Bone orthotropic remodeling as a thermodynamically-driven evolution

International audienceIn this contribution we present and discuss a model of bone remodeling set up in the framework of the theory of generalized continuum mechanics and first introduced by DiCarlo et al.[1]. Bone is described as an orthotropic body experiencing remodeling as a rotation of its microstruc-ture. Thus, the complete kinematic description of a material point is provided by its position in space and a rotation tensor describing the orientation of its microstructure. Material motion is driven by energetic considerations , namely by the application of the Clausius-Duhem inequality to the microstructured material. Within this framework of orthotropic re-modeling, some key features of the remodeling equilibrium configurations are deduced in the case of homogeneous strain or stress loading conditions. First, it is shown that remodeling equilibrium configurations correspond to energy extrema. Second, stability of the remodeling equilibrium configurations is assessed in terms of the local convexity of the strain and complementary energy functionals hence recovering some classical energy theorems. Eventually, it is shown that the remodeling equilibrium configurations are not only highly dependent on the loading conditions, but also on the material properties

When moles became diggers: Tegulariscaptor gen. nov., from the early Oligocene of south Germany, and the evolution of talpid fossoriality

The systematics of Geotrypus is among the most debated within Talpidae, but the recent development of quantitative methods for shape analyses allows us to provide a thorough reconsideration of Geotrypus spp. In the present study, we perform a systematic revision of the species Geotrypus minor from the early Oligocene of Germany using two-dimensional geometric morphometrics on the humerus, and cladistic analyses using two different character matrices. Our results suggest a distinct generic allocation for this species based on its unique humeral shape. Cladistic analyses reveal that G. minor has closer phylogenetic relationships with urotrichine shrew-moles than with other Geotrypus species or highly fossorial moles. Quantitative methods applied in this study support qualitative observations and fully justify a new generic allocation. In light of these results, Tegulariscaptor gen. nov. is proposed to encompass the material previously assigned to G. minor.http://zoobank.org/urn:lsid:zoobank.org:pub:8A839F1E-0EC8-4799-B3AE-1A4E54A95F0

Raptor talon shape and biomechanical performance are controlled by relative prey size but not by allometry

Most birds of prey (raptors), rely heavily on their talons for capturing prey. However, the relationship between talon shape and the ability to take prey is poorly understood. In this study we investigate whether raptor talons have evolved primarily in response to adaptive pressures exerted by different dietary demands, or if talon morphology is largely constrained by allometric or phylogenetic factors. We focus on the hallux talon and include 21 species in total varying greatly in body mass and feeding ecology, ranging from active predation on relatively large prey to obligate scavenging. To quantify the variation in talon shape and biomechanical performance within a phylogenetic framework, we combined three dimensional (3D) geometric morphometrics, finite element modelling and phylogenetic comparative methods. Our results indicate that relative prey size plays a key role in shaping the raptorial talon. Species that hunt larger prey are characterised by both distinct talon shape and mechanical performance when compared to species that predate smaller prey, even when accounting for phylogeny. In contrast to previous results of skull-based analysis, allometry had no significant effect. In conclusion, we found that raptor talon evolution has been strongly influenced by relative prey size, but not allometry and, that talon shape and mechanical performance are good indicators of feeding ecology

Trapped in the morphospace: The relationship between morphological integration and functional performance

The evolution of complex morphological structures can be characterized by the interplay between different anatomical regions evolving under functional integration in response to shared selective pressures. Using the highly derived humeral morphology of talpid moles as a model, here we test whether functional performance is linked to increased levels of evolutionary integration between humerus subunits and, if so, what the strength is of the relationship. Combining two-dimensional geometric morphometrics, phylogenetic comparative methods, and functional landscape modeling, we demonstrate that the high biomechanical performance of subterranean moles’ humeri is coupled with elevated levels of integration, whereas taxa with low-performance values show intermediate or low integration. Theoretical morphs occurring in high-performance areas of the functional landscape are not occupied by any species, and show a marked drop in covariation levels, suggesting the existence of a strong relationship between integration and performance in the evolution of talpid moles’ humeri. We argue that the relative temporal invariance of the subterranean environment may have contributed to stabilize humeral morphology, trapping subterranean moles in a narrow region of the landscape and impeding any attempt to reposition on a new ascending slope

Unexpectedly uneven distribution of functional trade-offs explains cranial morphological diversity in carnivores

Functional trade-offs can affect patterns of morphological and ecological evolution as well as the magnitude of morphological changes through evolutionary time. Using morpho-functional landscape modelling on the cranium of 132 carnivore species, we focused on the macroevolutionary effects of the trade-off between bite force and bite velocity. Here, we show that rates of evolution in form (morphology) are decoupled from rates of evolution in function. Further, we found theoretical morphologies optimising for velocity to be more diverse, while a much smaller phenotypic space was occupied by shapes optimising force. This pattern of differential representation of different functions in theoretical morphological space was highly correlated with patterns of actual morphological disparity. We hypothesise that many-to-one mapping of cranium shape on function may prevent the detection of direct relationships between form and function. As comparatively only few morphologies optimise bite force, species optimising this function may be less abundant because they are less likely to evolve. This, in turn, may explain why certain clades are less variable than others. Given the ubiquity of functional trade-offs in biological systems, these patterns may be general and may help to explain the unevenness of morphological and functional diversity across the tree of life

Pequeños vertebrados del relleno kárstico del Pleistoceno Superior de Avetrana (Apulia, Sur de Italia)

The fossiliferous deposit (karst cavity) in La Grave, a locality near the small town of Avetrana (Taranto, south­ern Italy), has yielded numerous fossils of vertebrates. The remains of large mammals have been the subject of several studies. This paper examines the remains of small vertebrates and identifies four taxa of amphibians (Bufo bufo, Bufotes gr. B. viridis, Hyla gr. H. Arborea and Rana (s.l.) sp.), four taxa of reptiles (Testudo hermanni, Podarcis sp., Zamenis gr. Z. longissimus, Natrix natrix), and nine taxa of small mammals (Erinaceus europaeus, Crocidura suaveolens, Arvicola italicus, Microtus (Terricola) savii, Microtus (Microtus) arvalis, Apodemus gr. A. sylvaticus - A. flavicollis, Hystrix (Acanthion) vinogradovi, Oryctolagus cuniculus and Lepus corsicanus). From a biochronological point of view, the data on small and large vertebrates indicate an age between the beginning of the Late Pleistocene (MIS 5e) and the central part of MIS 3. The most recent fossiliferous layer (bed 8) is likely to have been deposited during a cooler period when compared to the previous layers.The data from small fossil vertebrates combined with those emerging from the large mammals and birds evidence the presence, near the karstic cavity, of open spaces (prairies) with pools of water, bordered by wooded areas and, not far, the presence of a rocky coastline.El depósito (cavidad kárstica) de La Grave, localidad cercana a la pequeña ciudad de Avetrana (Tarento, Italia meridional), ha dado lugar a numerosos fósiles de vertebrados. Los restos de grandes mamíferos han sido objeto de varios estudios. En este trabajo se examinan los restos de pequeños vertebrados y se identifican cuatro taxones de anfibios (Bufo bufo, Bufotes gr. B. viridis, Hyla gr. H. Arborea and Rana (s.l.) sp.), cuatro de reptiles (Testudo hermanni, Podarcis sp., Zamenis gr. Z. longissimus, Natrix natrix), y nueve de pequeños mamíferos (Erinaceus europaeus, Crocidura suaveolens, Arvicola italicus, Microtus (Terricola) savii, Microtus (Microtus) arva­lis, Apodemus gr. A. sylvaticus - A. flavicollis, Hystrix (Acanthion) vinogradovi, Oryctolagus cuniculus and Lepus corsicanus). Desde un punto de vista biocronológico, los datos sobre los vertebrados pequeños y grandes indican una edad entre el comienzo del Pleistoceno tardío (MIS 5e) y la parte central del MIS 3. Es probable que el estrato fosilífero más reciente (nivel 8) se haya depositado durante un período más frío en comparación con las capas anteriores. Los datos de pequeños vertebrados fósiles combinados con los que proceden de los grandes mamífe­ros y aves evidencian la presencia, cerca de la cavidad kárstica, de espacios abiertos (praderas) con charcos de agua, bordeados por zonas boscosas y, no muy lejos, la presencia de una costa rocosa

Trapped in the morphospace: The relationship between morphological integration and functional performance

The evolution of complex morphological structures can be characterized by the interplay between different anatomical regions evolving under functional integration in response to shared selective pressures. Using the highly derived humeral morphology of talpid moles as a model, here we test whether functional performance is linked to increased levels of evolutionary integration between humerus subunits and, if so, what the strength is of the relationship. Combining two-dimensional geometric morphometrics, phylogenetic comparative methods, and functional landscape modeling, we demonstrate that the high biomechanical performance of subterranean moles’ humeri is coupled with elevated levels of integration, whereas taxa with low-performance values show intermediate or low integration. Theoretical morphs occurring in high-performance areas of the functional landscape are not occupied by any species, and show a marked drop in covariation levels, suggesting the existence of a strong relationship between integration and performance in the evolution of talpid moles’ humeri. We argue that the relative temporal invariance of the subterranean environment may have contributed to stabilize humeral morphology, trapping subterranean moles in a narrow region of the landscape and impeding any attempt to reposition on a new ascending slope

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

RRmorph—a new R package to map phenotypic evolutionary rates and patterns on 3D meshes

The study of evolutionary rates and patterns is the key to understand how natural selection shaped the current and past diversity of phenotypes. Phylogenetic comparative methods offer an array of solutions to undertake this challenging task, and help understanding phenotypic variation in full in most circumstances. However, complex, three-dimensional structures such as the skull and the brain serve disparate goals, and different portions of these phenotypes often fulfil different functions, making it hard to understand which parts truly were recruited by natural selection. In the recent past, we developed tools apt to chart evolutionary rate and patterns directly on three-dimensional shapes, according to their magnitude and direction. Here, we present further developments of these tools, which now allow to restitute the mapping of rates and patterns with full biological realism. The tools are condensed in a new R software package.Evolutionary rates embody the velocity of evolution. Parcellating different velocities across the phenotype is difficult. RRmorph resolves this conundrum by charting evolutionary patterns on 3D shapes, according to their magnitude and direction