James M. Neenan

The Braincase and Inner Ear of Placodus gigas (Sauropterygia, Placodontia)—A New Reconstruction Based on Micro-Computed Tomographic Data

ABSTRACT Placodus gigas is an unarmored placodont marine reptile (Diapsida, Sauropterygia) known from the Middle Triassic of Europe, most commonly found in the shallow marine carbonate facies of the German Muschelkalk (lower Anisian to middle Ladinian, approximately 243–235 Ma). Generally, the morphology of the skull is well understood, with the exception of the braincase, which is partially obscured by dermatocranial bone. Two skulls that display well-preserved and intact chondrocranial elements were scanned using industrial micro-computed tomography (µCT), thus revealing parts of the braincase that were previously obscured and allowing a new three-dimensional reconstruction of the region to be constructed. This includes a complete osteological description, the first reconstruction of a sauropterygian vestibular apparatus, and a new virtual cranial endocast. The morphology of the braincase and sphenoid region has been revised, revealing the position of the hypophyseal pit. The enigmatic ‘alisphenoid bridge’ has been reinterpreted as being a dorsally extended dorsum sellae. The vestibular apparatus is shown to have strongly dorsoventrally compressed vertical semicircular canals, a commonly observed morphology of other marine reptiles, and an ‘alert’ head position of about 20°, indicating a highly aquatic lifestyle. Because placodonts are the sister group to all other sauropterygians, these new data are of great comparative and phylogenetic significance, providing insight into some of the morphological and functional changes that occurred on the stem leading to the more derived sauropterygians.

Feeding biomechanics in Acanthostega and across the fish–tetrapod transition

Acanthostega is one of the earliest and most primitive limbed vertebrates. Its numerous fish-like features indicate a primarily aquatic lifestyle, yet cranial suture morphology suggests that its skull is more similar to those of terrestrial taxa. Here, we apply geometric morphometrics and two-dimensional finite-element analysis to the lower jaws of Acanthostega and 22 other tetrapodomorph taxa in order to quantify morphological and functional changes across the fish–tetrapod transition. The jaw of Acanthostega is similar to that of certain tetrapodomorph fish and transitional Devonian taxa both morphologically (as indicated by its proximity to those taxa in morphospace) and functionally (as indicated by the distribution of stress values and relative magnitude of bite force). Our results suggest a slow tempo of morphological and biomechanical changes in the transition from Devonian tetrapod jaws to aquatic/semi-aquatic Carboniferous tetrapod jaws. We conclude that Acanthostega retained a primitively aquatic lifestyle and did not possess cranial adaptations for terrestrial feeding.

Unique method of tooth replacement in durophagous placodont marine reptiles, with new data on the dentition of Chinese taxa

The placodonts of the Triassic period (~252–201 mya) represent one of the earliest and most extreme specialisations to a durophagous diet of any known reptile group. Exceptionally enlarged crushing tooth plates on the maxilla, dentary and palatine cooperated to form functional crushing areas in the buccal cavity. However, the extreme size of these teeth, combined with the unusual way they occluded, constrained how replacement occurred. Using an extensive micro‐computed tomographic dataset of 11 specimens that span all geographic regions and placodont morphotypes, tooth replacement patterns were investigated. In addition, the previously undescribed dental morphologies and formulae of Chinese taxa are described for the first time and incorporated into the analysis. Placodonts have a unique tooth replacement pattern and results follow a phylogenetic trend. The plesiomorphic Placodus species show many replacement teeth at various stages of growth, with little or no discernible pattern. On the other hand, the more derived cyamodontoids tend to have fewer replacement teeth growing at any one time, replacing teeth unilaterally and/or in functional units, thus maintaining at least one functional crushing area at all times. The highly derived placochelyids have fewer teeth and, as a result, only have one or two replacement teeth in the upper jaw. This supports previous suggestions that these taxa had an alternative diet to other placodonts. Importantly, all specimens show at least one replacement tooth growing at the most posterior palatine tooth plates, indicating increased wear at this point and thus the most efficient functional crushing area.

Revised paleoecology of placodonts – with a comment on ‘The shallow marine placodont Cyamodus of the central European Germanic Basin: its evolution, paleobiogeography and paleoecology’ by C.G. Diedrich (Historical Biology, iFirst article, 2011, 1–19, doi: 10.1080/08912963.2011.575938)

A recent article published by Diedrich (2011a, Hist Biol. iFirst online, 1–19, doi: 10.1080/08912963.2011.575938) aspired to provide a complete revision of the known material of the placodont genus Cyamodus Meyer, 1863 from the Germanic Basin of central Europe. It is the latest in a series of similar articles by the same author (see Diedrich 2010, Palaeogeogr Palaeoclimatol Palaeoecol. 285(3–4):287–306; 2011b, Nat Sci. 3(1):9–27 for overview) focussing on the European members of the Placodontia (Reptilia: Sauropterygia), a diverse group of enigmatic marine reptiles known from Triassic shallow marine deposits. In a similar fashion to some previous works by Diedrich (see Tintori 2011, Palaeogeogr Palaeoclimatol Palaeoecol. 300(1–4):205–207 for similar points of criticism), this newest article demonstrates a narrow scope of presenting and discussing data, including omitted articles relevant to the topic, and over-interpretation of results, all with the aim of embedding the idea of placodonts being herbivorous Triassic ‘sea cows’ feeding on macroalgae (Diedrich 2010, 2011b). The present contribution is intended to clarify mistakes and misinterpretations made by Diedrich (2011a), to incorporate vital citations previously omitted which allow alternative interpretations, and to put the paper into perspective by including a more general evolutionary and paleoecological overview of the remaining placodonts.

Growth patterns and life-history strategies in Placodontia (Diapsida: Sauropterygia).

Placodontia is a clade of durophagous, near shore marine reptiles from Triassic sediments of modern-day Europe, Middle East and China. Although much is known about their primary anatomy and palaeoecology, relatively little has been published regarding their life history, i.e. ageing, maturation and growth. Here, growth records derived from long bone histological data of placodont individuals are described and modelled to assess placodont growth and life-history strategies. Growth modelling methods are used to confirm traits documented in the growth record (age at onset of sexual maturity, age when asymptotic length was achieved, age at death, maximum longevity) and also to estimate undocumented traits. Based on these growth models, generalized estimates of these traits are established for each taxon. Overall differences in bone tissue types and resulting growth curves indicate different growth patterns and life-history strategies between different taxa of Placodontia. Psephoderma and Paraplacodus grew with lamellar-zonal bone tissue type and show growth patterns as seen in modern reptiles. Placodontia indet. aff. Cyamodus and some Placodontia indet. show a unique combination of fibrolamellar bone tissue regularly stratified by growth marks, a pattern absent in modern sauropsids. The bone tissue type of Placodontia indet. aff. Cyamodus and Placodontia indet. indicates a significantly increased basal metabolic rate when compared with modern reptiles. Double lines of arrested growth, non-annual rest lines in annuli, and subcycles that stratify zones suggest high dependence of placodont growth on endogenous and exogenous factors. Histological and modelled differences within taxa point to high individual developmental plasticity but sexual dimorphism in growth patterns and the presence of different taxa in the sample cannot be ruled out.

New specimen of Psephoderma alpinum (Sauropterygia, Placodontia) from the Late Triassic of Schesaplana Mountain, Graubunden, Switzerland

Psephoderma alpinum is an armoured, durophagous placodont known from the alpine Late Triassic. Here we present a new, well-preserved isolated skull discovered in the Alplihorn Member (Late Norian–Early Rhaetian) of the Kössen Formation, Schesaplana Mountain, which straddles the Swiss/Austrian border. Micro-computed tomographic (µCT) scanning was used to create an accurate osteological reconstruction of the specimen, the first time this has been conducted for Psephoderma. We thus clarify disputed anatomical features from previous descriptions, such as a lack of a lacrimal and a pineal foramen that is enclosed by the parietal. We also present the first description based on µCT data of the lateral braincase wall, sphenoid region and some cranial nerve canals for Psephoderma, with the location of the hypophyseal seat, cerebral carotid foramina, dorsum sellae, prootic foramen, lacrimal foramen, as well as all dental foramina being described. This specimen represents the first skull of Psephoderma recovered in Switzerland, and features such as poorly-sutured braincase elements and its relatively small size compared to other known specimens may indicate that it was a sub-adult.

An enigmatic aquatic snake from the Cenomanian of Northern South America

We report the first record of a snake from the Cretaceous of northern South America. The remains come from the La Luna Formation (La Aguada Member, Cenomanian of Venezuela) and consist of several vertebrae, which belong to the precloacal region of the vertebral column. Comparisons to extant and extinct snakes show that the remains represent a new taxon, Lunaophis aquaticus gen. et sp nov. An aquatic mode of life is supported by the ventral position of the ribs, indicating a laterally compressed body. The systematic relationships of this new taxon are difficult to determine due to the scarcity of fossil material; it is, however, a representative of an early lineage of snakes that exploited tropical marine pelagic environments, as reflected by the depositional conditions of the La Aguada Member. Lunaophis is also the first aquatic snake from the Cenomanian found outside of the African and European Tethyan and Boreal Zones.

A new, exceptionally preserved juvenile specimen of Eusaurosphargis dalsassoi (Diapsida) and implications for Mesozoic marine diapsid phylogeny

Recently it was suggested that the phylogenetic clustering of Mesozoic marine reptile lineages, such as thalattosaurs, the very successful fish-shaped ichthyosaurs and sauropterygians (including plesiosaurs), among others, in a so-called ‘superclade’ is an artefact linked to convergent evolution of morphological characters associated with a shared marine lifestyle. Accordingly, partial ‘un-scoring’ of the problematic phylogenetic characters was proposed. Here we report a new, exceptionally preserved and mostly articulated juvenile skeleton of the diapsid reptile, Eusaurosphargis dalsassoi, a species previously recovered within the marine reptile ‘superclade’, for which we now provide a revised diagnosis. Using micro-computed tomography, we show that besides having a deep skull with a short and broad rostrum, the most outstanding feature of the new specimen is extensive, complex body armour, mostly preserved in situ, along its vertebrae, ribs, and forelimbs, as well as a row of flat, keeled ventrolateral osteoderms associated with the gastralia. As a whole, the anatomical features support an essentially terrestrial lifestyle of the animal. A review of the proposed partial character ‘un-scoring’ using three published data matrices indicate that this approach is flawed and should be avoided, and that within the marine reptile ‘superclade’ E. dalsassoi potentially is the sister taxon of Sauropterygia.

Bony labyrinth morphology clarifies the origin and evolution of deer

Deer are an iconic group of large mammals that originated in the Early Miocene of Eurasia (ca. 19 Ma). While there is some consensus on key relationships among their members, on the basis of molecular- or morphology-based analyses, or combined approaches, many questions remain, and the bony labyrinth has shown considerable potential for the phylogenetics of this and other groups. Here we examine its shape in 29 species of living and fossil deer using 3D geometric morphometrics and cladistics. We clarify several issues of the origin and evolution of cervids. Our results give new age estimates at different nodes of the tree and provide for the first time a clear distinction of stem and crown Cervidae. We unambiguously attribute the fossil Euprox furcatus (13.8 Ma) to crown Cervidae, pushing back the origin of crown deer to (at least) 4 Ma. Furthermore, we show that Capreolinae are more variable in bony labyrinth shape than Cervinae and confirm for the first time the monophyly of the Old World Capreolinae (including the Chinese water deer Hydropotes) based on morphological characters only. Finally, we provide evidence to support the sister group relationship of Megaloceros giganteus with the fallow deer Dama.

Tooth occlusal morphology in the durophagous marine reptiles, Placodontia (Reptilia: Sauropterygia)

Abstract. Placodontia were a group of marine reptiles that lived in shallow nearshore environments during the Triassic. Based on tooth morphology it has been inferred that they were durophagous, but tooth morphology differs among species: placodontoid placodonts have teeth described as hemispherical, and the teeth of more highly nested taxa within the cyamodontoid placodonts have been described as flat. In contrast, the sister taxon to the placodonts, Palatodonta bleekeri, like many other marine reptiles, has tall pointed teeth for eating soft-bodied prey. The goals of this paper are to quantify these different tooth morphologies and compare tooth shape among taxa and with a functionally “optimal” tooth. To quantify tooth morphology we measured the radius of curvature (RoC) of the occlusal surface by fitting spheres to 3D surface scans or computed microtomographic scans. Large RoCs correspond to flatter teeth, while teeth with smaller RoCs are pointier; positive RoCs have convex occlusal surfaces, and a negative RoC indicates that the occlusal surface of the tooth is concave. We found the placodontoid taxa have teeth with smaller RoCs than more highly nested taxa, and palatine teeth tend to be flatter and closer to the optimal morphology than maxillary teeth. Within one well-nested clade, the placochelyids, the rearmost palatine teeth have a more complex morphology than the predicted optimal tooth, with an overall concave occlusal surface with a small, medial cusp. These findings are in keeping with the hypothesis that placodonts were specialized durophagous predators with teeth modified to break hard prey items while resisting tooth failure.