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20 pages; International audience; The Southern Ocean (SO) is among the regions on Earth that are undergoing regionally the fastest environmental changes. The unique ecological features of its marine life make it particularly vulnerable to the multiple effects of climate change. A network of Marine Protected Areas (MPAs) has started to be implemented in the SO to protect marine ecosystems. However, considering future predictions of the Intergovernmental Panel on Climate Change (IPCC), the relevance of current, static, MPAs may be questioned under future scenarios. In this context, the ecoregionalization approach can prove promising in identifying well‐delimited regions of common species composition and environmental settings. These so‐called ecoregions are expected to show similar biotic responses to environmental changes and can be used to define priority areas for the designation of new MPAs and the update of their current delimitation. In the present work, a benthic ecoregionalization of the entire SO is proposed for the first time based on abiotic environmental parameters and the distribution of echinoid fauna, a diversified and common member of Antarctic benthic ecosystems. A novel two‐step approach was developed combining species distribution modeling with Random Forest and Gaussian Mixture modeling from species probabilities to define current ecoregions and predict future ecoregions under IPCC scenarios RCP 4.5 and 8.5. The ecological representativity of current and proposed MPAs of the SO is discussed with regard to the modeled benthic ecoregions. In all, 12 benthic ecoregions were determined under present conditions, they are representative of major biogeographic patterns already described. Our results show that the most dramatic changes can be expected along the Antarctic Peninsula, in East Antarctica and the sub‐Antarctic islands under both IPCC scenarios. Our results advocate for a dynamic definition of MPAs, they also argue for improving the representativity of Antarctic ecoregions in proposed MPAs and support current proposals of Conservation of Antarctic Marine Living Resources for the creation of Antarctic MPAs.

17 pages; International audience; Modularity and integration are variational properties expressed at various levels of the biological hierarchy. Mismatches among these levels, for example developmental modules that are integrated in a functional unit, could be informative of how evolutionary processes and trade‐offs have shaped organismal morphologies as well as clade diversification. In the present study, we explored the full, integrated and modular spaces of two developmental modules in phacopid trilobites, the cephalon and the pygidium, and highlight some differences among them. Such contrasts reveal firstly that evolutionary processes operating in the modular spaces are stronger in the cephalon, probably due to a complex regime of selection related to the numerous functions ensured by this module. Secondly, we demonstrate that the same pattern of covariation is shared among species, which also differentiate along this common functional integration. This common pattern might be the result of stabilizing selection acting on the enrolment and implying a coordinate variation between the cephalon and the pygidium in a certain direction of the morphospace. Finally, we noticed that Austerops legrandi differs slightly from other species in that its integration is partly restructured in the way the two modules interact. Such a divergence can result from the involvement of the cephalon in several vital functions that may have constrained the response of the features involved in enrolment and reorganized the covariation of the pygidium with the cephalon. Therefore, it is possible that important evolutionary trade‐offs between enrolment and other functions on the cephalon might have partly shaped the diversification of trilobites.

15 pages; International audience; Lemmus and Myopus are two lemming species with distinct habitat requirements but which show very similar dental morphologies. They are thus extremely difficult to distinguish from one another in the fossil record on the basis of their dental remains, leading to poor understanding of the palaeobiogeographical evolution of Myopus as well as inaccurate palaeoenvrionmental reconstructions. Currently, the presence of Myopus in the fossil register from the Pleistocene is still debated and no firm occurrence of this lemming in western Europe has yet been confirmed for the Late Pleistocene. In this paper, we used geometric morphometrics on modern material to establish morphological differences between Lemmus and Myopus teeth (first lower and third upper molars). Morphological data were then used to build a robust linear discriminant model able to confidently classify isolated teeth of these two genera, and finally, linear discriminant models were used on fossil remains of Lemmus/Myopus from two Late Pleistocene archaeological/palaeontological sites (Grotte des Gorges and Gully Cave). This study demonstrates, for the first time, the presence of Myopus schisticolor in west European Late Pleistocene sites between the end of Marine Isotope Stage 3 and the beginning of the Holocene, during climatic events that favoured the development of taiga forest of birch and pine in these regions.

25 pages; International audience; The Mediterranean Region is a biodiversity/endemism hotspot whose freshwater fauna remains largely unexplored. Our integrative study challenges the taxonomic status of two freshwater palaemonid shrimps, Palaemon antennarius and Palaemon minos. Three molecular operational taxonomic units (MOTUs) were defined based on 352 cytochrome oxidase subunit I (COI) sequences and 88 haplotypes. Two belonged to P. antennarius: one inhabiting the Apennine Peninsula and Sicily, and the other from the Balkan Peninsula. Palaemon minos was the third MOTU, found on Crete. The Balkan MOTU of P. antennarius was genetically closer to P. minos than to the other conspecific MOTU. Data from a nuclear marker (Histone 3) is congruent with such a pattern. The carapace shape variation (based on 180 individuals) was mainly explained by the geographical distribution. Balkan and Cretan groups were clearly recovered, while other samples clustered along a shape gradient from Sicily, through the Apennine Peninsula to the Balkans. Our results show that, for taxonomic consistency, the MOTU inhabiting the Balkan Peninsula should be either described as a new separate species or synonymised with P. minos. The third possible option would be treating all the populations as part of P. antennarius. Geometric morphometrics supports the first option, phylogenetic reconstructions point to the second one, yet the low genetic divergence favours the third one, illustrating that even emblematic taxa such as shrimps require an in-depth integrative approach.

18 pages; International audience; Mammal remains, preserved in archaeological and palaeontological deposits, are commonly used to reconstruct past terrestrial climates and environments. Here we propose new species-specific models for Bioclimatic Analysis, a palaeoclimatic method based on a climatic restriction index for each mammal species, discriminant analysis, and multiple linear regressions. Our new models are based on small mammal associations, particularly insectivores and rodents, from Quaternary paleoarctic contexts. A dataset including new localities and an updated taxonomy was constructed in order to develop two approaches, the first using only Rodentia, the second based on associations including both Rodentia and Eulipotyphla. Both approaches proved to be reliable for inferring both climate zone and quantifying temperature, precipitation, and seasonality. Rarefaction analysis revealed these new models to be reliable even when a substantial percentage of species from the original palaeocommunity was absent from the fossil site. Application of these new models to small mammal associations from two sequences (Balma de l'Abeurador, France and El Mirón, Spain) spanning from the Last Glacial Maximum to the Holocene are consistent with the primary climatic changes recorded by regional Pyrenean proxies and showed an increase in mean annual temperature of between 3 and 5 °C.

18 pages; International audience; The increase in species range size towards high latitudes, known as Rapoport's rule, remains one of the most debated and poorly understood macroecological patterns. Numerous studies have challenged both its universality and the main mechanism originally proposed to explain it: the climatic variability hypothesis. Here, we study this pattern using a group of fossil marine organisms: the early Pliensbachian ammonites of the western Tethys. We further take into account the influence of the marked provincialism prevailing at that time, with a Mediterranean province (MED) and a north‐west European province (NWE) located on each side of a latitudinally oriented palaeobiogeographical barrier. We find that only species from the NWE province display a Rapoport effect, whereas species from the more tropical MED province show a boundary effect and have larger range sizes on average. This dual pattern can be explained by an alternative climatic variability hypothesis that better captures latitudinal seasonal variations and outlines the influence of the intertropical zone, characterized by stable and homogeneous climate that allows species to disperse over very large areas, regardless of their thermal tolerance. Accordingly, the NWE province probably displayed a gradient of seasonal climatic variations which caused the emergence of a Rapoport effect, whereas the MED province was probably located in the intertropical zone where no gradient in species range size is expected. Our multiscale approach further shows that the Rapoport effect is scale‐dependent and may be labile through time. This probably explains the conflicting results of previous studies carried out at various spatiotemporal scales.

16 pages; International audience; Significant environmental changes have already been documented in the Southern Ocean (e.g. seawater temperature increase and salinity drop) but its marine life is still incompletely known given the heterogeneousnature of biogeographic data. However, to establish sustainable conservation areas, understandingspecies and communities distribution patterns is critical. For this purpose, the ecoregionalization approachcan prove useful by identifying spatially explicit and well-delimited regions of common species compositionand environmental settings. Such regions are expected to have similar biotic responses to environmentalchanges and can be used to define priorities for the designation of Marine Protected Areas. In the presentwork, a benthic ecoregionalization of the Southern Ocean is proposed based on echinoids distribution dataand abiotic environmental parameters. Echinoids are widely distributed in the Southern Ocean, they are taxonomicallyand ecologically well diversified and documented. Given the heterogeneity of the sampling effort,predictive spatial models were produced to fill the gaps in between species distribution data. A first procedurewas developed using Gaussian Mixture Models (GMM) to combine individual species models into ecoregions.A second, integrative procedure was implemented using the Generalized Dissimilarity Models (GDM)to model and assemble species distributions. Both procedures were compared to propose benthic ecoregionsat the scale of the entire Southern Ocean.

The Early Aptian encountered several crises in neritic and pelagic carbonate production, major perturbations in the carbon cycle, and an oceanic anoxic event (OAE1a). Yet the causal links between these perturbations and climate changes remain poorly understood, partly because temperature records spanning the Early Aptian interval are still scant. We present new δ18O data from well-preserved bivalves from a carbonate platform of the Galve subbasin (Spain) that document a major cooling event postdating most of OAE1a. Our data show that cooling postdates the global platform demise and cannot have triggered this event that occurred during the warmest interval. The warmest temperatures coincide with the time equivalent of OAE1a and with platform biotic assemblages dominated by microbialites at Aliaga as well as on other Tethyan platforms. Coral-dominated assemblages then replace microbialites during the subsequent cooling. Nannoconids are absent during most of the time equivalent of the OAE1a, probably related to the well-known crisis affecting this group. Yet they present a transient recovery in the upper part of this interval with an increase in both size and abundance during the cool interval portion that postdates OAE1a. An evolution toward cooler and drier climatic conditions may have induced the regional change from microbial to coral assemblages as well as nannoconids size and abundance increase by limiting continent-derived input of nutrients.

18 pages; International audience; Accurate species identification in fossil remains is a complex task but is a key component for developing good inferences on many, if not all, fundamental questions in macroecology and macroevolution. In the Quaternary, arvicolines are very abundant remains in archeological and paleontological sites in Western Europe and their identification is often based on the first lower molar. The common vole Microtus arvalis (Pallas, 1778) and the field vole Microtus agrestis (Linnaeus, 1761) are commonly found in those deposits. These two species are genetically and ecologically divergent. Nonetheless, their lower molars, on which species identification is done, exhibit a large morphological variation that can potentially lead to some confusion and misinterpretation. Moreover, molecular data suggest that present-day M. agrestis populations are a complex of divergent lineages, some of them being recognized nowadays as valid species. On the basis of extant populations representing a large part of the present-day geographical distribution of these two species, we developed a classification model based on geometric morphometrics of the first lower molar. Our statistical model was then applied on four fossil sites selected to evaluate the relevance of taxonomic determination found in species lists. The model using landmarks describing the overall shape of the first lower molar classifies the two species with the smallest prediction error together with very high individual posterior probabilities. The obtained classification is much better than those arising from shapes of any specific molar part such as the anterior loop, asymmetry or peculiar triangle shape. Discrepancies with expert classification on fossils suggest that existing faunal lists should always be considered cautiously for these two species. Our morphometric model provides a first step towards a rationalized way of revising past collections and expertise for future small mammal assemblages. It will thus help us better understand the paleobiogeographical expansion of these two key species in Quaternary faunas.

19 pages; International audience; Evolutionary radiations are fascinating phenomena corresponding to a dramatic diversification of taxa and a burst of cladogenesis over short periods of time. Most evolutionary radiations have long been regarded as adaptive but this has seldom been demonstrated with large-scale comparative datasets including fossil data. Originating in the Early Jurassic, irregular echinoids are emblematic of the spectacular diversification of mobile marine faunas during the Mesozoic Marine Revolution. They diversified as they colonized various habitats, and now constitute the main component of echinoid fauna in modern seas. The evolutionary radiation of irregular echinoids has long been considered as adaptive but this hypothesis has never been tested. In the present work we analyze the evolution of echinoid species richness and morphological disparity over 37 million years based on an extensive fossil dataset. Our results demonstrate that morphological and functional diversifications in certain clades of irregular echinoids were exceptionally high compared to other clades and that they were associated with the evolution of new modes of life and so can be defined as adaptive radiations. The role played by ecological opportunities in the diversification of these clades was critical, with the evolution of the infaunal mode of life promoting the adaptive radiation of irregular echinoids.

12 pages; International audience; AimTo test for the phylogenetic conservatism of geographic range size and to explore the effect of the environment on this potential conservatism.LocationThe western Tethys Ocean and its surroundings (present-day Europe, the Middle East and North Africa) during the early Pliensbachian (Early Jurassic).MethodsUsing 104 localities and 1,765 occurrences of ammonite species, we estimated geographic range sizes using the extent of occurrence and the latitudinal range. The phylogenetic conservatism of range sizes was tested using Moran's I index which measures phylogenetic autocorrelation, and Pagel's λ which indicates whether a phylogeny correctly predicts covariance patterns among taxa on a given trait according to a Brownian evolution model. We conducted these analyses for two neighbouring provinces with contrasting environmental features (Mediterranean and Northwest European). We also explored scale effects by considering the whole western Tethys and two temporal resolutions (chronozone and sub-stage).ResultsA marked difference in phylogenetic signal is observed between Mediterranean (MED) and Northwest European (NWE) species; the range size of MED species is more frequently phylogenetically conserved than that of NWE species. No phylogenetic conservatism of species range size is observed during the last chronozone of the early Pliensbachian which is characterized by numerous palaeoclimatic and palaeoenvironmental changes.Main conclusionsSpecies range size may be partly determined by phylogeny, but this phylogenetic conservatism is modulated by spatio-temporal environmental stability. The phylogenetic signal of species range size may be labile through time within the same lineage and may differ between contemporaneous species of the same group. This lability stems from the fact that species range size results from a complex interplay of intrinsic and extrinsic factors.

14 pages; International audience; The Early Jurassic was broadly a greenhouse climate period that was punctuated by short warm and cold climatic events, positive and negative excursions of carbon isotopes, and episodes of enhanced organic matter burial. Clay minerals from Pliensbachian sediments recovered from two boreholes in the Paris Basin, are used here as proxies of detrital supplies, runoff conditions, and palaeoceanographic changes. The combined use of these minerals with stable isotope data (C-O) from bulk carbonates and organic matter allows palaeoclimatic reconstructions to be refined for the Pliensbachian. Kaolinite/illite ratio is discussed as a reliable proxy of the hydrological cycle and runoff from landmasses. Three periods of enhanced runoff are recognised within the Pliensbachian. The first one at the Sinemurian-Pliensbachian transition shows a significant increase of kaolinite concomitant with the negative carbon isotope excursion at the so-called Sinemurian Pliensbachian Boundary Event (SPBE). The Early/Late Pliensbachian transition was also characterised by more humid conditions. This warm interval is associated with a major change in oceanic circulation during the Davoei Zone, likely triggered by sea-level rise; the newly created palaeogeography, notably the flooding of the London-Brabant Massif, allowed boreal detrital supplies, including kaolinite and chlorite, to be exported to the Paris Basin. The last event of enhanced runoff occurred during the late Pliensbachian (Subnodosus Subzone of the Margaritatus Zone), which occurred also during a warm period, favouring organic matter production and preservation. Our study highlights the major role of the London Brabant Massif in influencing oceanic circulation of the NW European area, as a topographic barrier (emerged lands) during periods of lowstand sea-level and its flooding during period of high sea-level. This massif was the unique source of smectite in the Paris Basin. Two episodes of smectite-rich sedimentation (‘smectite events’), coincide with regressive intervals, indicating emersion of the London Brabant Massif and thus suggesting that an amplitude of sea-level change high enough to be linked to glacio-eustasy. This mechanism is consistent with sedimentological and geochemical evidences of continental ice growth notably during the Latest Pliensbachian (Spinatum Zone), and possibly during the Early Pliensbachian (late Jamesoni/early Ibex Zones).

The evolution of mammalian dentition is constrained by functional necessity and by the non-independence of morphological structures. Efficient chewing implies coherent tooth coordination from development to motion, involving covariation patterns (integration) within dental parts. Using geometric morphometrics, we investigate the modular organization of the highly derived vole dentition. Integration patterns between and within the upper and lower molar rows are analyzed to identify potential modules and their origins (functional and developmental). Results support an integrated adult dentition pattern for both developmental and functional aspects. The integration patterns between opposing molar pairs suggest a transient role for the second upper and lower molars during the chewing motion. Upper and lower molar rows form coherent units but the relative integration of molar pairs is in contradiction with existing developmental models. Emphasis on the first three cusps to grow leads to a very different integration pattern, which would be congruent with developmental models. The early developmental architecture of traits is masked by later stages of growth, but may still be deciphered from the adult phenotype, if careful attention is paid to relevant features.

The characteristics of dissolved organic matter (DOM) in soils are often determined through laboratory experiments. Many different protocols can be used to extract organic matter from soil. In this study, we used five air-dried soils to compare three extraction methods for water-extractable organic matter (WEOM) as follows: (i) pressurised hot-water-extractable organic carbon (PH-WEOC), a percolation at high pressure and temperature; (ii) water-extractable organic carbon (WEOC), a 1-hour end-over shaking; and (iii) leaching-extractable organic carbon (LEOC), a leaching of soil columns at ambient conditions. We quantified the extraction yield of organic carbon; the quality of WEOM was characterized by UV absorbance, potential biodegradability (48-day incubation) and parallel factor analysis (PARAFAC) modelling of fluorescence excitation emission matrices (FEEMs). Biodegradation of dissolved organic carbon (DOC) was described by two pools of organic C. The proportions of labile and stable DOC pools differed only slightly between the WEOC and LEOC methods, while PH-WEOC contains more stable DOC. The mineralization rate constants of both labile and stable DOC pools were similar for the three methods. The FEEMs were decomposed into three components: two humic-like fluorophores and a tryptophan-like fluorophore. The effect of extraction method was poorly discriminant and the most similar procedures were PH-WEOC and LEOC while WEOC extracts were depleted in humic-like fluorophores. This study demonstrates that WEOM quality is primarily determined by soil characteristics and that the extraction method has a smaller, but still significant, impact on WEOM quality. Furthermore, we observed considerable interaction between extraction procedure and soil type, showing that method-induced differences in WEOM quality vary with soil characteristics.

The genusNeseuretusHicks, 1873 is the most abundant trilobite of the Ordovician siltite succession of the Andouillé and Traveusot Formations in the French Armorican massif. The systematics of some species ofNeseuretusis still unclear. Armorican and Iberian domains formed part of a distinctive paleobiogeographical province in the Ordovician and, while fiveNeseuretusspecies were defined in Iberia that follow each other through time, from the Middle to the Upper Ordovician, only oneNeseuretusspecies,N. tristani, has been identified in the Armorican massif so far. The discovery of new fossil deposits in the Ménez-Belair syncline has led to the identification and re-description of threeNeseuretustaxa:Neseuretus avusHamman, 1977 from the early-middle Darriwilian,Neseuretus tristani(Brongniart in Desmarest, 1817) from the late–middle Darriwilian to late Darriwilian andNeseuretus tardus(Hammann, 1983) from the Darriwilian–Sandbian boundary and early Sandbian. Morphological characters of the cephalon and pygidium were determined that can definitively distinguish the three identified taxa. These results stem from both traditional and geometric (outline and landmark-based) morphometric analyses. The three taxa follow each other through time and constitute a morphological sequence that contributes to improving the stratigraphy of the Middle and Upper Ordovician.

17 pages; International audience; In biogeography, the similarity distance decay (SDD) relationship refers to the decrease in compositional similarity between communities with geographical distance. Although representing one of the most widely used relationships in biogeography, a review of the literature reveals that: (1) SDD is influenced by both spatial extent and sample size; (2) the potential effect of the phylogenetic level has yet to be tested; (3) the effect of a marked biogeographical structuring upon SDD patterns is largely unknown; and (4) the SDD relationship is usually explored with modern, mainly terrestrial organisms, whereas fossil taxa are seldom used in that perspective. Using this relationship, we explore the long-distance dispersal of the Early Jurassic (early Pliensbachian, c. 190.8 Ma to 187.6 Ma) ammonites of the western Tethys and adjacent areas, in a context of marked provincialism. We show that the long-distance dispersal of these ammonites is not related to shell size and shape, but rather to the environmental characteristics of the province to which they belong. This suggests that their long-distance dispersal may have been essentially driven by passive planktonic drift during early juvenile, post-hatching stages. Furthermore, it seems that the SDD relationship is not always an appropriate method to characterize the existence of a biogeographical structuring. We conducted SDD analyses at various spatial, sampling and phylogenetic scales in order to evaluate their sensitivity to scale effects. This multi-scale approach indicates that the sampling scale may influence SDD rates in an unpredictable way and that the phylogenetic level has a major impact on SDD patterns.

11 pages; International audience; Campanian sediments from two French sedimentary basins were studied, using clay mineralogy and stable isotope (δ13C and δ18O) geochemistry, in order to investigate the Late Campanian Event. The clay fraction of the Campanian sediments from the Tercis-les-Bains section (Aquitaine Basin) and from the Poigny borehole (Paris Basin) is mainly composed of smectite. This background sedimentation was, however, interrupted during the Upper Campanian in the two basins by a substantial increase in detrital inputs, including illite, kaolinite, and chlorite at Tercis-les-Bains, and illite at Poigny. This detrital event, resulting from the enhanced erosion of nearby continental areas triggered by increasing runoff, has also been recognized in the Tethys and South Atlantic oceans. It coincided with a global negative carbon isotope excursion, the Late Campanian Event (LCE). Carbon isotope stratigraphy was used to correlate the two basins with previously studied sections from distant areas. Spectral analysis of the bulk δ13C from Tercis-les-Bains suggests a duration of ca. 400 kyr for a pre-LCE negative excursion and ca. 800–900 kyr for the LCE sensu stricto. The detrital event, as characterized by clay mineralogy, spans the interval that comprises the pre-LCE and the LCE, with a duration of 1.3 Myr. Intensification of continental erosion during the LCE may have resulted either from the Late Campanian polyplocum regression and/or from a regional tectonic pulse that triggered the emersion of previous submerged shelf areas and the increase of silicate erosion. As the LCE seems to be recorded at a large geographic scale, it is proposed here that enhanced chemical weathering and an associated decrease in atmospheric pCO2 levels could have contributed to the long-term Late Cretaceous cooling trend.

Describing patterns of connectivity among organs is essential for identifying anatomical homologies among taxa. It is also critical for revealing morphogenetic processes and the associated constraints that control the morphological diversification of clades. This is particularly relevant for studies of organisms with skeletons made of discrete elements such as arthropods, vertebrates, and echinoderms. Nonetheless, relatively few studies devoted to morphological disparity have considered connectivity patterns as a level of morphological organization or developed comparative frameworks with proper tools. Here, we analyze connectivity patterns among apical plates in Atelostomata, the most diversified clade among irregular echinoids. The clade comprises approximately 1600 fossil and Recent species (e.g., 25% of post-Paleozoic species of echinoids) and shows high levels of morphological disparity. Plate connectivity patterns were analyzed using tools and statistics of graph theory. To describe and explore the diversity of connectivity patterns among plates, we symbolized each pattern as a graph in which plates are coded as nodes that are connected pairwise by edges. We then generated a comparative framework as a morphospace of connections, in which the disparity of plate patterns observed in nature was mapped and analyzed. Main results show that apical plate patterns are both highly disparate between and within atelostomate groups and limited in number; overall, they also constitute small, compact, and simple structures compared to possible random patterns. Main traits of the evolution of apical plate patterns reveal the existence of strong morphogenetic constraints that are phylogenetically determined. In contrast, evolutionary radiations within atelostomates were accompanied by a clear increase in disparity, suggesting a release of some constraints at the origin of clades.

10 pages; International audience; For archaeologists, metallic artifacts are key materials to assess Middle Bronze Age production areas and cultural exchanges. Here, a set of 629 bronze palstaves excavated in northern France, belonging to Breton and Norman typological groups, was treated by (open) outline-based morphometrics with orthogonal polynomial regression. Using robust statistics developed for outlier detection, these Norman and Breton palstave outlines can be divided into two groups: those for which the shape fluctuates close to the standard shape, called "congruent" axes, and those which are far enough from this standard to be considered as "non-congruent", although they possess most of the features of the typological group. The highest density of discovery (whether congruent and non-congruent in shape) is in the extreme east of Brittany for the Breton axes, while the Norman axes are concentrated in northern Normandy, hence the choice of names. However, the distribution of congruent and non-congruent artifacts appears to be spatially dependent for the Norman group, and to a lesser extent for the Breton group, as there are proportionally more congruent specimens inside the supposed production areas than outside. This contradicts the generally accepted archaeological scheme which hypothesizes that all axes in a group originate from the same production center, and that some items were exported from there to supply neighboring regions. Other minor production centers probably existed, copying the original model with greater shape variation.

10 pages; International audience; Tooth number in rodents is an example of reduction in evolution. All rodents have a toothless diastema lacking canine and most premolars present in most other mammals. Whereas some rodent lineages retained one premolar (p4), many others lost it during evolution. Recently, an 'inhibitory cascade' developmental model (IC) has been used to predict how the first molar (m1) influences the number and relative sizes of the following distal molars (m2 and m3). The model does not, however, consider the presence of premolars, and here we examine whether the premolar could influence and constrain molar proportions during development and evolution. By investigating a large data set of both extinct and extant rodent families over more than 40 million years, we show that the basal phenotype is characterized by the presence of premolars together with equally sized molars. More recent rodent families, with and without premolar, show more unequal molar sizes. Analysing molar areas, we demonstrated that (i) rodents harbour almost all the molar proportions known in mammals, and the IC model can explain about 80% of taxa in our data set; (ii) proportions of molars are influenced by the presence or absence of p4; and (iii) the most variable teeth in the dental row are m1 and m3, whether p4 is present or not. Moreover, m1 can represent up to half of the total molar area when p4 is absent. We hypothesize that p4 loss during evolution released the constraint on m1 development, resulting in a more variable size of m1 and thereby having an indirect effect on the evolution of the whole molar row.

7 pages; International audience; Conservation biologists and palaeontologists are increasingly investigating the phylogenetic distribution of extinctions and its evolutionary consequences. However, the dearth of palaeontological studies on that subject and the lack of methodological consensus hamper our understanding of that major evolutionary phenomenon. Here we address this issue by (i) reviewing the approaches used to quantify the phylogenetic selectivity of extinctions and extinction risks; (ii) investigating with a high-resolution dataset whether extinctions and survivals were phylogenetically clustered among early Pliensbachian (Early Jurassic) ammonites; (iii) exploring the phylogenetic and temporal maintenance of this signal. We found that ammonite extinctions were significantly clumped phylogenetically, a pattern that prevailed throughout the 6.6 Myr-long early Pliensbachian interval. Such a phylogenetic conservatism did not alter - or may even have promoted - the evolutionary success of this major cephalopod clade. However, the comparison of phylogenetic autocorrelation among studies remains problematic because the notion of phylogenetic conservatism is scale-dependent and the intensity of the signal is sensitive to temporal resolution. We recommend a combined use of Moran's I, Pearson's ϕ and Fritz and Purvis' D statistics because they highlight different facets of the phylogenetic pattern of extinctions and/or survivals.

Understanding the factors that determine the distribution of taxa at various spatial scales is a crucial challenge in the context of global climate change. This holds particularly true for polar marine biota that are composed of both highly adapted and vulnerable faunas. We analysed the distribution of 2 Antarctic echinoid species, Sterechinus antarcticus and S. neumayeri, at the scale of the entire Southern Ocean using 2 niche modelling procedures. The performance of distribution models was tested with regard to the known ecology of the species. The respective contributions of environmental parameters are discussed along with the putative roles played by biotic interactions and biogeographic processes. Depth was the parameter that contributed most to both distribution models, whereas sea ice coverage and sea surface temperature had significant contributions for S. neumayeri only. Suitability maps of the 2 species were mostly similar, with a few notable differences. The Campbell Plateau and Tasmania were predicted as suitable areas for S. antarcticus only, while S. neumayeri was restricted to the south of the Antarctic Polar Front. However, numerous sampling data attest that S. antarcticus is absent from the Campbell Plateau and from Tasmania. Different hypotheses are formulated to explain the mismatch between observed and modelled distribution data. They stress the putative roles played by both oceanographic barriers to dispersal (Antarctic Polar Front), biotic factors (species exclusion patterns) and biogeographic processes (ongoing dispersal).

Developments of future scenarios of Antarctic ecosystems are still in their infancy, whilst predictions of the physical environment are recognized as being of global relevance and corresponding models are under continuous development. However, in the context of environmental change simulations of the future of the Antarctic biosphere are increasingly demanded by decision makers and the public, and are of fundamental scientific interest. This paper briefly reviews existing predictive models applied to Antarctic ecosystems before providing a conceptual framework for the further development of spatially and temporally explicit ecosystem models. The concept suggests how to improve approaches to relating species’ habitat description to the physical environment, for which a case study on sea urchins is presented. In addition, the concept integrates existing and new ideas to consider dynamic components, particularly information on the natural history of key species, from physiological experiments and biomolecular analyses. Thereby, we identify and critically discuss gaps in knowledge and methodological limitations. These refer to process understanding of biological complexity, the need for high spatial resolution oceanographic data from the entire water column, and the use of data from biomolecular analyses in support of such ecological approaches. Our goal is to motivate the research community to contribute data and knowledge to a holistic, Antarctic-specific, macroecological framework. Such a framework will facilitate the integration of theoretical and empirical work in Antarctica, improving our mechanistic understanding of this globally influential ecoregion, and supporting actions to secure this biodiversity hotspot and its ecosystem services.Keywords: Environmental change; integrative modelling framework; spatially and temporally explicit modelling macroecology; biodiversity; habitat suitability models(Published: 4 May 2012)Citation: Polar Research 2012, 31, 11091, http://dx.doi.org/10.3402/polar.v31i0.11091 To access the supplementary material to this article: Supplementary Tables S1, S2, please see Supplementary Files in the column to the right (under Article Tools).

4 pages; International audience; The tropical, warm, and equable climate of the Jurassic world is regularly challenged by geoscientists, especially since oxygen isotopes ( 18O) of fossil hardparts have been used to reconstruct the paleotemperature history of seawater. By applying the innovative “SiZer” (significant zero crossings of the derivatives) statistical approach to a newly compiled 18O database for the Jurassic, we demonstrate the occurrence of major and multiscale 18O changes mainly related to climate disturbances. For the first time, two long-term anomalies in 18O are identified during the Toarcian and the Late Jurassic, in conjunction with intensive volcanism in large igneous provinces. These results support a strong influence of repeated volcanic pulses on the modulation of pCO2, temperatures, and polar ice cap volumes over protracted periods. At shorter time scales, 13 relatively rapid (0.5–1 m.y.) and significant warming and cooling events are identified, the causes of which include transient fluctuations in greenhouse gas concentrations related to still-debated mechanisms.

10 pages; International audience; Disparity (shape diversity) is a key aspect of biodiversity, both present and past. The shapes of organisms are usually quantified by means of morphometrics. In this article, after a short review of recent advances and applications of morphometric methods, examples are presented as an overview of morphometric studies undertaken at the Biogéosciences research unit of Burgundy University. They concern both works on shape differentiation and evolution of disparity through time, and work aiming to infer, from the shapes of the organisms, any developmental stresses, constraints or processes which could explain in part the resulting disparity. The objective is to demonstrate how useful morphometrics can be for research in evolutionary and developmental biology.

The Pliensbachian–Toarcian interval was marked by major environmental disturbances and by a second-order mass extinction. Here, we reappraise the taxonomic, spatiotemporal and selective dynamics of extinctions over the whole interval, by analysing a high-resolution dataset of 772 ammonite species from NW Tethyan and Arctic domains. On average, 40–65% of ammonite species disappeared during each subchronozone, but higher extinction pulses (reaching 70–90%) prevailed from the Margaritatus to the Dispansum Chronozone. The main extinctions, corresponding to the Gibbosus, Pliensbachian–Toarcian boundary, Semicelatum, Bifrons–Variabilis, and Dispansum events, differed in their dynamics, suggesting episodes of ecological stress related to climate change, regression, disturbance in the carbon cycle or anoxia. The multi-pulsed volcanic activity in the Karoo–Ferrar province could well have triggered these ecological changes. In addition, ammonites experienced a morphological bottleneck during the Gibbosus event, 1 Ma before the Early Toarcian diversity collapse. Typically, drops in richness were related both to high extinctions and to declines in origination rates. This feature could result from strengthened ecological stresses related to the temporal overlap of environmental disturbances. After the Early Toarcian crisis, the recovery of ammonites was rapid (2 Ma) and probably influenced by a coeval marine transgression. Supplementary material: Figures showing a comparison of extinction and origination patterns based on different datasets, and variations of the morphospace occupation are available at http://www.geolsoc.org.uk/SUP18381.

14 pages.; International audience; The study of mammalian evolution is often based on insights into the evolution of teeth. Developmental studies may attempt to address the mechanisms that guide evolutionary changes. One example is the new developmental model proposed by Kavanagh et al. (2007), which provides a high-level testable model to predict mammalian tooth evolution. It is constructed on an inhibitory cascade model based on a dynamic balance of activators and inhibitors, regulating differences in molar size along the lower dental row. Nevertheless, molar sizes in some mammals differ from this inhibitory cascade model, in particular in voles. The aim of this study is to point out arvicoline and murine differences within this model and to suggest an alternative model. Here we demonstrate that the inhibitory cascade is not followed, due to the arvicoline's greatly elongated first lower molar. We broaden the scope of the macroevolutionary model by projecting a time scale on to the developmental model. We demonstrate that arvicoline evolution is rather characterized by a large gap from the oldest vole to more recent genera, with the rapid acquisition of a large first lower molar contemporaneous to their radiation. Our study provides alternative evolutionary hypotheses for mammals with different trajectories of development.

10 pages; International audience; Knowledge of mammalian tooth formation is increasing, through numerous genetic and developmental studies. The prevalence of teeth in fossil remains has led to an intensive description of evolutionary patterns within and among lineages based on tooth morphology. The extent to which developmental processes have influenced tooth morphologies and therefore the role of these processes in these evolutionary patterns are nonetheless challenging. Recent methodological advances have been proposed allowing the inference of developmental processes from adult morphologies and the characterization of the degree of developmental integration/modularity of morphological traits by studying the patterns of variation within and among individuals. This study focuses on the geometric shape of the lower molars of the vole species Microtus arvalis. Our results suggest (i) quasi-independence of each molar at the developmental level (developmental modules), even slightly stronger for the third molar supporting some genetic and developmental hypotheses and (ii) more pervasive integration processes among molars at the morphological level.

Most animals have a body plan that is bilaterally symmetric, at least externally. In some species, however, directional asymmetry (DA) occurs, where there is a preferential ‐ 32 ‐ direction of asymmetries, with one side that differs systematically from the other. Although it was first thought tat DA is relatively rare, subtle DA has been found in all studies that examined asymmetries with geometric morphometric techniques. Until now all the studies have looked at only one structure or at the comparison between modules. Here we analysed directional asymmetries of several different parts showing matching and object symmetries in Paussus favieri (Coleoptera, Carabidae, Paussini). Because beetles of the tribe Paussini are parasites of ants they are of great interest because of the striking myrmecophilous adaptations. Most of the species are extremely rare and localized, and their biology and systematics are still poorly known. We used geometric morphometric techniques to analyze the variation of antennae (both left and right), head, pronotum and abdomen in P. favieri (a relatively common species) from different European and Maghrebian localities. A clear and high evidence of a pattern of directional asymmetry emerged from the analyses in all parts examined. In particular principal component analysis revealed a pattern of variation associated with DA, higher in the antennae, but still present in all other body parts except of the abdomen. In all analyses differences between populations emerged as well. Finally, Procrustes ANOVA provides similar results. For size DA was statistically significant in all body parts except the abdomen. For shape DA was significant only in the antennae. From our analyses antennae have emerged as the part most under the DA, with a high shape and size variability. Moreover differences between populations emerged, with Moroccan population showing the highest differences. We conclude that multiple body parts have directional asymmetry, and the differences find between populations show that directional asymmetry is evolving.