Michael J. Polcyn

Dallasaurus turneri, a new primitive mosasauroid from the Middle Turonian of Texas and comments on the phylogeny of Mosasauridae (Squamata)

A new genus and species of primitive-limbed mosasauroid, Dallasaurus turneri, from the Middle Turonian (~92 Ma) of north-central Texas, is described on the basis of two incomplete skeletons. The new taxon retains plesiomorphic characters such as facultatively terrestrial limbs (plesiopedal) but also exhibits certain characters shared with derived mosasaurs of the subfamily Mosasaurinae. In phylogenetic analysis, the new taxon reconstructs as the basal member of that clade. Other plesiopedal taxa previously included in the family Aigialosauridae reconstruct in basal positions within three different, major clades that include members that achieved the derived (hydropedal) fin-like limb condition. In addition, Opetiosaurus and Aigialosaurus reconstruct as successive outgroup taxa to all other mosasauroids, thereby demonstrating the paraphyletic nature of the current concept of Aigialosauridae. Interpretation of our phylogenetic analysis suggests that three different lineages of mosasauroids independently achieved the derived mosasaur body plan, including fully marine limb modifications culminating in the development of flippers or paddle-like appendages. The inclusion of plesiopedal forms within lineages of well-established hydropedal clades requires a reorganisation of our concepts of Mosasauridae to include these basal forms. In order to avoid continued use of the paraphyletic taxon ‘Aigialosauridae’ as currently defined, we recommend ‘Aigialosauridae’ as a formal taxonomic name be used only for inclusion of Aigialosaurus dalmaticus and potential members of its own independent lineage. This also avoids the implicit polyphyletic use of ‘Mosasauridae’ the diagnosis of Mosasauridae should be modified to exclude limb characters that discriminate between more terrestrial versus more aquatic adaptations.

Russellosaurus coheni n. gen., n. sp., a 92 million-year-old mosasaur from Texas (USA), and the definition of the parafamily Russellosaurina

A new mosasaur, Russellosaurus coheni , is described from the Collignoniceras woollgari Zone (lower Middle Turonian) at Cedar Hill, Dallas County, Texas. At approximately 92 Ma, it is the oldest well-preserved mosasaur skull from North America. It possesses characters diagnostic of Plioplatecarpinae but retains numerous plesiomorphies as well. Phylogenetic analysis indicates a close relationship with Yaguarasaurus columbianus , and these two, together with Tethysaurus nopcsai , form a clade that occupies a position basal to the divergence of the subfamilies Tylosaurinae and Plioplatecarpinae. Russellosaurus coheni is proposed as the nominal taxon of a new mosasaur clade, parafamily taxon novum Russellosaurina, which includes Plioplatecarpinae, Tylosaurinae, their common ancestor and all descendants. Tethysaurus retains a plesiopedal limb and girdle morphology, and along with Russellosaurus and Yaguarasaurus , cranial plesiomorphies. Dallasaurus turneri , a temporally and geographically sympatric plesiopedal mosasaur, occupies a basal position within Mosasaurinae. This phyletic arrangement confirms that marine adaptations, such as development of paddle-like limbs, occurred independently in at least two lineages of mosasaurs, once within Mosasaurinae and once within Russellosaurina.

THE ANATOMY AND RELATIONSHIPS OF HAASIOPHIS TERRASANCTUS, A FOSSIL SNAKE WITH WELL-DEVELOPED HIND LIMBS FROM THE MID-CRETACEOUS OF THE MIDDLE EAST

Abstract The fossil snake species Haasiophis terrasanctus Tchernov, Rieppel, Zaher, Polcyn, and Jacobs, 2000, from the early Upper Cretaceous of the Middle East, is described and illustrated, following a review of the current debate on snake relationships and origins. The description and discussion presented here adds important detail to the knowledge of this taxon and its phylogenetic significance beyond the limited account presented in the original description of Haasiophis. The species is remarkable in that it shows the skull of a relatively advanced (i.e., macrostomatan) snake, yet preserves well-developed hind limbs. The hind limb includes a femur, tibia, fibula, astragalus, calcaneum, distal tarsal four, and remains of four metatarsals and two phalanges. Haasiophis cannot be considered a juvenile specimen of Pachyrhachis. The implications of the presence of well-developed hind limbs in Haasiophis, Pachyrhachis, and Podophis for the cladistic analysis of the phylogenetic interrelationships of these fossil snakes is discussed. The presence of well-developed hind limbs in Pachyrhachis and Haasiophis also creates methodological problems for the cladistic analysis of the phylogenetic relationships of these fossil snakes. Scenarios of snake origins are reviewed and found to be deficient in the absence of a well-corroborated hypothesis of snake relationships within Squamata.

Skin pigmentation provides evidence of convergent melanism in extinct marine reptiles

Throughout the animal kingdom, adaptive colouration serves critical functions ranging from inconspicuous camouflage to ostentatious sexual display, and can provide important information about the environment and biology of a particular organism. The most ubiquitous and abundant pigment, melanin, also has a diverse range of non-visual roles, including thermoregulation in ectotherms. However, little is known about the functional evolution of this important biochrome through deep time, owing to our limited ability to unambiguously identify traces of it in the fossil record. Here we present direct chemical evidence of pigmentation in fossilized skin, from three distantly related marine reptiles: a leatherback turtle, a mosasaur and an ichthyosaur. We demonstrate that dark traces of soft tissue in these fossils are dominated by molecularly preserved eumelanin, in intimate association with fossilized melanosomes. In addition, we suggest that contrary to the countershading of many pelagic animals, at least some ichthyosaurs were uniformly dark-coloured in life. Our analyses expand current knowledge of pigmentation in fossil integument beyond that of feathers, allowing for the reconstruction of colour over much greater ranges of extinct taxa and anatomy. In turn, our results provide evidence of convergent melanism in three disparate lineages of secondarily aquatic tetrapods. Based on extant marine analogues, we propose that the benefits of thermoregulation and/or crypsis are likely to have contributed to this melanisation, with the former having implications for the ability of each group to exploit cold environments.

Angolatitan adamastor, a new sauropod dinosaur and the first record from Angola

A forelimb of a new sauropod dinosaur (Angolatitan adamastor n. gen. et sp.) from the Late Turonian of Iembe (Bengo Province) represents the first dinosaur discovery in Angola, and is one of the few occurrences of sauropod dinosaurs in sub-Saharan Africa collected with good chronological controls. The marginal marine sediments yielding the specimen are reported to be late Turonian in age and, thus it represents a non-titanosaurian sauropod in sub-Saharan Africa at a time taken to be dominated by titanosaurian forms. Moreover, Angolatitan adamastor is the only basal Somphospondyli known in the Late Cretaceous which implies in the existence of relict forms in Africa.

A 17-My-old whale constrains onset of uplift and climate change in east Africa

Significance An enigmatic fossil representing the deep-diving, open-ocean whale family Ziphiidae found 740 km inland and at 620 m elevation in West Turkana, Kenya was rediscovered after it went missing for more than 30 years. This stranded whale fossil provides the first constraint on the initiation of east African uplift from near sea level at 17 Ma, limiting the timing and initial elevation of environmental change indicated by geodynamic and climatic modeling, paleosols, isotopes, paleobotany, and the mammalian fossil record. At 17 Ma, elevation was low, rainfall was high, vegetation was forested, and mammalian communities contained immigrants and native African species, including diverse primates. Uplift resulted in increasing aridity and open habitats that drove human evolution. Timing and magnitude of surface uplift are key to understanding the impact of crustal deformation and topographic growth on atmospheric circulation, environmental conditions, and surface processes. Uplift of the East African Plateau is linked to mantle processes, but paleoaltimetry data are too scarce to constrain plateau evolution and subsequent vertical motions associated with rifting. Here, we assess the paleotopographic implications of a beaked whale fossil (Ziphiidae) from the Turkana region of Kenya found 740 km inland from the present-day coastline of the Indian Ocean at an elevation of 620 m. The specimen is ∼17 My old and represents the oldest derived beaked whale known, consistent with molecular estimates of the emergence of modern strap-toothed whales (Mesoplodon). The whale traveled from the Indian Ocean inland along an eastward-directed drainage system controlled by the Cretaceous Anza Graben and was stranded slightly above sea level. Surface uplift from near sea level coincides with paleoclimatic change from a humid environment to highly variable and much drier conditions, which altered biotic communities and drove evolution in east Africa, including that of primates.

Microanatomical and Histological Features in the Long Bones of Mosasaurine Mosasaurs (Reptilia, Squamata) - Implications for Aquatic Adaptation and Growth Rates

Background During their evolution in the Late Cretaceous, mosasauroids attained a worldwide distribution, accompanied by a marked increase in body size and open ocean adaptations. This transition from land-dwellers to highly marine-adapted forms is readily apparent not only at the gross anatomic level but also in their inner bone architecture, which underwent profound modifications. Methodology/Principal Findings The present contribution describes, both qualitatively and quantitatively, the internal organization (microanatomy) and tissue types and characteristics (histology) of propodial and epipodial bones in one lineage of mosasauroids; i.e., the subfamily Mosasaurinae. By using microanatomical and histological data from limb bones in combination with recently acquired knowledge on the inner structure of ribs and vertebrae, and through comparisons with extant squamates and semi-aquatic to fully marine amniotes, we infer possible implications on mosasaurine evolution, aquatic adaptation, growth rates, and basal metabolic rates. Notably, we observe the occurrence of an unusual type of parallel-fibered bone, with large and randomly shaped osteocyte lacunae (otherwise typical of fibrous bone) and particular microanatomical features in Dallasaurus, which displays, rather than a spongious inner organization, bone mass increase in its humeri and a tubular organization in its femora and ribs. Conclusions/Significance The dominance of an unusual type of parallel-fibered bone suggests growth rates and, by extension, basal metabolic rates intermediate between that of the extant leatherback turtle, Dermochelys, and those suggested for plesiosaur and ichthyosaur reptiles. Moreover, the microanatomical features of the relatively primitive genus Dallasaurus differ from those of more derived mosasaurines, indicating an intermediate stage of adaptation for a marine existence. The more complete image of the various microanatomical trends observed in mosasaurine skeletal elements supports the evolutionary convergence between this lineage of secondarily aquatically adapted squamates and cetaceans in the ecological transition from a coastal to a pelagic lifestyle.

Microspectroscopic evidence of cretaceous bone proteins.

Low concentrations of the structural protein collagen have recently been reported in dinosaur fossils based primarily on mass spectrometric analyses of whole bone extracts. However, direct spectroscopic characterization of isolated fibrous bone tissues, a crucial test of hypotheses of biomolecular preservation over deep time, has not been performed. Here, we demonstrate that endogenous proteinaceous molecules are retained in a humerus from a Late Cretaceous mosasaur (an extinct giant marine lizard). In situ immunofluorescence of demineralized bone extracts shows reactivity to antibodies raised against type I collagen, and amino acid analyses of soluble proteins extracted from the bone exhibit a composition indicative of structural proteins or their breakdown products. These data are corroborated by synchrotron radiation-based infrared microspectroscopic studies demonstrating that amino acid containing matter is located in bone matrix fibrils that express imprints of the characteristic 67 nm D-periodicity typical of collagen. Moreover, the fibrils differ significantly in spectral signature from those of potential modern bacterial contaminants, such as biofilms and collagen-like proteins. Thus, the preservation of primary soft tissues and biomolecules is not limited to large-sized bones buried in fluvial sandstone environments, but also occurs in relatively small-sized skeletal elements deposited in marine sediments.

Landlubbers to leviathans: evolution of swimming in mosasaurine mosasaurs

Abstract Incremental stages of major evolutionary transitions within a single animal lineage are rarely observed in the fossil record. However, the extraordinarily complete sequence of well preserved material spanning the 27-Myr existence of the marine squamate subfamily Mosasaurinae provides a unique exception. By comparison with extant and extinct analogs, the tail morphology of four mosasaurine genera is examined, revealing a pattern of evolution that begins with the generalized varanoid anatomy and culminates in a high-aspect-ratio fin, similar to that of sharks. However, unlike the epicercal caudal fluke of selachians in which the tail bends dorsocaudally, derived mosasaurs develop a hypocercal tail with a ventrocaudal bend. Progressive caudal regionalization, reduced intervertebral mobility, increased tail depth due to a marked downturn of the posterior caudal segment, and the development of finlike paired appendages reveal a pattern of adaptation toward an optimized marine existence. This change in morphology reflects a transition from anguilliform or sub-carangiform locomotion to carangiform locomotion, and indicates a progressive shift from nearshore dwellers to pelagic cruisers—a change in foraging habitat independently corroborated by paleobiogeographic, stable isotope, osteohistological, and paleopathological data. Evolutionary patterns similar to those observed in mosasaurine mosasaurs are seen in other secondarily aquatically adapted amniotes, notably metriorhynchid crocodyliforms, cetaceans, and ichthyosaurs, and may be explained by developmental modularity governing the observed phenotypic expression.

Soft tissue preservation in a fossil marine lizard with a bilobed tail fin

Mosasaurs are secondarily aquatic squamates that became the dominant marine reptiles in the Late Cretaceous about 98-66 million years ago. Although early members of the group possessed body shapes similar to extant monitor lizards, derived forms have traditionally been portrayed as long, sleek animals with broadened, yet ultimately tapering tails. Here we report an extraordinary mosasaur fossil from the Maastrichtian of Harrana in central Jordan, which preserves soft tissues, including high fidelity outlines of a caudal fluke and flippers. This specimen provides the first indisputable evidence that derived mosasaurs were propelled by hypocercal tail fins, a hypothesis that was previously based on comparative skeletal anatomy alone. Ecomorphological comparisons suggest that derived mosasaurs were similar to pelagic sharks in terms of swimming performance, a finding that significantly expands our understanding of the level of aquatic adaptation achieved by these seagoing lizards.