Ricardo N. Martínez

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.

Small-molecule p21-activated kinase inhibitor PF-3758309 is a potent inhibitor of oncogenic signaling and tumor growth

Despite abundant evidence that aberrant Rho-family GTPase activation contributes to most steps of cancer initiation and progression, there is a dearth of inhibitors of their effectors (e.g., p21-activated kinases). Through high-throughput screening and structure-based design, we identify PF-3758309, a potent (Kd = 2.7 nM), ATP-competitive, pyrrolopyrazole inhibitor of PAK4. In cells, PF-3758309 inhibits phosphorylation of the PAK4 substrate GEF-H1 (IC50 = 1.3 nM) and anchorage-independent growth of a panel of tumor cell lines (IC50 = 4.7 ± 3 nM). The molecular underpinnings of PF-3758309 biological effects were characterized using an integration of traditional and emerging technologies. Crystallographic characterization of the PF-3758309/PAK4 complex defined determinants of potency and kinase selectivity. Global high-content cellular analysis confirms that PF-3758309 modulates known PAK4-dependent signaling nodes and identifies unexpected links to additional pathways (e.g., p53). In tumor models, PF-3758309 inhibits PAK4-dependent pathways in proteomic studies and regulates functional activities related to cell proliferation and survival. PF-3758309 blocks the growth of multiple human tumor xenografts, with a plasma EC50 value of 0.4 nM in the most sensitive model. This study defines PAK4-related pathways, provides additional support for PAK4 as a therapeutic target with a unique combination of functions (apoptotic, cytoskeletal, cell-cycle), and identifies a potent, orally available small-molecule PAK inhibitor with significant promise for the treatment of human cancers.

A Basal Dinosaur from the Dawn of the Dinosaur Era in Southwestern Pangaea

Two hundred thirty million years ago, in what is now Argentina, dinosaurs could be found as the dominant carnivores or as small herbivores. Upper Triassic rocks in northwestern Argentina preserve the most complete record of dinosaurs before their rise to dominance in the Early Jurassic. Here, we describe a previously unidentified basal theropod, reassess its contemporary Eoraptor as a basal sauropodomorph, divide the faunal record of the Ischigualasto Formation with biozones, and bracket the formation with 40Ar/39Ar ages. Some 230 million years ago in the Late Triassic (mid Carnian), the earliest dinosaurs were the dominant terrestrial carnivores and small herbivores in southwestern Pangaea. The extinction of nondinosaurian herbivores is sequential and is not linked to an increase in dinosaurian diversity, which weakens the predominant scenario for dinosaurian ascendancy as opportunistic replacement.

Evidence for Avian Intrathoracic Air Sacs in a New Predatory Dinosaur from Argentina

Background Living birds possess a unique heterogeneous pulmonary system composed of a rigid, dorsally-anchored lung and several compliant air sacs that operate as bellows, driving inspired air through the lung. Evidence from the fossil record for the origin and evolution of this system is extremely limited, because lungs do not fossilize and because the bellow-like air sacs in living birds only rarely penetrate (pneumatize) skeletal bone and thus leave a record of their presence. Methodology/Principal Findings We describe a new predatory dinosaur from Upper Cretaceous rocks in Argentina, Aerosteon riocoloradensis gen. et sp. nov., that exhibits extreme pneumatization of skeletal bone, including pneumatic hollowing of the furcula and ilium. In living birds, these two bones are pneumatized by diverticulae of air sacs (clavicular, abdominal) that are involved in pulmonary ventilation. We also describe several pneumatized gastralia (“stomach ribs”), which suggest that diverticulae of the air sac system were present in surface tissues of the thorax. Conclusions/Significance We present a four-phase model for the evolution of avian air sacs and costosternal-driven lung ventilation based on the known fossil record of theropod dinosaurs and osteological correlates in extant birds: (1) Phase I—Elaboration of paraxial cervical air sacs in basal theropods no later than the earliest Late Triassic. (2) Phase II—Differentiation of avian ventilatory air sacs, including both cranial (clavicular air sac) and caudal (abdominal air sac) divisions, in basal tetanurans during the Jurassic. A heterogeneous respiratory tract with compliant air sacs, in turn, suggests the presence of rigid, dorsally attached lungs with flow-through ventilation. (3) Phase III—Evolution of a primitive costosternal pump in maniraptoriform theropods before the close of the Jurassic. (4) Phase IV—Evolution of an advanced costosternal pump in maniraptoran theropods before the close of the Jurassic. In addition, we conclude: (5) The advent of avian unidirectional lung ventilation is not possible to pinpoint, as osteological correlates have yet to be identified for uni- or bidirectional lung ventilation. (6) The origin and evolution of avian air sacs may have been driven by one or more of the following three factors: flow-through lung ventilation, locomotory balance, and/or thermal regulation.

A basal sauropodomorph (Dinosauria: Saurischia) from the Ischigualasto Formation (Triassic, Carnian) and the early evolution of Sauropodomorpha

Background The earliest dinosaurs are from the early Late Triassic (Carnian) of South America. By the Carnian the main clades Saurischia and Ornithischia were already established, and the presence of the most primitive known sauropodomorph Saturnalia suggests also that Saurischia had already diverged into Theropoda and Sauropodomorpha. Knowledge of Carnian sauropodomorphs has been restricted to this single species. Methodology/Principal Findings We describe a new small sauropodomorph dinosaur from the Ischigualsto Formation (Carnian) in northwest Argentina, Panphagia protos gen. et sp. nov., on the basis of a partial skeleton. The genus and species are characterized by an anteroposteriorly elongated fossa on the base of the anteroventral process of the nasal; wide lateral flange on the quadrate with a large foramen; deep groove on the lateral surface of the lower jaw surrounded by prominent dorsal and ventral ridges; bifurcated posteroventral process of the dentary; long retroarticular process transversally wider than the articular area for the quadrate; oval scars on the lateral surface of the posterior border of the centra of cervical vertebrae; distinct prominences on the neural arc of the anterior cervical vertebra; distal end of the scapular blade nearly three times wider than the neck; scapular blade with an expanded posterodistal corner; and medial lamina of brevis fossa twice as wide as the iliac spine. Conclusions/Significance We regard Panphagia as the most basal sauropodomorph, which shares the following apomorphies with Saturnalia and more derived sauropodomorphs: basally constricted crowns; lanceolate crowns; teeth of the anterior quarter of the dentary higher than the others; and short posterolateral flange of distal tibia. The presence of Panphagia at the base of the early Carnian Ischigualasto Formation suggests an earlier origin of Sauropodomorpha during the Middle Triassic.

Adeopapposaurus mognai, gen. et sp. nov. (Dinosauria: Sauropodomorpha), with Comments on Adaptations of Basal Sauropodomorpha

ABSTRACT Prosauropods are basal sauropodomorphs that were the major terrestrial faunal components from the Norian until their extinction during the Toarcian. Their status as a natural group is debatable. In the present work I describe Adeopapposaurus mognai, a new sauropodomorph from the Cañón del Colorado Formation, in northwestern Argentina. Diagnostic autapomorphies and combination of characters of Adeopapposaurus include a series of large foramina in a sub-vertical row on the lateral surface of the premaxilla; strongly rugose depression bordered by a protuberance with a series of foramina in a sub-vertical row, on the lateral surface of the anterior end of the dentary; eleven anteroposteriorly elongated cervical vertebrae and thirteen dorsal vertebrae with neural arches taller than the respective centra. Phylogenetically Adeopapposaurus is resolved as the sister group to Massospondylus; differing from the latter based on differences in mandible and premaxilla and addition of one dorsal vertebra to the neck. The specimens described here reveal numerous herbivorous adaptations, including the presence of a highly vascularized bony plate in the premaxilla and dentary, which indicates that it had a horny beak.

Osteology of Eoraptor lunensis (Dinosauria, Sauropodomorpha)

ABSTRACT We describe the basal sauropodomorph Eoraptor lunensis, based on the nearly complete holotypic skeleton and referred specimens, all of which were discovered in the Cancha de Bochas Member of the Ischigualasto Formation in northwestern Argentina. The lightly built skull has a slightly enlarged external naris and a spacious antorbital fossa with a prominent, everted dorsal margin and internal wall lacking any pneumatic extensions into surrounding bones. The tall quadrate is lapped along its anterior margin by the long, slender ventral process of the squamosal, and the lower jaw has a mid-mandibular joint between a tongue-shaped splenial process and a trough in the angular. All but the posterior-most maxillary and dentary crowns have a basal constriction, and the marginal denticles are larger and oriented more vertically than in typical theropod serrations. Rows of rudimentary palatal teeth are present on the pterygoid. Vertebral centra are hollow, although not demonstrably pneumatized, and all long bones have hollow shafts. The radius and ulna are more robust, the manus proportionately shorter, and the manual unguals less recurved than in the contemporaneous basal theropod Eodromaeus murphi. An outstanding feature of the manus of Eoraptor is the twisted shaft of the first phalanx of the pollex, which deflects medially the tip of the ungual as in basal sauropodomorphs. The long bones of the hind limb have more robust shafts than those of Eodromaeus, although in both genera the tibia remains slightly longer than the femur.

A new carnivorous cynodont from the Ischigualasto Formation (Late Triassic, Argentina), with comments on eucynodont phylogeny

ABSTRACT The nearly complete skull of a recently discovered carnivorous eucynodont, Ecteninion lunensis, gen. et sp. nov., is described. This is the first new genus of carnivorous cynodont reported from the Late Triassic (middle Carnian) Ischigualasto Formation of northwestern Argentina, and is unique to the formation. Diagnostic features include partially differentiated cheek teeth that overlap in an imbricate pattern, elongate braincase and parietal crest, pronounced lateral expansion of the braincase, sagittal crest that overhangs the occipital plate in dorsal view, gracile postorbital bars, short osseous secondary palate, maxilla excluded from the border of the subtemporal fenestra, palatine and orbitosphenoid that contribute to the medial orbital wall, post-temporal foramen enclosed within the tabular, squamosals that nearly meet at the posterior end of the sagittal crest, narrow and deep groove for the external auditory meatus, jaw articulation located at the anterior base of the zygomatic arches, a...

Vertebrate succession in the Ischigualasto Formation

ABSTRACT The Upper Triassic (Carnian—Norian) Ischigualasto Formation has yielded a diverse vertebrate fauna that records the initial phase of dinosaur evolution. Radioisotopic dates from ash layers within the formation provide a chrono-stratigraphic framework, and stratigraphic and sedimetological studies have subdivided the formation into four members and three abundance-based biozones. We describe two new basal dinosauromorphs, an unnamed lagerpetid and a new silesaurid, Ignotosaurus fragilis, gen. et sp. nov., which increase to 29 the number of vertebrates in the Ischigualasto fauna. We provide a census of 848 fossil specimens representing 26 vertebrate taxa logged to stratigraphic intervals of 50 m. This temporally calibrated census shows that abundance and taxonomic diversity within the Ischigualasto Formation does not change suddenly but rather appears to gradually decline in response to climatic deterioration. The only abrupt shift in faunal composition occurs at the end of the second of three biozones, when the abundant cynodont Exaeretodon is replaced by the rare dicynodont Jachaleria.

Age constraints on the dispersal of dinosaurs in the Late Triassic from magnetochronology of the Los Colorados Formation (Argentina)

Significance Uncertainties in reported 40Ar/39Ar dates from the Ischigualasto Formation of Argentina allow its dinosaur-bearing fauna to be Norian in age and possibly contemporaneous with some of the older U-Pb dated dinosaur-bearing units in the Chinle Formation of the American Southwest. Our magnetochronology of the previously undated Los Colorados Formation, which also contains a diverse dinosaur assemblage, constrains its age to the interval from 227 to 213 Ma (Norian) and thereby largely restricts the underlying Ischigualasto Formation to the Carnian. Rise of early dinosaurs was thus diachronous across the Americas with their dispersal from the austral temperate belt blocked until later in the Norian. The breakout may have resulted from critically lowered climatic barriers associated with decreasing atmospheric pCO2 levels. A measured magnetozone sequence defined by 24 sampling sites with normal polarity and 28 sites with reverse polarity characteristic magnetizations was established for the heretofore poorly age-constrained Los Colorados Formation and its dinosaur-bearing vertebrate fauna in the Ischigualasto–Villa Union continental rift basin of Argentina. The polarity pattern in this ∼600-m-thick red-bed section can be correlated to Chrons E7r to E15n of the Newark astrochronological polarity time scale. This represents a time interval from 227 to 213 Ma, indicating that the Los Colorados Formation is predominantly Norian in age, ending more than 11 My before the onset of the Jurassic. The magnetochronology confirms that the underlying Ischigualasto Formation and its vertebrate assemblages including some of the earliest known dinosaurs are of Carnian age. The oldest dated occurrences of vertebrate assemblages with dinosaurs in North America (Chinle Formation) are younger (Norian), and thus the rise of dinosaurs was diachronous across the Americas. Paleogeography of the Ischigualasto and Los Colorados Formations indicates prolonged residence in the austral temperate humid belt where a provincial vertebrate fauna with early dinosaurs may have incubated. Faunal dispersal across the Pangean supercontinent in the development of more cosmopolitan vertebrate assemblages later in the Norian may have been in response to reduced contrasts between climate zones and lowered barriers resulting from decreasing atmospheric pCO2 levels.