Jérémie Bardin

Pierre and Marie Curie University - Paris 6 | UPMC · Centre de Recherche sur la Paléobiodiversité et les Paléoenvironnements (CR2P)

Accurate modeling of the complexity of morphological evolution is crucial for morphological phylogenetics and for performing tests on a wide variety of evolutionary scenarios. In this context, morphological integration and the problem of correlated categorical characters represent a major challenge. In particular, the magnitude and implications of correlations among serially homologous structures such as teeth have been much debated but were never tested statistically within a broad phylogenetic context. Here, we present a large-scale empirical study analyzing the serial variation of cingular crests on successive molars (M1, M2 and M3) of 274 placental species in a phylogenetic context. Both likelihood analyses and analysis of phylogenetic co-distributions demonstrated highly correlated evolution in the entire sample and thus the non-independence of these serial features at a macroevolutionary scale. Likelihood analyses show that their serial variation should be better scored within a single composite character model with constrained paths for transitions enabling simultaneous changes on all three molars, which suggests a strong developmental or genetic integration. These results are congruent with current molecular and developmental knowledge related to dental morphological variation and call into question the frequent use of separate characters scored on serially homologous structures of the dentition in phylogenetic analyses. Overall, they provide long-overdue and clear empirical evidence that in-depth studies of patterns of integration constitute an essential step towards more realistic character construction and modeling. This approach is critical for more accurate morphological phylogenetics and, more generally, for testing macroevolutionary scenarios on groups of correlated characters.

Ammonite phylogeny has mainly been established based on a stratigraphic approach, with cladistics underconsidered. The main arguments against the use of cladistics are the supposed large amount of homoplasy and the small number of characters. Resolving the phylogeny of the Hildoceratidae (Early Jurassic) is especially challenging because of its large diversity and disparity. Many forms that have not been determined as closely related in previous studies exhibit very similar shapes. Moreover, some groups are morphologically very different, adding difficulties to building a unique coding scheme at a low taxonomic resolution (i.e. species). Here we propose an integrated coding scheme of the peristome shape and the ornamentation, allowing an increased level of comparison. The shape of the peristome is used as a new reference to locate ornamental features and propose new homology hypotheses. In total, 105 taxa have been analysed for 47 characters. We code continuous characters by their means and ranges ± one standard deviation. We test two weighting schemes: equal weights standardized by unit range and implied weighting with several concavity constants. This work has led to redefinition of the phylogenetic inclusivenesses of all the hildoceratid subfamilies. The new coding scheme based on peristome shapes provides the fewest homoplastic characters. The schemes appear promising to improve phylogenetic analyses in ammonoids as well as molluscs as a whole by creating a general coding framework.

Palaeontological data are key elements for inferring ancestral character states and the assembly of character complexes, but cephalopod fossils preserving soft tissues are very rare. The exceptionally well-preserved, unique specimen of Jurassic Proteroctopus ribeti Fischer & Riou from the Lagerstätte of La-Voulte-sur-Rhône (c. 165 Ma, France) is one of the few fossil octopod related taxa, but is rarely considered in evolutionary studies. In this paper, we used synchrotron microtomography to reappraise its external characters and for the first time, to reveal its internal structures. A unique character association is found with two fins, head fused to the body, eight well-developed arms with cirri and two rows of oblique suckers, a gladius and absence of an ink sac. The phylogenetic analysis indicates that Proteroctopus is a basal member of the Vampyropoda. However, this result should be interpreted with caution due to the number of unknown character states in the matrix. Contrary to previous assumptions, the phylogenetic position of Proteroctopus, as well as its stratigraphic occurrence, suggest that the arrangement of biserial suckers may be the ancestral condition in Vampyropoda.

Ontogeny is rarely included in phylogenetic analyses of morphological data. When used, the ontogenetic information is reduced to one character for two or three different ontogenetic stages. Several examples show that current methods miss a major part of the information. We here propose a new method for including the ontogenetic dimension in coding schemes of phylogenetic analyses. Our goal was to maximise the phylogenetic information extracted from ontogenetic trajectories. For discrete features, we recommend including precise timings of transformation(s) from one state to another in the ontogenetic trajectories. For continuously varying features, growth laws are modelled on raw data using least-square regressions. Then, parameters of models are included in the coding scheme as continuous characters. This method is employed to reconstruct phylogenetic relationships using the ammonite family Amaltheidae as a test subject. Based on the same data set, a second analysis has been performed only for characters of the adult stage. Comparisons of retention index, bootstrap support and stratigraphic congruence between the two analyses show that the inclusion of ontogeny yields better phylogenetic reconstruction. Morphological traits in ammonites which are usually the most homoplastic show a better fit to most parsimonious trees by including the ontogenetic dimension. In several cases, growth rates and patterns of growth have better fit to phylogeny than adult shapes, implying that paths of ontogeny can be more relevant than its products.

Among some modern and recent fossil species of planktonic foraminifera, the proportion of left- to right-coiled shells in a population appears to be temperature-dependent; the relative abundance of each morphotype reflecting ecological preferences. A similar relationship is identified among Muricohedbergella delrioensis (Carsey, 1926) at the Cenomanian-Turonian stage boundary in mid-latitude sites of the Western Interior Seaway, including the Pueblo type section. The increase of sinistral M. delrioensis in the assemblage is related to higher δ18O carb values and decrease in inner porosity which suggests that changes in the coiling direction in this morphospecies could represent a new proxy for constraining Sea Surface Temperature (SST) variations. As surface dwellers, muricohedbergellids were not affected by Oceanic Anoxic Event (OAE2) and provide a continuous paleoclimatic signal throughout the Cenomanian-Turonian boundary interval. Furthermore, genetic evidence obtained from extant foraminifera indicates that shifts of coiling ratios in planktonic foraminifera species can express the signature of distinct genetic types, which are revealed through their opposite coiling directions. Coiling direction could be a genetic trait, implying that cryptic species may occur in the Mesozoic.

Ammonite phylogeny has mainly been established based on a stratigraphic approach, with cladistics underconsidered. The main arguments against the use of cladistics are the supposed large amount of homoplasy and the small number of characters. Resolving the phylogeny of the Hildoceratidae (Early Jurassic) is especially challenging because of its large diversity and disparity. Many forms that have not been determined as closely related in previous studies exhibit very similar shapes. Moreover, some groups are morphologically very different, adding difficulties to building a unique coding scheme at a low taxonomic resolution (i.e. species). Here we propose an integrated coding scheme of the peristome shape and the ornamentation, allowing an increased level of comparison. The shape of the peristome is used as a new reference to locate ornamental features and propose new homology hypotheses. In total, 105 taxa have been analysed for 47 characters. We code continuous characters by their means and ranges ± one standard deviation. We test two weighting schemes: equal weights standardized by unit range and implied weighting with several concavity constants. This work has led to redefinition of the phylogenetic inclusivenesses of all the hildoceratid subfamilies. The new coding scheme based on peristome shapes provides the fewest homoplastic characters. The schemes appear promising to improve phylogenetic analyses in ammonoids as well as molluscs as a whole by creating a general coding framework.

Cladistics appears as one of the most useful method to reconstruct phylogeny of fossil taxa. However, ammonoids workers tend to sulk this method. The capital step of cladistic analysis is the recognition of homology hypothesis as clue to reconstruct monophyletic clades based on the sharing of derived traits. Previous authors have suggested that coding schemes are usually direct transcription of original taxa description. However, establishing a list of characters (i.e. a matrix taxa/characters) is a very different work compared to a compilation of diagnoses. How morphology is coded in ammo- noids? How coding schemes are influenced by traditional descriptions/characters? Here, we review all cladistic analyses of ammonoids published in the literature to compare characters and the way authors have dealt with the treatment of continuous characters, polymorphism and ontogeny. Several barriers are usually invoked to justify that cladistics cannot be applied to reconstruct ammonoids phylogenies. We show that an appropriate use of improvements both on ammonoids' knowledge and cladistics methodology may overcome limitations usually invoked to perform cladistic analysis on ammonoids.

A rich ammonite fauna from the Lower Toarcian (Lower Jurassic) of the South Riffian ridges (northern Morocco), collected bed-by-bed from 14 new outcrops, is described. The fauna is nearly exclusively composed of already-known taxa, allowing correlation from the chronozone down to the zonule level. Fourteen species are described, among them the new species Dactylioceras (Dactylioceras) laticostatum sp. nov. The stratigraphical interval studied spans the first chronozone of the Toarcian, the Polymorphum Chronozone, but it remains unclear whether the Toarcian Oceanic Anoxic Event is represented in the studied succession. The Early Toarcian is mainly characterized by representatives of the family Dactylioceratidae, which are abundant but commonly crushed or deformed. Two new descriptors are proposed to describe the shell shape of crushed or incomplete specimens and have been used together with the classical ones for the taxonomic treatment of this group. A hypothesis about dimorphism in Dactylioceras (Orthodactylites) semicelatum (Simpson, 1843) and Dactylioceras (Dactylioceras) laticostatum sp. nov. is proposed.

Abstract: A belemnite fauna collected in the lowermost Toarcian succession which crops out near Moulay Idriss (northern Morocco) is studied in this paper. This is the first palaeontological study of Early Toarcian belemnites from Northern Africa, i.e. the north-eastern margin of the Gondwana, in connexion with Tethys. Four species of the family Passaloteuthidae Naef, 1922 have been identified: Passaloteuthis bisulcata (de Blainville, 1827), Pseudohastites longiformis (Blake, 1876), Parapassaloteuthis zieteni (Mayer-Eymar, 1884) and Parapassaloteuthis sp. A. All species have been collected in lowermost Toarcian beds dated to the ammonite Polymorphum Chronozone (=Tenuicostatum Chronozone), which coincides with the belemnite Passaloteuthis bisulcata biozone. The discovery of a syntype of this zonal index is discussed. The identified species are common with Europe, thus suggesting that the onset of the belemnite provincialism in the Toarcian could post-date the earliest Toarcian Polymorphum – Tenuicostatum Chronozone. However, records of Early Jurassic belemnites are still too sparse to recognize the establishment of provincialism and the timing of its onset. Key words: belemnites, systematics, biostratigraphy, biogeography, Jurassic, Toarcian, Morocco

Since the introduction of the cladistic method in systematics, continuous characters have been integrated into analyses but no methods for their treatment have received unanimous support. Some methods require a large number of character states to discretise continuous characters in order to keep the maximum level of information about taxa differences within the coding scheme. Our objective was to assess the impact of increasing character state number on the outcomes of phylogenetic analyses. Analysis of a variety of simulated data sets shows that these methods for coding continuous characters can lead to the generation of well-resolved trees that do not reflect phylogenetic signal. We call this phenomenon the flattening of the tree-length distribution; it is influenced by both the relative quantity of continuous characters in relation to discrete characters, and the number of characters in relation to the number of taxa. Bootstrap tests provide one method to avoid this potential bias.

The present paper deals with the systematic description of a rich and well-preserved ammonite fauna of the Late Pliensbachian, collected in two outcrops near the "Lac de Charmes" (Haute-Marne, France). In terms of ammonite zonation, the studied faunas indicate the Margaritatus and Spinatum Chronozones. Most of their subdivisions (subchronozones and zonules) have been recognized for the first time in this region. We propose a correlation with zonations established in close areas (Germany, Southern France). As already suggested on the litera-ture, the possibility to further subdivide the Solare zonule in finner intervals is confirmed by our data. Within this zonule, we recognize a first interval with only Pleuroceras solare (Phillips, 1829), and a second one where this species oc-curs with several subspecies and where Pleuroceras spinatum (Bruguière, 1789) appears. Our results also confirm the possibility to subdivide more finely the Solare zonule firstly suggested by some authors. We recognize a first part with only Pleuroceras solare and a second one where this species present various sub-species and where Pleuroceras spinatum appears. Some points of view are given about Amaltheus gr. margaritatus Montfort, 1808, Pleuroceras gr. solare (Phillips, 1829) and the species Pleuroceras transiens (Frentzen, 1934).