T. Kotsakis

The Gavialis--Tomistoma debate: the contribution of skull ontogenetic allometry and growth trajectories to the study of crocodylian relationships

SUMMARY The phylogenetic placement of Tomistoma and Gavialis crocodiles depends largely upon whether molecular or morphological data are utilized. Molecular analyses consider them as sister taxa, whereas morphological/paleontological analyses set Gavialis apart from Tomistoma and other crocodylian species. Here skull allometric trajectories of Tomistoma and Gavialis were contrasted with those of two longirostral crocodylian taxa, Crocodylus acutus and Mecistops cataphractus, to examine similarities in growth trajectories in light of this phylogenetic controversy. Entire skull shape and its two main modules, rostrum and postrostrum, were analyzed separately. We tested differences for both multivariate angles between trajectories and for shape differences at early and late stages of development. Based on a multivariate regression of shape data and size, Tomistoma seems to possess a peculiar rate of growth in comparison to the remaining taxa. However, its morphology at both juvenile and adult sizes is always closer to those of Brevirostres crocodylians, for the entire head shape, as well as the shape of the postrostrum and rostrum. By contrast, the allometric trajectory of Gavialis always begins and ends in a unique region of the multidimensional morphospace. These findings concur with a morphological hypothesis that places Gavialis separate from Brevirostres, and Tomistoma closer to other crocodylids, and provides an additional, and independent, data set to inform on this ongoing phylogenetic discussion.

The role of post-natal ontogeny in the evolution of phenotypic diversity in Podarcis lizards.

Understanding the role of the developmental pathways in shaping phenotypic diversity allows appreciating in full the processes influencing and constraining morphological change. Podarcis lizards demonstrate extraordinary morphological variability that likely originated in short evolutionary time. Using geometric morphometrics and a broad suite of statistical tests, we explored the role of developmental mechanisms such as growth rate change, ontogenetic divergence/convergence/parallelism as well as morphological expression of heterochronic processes in mediating the formation of their phenotypic diversity during the post‐natal ontogeny. We identified hypermorphosis – the prolongation of growth along the same trajectory – as the process responsible for both intersexual and interspecific morphological differentiation. Albeit the common allometric pattern observed in both sexes of any species constrains and canalizes their cephalic scales variation in a fixed portion of the phenotypic space, the extended growth experienced by males and some species allows them to achieve peramorphic morphologies. Conversely, the intrasexual phenotypic diversity is accounted for by non‐allometric processes that drive the extensive morphological dispersion throughout their ontogenetic trajectories. This study suggests a model of how simple heterochronic perturbations can produce phenotypic variation, and thus potential for further evolutionary change, even within a strictly constrained developmental pathway.

Bite of the Cats: Relationships between Functional Integration and Mechanical Performance as Revealed by Mandible Geometry

Cat-like carnivorous mammals represent a relatively homogeneous group of species whose morphology appears constrained by exclusive adaptations for meat eating. We present the most comprehensive data set of extant and extinct cat-like species to test for evolutionary transformations in size, shape and mechanical performance, that is, von Mises stress and surface traction, of the mandible. Size and shape were both quantified by means of geometric morphometrics, whereas mechanical performance was assessed applying finite element models to 2D geometry of the mandible. Additionally, we present the first almost complete composite phylogeny of cat-like carnivorans for which well-preserved mandibles are known, including representatives of 35 extant and 59 extinct species of Felidae, Nimravidae, and Barbourofelidae. This phylogeny was used to test morphological differentiation, allometry, and covariation of mandible parts within and among clades. After taking phylogeny into account, we found that both allometry and mechanical variables exhibit a significant impact on mandible shape. We also tested whether mechanical performance was linked to morphological integration. Mechanical stress at the coronoid process is higher in sabertoothed cats than in any other clade. This is strongly related to the high degree of covariation within modules of sabertooths mandibles. We found significant correlation between integration at the clade level and per-clade averaged stress values, on both original data and by partialling out interclade allometry from shapes when calculating integration. This suggests a strong interaction between natural selection and the evolution of developmental and functional modules at the clade level.

The Pleistocene deer hunter of Sardinia

The Pleistocene mammalian fauna of Sardinia isendemic and consists of only a few taxa, indicating island conditions at the time. A sharp faunal break in the Middle Pleistocene separates an older fauna with Nesogoral from a younger one (Middle/Upper Pleistocene) which contains mainly deer (Megaceros cazioti), and ochotonid (Prolagus sardus), a small canid (Cynotherium sardous), and Tyrrhenicola. Commonly paleoecological conditions on islands have led to the development of faunas with dwarfed species, showing a low gear locomotion due to lack of large carnivores. In this aspect the younger Pleistocene fauna of Sardinia is different. A deer with the proportion of mainland species and a small canid were found. This is suggestive of the presence of a large predator. Indeed, together with this fauna a human temporal bone, showing some distinctive characters, was found in Grotta di Corbeddu, Oliena, Sardinia. These findings gave rise to further excavations in the same cave. A paleolithic settlement was discovered and two types of bone artefact could be distinguished. 14C dating revealed an age of 13590±140 yrs. B.P. These results suggest the development of an endemichuman culture, adapted to the restricted island environment and to the monotonous prey of deer and ochotonids, different from any known culture of that age.

Testing Convergent and Parallel Adaptations in Talpids Humeral Mechanical Performance by means of Geometric Morphometrics and Finite Element Analysis

The shape and mechanical performance in Talpidae humeri were studied by means of Geometric Morphometrics and Finite Element Analysis, including both extinct and extant taxa. The aim of this study was to test whether the ability to dig, quantified by humerus mechanical performance, was characterized by convergent or parallel adaptations in different clades of complex tunnel digger within Talpidae, that is, Talpinae+Condylura (monophyletic) and some complex tunnel diggers not belonging to this clade. Our results suggest that the pattern underlying Talpidae humerus evolution is evolutionary parallelism. However, this insight changed to true convergence when we tested an alternative phylogeny based on molecular data, with Condylura moved to a more basal phylogenetic position. Shape and performance analyses, as well as specific comparative methods, provided strong evidence that the ability to dig complex tunnels reached a functional optimum in distantly related taxa. This was also confirmed by the lower phenotypic variance in complex tunnel digger taxa, compared to non‐complex tunnel diggers. Evolutionary rates of phenotypic change showed a smooth deceleration in correspondence with the most recent common ancestor of the Talpinae+Condylura clade. J. Morphol. 2012.

Testing evolutionary stasis and trends in first lower molar shape of extinct Italian populations of Terricola savii (Arvicolidae, Rodentia) by means of geometric morphometrics

Extinct populations of Terricola savii have been investigated in order to analyse evolutionary stasis and correlation of first lower molar shape with climatic proxies by means of geometric morphometrics. Evolutionary stasis, its recognition and explanation are central topics in evolutionary paleobiology. In this study, tooth shape variation of the arvicolid T. savii has been analysed through time. In addition to explicit multivariate tests of stasis based on landmark and semi‐landmark geometric morphometrics, first lower molar M1 shape has been decomposed in orthogonal axes of variation and tested for correlation with climate changes. Multivariate tests were consistent with evolutionary stasis. Yet, according to univariate tests, the dominant dimension of shape variation shows a temporal trend well correlated with a climatic proxy, i.e. δ18O. The remaining variation does not show any trend. Adaptation to current climatic condition might occur even without affecting shape as a whole. Phenotypic plasticity of this species could be invoked to explain evolutionary stasis, as a long time pattern.

Is torosaurus triceratops? Geometric morphometric evidence of late maastrichtian ceratopsid dinosaurs.

Background Recent assessments of morphological changes in the frill during ontogeny hypothesized that the late Maastrichtian horned dinosaur Torosaurus represents the “old adult” of Triceratops, although acceptance of this finding has been disputed on several lines of evidence. Methodology/Principal Findings Examining the cranial morphology of 28 skulls in lateral view and 36 squamosals of Nedoceratops hatcheri, Triceratops spp. and Torosaurus spp. by means of landmark-based geometric morphometrics, we compared ontogenetic trajectories among these taxa. Principal Component Analysis and cluster analysis confirmed different cranial morphologies. Torosaurus shape space is well separated from Triceratops, whereas Triceratops horridus and Triceratops prorsus partially overlap within Triceratops shape space. Linear regressions between shape and size suggest different ontogenetic trajectories among these taxa. Results support the “traditional” taxonomic status of Torosaurus. We hypothesize that ontogeny drives cranial morphology with different patterns between Torosaurus and Triceratops. Conclusions/Significance Torosaurus is a distinct and valid taxon. Whether looking at entire skulls, skulls without the frill, frills alone, or squamosals, Torosaurus has different morphologies and distinct allometric trajectories compared to Triceratops. This new approach confirms the taxonomic status of Torosaurus as well as the comparatively low diversity of ceratopsids at the end of the Maastrichtian in North America.

Morphological integration and functional modularity in the crocodilian skull.

We explored the morphological organization of the skull within Crocodylidae, analyzing functional and phylogenetic interactions between its 2 constituent functional modules: the rostrum and the postrostrum. We used geometric morphometrics to identify localized shape changes, focusing on the differences between the major clades of the crown-group Crocodylia: Alligatoridae and Crocodylidae. We used published bite performance data to correlate rostral function with postrostral morphology. The skull modules appear more integrated within Alligatoridae than within Crocodyliade. Phylogenetic effects on shape variation are more evident in Alligatoridae than in Crocodylidae, where functional parameters concerning the rostral morphology are proportionally more important than phylogeny. Long-snouted species are characterized by low structural performance, which is significantly associated with a reduction of the pterygoid-quadrate cranial nipper, suggesting that the nipper is important for the ingestion of large food items in generalist species. This functional association is coupled with a significant evolutionary allometry at the clade level, while Alligatoridae and Crocodylidae show different degrees of evolutionary allometry for their entire shape and rostrum. The postrostrum is more conservative than the rostrum in terms of morphospace occupation, evolutionary allometry and phylogenetic signal.

Digging adaptation in insectivorous subterranean eutherians. The enigma of Mesoscalops montanensis unveiled by geometric morphometrics and finite element analysis

The enigmatic Early Miocene fossorial mammal Mesoscalops montanensis shows one of the most modified humeri among terrestrial mammals. It has been suggested, on qualitative considerations, that this species has no extant homologues for humerus kinematics and that, functionally, the closest extant group is represented by Chrysochloridae. We combine here three dimensional geometric morphometrics, finite element analysis and phylogenetic comparative methods to explore the shape and mechanical stress states of Mesoscalops montanensis as well as of extant and extinct Talpidae and Chrysochloridae under realistic digging simulations. Evolutionary convergence analyses reveal that the shape of Mesoscalops montanensis represents a unique morphology in the context of fossorial mammals and that its functional performance, albeit superficially similar to that of extant Chrysochloridae, still represents a nonconvergent optimum for adaptation to digging. J. Morphol. 276:1157–1171, 2015.

New Systematic Insights about Plio-Pleistocene Moles from Poland

The generic attribution of the Plio-Pleistocene Polish moles ?Neurotrichus polonicus and ?Neurotrichus skoczeni has been questioned several times in the past. The fossil material belonging to ?Neurotrichus polonicus and ?Neurotrichus skoczeni is re-evaluated here and a new diagnosis is provided on the basis of qualitative considerations. In addition, a Geometric Morphometric analysis of the humerus has been performed including both extant and extinct Neurotrichini and Urotrichini taxa for comparison. Our results proved the unique morphology of the Polish material suggesting a distinct taxonomic state. The morphological variations evidenced by the humeral shape analysis agree with the observed qualitative differences and support a new generic allocation. The new genus Rzebikia gen. nov. is proposed for all the material previoulsly ascribed to ?Neurotrichus polonicus and ?Neurotrichus skoczeni.