The development of X‐ray tomography in the last decade has led to a revolution in palaeontology by providing a means of imaging 3D fossils. In turn, imaging of flat fossils has strongly benefitted from critical improvement of synchrotron X‐ray fluorescence (XRF). The latter, which allows the mapping of 2D distributions of major‐to‐trace elements over decimetre‐scale objects, usually targets metals with atomic number (Z) up to strontium (K‐shell emission lines); in the same energy domain, L‐shell emission lines of heavier elements (particularly lead) can also be analysed. A fluorescence signal from strontium can escape from a depth of a few 100 μm in fossils, thereby revealing, due to its substitution for calcium in calcium phosphates (apatite group minerals and bone), hidden fossil bone or phosphatized remains. Nonetheless, strontium similarly substitutes for calcium in calcium carbonates, resulting in the absence of contrast when fossils are preserved in limestone. Here, we show that this issue can easily be overcome by using X‐rays slightly higher in energy (17.2 keV and above) to excite and detect yttrium fluorescence. Together with lanthanides (collectively known as the rare earth elements), yttrium preferentially substitutes for calcium in calcium phosphates, offering anatomical contrasts for a wider range of fossils. This is demonstrated here for three fossils (vertebrate and invertebrate) from different ages and depositional environments. We then discuss the different chemical behaviours of strontium and yttrium in calcium phosphates and carbonates. Although yttrium is also found in other (rarer) minerals, its mapping using synchrotron XRF could be used as a proxy to pinpoint calcium phosphates in fossils and other geological materials.
Polychelidan lobsters are fascinating crustaceans that were known as fossils before being discovered in the deep-sea. They differ from other crustaceans by having four to five pairs of claws. Although recent palaeontological studies have clarified the systematics and phylogeny of the group, the biology of extant polychelidans and--first of all--their anatomy are poorly documented. Numerous aspects of the evolutionary history of the group remain obscure, in particular, how and when polychelidans colonized the deep-sea and became restricted to it. Surprisingly, the biology of extant polychelidans and the anatomy of all species, fossil and recent, are poorly documented. Here, X-ray microtomography (XTM), applied to an exceptionally well-preserved specimen from the La Voulte Lagerstätte, reveals for the first time vital aspects of the external and internal morphology of Voulteryon parvulus (Eryonidae), a 165-million-year-old polychelidan: 1) its mouthparts (maxillae and maxillipeds), 2) its digestive tract and 3) its reproductive organs. Comparisons with dissected specimens clearly identify this specimen as a female with mature ovaries. This set of new information offers new insights into the feeding and reproductive habits of Mesozoic polychelidans. Contrasting with other Jurassic polychelidans that lived in shallow-water environments, V. parvulus spawned in, and probably inhabited, relatively deep-water environments, as do the survivors of the group.
Konservat-Lagerst€ atten are seen as snapshots of past biodiversity for a given location and time. However, processes leading to the exceptional morphological preservation of fossils in these deposits remain incompletely understood. This results in a deficient assessment of taphonomic biases and limits the robustness/relevance of palaeobiological reconstructions. Here, we report the mineralogical characterization of crustacean fossils preserved within carbonate-rich concretions from the Jurassic Konservat-Lagerst€ atte of La Voulte-sur-Rhône (Ard eche, France). The combination of SEM-EDS, TEM, synchrotron-based XRF, XRD and XANES allows the mineralogical phases composing these fossils (i.e. fluorapatite, Fe-sulfides (pyrite, pyrrhotite) and Mg-calcite) and the surrounding matrix (i.e. Mg-calcite, clays and detrital silicates) to be identified. Fluorapatite and pyrite (and pyrrhotite) precipitated during decay under anoxic conditions, replacing delicate organic structures and preserving anatomical details. These mineral structures were subsequently consolidated by a Mg-calcite cement. Of note, histologically similar tissues were replaced by the same mineral phases, confirming that fossilization (in La Voulte) occurred rapidly enough to be influenced by tissue composition. Altogether, the present study shows that exceptional preservation requires fast biodegradation, thereby confirming recent experimental evidence.
Exceptional and extremely rare preservation of soft parts, eyes, or syn-vivo associations provide crucial palaeoecological information on fossil-rich deposits. Here we present exceptionally preserved specimens of the polychelidan lobster Voulteryon parvulus , from the Jurassic of La Voulte-sur-Rhône Fossil-Lagerstätte, France, bearing eyes with hexagonal and square facets, ovaries, and a unique association with epibiont thecideoid brachiopods, giving insights onto the palaeoenvironment of this Lagerstätte. The eyes, mostly covered in hexagonal facets are interpreted as either apposition eyes (poorly adapted to low-light environment) or, less likely, as refractive or parabolic superposition eyes (compatible with dysphotic palaeoenvironments). The interpretation that V. parvulus had apposition eyes suggests an allochthonous, shallow water origin. However, the presence of thecideoid brachiopod ectosymbionts on its carapace, usually associated to dim-light paleoenvironments and/or rock crevices, suggests that V. parvulus lived in a dim-light setting. This would support the less parsimonious interpretation that V. parvulus had superposition eyes. If we accept the hypothesis that V. parvulus had apposition eyes, since the La Voulte palaeoenvironment is considered deep water and had a soft substrate, V. parvulus could have moved into the La Voulte Lagerstätte setting. If this is the case, La Voulte biota would record a combination of multiple palaeoenvironments.
Teredinidae are obligate xylophagous bivalves that colonize drift wood. They display a highly derived anatomy with a reduced shell; most of their body consists of soft tissues. Consequently, fossil teredinids mostly correspond to burrows, shells or small terminal aragonite structures called ‘pallets’. We report, from mid‐Cretaceous logs of the Envigne Valley, France, exceptionally preserved wood‐boring bivalves with silicified soft parts. After characterizing the wood, we report both the molluscs’ anatomy and their distribution inside the wood (using computed tomography). The 3D‐reconstructions reveal rarely preserved soft tissues (mantle, respiratory siphons, visceral pouch) but surprisingly no mineralized pallets. Envigne shipworms display neat dorsal condyles and a vermiform body plan, making them the oldest known Teredinidae. To document the evolutionary or taphonomic origin of this lack of pallet, the mineralization of these bivalves was investigated, and it is concluded that pallets could have dissolved prior to other carbonate components. From a survey of the fossil wood‐boring bivalves and their characters, we provide a time scaled origin of the main pholadoid clades supported in recent phylogenies. Since we found no correlation between the presence of pallets in Pholadoidea and their occurrence in the stratigraphic record, we suggest that their absence during the Mesozoic could be the result of a taphonomic bias related to the ‘calcite vs aragonite seas’ paradigm. The Cenomanian Envigne Valley was an estuarine to intertidal environment in which the Teredinidae appear to have selected their wood habitat on the basis of size but not type.
Crustacean eggs are rare in the fossil record. Here we report the exquisite preservation of a fossil polychelidan embedded within an unbroken nodule from the Middle Jurassic La Voulte-sur-Rhône Lagerstätte (France) and found with hundreds of eggs attached to the pleon. This specimen belongs to a new species, Palaeopolycheles nantosueltae sp. nov. and offers unique clues to discuss the evolution of brooding behaviour in polychelidan lobsters. In contrast to their development, which now relies on a long-lived planktic larval stage that probably did not exist in the early evolutionary steps of the group, the brood size of polychelidan lobsters seems to have remained unchanged and comparatively small since the Jurassic. This finding is at odds with reproductive strategies in other lobster groups, in which a long-lived planktic larval stage is associated with a large brood size.
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