Sylvain Charbonnier

Ecological Significance of the Arthropod Fauna from the Jurassic (Callovian) La Voulte Lagerstätte

The La Voulte Lagerstätte is remarkable for its unique soft-bodied fauna (e.g., worms, coleoid squids) and its exceptionally preserved arthropods mainly found in small sideritic concretions. This arthropod fauna includes 30 different species assigned to the crustaceans, the thylacocephalans and the pycnogonids. Crustaceans are the most diversified group with 23 species distributed in a dozen families. Quantitative analyses based on 388 nodules reveals four dominant groups: (i) the enigmatic thylacocephalan arthropods (33%), (ii) the Solenoceridae shrimps (22%), (iii) the Coleiidae crustaceans (15%), and (iv) the Penaeidae shrimps (10%). Converging lines of evidence from depositional environment and modern analogues, indicate that this arthropod fauna probably inhabited a deep water setting most probably exceeding 200 m (= bathyal zone) under dysphotic or aphotic conditions. This new set of data sheds new light on the deep-sea colonisation by animal communities in the Mesozoic.

Tomographic reconstruction of neopterous Carboniferous insect nymphs

Two new polyneopteran insect nymphs from the Montceau-les-Mines Lagerstätte of France are presented. Both are preserved in three dimensions, and are imaged with the aid of X-ray micro-tomography, allowing their morphology to be recovered in unprecedented detail. One–Anebos phrixos gen. et sp. nov.–is of uncertain affinities, and preserves portions of the antennae and eyes, coupled with a heavily spined habitus. The other is a roachoid with long antennae and chewing mouthparts very similar in form to the most generalized mandibulate mouthparts of extant orthopteroid insects. Computer reconstructions reveal limbs in both specimens, allowing identification of the segments and annulation in the tarsus, while poorly developed thoracic wing pads suggest both are young instars. This work describes the morphologically best-known Palaeozoic insect nymphs, allowing a better understanding of the juveniles’ palaeobiology and palaeoecology. We also consider the validity of evidence from Palaeozoic juvenile insects in wing origin theories. The study of juvenile Palaeozoic insects is currently a neglected field, yet these fossils provide direct evidence on the evolution of insect development. It is hoped this study will stimulate a renewed interest in such work.

Unexpected Early Triassic marine ecosystem and the rise of the Modern evolutionary fauna.

A new marine fossil assemblage from the Early Triassic shows unexpected phyletic diversity and functional complexity. In the wake of the end-Permian mass extinction, the Early Triassic (~251.9 to 247 million years ago) is portrayed as an environmentally unstable interval characterized by several biotic crises and heavily depauperate marine benthic ecosystems. We describe a new fossil assemblage—the Paris Biota—from the earliest Spathian (middle Olenekian, ~250.6 million years ago) of the Bear Lake area, southeastern Idaho, USA. This highly diversified assemblage documents a remarkably complex marine ecosystem including at least seven phyla and 20 distinct metazoan orders, along with algae. Most unexpectedly, it combines early Paleozoic and middle Mesozoic taxa previously unknown from the Triassic strata, among which are primitive Cambrian-Ordovician leptomitid sponges (a 200–million year Lazarus taxon) and gladius-bearing coleoid cephalopods, a poorly documented group before the Jurassic (~50 million years after the Early Triassic). Additionally, the crinoid and ophiuroid specimens show derived anatomical characters that were thought to have evolved much later. Unlike previous works that suggested a sluggish postcrisis recovery and a low diversity for the Early Triassic benthic organisms, the unexpected composition of this exceptional assemblage points toward an early and rapid post-Permian diversification for these clades. Overall, it illustrates a phylogenetically diverse, functionally complex, and trophically multileveled marine ecosystem, from primary producers up to top predators and potential scavengers. Hence, the Paris Biota highlights the key evolutionary position of Early Triassic fossil ecosystems in the transition from the Paleozoic to the Modern marine evolutionary fauna at the dawn of the Mesozoic era.

A 365-Million-Year-Old Freshwater Community Reveals Morphological and Ecological Stasis in Branchiopod Crustaceans

Branchiopod crustaceans are represented by fairy, tadpole, and clam shrimps (Anostraca, Notostraca, Laevicaudata, Spinicaudata), which typically inhabit temporary freshwater bodies, and water fleas (Cladoceromorpha), which live in all kinds of freshwater and occasionally marine environments [1, 2]. The earliest branchiopods occur in the Cambrian, where they are represented by complete body fossils from Sweden such as Rehbachiella kinnekullensis [3] and isolated mandibles preserved as small carbonaceous fossils [4-6] from Canada. The earliest known continental branchiopods are associated with hot spring environments [7] represented by the Early Devonian Rhynie Chert of Scotland (410 million years ago) and include possible stem-group or crown-group Anostraca, Notostraca, and clam shrimps or Cladoceromorpha [8-10], which differ morphologically from their modern counterparts [1, 2, 11]. Here we report the discovery of an ephemeral pool branchiopod community from the 365-million-year-old Strud locality of Belgium. It is characterized by new anostracans and spinicaudatans, closely resembling extant species, and the earliest notostracan, Strudops goldenbergi [12]. These branchiopods released resting eggs into the sediment in a manner similar to their modern representatives [1, 2]. We infer that this reproductive strategy was critical to overcoming environmental constraints such as seasonal desiccation imposed by living on land. The pioneer colonization of ephemeral freshwater pools by branchiopods in the Devonian was followed by remarkable ecological and morphological stasis that persists to the present day.

Diversity and Paleoenvironment of the Flora From the Nodules of the Montceau-Les-Mines Biota (Late Carboniferous, France)

Abstract The flora from the Montceau-les-Mines Lagerstätte (Massif Central, France, Late Pennsylvanian) is preserved in small sideritic concretions and was studied in three locations in the (1) Saint-Louis, (2) Saint-François, and (3) Sainte-Hélène opencast mines. Qualitative and quantitative analyses of plant diversity and floristic composition in 6812 nodules indicate substantial variations in the floral composition of these opencast mines. More than 50 taxa are recognized and belong to groups typical of the Late Pennsylvanian flora (lycopsids, sphenopsids, tree ferns, and pteridosperms). Arborescent sphenopsids and tree ferns were the major components at Saint-Louis, whereas the flora from Saint-François consisted mainly of pteridosperms; the one from Sainte-Hélène has a more balanced composition. Taphonomic and sedimentological data show that the flora contained in the nodules was hypoautochthonous to parautochthonous. The Montceau Basin displayed a mosaic of paleoenvironments (e.g., deltaic lacustrine, paludal to fluvial) which favored colonization by plants and animals.

Phylogeny of fossil and extant glypheid and litogastrid lobsters (Crustacea, Decapoda) as revealed by morphological characters

A phylogenetic analysis of a total of 31 species: 27 fossil species from seven families (Glypheidae, Litogastridae, Mecochiridae, Pemphicidae, Erymidae, Clytiopsidae, Chimaerastacidae), and four extant species from three families (Glypheidae, Nephropidae, Stenopodidae) is proposed. Most of the genera considered are coded exclusively based upon their type species and, as much as possible, based upon the type specimens. The cladistic analysis demonstrates that the glypheidean lobsters (infraorder Glypheidea) form a monophyletic group including two superfamilies: Glypheoidea and Pemphicoidea new status. Glypheoidea includes three families: Glypheidae, Mecochiridae and Litogastridae. Litogastridae is the sister group of the clade Glypheidae + Mecochiridae. Pemphicoidea includes a single family: Pemphicidae. A new classification of Glypheidea is proposed and currently known genera are rearranged based upon the phylogenetic analysis.

Sea Stars from Middle Jurassic Lagerstätte of La Voulte-sur-Rhône (Ardèche, France)

Abstract The Callovian Lagerstätte of La Voulte-sur-Rhône represents deep (bathyal) environments at the bottom of a submarine rocky slope. Three sea star body fossils partially replaced with pyrite were collected from clay beds. A small, stellate specimen is assigned to Terminaster cancriformis, a species already known from Jurassic clay deposits of Germany, Switzerland and England. The morphology of Terminaster illustrates mostly plesiomorphic features for post-Paleozoic sea stars, and the genus may be considered either as a paedomorphic Zoroasteridae or a stem group Forcipulatacea. Two specimens of a large multiarmed sea star are assigned to Decacuminaster n. gen. solaris n. sp., characterized by a lack of abactinal plates and an axial skeleton typical of the Velatida. The two genera Xandarodaster and Plesiastropecten are close to Decacuminaster and must be transferred to the Velatida. In modern oceans, the association of a multiarmed Velatida and a zoroasterid-like species would be typical of bathyal/abyssal environments, which suggests that these groups explored deep sea environments early in their history.

New sea spiders from the Jurassic La Voulte-sur-Rhône Lagerstätte

The diverse and exceptionally well-preserved pycnogonids described herein from the Middle Jurassic La Voulte Lagerstätte fill a 400 Myr gap of knowledge in the evolutionary history of this enigmatic group of marine arthropods. They reveal very close morphological and functional (locomotion, feeding) similarities with present-day pycnogonids and, by contrast, marked differences with all Palaeozoic representatives of the group. This suggests a relatively recent, possibly Mesozoic origin for at least three major extant lineages of pycnogonids (Ammotheidae, Colossendeidae, Endeidae). Combined evidence from depositional environment, faunal associates and recent analogues indicate that the La Voulte pycnogonids probably lived in the upper bathyal zone (ca 200 m). Our results point to a remarkable morphological and ecological stability of this arthropod group over at least 160 Myr and suggest that the colonization of the deep sea by pycnogonids occurred before the Jurassic.

Exceptional preservation of eye structure in arthropod visual predators from the Middle Jurassic

Vision has revolutionized the way animals explore their environment and interact with each other and rapidly became a major driving force in animal evolution. However, direct evidence of how ancient animals could perceive their environment is extremely difficult to obtain because internal eye structures are almost never fossilized. Here, we reconstruct with unprecedented resolution the three-dimensional structure of the huge compound eye of a 160-million-year-old thylacocephalan arthropod from the La Voulte exceptional fossil biota in SE France. This arthropod had about 18,000 lenses on each eye, which is a record among extinct and extant arthropods and is surpassed only by modern dragonflies. Combined information about its eyes, internal organs and gut contents obtained by X-ray microtomography lead to the conclusion that this thylacocephalan arthropod was a visual hunter probably adapted to illuminated environments, thus contradicting the hypothesis that La Voulte was a deep-water environment.

Trace elemental imaging of rare earth elements discriminates tissues at microscale in flat fossils.

The interpretation of flattened fossils remains a major challenge due to compression of their complex anatomies during fossilization, making critical anatomical features invisible or hardly discernible. Key features are often hidden under greatly preserved decay prone tissues, or an unpreparable sedimentary matrix. A method offering access to such anatomical features is of paramount interest to resolve taxonomic affinities and to study fossils after a least possible invasive preparation. Unfortunately, the widely-used X-ray micro-computed tomography, for visualizing hidden or internal structures of a broad range of fossils, is generally inapplicable to flattened specimens, due to the very high differential absorbance in distinct directions. Here we show that synchrotron X-ray fluorescence spectral raster-scanning coupled to spectral decomposition or a much faster Kullback-Leibler divergence based statistical analysis provides microscale visualization of tissues. We imaged exceptionally well-preserved fossils from the Late Cretaceous without needing any prior delicate preparation. The contrasting elemental distributions greatly improved the discrimination of skeletal elements material from both the sedimentary matrix and fossilized soft tissues. Aside content in alkaline earth elements and phosphorus, a critical parameter for tissue discrimination is the distinct amounts of rare earth elements. Local quantification of rare earths may open new avenues for fossil description but also in paleoenvironmental and taphonomical studies.