Metrics details
freshwater crabs (Decapoda: Brachyura) are among the most diverse decapod crustaceans
their fossil record is extremely limited: only Potamidae
Potamonautidae and Trichodactylidae are reported up to the Eocene of the Neotropics so far
This work documents unusually large decapod claws from the Upper Cretaceous (Campanian) continental deposits of Velaux and vicinity (southern France)
in close association with large vertebrate remains
In addition to (1) the systematic assignment of these claws
the study addresses (2) the salinity trends in the deposit environment from its faunal assemblage and the elementary chemical patterns of fossils
and (3) the likely scenario for their auto/allochthony in the Velaux fluvial system
a group of “true” crabs nowadays linked to marine systems
the claw taphonomy and the carbonates Y/Ho signatures support their ancient freshwater/terrestrial ecology
making them the oldest reported continental brachyurans and extending the presence of crabs in freshwater environments by 40 Ma
Either as primary or as secondary freshwater crabs
the occurrence of these portunoids in Velaux is an evidence for the independent colonizations of continental environments by multiple brachyuran clades over time
The study aims at (1) proposing a systematic assignment for these claws
(2) characterizing the actual salinity trend of their depositional environment
based on the channel fauna assemblage and elementary chemical patterns of fossils and (3) identifying the relevant taphonomic scenario for the presence of crab claws within a fluvial system
As all these approaches support a freshwater or terrestrial signature for the paleoenvironment of these large-clawed brachyurans; we then discuss the implications for presumed multiple invasions of freshwater habitats by crustacean decapods over time
1) consists of seven (partial) claws and associated vertebrate remains collected from sequence 2 of the sedimentary succession of Velaux-La Bastide Neuve locality and one from the close locality of Rognac-Les Frégates (about four km from Velaux
Specimens are housed in the palaeontological collections of Velaux municipal institutions (Musée du Moulin Seigneurial/Velaux-La Bastide Neuve: MMS/VBN.00.004
12.A.003) and of the Muséum d'Histoire naturelle d'Aix-en-Provence
the Dinocarcinus material is not preserved sufficiently enough to allow detailed comparison of the cheliped dentition of various portunoid or xanthoid crabs
its taxonomic features (as mentioned above) point out affinities with portunoids
although this attribution has to be considered as preliminary
Dinocarcinus velauciensis is kept in open nomenclature
Portunoidea sensu lato (see above) Dinocarcinus n. gen. Van Bakel, Hyžný, Valentin & Robin Figs. 1, 2 and 3.
Dinocarcinus velauciensis Van Bakel, Hyžný, Valentin & Robin n. gen., n. sp. (a,b) Illustrative features of holotype MMS/VBN.00.004 (complete chela). (d) = dactylus; ff = fixed finger; pr = propodus; pr/d ar = propodus/dactylus articulation; t1-4 = dactylus teeth; T1-4 = fixed finger teeth;? = questionable limits. Scale bars = 1 cm. Drawing. B. van Bakel, photograph. L. Cazes.
Isolated claws found scattered in sediment
outer and occlusal views of dactyli showing teeth arrangement
marginal and occlusal two views showing teeth arrangement
Etymology: Denoting the actual association with dinosaur (ornithopodan) remains
Fixed finger dorsal surface with single “pitted groove”
the claw morphology of Dinocarcinus velauciensis n
does not match that of the Dakoticancroidea
are highly comparable with those of Dinocarcinus n
however they show a bulbous proximal crushing tooth on the dactylus of the major claw
The claw morphology of Dinocarcinus n. gen. shows few diagnostic characters for superfamily level assignment (Portunoidea), namely heavily calcified claws, a grooved fixed finger, molariform teeth, palm, and fingers subequal in length, and blunt, non-hooked fingertips (Fig. 2)
More accurate assignation is not possible at this point
An early diverging position within Portunoidea is possible considering morphology
and similar families occurring at that time
Dinocarcinus velauciensis Van Bakel, Hyžný, Valentin & Robin n. sp. Figures 1, 2 and 3
Type material: Holotype: MMS/VBN.00.004; Paratypes 1–4: MMS/VBN.02.94, 09.132d, 12.A.003, 12.A.006 (see Table 1)
Description: Only claws known; claw very large (approximately 85 mm for holotype MMS.VBN.00.004)
Upper (cutting) margin of fixed finger straight
Surface of molariform proximal tooth bulbous on fixed finger
Fixed finger dorsal surface with a single “pitted groove”
Dactylus upper and lower margin curved in dorsal view; dactylus cross-section flatter than propodus cross-section
Dactylus more strongly curved in dorsal view than fixed finger
Dactylus margin at approximately 100 degrees at upper (cutting) margin of fixed finger
cuticle surface (as preserved) covered with microscopic dense
has been extracted from a comparable block
as well as a partial titanosaurid dorsal vertebra showing preserved ossified tendons
Distribution of the taxa remains (n = 308)
recovered from the sequence 2 and associated described depositional environments (see III
Taxonomic/ecological diversity in the sequence 2 of the channel)
based on Velaux-La Bastide Neuve excavation campaigns
with Ho removed from seawater twice as fast as Y owing to differences in surface complexation behaviour
Y and Ho absolute concentrations (which we could expect to interpret from the studied material) depend on biological and taphonomic factors of integration that simply cannot be estimated here
the salinity of the studied channel cannot be assessed from the chemistry
the observed Y/Ho ratio formally excludes a marine/costal pattern and seems to distinguish strongly from it
Distribution of the fossil assemblage in the deposit of Velaux-La Bastide Neuve
(a) Lower portion of the Velaux log (Seq 0–4)
(b) Schematic spatial extension of the Velaux channel showing the distribution of bonebeds 2 (B2) and 3 (B3) in section
(c) Top view of the distribution of claws/bones on the excavation site
(d) Bloc assemblage of claws (small arrow) among vertebrate remains
the authors question the allochthonous marine origin of the claws
The consistent autochthony of all other invertebrates in the channel (Unio
and the Y/Ho carbonates displayed by claws
limit our interpretation to the most parsimonious assertion: it is rather unlikely that claws were moved into the Velaux fluvial system
Decapod crustaceans inhabit virtually all water-influenced habitats
from streams and rivers to ponds and lakes
representatives of a number of originally exclusively marine decapod clades have successfully invaded freshwater and/or terrestrial habitats
the following listing of freshwater decapods is not meant to be exhaustive but rather illustrative of the independent colonization of freshwater habitats within this group of crustaceans
All primary freshwater crabs are entirely free from the need to enter seawater
As for the fossils from Velaux and surroundings newly described herein
it cannot be decided unequivocally whether representatives of Dinocarcinus n
were able to complete their life cycle in the freshwater habitat or/and had direct development
The sheer size and number of claw fragments
may prove that crabs from the Late Cretaceous of Velaux were not only an occasional element of the respective environment
but rather a natural part of the assemblage suggesting that they were fully adapted to freshwater environment
so it is more likely they were primary freshwater crabs
whereas the age of the African Potamonautidae clade was given with 75–73 Ma (Campanian)
From the fossil record of the modern freshwater families alone
such timing cannot be apprehended; however
the discovery of fossil crabs from Velaux-La Bastide Neuve illustrates that brachyuran crabs attempted to colonize freshwater habitats in the Old World at least from the Campanian onwards
Based on the material from Velaux and Rognac described herein
we suggest that the freshwater habitats of islands in the Tethyan epicontinental sea were colonized by marine portunoids during the Late Cretaceous
most secondary freshwater brachyurans have a marine larval development and would reach inland habitats more likely as adults
This might also have been the case for Dinocarcinus velauciensis
from the late Campanian of Southern France
a group of “true crab” that are nowadays intimately linked to marine systems
faunal assemblage and taphonomy of these fossils
as well as the Y/Ho ratio of their carbonates indicate an ancient freshwater or terrestrial ecology
This makes them the oldest freshwater/terrestrial brachyurans ever reported
extending the existence of freshwater crabs by 40 Ma
“true” crabs were associated to terrestrial vertebrates
Although they were likely well adapted to this environment
it cannot be decided whether Dinocarcinus was able to complete its life cycle in the freshwater habitat or/and had direct development
Its occurrence in the Late Cretaceous of Velaux-La Bastide Neuve
is an evidence for the independent colonizations of freshwater environments by multiple Brachyura clades over time
beside that of modern primary freshwater crabs (Potamoidea
It also supports the molecular clock estimation of an Early Cretaceous start for the radiation of Afrotropical freshwater crab taxa (just appearing in the Late Cretaceous)
with the evidence of brachyuran crabs colonizing freshwater habitats as early as the Campanian
The elementary composition of a brachyuran claw and of its surrounding matrix were investigated for their Y/Ho ratios
One gram of each was sampled on MMS.VBN.09.69e (claw/sediment)
the basis of the propodus embedded in the matrix was mechanically sampled to preserve the connection between the claw and the turtle plate
Samples were microground and analysed for their composition in minor elements (in µg/g) normalized to PAAS
using ICMPS at the Service d'Analyses des Roches et des Minéraux of the CRPG
All data needed to evaluate the conclusions in the paper are present in the paper and supplementary information
In Freshwater animal diversity assessment 275–286 (Springer
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Authors want to acknowledge Clément Jauvion for discussions on elementary ratio in marine/non marine sediments
Turini for photographs of the specimen from Rognac
and communication assistance from the Velaux Municipality (J.-P
the Service Départemental d'Incendie et de Secours (SDIS) 13
and numerous volunteers during the field campaigns in 2009 and 2012
This work was supported in part by the French Ministry of Culture and Communication (research grant VR1013 to the Palaios Association) and the Bouches-du-Rhône department (CD 13) (proposals MAPADGAC23112010-1 and MAPADGAC16012014-1-AAPC)
We wish to thank Sean Murphy for English review of this work
and an anonymous reviewer for comments on manuscript improvements and discussion on claws systematic assignement
Centre de Recherche en Paléontologie – Paris (CR2P
Département Origines & Évolution (CP38)
Paléoécosystèmes et Paléoprimatologie (PALEVOPRIM
Royal Belgian Institute of Natural Sciences
provided the geological and taphonomic context of claws location
collected the fossils and provided data on faunal assemblage for Velaux deposit
supported material imaging and provided taxonomic review and comments
The authors declare no competing interests
Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations
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At their training base in the town of Velaux
gas-run metallic structures are used to teach firefighters to navigate through extreme heat
a setup threatened by rising fuel and environmental costs
By Antoine Albertini
Training maneuvers are carried out by a water bomber helicopter on the technical platform of the Velaux (Bouches-du-Rhône) fire department training center
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Artist’s impression of Matheronodon provincialis
Matheronodon provincialis was a primitive cousin of the well-known European dinosaur Iguanodon
The ancient beast lived 70 million years ago (Late Cretaceous epoch) and was approximately 16 feet (5 m) long
The fossilized jawbone and three teeth of the new species were discovered at the site of Velaux-La Bastide Neuve
“Matheronodon provincialis had extremely enlarged teeth
up to 2.4 inches (6 cm) long and 2 inches (5 cm) wide
They operated like self-sharpening serrated scissors,” said Dr
a paleontologist at Free University of Brussels
“Its teeth have ridged surfaces but are only covered with a thick enamel layer on one side.”
“Because the enamel is more resistant to wear than the exposed dentine
The fossilized jawbone of Matheronodon provincialis
“The denture of rhabdodontid dinosaurs (Rhabdodontidae) had evolved in a different direction than that of their contemporaries
the hadrosaurs or duck-billed dinosaurs,” said Dr
a paleontologist at the Royal Belgian Institute of Natural Sciences
“Hadrosaurs had sophisticated dental ‘batteries’ formed by little teeth with which they could crush conifers.”
“Matheronodon provincialis and other Rhabdodontidae probably ate leaves of palm trees
which were abundant in Europe at that time.”
“They had to cut rather than crush the fiber-rich leaves
Research describing the new species is published online in the journal Scientific Reports
article number: 13098; doi: 10.1038/s41598-017-13160-2
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She said: "Laurent only saw the good side of life
"He'd have lived somewhere where there were lots of other students
We didn't know it was a city where there are murders every day."
in her victim impact statement read to the Old Bailey
Mrs Bonomo described the devastating effect the murder had on her close-knit family
She wrote from the family home of Velaux-en-Provence in the south of France where
She describes every second of her anguish in the hours following the killing
Every time she tries to call him she reaches his answerphone
she postponed as long as possible the moment she had to tell us so that we wouldn't panic in vain."
Marie thought Laurent might have had the idea of making a spur-of-the-moment trip over
"He was quite capable of jumping on the Eurostar to give Marie a big surprise
they were in permanent contact via texts and conversations at all hours
"Then this silence became more and more incomprehensible and heavy."
hoping each time I'd hear his voice full of irony saying
you imagined me cut to pieces at the bottom of a ditch?'
The next day she got a call from the consulate
We found two people.' "Me: They're dead?' Yes
but we still don't know who they are.' "Me
in the middle of this suffocating unreality
My terrible fear that he might have disappeared into a unfathomable limbo was no longer there." At first she believed he died in the fire but learned the truth of what happened after reading news-papers on the internet the next day
Mrs Bonomo says that despite the tragedy she still has faith that thanks to British justice "clear light will be shone on the deaths of our children"
She says: "I'm not capable of hating those who have committed these terrible crimes
which made me feel the same thing every time I woke
Mr Ferez's mother described his killers as "monsters"
nurse Françoise Villemont said: "No one can carry out a massacre for two games consoles
Mr Bonomo had taken part in a civil union ceremony with fellow student Marie Bertez
who has now formed a bond with Mr Ferez's girlfriend Magali Bonnans
Ms Bertez described her beloved "Lolo" as "my world"
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