicken) buried in clay-rich sediment exhibit heavy biofilm infiltration, while bones buried in sands and silts exhibit moderate levels. Crushed bones exhibit significantly lower biofilm infiltration than whole bone samples. Strong interactions between biostratinomic alteration and sediment size are also identified with respect to biofilm development. Sediments modeling crevasse splay deposits exhibit considerable variability; whole-bone crevasse splay samples exhibit higher frequencies of high-level biofilm infiltration, and crushed-bone samples in modeled crevasse splay deposits display relatively high frequencies of low-level biofilm infiltration. These results suggest that sediment size, depositional setting, and biostratinomic condition play key roles in biofilm infiltration in vertebrate remains, and may influence soft tissue preservation in fossil vertebrates.