Raymond R. Rogers

The Ischigualasto Tetrapod Assemblage (Late Triassic, Argentina) and 40Ar/39Ar Dating of Dinosaur Origins

40Ar/39Ar dating of sanidine from a bentonite interbedded in the Ischigualasto Formation of northwestern Argentina yielded a plateau age of 227.8 � 0.3 million years ago. This middle Carnian age is a direct calibration of the Ischigualasto tetrapod assemblage, which includes some of the best known early dinosaurs. This age shifts last appearances of Ischigualasto taxa back into the middle Carnian, diminishing the magnitude of the proposed late Carnian tetrapod extinction event. By 228 million years ago, the major dinosaurian lineages were established, and theropods were already important constituents of the carnivorous tetrapod guild in the Ischigualasto—Villa Uni�n Basin. Dinosaurs as a whole remained minor components of tetrapod faunas for at least another 10 million years.

Taphonomy of three dinosaur bone beds in the Upper Cretaceous Two Medicine Formation of northwestern Montana; evidence for drought-related mortality

Two bone beds, Canyon Bone Bed and Dino Ridge Quarry, have yielded the near-exclusive remains of a new species of Styracosaurus (Family Ceratopsidae); the third bone bed, Westside Quarry, is dominated by a new species of Prosaurolophus (Family Hadrosauridae). Evidence supporting a drought hypothesis includes: 1) a seasonal, semiarid paleoclimate, 2) associated caliche horizons, 3) aqueous depositional settings, 4) apparent age distributions characteristic of modern drought mortality (CBB and DRQ), and 5) the intraformational recurrence of low-diversity bone beds. Several alternative scenarios were considered, but drought proved most reasonable in light of the enhanced probability of preserving drought assemblages, and the species-selective and recurrent nature of modern drought mortality

LATE CRETACEOUS TERRESTRIAL VERTEBRATES FROM MADAGASCAR: IMPLICATIONS FOR LATIN AMERICAN BIOGEOGRAPHY1

Abstract The Mahajanga Basin Project, initiated in 1993 and centered in Upper Cretaceous strata of northwestern Madagascar, has resulted in the discovery of some of the most complete, well-preserved, and significant specimens of Late Cretaceous vertebrate animals from the Southern Hemisphere and indeed the world. Among the most important finds are various specimens of crocodyliforms, non-avian dinosaurs, and mammals; these finds have the potential to provide key insights into the biogeographic and paleogeographic history of Gondwana. Madagascar has been physically isolated from Africa for over 160 million years and from all other major landmasses for more than 85 million years. The closest known relatives of many of the Late Cretaceous Malagasy taxa are penecontemporaneous forms from South America (primarily Argentina) and India, thus documenting a previously unrecognized high level of cosmopolitanism among Gondwanan vertebrates near the end of the Cretaceous. The family-level taxa that are shared among Madagascar, South America, and the Indian subcontinent are not known from penecontemporaneous horizons in mainland Africa, but it cannot yet be confidently determined if this is due to differential extinction, poor sampling, true absence (i.e., the taxa were never present on Africa), or some combination thereof. Nonetheless, currently available geologic and paleontologic data are most consistent with the Africa-first model, suggesting that Africa was the first of the major Gondwanan landmasses to be fully isolated prior to the Albian/Cenomanian boundary, and that its terrestrial vertebrate faunas became progressively more provincial during the Cretaceous, while those on other Gondwanan landmasses remained relatively cosmopolitan until the later stages of the Late Cretaceous.

Stratigraphic Analysis of Upper Cretaceous Rocks in the Mahajanga Basin, Northwestern Madagascar: Implications for Ancient and Modern Faunas.

Upper Cretaceous strata of the Mahajanga Basin, northwestern Madagascar, yield some of the most significant and exquisitely preserved vertebrate fossils known from Gondwana. The sedimentology of these strata and their stratigraphic relations have been the focus of renewed geological investigations during the course of five expeditions since 1993. We here designate stratotypes and formalize the terrestrial Maevarano Formation, with three new members (Masorobe, Anembalemba, Miadana), and the overlying marine Berivotra Formation. The Maevarano Formation accumulated on a broad, semiarid alluvial plain bounded to the southeast by crystalline highlands and to the northwest by the Mozambique Channel. The Berivotra Formation was deposited in an open marine setting that evolved from a clastic‐ to a carbonate‐dominated shelf, resulting in deposition of the overlying Betsiboka limestone of Danian age. New stratigraphic data clearly indicate that the Maevarano Formation correlates, at least in part, with the Maastrichtian Berivotra Formation, and this in turn indicates that the most fossiliferous portions of the Maevarano Formation are Maastrichtian in age, rather than Campanian as previously reported. This revised age for the Maevarano vertebrate assemblage indicates that it is approximately contemporaneous with the vertebrate fauna recovered from the Deccan basalt volcano‐sedimentary sequence of India. The comparable age of these two faunas is significant because the faunas appear to be more similar to one another than either is to those from any other major Gondwanan landmass. The revised age of the Maevarano Formation, when considered in the light of our recent fossil discoveries, further indicates that the ancestral stocks of Madagascars overwhelmingly endemic modern vertebrate fauna arrived on the island in post‐Mesozoic times. The basal stocks of the modern vertebrate fauna are conspicuously absent in the Maevarano Formation. Finally, the revised age of the Maevarano Formation serves to expand our global perspective on the K/T event by clarifying the age of a diverse, and arguably the best preserved, sample of Gondwanan vertebrates from the terminal Cretaceous.

Paleoenvironment and Taphonomy of the Chañares Formation Tetrapod Assemblage (Middle Triassic), Northwestern Argentina: Spectacular Preservation in Volcanogenic Concretions

Abstract The enigmatic concretionary exposures that typify the Chañares Formation (Ladinian, northwestern Argentina) long have defied precise paleoenvironmental characterization. Recent work indicates that the formation accumulated in an alluvial-to-lacustrine setting within an active rift basin that received sedimentary detritus from surrounding highlands, as well as copious amounts of volcanic ash. Ash- flow sheets were emplaced presumably as secondary mass flows on alluvial surfaces characterized by small fluvial channels and shallow lakes. Thin bentonite beds intercalated in the Chañares Formation indicate that ash also accumulated via direct airfall, although this mode of emplacement accounts for a very small fraction of the overall section. A shift to widespread lacustrine deposition is recorded by the superjacent Los Rastros Formation, which preserves at least six shallowing-upward hemicycles, five of which commenced amidst explosive volcanic activity as evidenced by intercalated bentonite beds. Volcanism played an important role in the generation and preservation of the Chañares Formations exceptional tetrapod fossil record. This is especially true of the classic Los Chañares locality, where more than 100 individuals representing a diverse array of taxa (archosaurs, cynodonts, dicynodonts) are entombed in volcanogenic concretions with matrices of relic glass shards diagenetically replaced by calcite. Taphonomic attributes of the Los Chañares locality are consistent with the scenario of mass mortality, and several clues hint at the nature of the event. The killing agent was lethal to a variety of taxa, killed both adults and juveniles, and led to the concentration of taxa that under normal circumstances would tend to dissociate, such as carnivores and their potential prey. It also produced a counterintuitive bias against the preservation of large-bodied taxa, which may have been largely unsusceptible to the death event, or perhaps were excluded from the Los Chañares death assemblage via post-mortem sorting. The spatial arrangement of skeletal material in a small sample of concretions is consistent with the stranding of tetrapod carcasses along a strandline, and it is feasible that volcanism led to catastrophic flooding of the landscape via damming and/or diversion of local drainages. Uncompacted skeletal elements and relic outlines of glass shards indicate that carbonate concretions formed shortly after skeletal material was buried in reworked volcanic ash. The microbial decay of organic matter presumably catalyzed concretion diagenesis. There is no indication that bone hydroxyapatite diffused into the entombing glassy matrix and contributed to concretion formation. Bones entombed within early diagenetic concretions were safeguarded from subsequent destructive pedogenic and/or diagenetic processes, and were incorporated in exquisite quality into the fossil record.

CONTINENTAL INSECT BORINGS IN DINOSAUR BONE: EXAMPLES FROM THE LATE CRETACEOUS OF MADAGASCAR AND UTAH

Abstract Two new insect-related ichnogenera are reported in fossil dinosaur bones from Upper Cretaceous continental strata in Madagascar and Utah. Cubiculum ornatus n. igen. and isp. is described from numerous fossil bones in the Upper Cretaceous Maevarano Formation of northwestern Madagascar, and consists of hollow, ovoid chambers with concave flanks excavated into both spongy and compact bone. Traces similar in morphology to Cubiculum ornatus have been reported elsewhere in North America, Asia, Europe, and Africa in bones ranging in age from Jurassic to Pleistocene, and have been interpreted as pupal chambers constructed by carrion beetle larvae. Osteocallis mandibulus n. igen. and isp. is described in dinosaur bones from continental deposits of the Upper Cretaceous Maevarano Formation of Madagascar and the Upper Cretaceous Kaiparowits Formation of southern Utah. O. mandibulus consists of shallow, meandering surface trails, composed of numerous arcuate grooves, bored into compact (cortical) bone surfaces, and is tentatively interpreted as a feeding trace. Based on similar patterns of bioglyph preserved in both Cubiculum ornatus and Osteocallis mandibulus, the tracemaker is interpreted to be the same or similar for both borings. Given the recurrent association with animal remains, the tracemaker is furthermore presumed to be a necrophagous or osteophagous insect that used bone as a substrate for both reproduction (C. ornatus) and feeding (O. mandibulus).

Visualizing fossilization using laser ablation-inductively coupled plasma-mass spectrometry maps of trace elements in Late Cretaceous bones

Elemental maps generated by laser ablation–inductively coupled plasma–mass spectrometry (LA-ICP-MS) provide a previously unavailable high-resolution visualization of the complex physicochemical conditions operating within individual bones during the early stages of diagenesis and fossilization. A selection of LA-ICP-MS maps of bones collected from the Late Cretaceous of Montana (United States) and Madagascar graphically illustrate diverse paths to recrystallization, and reveal unique insights into geochemical aspects of taphonomic history. Some bones show distinct gradients in concentrations of rare earth elements and uranium, with highest concentrations at external bone margins. Others exhibit more intricate patterns of trace element uptake related to bone histology and its control on the flow paths of pore waters. Patterns of element uptake as revealed by LA-ICP-MS maps can be used to guide sampling strategies, and call into question previous studies that hinge upon localized bulk samples of fossilized bone tissue. LA-ICP-MS maps also allow for comparison of recrystallization rates among fossil bones, and afford a novel approach to identifying bones or regions of bones potentially suitable for extracting intact biogeochemical signals.

Multiple taxon–multiple locality approach to providing oxygen isotope evidence for warm-blooded theropod dinosaurs

Oxygen isotope ratios of fossil remains of coexisting taxa from several different localities can be used to help investigate dinosaur thermoregulation. Focusing on the Late Cretaceous, oxygen isotope ratios of crocodile tooth enamel from four separate localities exhibit less of a decrease with latitude than do ratios of tooth enamel from coexisting theropod dinosaurs. A shallower latitudinal gradient for crocodiles is consistent with how oxygen isotope ratios should vary for heterothermic animals having body temperatures coupled with their environments (“cold blooded”), while a steeper gradient for theropods is consistent with how these ratios should vary for homeothermic animals having constant body temperatures independent of their environments (“warm blooded”). This inferred homoethermy in theropods is likely due to higher rates of metabolic heat production relative to crocodiles and is not an artifact of body size.

Cannibalism in the Madagascan dinosaur Majungatholus atopus

Many lines of evidence have been brought to bear on the question of theropod feeding ecology, including functional and physiological considerations, morphological constraints, taphonomic associations, and telling—although rare—indications of direct ingestion. Tooth marks of theropods, although rarely described and generally left unassigned to a particular taxon, can provide unique clues into predator–prey interaction, and can also yield insights into the extent of carcass utilization. Here we describe a sample of tooth-marked dinosaur bone recovered from three well-documented localities in the Upper Cretaceous Maevarano Formation of Madagascar that provides insights into the feeding ecology of the abelisaurid theropod Majungatholus atopus. Intensely tooth-marked elements from multiple individuals show that Majungatholus defleshed dinosaur carcasses. Furthermore, Majungatholus clearly fed upon the remains of not only sauropods, but also conspecifics, and thus was a cannibal. Cannibalism is a common ecological strategy among extant carnivores, but until now the evidence in relation to carnivorous dinosaurs has been sparse and anecdotal.

Associations of Vertebrate Skeletal Concentrations and Discontinuity Surfaces in Terrestrial and Shallow Marine Records: A Test in the Cretaceous of Montana.

Although lags of bones and teeth are commonly cited criteria for marine unconformities, the consistency of the association of vertebrate fossils and discontinuity surfaces, as well as the taphonomic (postmortem) controls on this relationship, are poorly understood. A field test across fluvial, paralic, and shallow marine facies in the Campanian Two Medicine and Judith River formations of Montana indicates that the distribution of vertebrate skeletal concentrations is poorly correlated with the inferred durations of erosional and omissional hiatuses. Instead, vertebrate concentrations associated with discontinuities of all durations tend to be patchy and closely track the abundance of fossil material in underlying and lateral facies. Based on the analysis of 83 measured sections, we found first that erosional bases of channels and minor scours within channels yield vertebrate lags; tidally influenced fluvial deposits are more productive than are “upland” fluvial deposits. Second, erosional shoreface ravinements and their correlative transgressive marine flooding surfaces (fourth‐order sequence boundaries) have well‐developed vertebrate lags only along segments that cut across older shoreface deposits. Third, a nonerosional, widely traceable discontinuity, which is interpreted as the nonmarine extension of a 75.4‐Ma third‐order transgressive surface, is completely lacking in vertebrate concentrations. Despite being unfossiliferous itself, this discontinuity does mark a regional change in the richness of the vertebrate fossil record, with overlying beds characterized by a much greater abundance of skeletal material. Fourth, a laterally extensive set of erosional surfaces, embedded within multistory fluvial sandstone sheets, is the nonmarine extension of an 80‐Ma third‐order sequence boundary in the marine basin and lacks vertebrate concentrations. The strong dependence of vertebrate lag development on preexisting local sources of skeletal material rather than on the magnitude of the erosional vacuity or the duration of the hiatus contrasts with skeletal concentrations of invertebrates in marine successions, where exhumation is generally much less important than the production of new elements during the hiatus. These findings provide a guide to prospecting productive fossil horizons in terrestrial records and underscore fundamental differences in the ways in which bioclastic material accumulates in terrestrial and shallow marine settings, the qualities of paleobiologic data derived from such concentrations, and the relative reliabilities of skeletal material as cues to stratigraphically significant discontinuities.