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Bernissart and the iguanodons: Historical perspective and new investigations

Authors:
2
1.1. The Sainte-Barbe pit and mine buildings in
1878, at the time when the iguanodons were
discovered.
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Bernissart and the Iguanodons:
Historical Perspective and New Investigations
Pascal Godefroit*, Johan Yans, and Pierre Bultynck
The discovery of complete and articulated skeletons of Iguanodon at Bernis-
sart in  came at a time when the anatomy of dinosaurs was still poorly
understood, and thus considerable advances were made possible. Here we
briey describe, mainly from documents in the archives of the Royal Bel-
gian Institute of Natural Sciences, the circumstances of the discovery of the
Bernissart iguanodons. We also provide information about their prepara-
tion and mounting in laboratories, for exhibitions, and in early studies. We
also summarize the latest results of a multidisciplinary project dedicated to
the material collected in the cores drilled in – in and around the
Iguanodon Sinkhole at Bernissart.
The discovery of the rst Iguanodon fossils has become a legend in the
small world of paleontology. Around , Mary Ann Mantell accompanied
her husband, the physician Dr. Gideon Algernon Mantell, on his medical
rounds and by chance discovered large fossilized teeth. Her husband found
the teeth intriguing. With advice from Georges Cuvier, William Clift, and
William Daniel Conybeare, he described them and named them Iguan-
odon, “iguana tooth,” because of their supercial resemblance to those of
living iguanas (Mantell, ). Iguanodon was one the three founding mem-
bers of the Dinosauria—along with Megalosaurus and Hylaeosaurus
named by Richard Owen in .
For  years, little was known about Iguanodon and other dinosaurs.
Mantell imagined these antediluvian animals to be some kind of giant
lizards with elongated bodies and sprawling limbs (Benton, ). In ,
the sculptor Waterhouse Hawkins, following Owen’s advice, realized full-
size reconstructions of Iguanodon and Megalosaurus for the Crystal Palace
exhibition in London. Iguanodon was reconstructed as a rhinoceros-like
heavy quadruped with a large spike on its nose. These impressive monsters
invoked the rst public sensation over dinosaurs (Norman, ).
The rst partial dinosaur skeleton, named Hadrosaurus foulkii Leidy,
, was discovered in  in New Jersey. This skeleton was reconstructed
in a bipedal gait at the Academy of Natural Sciences of Philadelphia, but
many questions were still left unanswered about the general appearance
of dinosaurs.
Then,  years later, another Iguanodon discovery broke the scientic
world—and the dinosaur world—wide open (Forster, ). The discovery
of complete and articulated skeletons of Iguanodon at Bernissart in 
revealed for the rst time the anatomy of dinosaurs, and thus consider-
able advances were made possible, in combination with the remarkable
The Bernissart Iguanodons:
A Cornerstone in the
History of Paleontology
1
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Godefroit et al.
4
discoveries in the American Midwest described by Marsh and Cope
(Norman, ).
Many manuscripts and plans relating to the original excavations at
Bernissart are preserved in the paleontological archives of the , which
allow us to reconstruct the circumstances of the discovery of these fantastic
dinosaurs.
Institutional abbreviations. , The Natural History Museum,
London (formerly the British Museum [Natural History]), U.K.; ,
Royal Belgian Institute of Natural Sciences, Brussels (formerly ,
Musée royal d’Histoire naturelle de Belgique), Belgium.
Bernissart is a former coal-mining village in southwestern Belgium, situated
 km south of Mons and less than  km from the Franco-Belgian frontier. Pre-
industrial coal extraction began at Bernissart around  (Delguste, ).
In , Duke Emmanuel de Croÿ grouped together the different coal com-
panies in northern France into the powerful Anzin Company, which started
the industrial exploitation of the coal in the Bernissart area during the second
half of the eighteenth century (Delguste, ). In the nineteenth century,
the Bernissart Coal Board Limited Company dug ve coal pits on Bernissart
territory. The Négresse pit (no. , exploited from ) and Sainte-Barbe pit
(no. , exploited from ; Fig. .) were used for coal extraction and coupled
with the Moulin pit (no. , exploited from ) for ventilation. The Sainte-
Catherine pit (no. , exploited from ) was the third extraction pit and was
coupled with pit no.  (exploited from ?) for ventilation. The maximum
distance between pits  and  was about , m. With a depth of m, the
Sainte-Barbe pit was the deepest. In spite of a rather archaic technology, the
daily production for the three extraction pits was about  tons. However,
the ood problems were more important than in other coal mines from the
Mons area; steam pumps were used to extract the water.
On February , , miners digging a horizontal exploration gallery
m below ground level suddenly encountered, m to the south of
the Luronne seam, disturbed rocks, indicating that they were penetrating
inside a vertical cran—a local term meaning a pit formed by natural col-
lapse through the coal seams that was lled especially with clayey deposits
normally located above the coal measures.
On March , chief overseer Cyprien Ballez, engineer Léon Latinis,
and mine director Gustave Fagès went down into the Sainte-Barbe pit to
evaluate the situation. It was decided to traverse this cran and to rejoin the
coal seam on the other side. Overseer Motuelle and miners Jules Créteur
and Alphonse Blanchard were put in charge of continuing the exploration
gallery through the perturbed layers of the cran. On March , Ballez no-
ticed that the exploration gallery was still in the perturbed zone of the cran.
In March, the miners had already collected dinosaur remains: fragmen-
tary bones and teeth, which are labeled “remains of the rst Iguanodon,
March ” and are housed in the paleontological collections of the
. But they apparently paid little attention to these discoveries, believ-
ing that they were just fossil wood.
On April , the exploration team again entered nondisturbed but in-
clined formations. On April , Ballez and Latinis went down again together
The Discovery and Excavation
of the Bernissart Iguanodons
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Bernissart and the Iguanodons 5
in the exploration gallery. The engineer estimated that they had again
reached coal-bearing formations. Latinis’s explanations apparently did not
satisfy Fagès. Indeed, the mine manager decided to accompany the engi-
neer and the chief overseer in the exploration gallery on April . While
inspecting the deposits, Fagès found a long object with an oval cross sec-
tion and a brous texture. Latinis believed that it was a fossil oak branch.
Conversely, Fagès ironically asserted that it was a rib of Father Adam.
Miner Jules Créteur mentioned that he had already found a larger fossil,
and the team soon unearthed limb bones in the gallery. In the evening,
miners brought several fragments of these fossils to Café Dubruille. There
the local doctor, Lhoir, who also worked for the coal mine, burned one of
the fragments and conrmed that the fossils collected by the miners were
bones, not wood. Many new fossils were discovered by the miners in the
night of April –. On April , Fagès ordered Ballez to bring all the frag-
ments of bones that the miners had collected to the surface and to lock up
the end of the gallery.
On Sunday, April , Latinis was commissioned to go to Mons to show
the fossils to the well-known geologist François-Léopold Cornet. But Cor-
net was not home. Latinis thus left the fossils to his young son, Jules (a
future renowned geologist), and asked him to tell his father that these bones
had been found in the Sainte-Barbe pit at Bernissart.
On April , F.-L. Cornet came to Bernissart and briey discussed the
Bernissart discovery with Latinis. He could not meet Fagès, who was with
Ballez in the Sainte-Catherine pit. On April , Cornet told the zoologist
Pierre-Joseph Van Beneden, professor of paleontology at Leuven University,
that Latinis, who was a former student of Van Beneden, had discovered fos-
sil bones at Bernissart, and he sent him some of the bones that Latinis had
left with his son. Van Beneden quickly identied the teeth as belonging
to the dinosaur Iguanodon, previously described from Wealden deposits
in England.
On April , Fagès went to Mons to meet the chief mining engineer,
Gustave Arnould, who immediately sent a telegram to Edouard Dupont,
director of the Musée royal d’Histoire naturelle de Belgique () at
Brussels to inform him of the important discovery at m below ground
level in the Sainte-Barbe Pit (Fig. .).
On Saturday, April , Louis De Pauw, head preparer at the  and
a man who already had extensive experience in the excavation and prepa-
ration of fossil vertebrates, met Arnould at Blaton. They went together to
Bernissart. Fagès showed them the bones recently found in the gallery; De
Pauw recognized two ungual phalanges and one vertebral centrum. It was
then decided to go down together into the fossiliferous gallery. De Pauw
() reported that the walls of the exploration gallery were completely
covered by fossil bones, plants, and shes. Ballez, Motuelle, Créteur, and
Blanchard soon unearthed a complete hind limb that they transported on
a plank covered with straw. But after a -m walk, the bones began to
disintegrate on contact with the fresh air of the mine galleries. De Pauw
protected the biggest remaining fragment with his own clothes, and Ballez
and Motuelle brought the fossils to the surface. De Pauw realized that the
presence of pyrite inside the bones was one of the biggest problems that
they had to face if they were to unearth the fossils from the Sainte-Barbe pit.
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Godefroit et al.
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He packed the collected bones in a box full of sawdust and brought them
back to Brussels. In  workshops, he succeeded in solidifying the limb
bones from Bernissart with gelatin. In the meantime, Latinis prepared 
more boxes full of fossils at Bernissart.
Fagès quickly gathered together the board of directors of the Bernis-
sart Coal Board Limited Company. They decided to donate the fossils
discovered in the Sainte-Barbe pit to the Belgian state and to notify
Charles Delcour, minister of the interior, and Edouard Dupont, direc-
tor of the , about this decision. But the excavations could not
immediately begin because the  team was busy preparing for the
Paris World’s Fair.
De Pauw settled in Bernissart on May , and the excavations be-
gan on Wednesday, May . The excavation team included one warder
(M.Sonnet) and one molder (A.Vandepoel) from the , six miners
(J.Créteur, A.Blanchard, J.Gérard, E.Saudemont, D.Lesplingart, and
Dieudonné), and the overseers Ballez, Mortuelle, and Pierrard. Every day
from : in the morning until : in the afternoon, the team went down
into the Sainte-Barbe pit. The excavation method De Pauw created proved
to be efcient and is still used today during paleontological excavations.
Each Iguanodon skeleton was split into pieces. The exposed bones were
rst covered by wet paper or liquid clay and coated by a layer of plaster of
Paris. The fossils were then undercut in a bed of matrix and the reverse
side plastered. The block was then reinforced with either strips of wood or
1.2. Telegram of April 12, 1878. Translation:
“Important discovery of bones in coalfield fault
Bernissart decomposing due to pyrite. Send De
Pauw tomorrow to arrive Mons station 8 a.m.
Shall be there. Urgent. Gustave Arnaut.”
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Bernissart and the Iguanodons 7
1.3. A, Drawing by G. Lavalette in 1883 of
specimen “L” (RBINS R56) of Iguanodon bernis-
sartensis, as discovered in the Sainte-Barbe pit.
B,Drawing by G. Lavalette in 1882 of specimen
“T” (RBINS R57) of Mantellisaurus atherfieldensis,
as discovered in Sainte-Barbe pit. C, Sketch of the
assemblage of plaster blocks containing pieces
into which specimen “T” (RBINS R57) was divided
for raising to the surface. Block “1T” contains
the skull and “5T,” the end of the tail of this
individual.
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Godefroit et al.
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steel, then coated with a second layer of plaster. After being sketched and
cataloged (Fig. .), the blocks were carried to the surface. Every afternoon,
from :, the team collected fossils on the coal tip in sediments previously
extracted from the pit (De Pauw, ).
In August , a big earthquake blocked the excavation team for 
hours in the gallery  m below ground level. This gallery was subse-
quently ooded, and on Tuesday, October , the team was forced to aban-
don their work for several months. The tools and the last fossiliferous blocks
had to be left behind in the ooded galleries. At that time, ve skeletons of
Iguanodon had already been discovered, although only that of “A” (
--) had been excavated completely.
Between October  and April , individual “A” was prepared
and mounted in the museum workshop at the St. Georges Chapel of Nas-
sau Palace. The front part of this specimen had been destroyed during the
original gallery excavations. This was one of the earliest mounted skeletons
of associated dinosaur remains (Norman, ; Fig. .).
In the meantime, Antoine Sohier replaced Latinis as engineer in the
Bernissart Coal Board Limited Company and received the task of repair-
ing the damaged galleries and replacing the old wooden shaft lining of the
Sainte-Barbe pit with a cast iron one (see Sohier, ). Since the discovery
of the rst fossils in the Sainte-Barbe pit, the relations between Fagès and
Latinis were characterized by conict. Latinis was regularly dressed down
because he did not regularly inspect the galleries. Latinis was apparently
absent without leave when the gallery collapsed after the earthquake, and
Fagès held him responsible for the collapse.
The excavations restarted on May , . De Pauw was accompanied
by four members of the  team (M.Sonnet, A.Collard, and A. and
L.Vandepoel) and by the same miners as in . J.Créteur was the rst
to nd the abandoned tools and blocks in the gallery at −m (Fig. .A).
The excavations proceeded with great success, resulting in the removal of
 more or less complete and four partial skeletons of iguanodontids, two
Bernissartia (a dwarf crocodile) skeletons, one “Goniopholis” (larger croco-
dile) skeleton, two turtles, and innumerable shes and plant remains. From
this rst concentration of fossils, the gallery at the -m level was extended
horizontally for about m in an east–southeast direction across the cran,
passing through an area where the stratied sediments were almost hori-
zontal but apparently devoid of large vertebrate remains. On October ,
, another Goniopholis specimen was discovered at about m from the
entrance of the cran. A further eight well-preserved Iguanodon skeletons
were discovered between  and m from the entrance before reaching
its opposite side (Fig. .B).
In , a new horizontal gallery was dug at a depth of −m. The
miners also encountered fossiliferous clays, but the diameter of the cran
was extremely restricted (approximately m) at this level. Three more ar-
ticulated skeletons were recovered from this third series of excavations (Fig.
.C). The clayey layers had completely disappeared m below.
After three years of excavations at Bernissart, about  blocks, weigh-
ing a total of more than  tonnes, were transported to Brussels in furniture
removal vans, each of  tonnes’ capacity.
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Bernissart and the Iguanodons 9
The excavations at Bernissart were particularly expensive for the Bel-
gian state (about , francs in modern currency), and the government
had already allocated two extraordinary grants. In , the expenses in-
volved by this enterprise were considered too high by the Belgian govern-
ment, and the excavations were stopped. Members of parliament suggested
that an Iguanodon skeleton should be sold abroad to defray expenses, but
public outcry prevented this transaction.
During World War I, the German occupation authorities decided to
start new excavations at Bernissart (see Roolf, Chapter  in this book). The
plans, revealed in documents captured after the liberation of Belgium,
indicated that a new gallery was to be excavated at −m. The explora-
tion gallery was stopped on October , , m in front of the border of
1.4. Mounting, in 1878, of the first Iguanodon
specimen (specimen “A,” RBINS VERT-5144-1716)
in the St. Georges Chapel, or Nassau Chapel,
assembly workshop of the MRHNB. This room is
now an exhibition hall in the Albert I Royal Library,
Brussels. To the left of the iguanodon’s hind limb
can be seen the skeletons of a kangaroo and
a cassowary, used as models in assembling the
skeleton.
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Godefroit et al.
10
the cran. Unfortunately, the newly excavated tunnel collapsed when the
occupying forces withdrew.
On January , , Albert Anciaux, then the director general of the
colliery at Bernissart, sent a letter to Gustave Gilson (the director of the
 after Edouard Dupont), complaining that the costs of the aborted
excavation in Bernissart between  and , which were entirely borne
by the colliery owners, amounted to , francs. The Belgian government
reimbursed these expenses in May  (Gosselin, ).
1.5. Plan views of the excavations at Bernissart,
with the skeletons restored in their original loca-
tions (adapted by Norman 1986 from original
archived documents in RBINS). Not all the letter/
number-coded individuals are now identifiable in
the collections. A, First series at −322m; B, sec-
ond series at −322m on the east–southeast side
of the cran; C, third series at −356m. Abbrevia-
tions: Be, the small crocodile Bernissartia; Go, the
larger crocodile “Goniopholis”; ch, turtles.
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Bernissart and the Iguanodons 11
After the war, the extraction pits were put again into exploitation at
Bernissart. But coal extraction at Bernissart was no longer nancially viable,
even as the mining activities at the neighboring Harchies colliery became
highly protable.
On September , Anciaux informed Gilson that for reasons of econ-
omy, the water pumps and ventilators at the Sainte-Barbe pit would had to
be removed and the pit abandoned, unless funds could be found elsewhere.
The director of the Bernissart colliery proposed two solutions to maintain
the paleontological research activities in Sainte-Barbe pit and estimated
the annual maintenance costs: ,, or , francs, depending on
the solution that was chosen. G. Gilson approached the Belgian govern-
ment and private sponsors to nd nancial support. Despite national and
international appeals, the Sainte-Barbe pit was denitively closed at the
end of October .
At the beginning of the s, Jules Destrée, then minister without
portfolio and senator, again requested that the Belgian government release
 million francs to restart excavations at Bernissart. The moment was badly
chosen, thanks to the dramatic consequences of the  stock market crash
on the Belgian economy, particularly on the collieries.
According to Gosselin (), the German occupation authorities
again tried to start excavations at Bernissart between  and . They
took maps and documents necessary for a new exploitation of the fossilifer-
ous site away from André Capart (then a section director at the ) and
De Pauw’s son. However, we did not nd any document in the archives of
the  that could corroborate this hypothesis.
From  onward, once the excavations at Bernissart had ceased, museum
preparation proceeded rapidly. Once the Iguanodon blocks arrived in Brus-
sels, they were stored in the museum workshop, housed in the St. Georges
Chapel of Nassau Palace, now preserved as an exhibition hall in the Albert
I Royal Library. De Pauw () described in detail the preparation of the
Iguanodon skeletons. The plastered blocks were exposed on their upper
surface (the surface containing the bones exposed in the gallery) by re-
moving the protective casing of plaster of Paris. Then a wall of plaster was
constructed around the block and a hot glue mixture, diluted with alcohol
and saturated with arsenic, was poured on top. De Pauw believed that the
arsenic was able to “kill” the pyrite. Excess glue mixture was cleaned off
and the block hardened in a drying room. The reverse side of the block
was then prepared with a cold chisel to remove the plaster and the matrix,
and the glue mixture was applied on this side. The pyrite was systematically
curetted from the bones. Some vertebrae contained more than  kg of py-
rite. The remaining cavities were lled with carton-pierre, a stable mixture
of paper, glue, and talc.
It was decided to mount the best preserved Iguanodon specimens
in a lifelike gait. In , the rst complete specimen (individual “Q,
 R, the holotype of Iguanodon bernissartensis) was assembled and
mounted by L. De Pauw and his team in the St. Georges Chapel. The
bones were suspended from scaffolding by ropes that could be adjusted
so as to obtain the most lifelike position for the complete skeleton, which
Preparation, Mounting,
and Exhibition of the
Bernissart Iguanodons
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Godefroit et al.
12
was then supported by an iron framework (Fig. .). This rst mounted
specimen was publicly exhibited in  in a glass cage constructed in the
interior court of Nassau Palace. In , the cage was lengthened to accom-
modate a second specimen (individual “T,”  R, the only complete
specimen of Mantellisaurus athereldensis) and a selection of fossils of the
Bernissart ora and fauna (Fig. .).
But the Nassau Palace chapel quickly became too small for the storage,
preparation, mounting, and exhibition of these numerous and bulky skel-
etons. In , the iguanodons were transported to a new location: the Royal
Museum of Natural History in Leopold Park. In , ve specimens were
mounted in a glass cage close to the entrance of the museum. From 
onward, the whole Bernissart exhibition was permanently installed in the
newly constructed Janlet Wing of the . Eleven complete specimens
1.6. Mounting in 1882 of the first complete
Iguanodon specimen (specimen “Q,” RBINSR51,
the holotype of I. bernissartensis) in the St.
Georges Chapel. The bearded figure closest to the
specimen is L.De Pauw.
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Bernissart and the Iguanodons 13
were exhibited in a lifelike gait, while  more or less complete and eight
fragmentary individuals were presented as an en gisement display (Fig. .).
Between  and , the Iguanodon skeletons were dismantled and
treated because  years of changes in temperature and humidity had dam-
aged them. The bones were soaked in a mixture of alcohol and shellac, a
natural lacquer secreted by coccid insects. The specimens were installed
into two large glass cages to stabilize the temperature and humidity of their
environment (Fig. .).
During World War II, all the specimens were again dismantled and
stocked in the cellars of the museum, for fear of aerial bombings. But the
humidity was too much for these fragile fossils, which were again mounted
in the exhibition hall before the end of the war (Bultynck, ).
From  to , the s Janlet Wing was renovated. On this
occasion, the iguanodon skeletons were again completely restored. All
the bones were reinforced by a solution of synthetic polyvinyl acetate in
acetone and alcohol (known by the trade name Mowilith). New glass cages
were constructed to protect the skeletons (Fig. .).
From the beginning, E. Dupont and G. Fagès had friendly relations. Du-
pont expressed his gratitude to Fagès, who had accepted the care of the
Bernissart fossils for the , and Dupont did everything in his power to
The Study of the
Bernissart Iguanodons
1.7. First two complete specimens of Bernissart
iguanodons exhibited in the interior court of the
Nassau Palace, Brussels, in 1884. Left, holotype
specimen of Iguanodon bernissartensis (individual
“Q,” RBINS R51); right, complete specimen of
Mantellisaurus atherfieldensis (individual “T,”
RBINS R57).
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Godefroit et al.
14
atter Fagès. In , he suggested to the minister of the interior that Fagès
be decorated with the Order of Leopold, the highest distinction in Belgium.
Dupont even invited Fagès and his wife to celebrate Christmas  in his
home! (They politely refused.) The explanatory label at the feet of the rst
mounted specimen in the interior court of Nassau Palace mentioned that
it was “discovered in  in Bernissart colliery by M.Fagès, director of the
society.” This label irritated P.-J. Van Beneden, who thought that he ought
to be credited with rst discovering the iguanodons because it was he who
had rst identied the fossils as belonging to the genus Iguanodon and who
had published the rst scientic note about these dinosaurs (Van Beneden,
). It was the start of an epic, although completely futile, dispute be-
tween Van Beneden and Dupont over the authorship of the Bernissart
iguanodons during noisy sessions of the Academy of Sciences in . As
a consequence of these disputes, Dupont insisted that Van Beneden give
the handful of bones that Cornet had sent to him in April  back to the
. Of course, Van Beneden refused, and the relations between the
director of the  and the professor at Leuven University continued
to deteriorate.
A third contentious point, again involving P. J. Van Beneden, con-
cerned the species that had been discovered at Bernissart: did it belong to a
new species or to Iguanodon mantelli, already described from disarticulated
specimens discovered in England? Just after the discovery of the Bernis-
sart iguanodons, Dupont had asked the young naturalist Georges Albert
Boulenger to study these specimens. In , Boulenger presented his rst
1.8. The Bernissart iguanodons, mounted in life-
like gait, in the Janlet Wing of the MRHNB in the
early 1930s.
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Bernissart and the Iguanodons 15
results to the Belgian Academy of Sciences, Letters, and Fine Arts. He de-
scribed the anatomy of the pelvis of these dinosaurs and proposed that the
greater number of sacral vertebrae (six) in the Bernissart form, as opposed
to the ve sacral vertebrae in I.mantelli, merited the establishment of a
new species that he named Iguanodon bernissartensis. Unfortunately, this
paper was refused publication, although a brief, highly critical review of
Boulenger’s paper was published by Van Beneden (), then president of
the science section of the academy, who claimed that observed anatomical
differences were most probably attributable to sexual dimorphism and that
the Bernissart iguanodons belonged to Iguanodon mantelli. Shortly after-
ward, in , Boulenger accepted a post at the British Museum (Natural
History), and in , study of the Bernissart iguanodons was entrusted
to Louis Dollo, a mining engineer of French origin who eventually be-
came a Belgian citizen and who entirely devoted his career to vertebrate
paleontology at the .
Between  and , Dollo (a, b, a, b, c, ,
a, b, , , ) published many preliminary notes on the
Bernissart fauna, especially on Iguanodon. While studying in detail several
parts of the Iguanodon skeleton, Dollo began to adopt a forensic approach
to understanding these fossils. He developed a new style of paleontology
that became known as paleobiology—paleontology expanded to investigate
1.9. The Bernissart iguanodons, presented in an
en gisement display in the Janlet Wing of the
RBINS in the early 2000s.
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Godefroit et al.
16
the biology, and by implication the ecology and behavior, of extinct crea-
tures. The rst paper (Dollo, a) examined the basis for the creation
of the new species, I.bernissartensis, as distinct from I.mantelli. Dollo
established an overall similarity in anatomy between the smaller and more
gracile species from Bernissart ( R) and the remains of the “Mantel-
piece” ( ), and therefore by convention identied  R
as Iguanodon mantelli (Norman, ). With respect to the larger species,
Dollo (a) circumvented the problems of sexual dimorphism in the
sacral count by demonstrating a wider range of additional anatomic differ-
ences: skull proportions, size of narial openings, shape of the orbit, size and
shape of the infratemporal openings, shape of scapular blade, completeness
of external coracoid foramen and overall shape of the coracoid, size of the
humerus, proportions of the manus and pollex, and shape of the anterior
pubic blade. Dollo nally concluded that they merited being considered
a separate species.
Dollo’s nal contribution to the Iguanodon story was published in
 as a synthetic study to honor the centenary of Mantell’s original pa-
per. He identied Iguanodon as an ecological equivalent of the giraffe. Its
kangaroo-like posture enabled it to reach high into the trees to gather its
fodder, which it was able to draw into its mouth with its long, muscular
tongue. The sharp beak was used to nip off tough stems, while the teeth
served to pulp the food before it was swallowed. This image of Iguanodon
as a gigantic kangaroo-style creature, as depicted by Dollo, has become
1.10. A new cage for the Bernissart iguanodons
in 2007.
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Bernissart and the Iguanodons 17
iconic during more than  years and was reinforced by the distribution of
full-size replicas of mounted skeletons of Iguanodon from Brussels to many
of the great museums around the world (Norman, ).
In , British paleontologist D. Norman published a monograph on
Iguanodon bernissartensis. He described the skeleton with the precision
required nowadays. Functional analysis of the skeleton revealed that the
vertebral column, stiffened by a network of ossied tendons, was held more
or less horizontal while the animal was walking or running. Norman also
believed that I.bernissartensis was mainly quadrupedal. The structure of
the pectoral girdle, the ratios of the forelimb and hind limb lengths, the
strongly fused carpal bones, and the presence of hooike unguals on the
middle three digits of the hand suggested that the adult of I.bernissartensis
spent most of its time in a quadrupedal posture, although juveniles had a
predominantly bipedal mode of life.
In , Norman described the small Iguanodon species from Bernis-
sart and concluded that it belonged to Iguanodon athereldensis Hooley,
, a species previously described from the Wealden Beds of the Isle of
Wight. Moreover, he stressed that the former name for it, Iguanodon man-
telli, is a nomen dubium as a result of the fragmentary preservation of the
type material of that species.
On the occasion of the mounting of an Iguanodon bernissartensis cast
in a quadrupedal position at the  in , Bultynck discussed in a short
paper the posture and gait of this species.
It is also worth mentioning that many specialists undertook the study
of the other fossils found at Bernissart: C.E. Bertrand () and G. Poinar
Jr. and A.J. Boucot () for coprolites, A. Lameere and G. Severin ()
for insects, R.H. Traquair () and L. Taverne (, , ) for shes,
Buffetaut () and M.A. Norell and J.M. Clark () for the crocodile
Bernissartia fagesii, and A. C. Seward (), K.L. Alvin (, , ,
), and F. Stockmans () for plants.
In –, three new boreholes were drilled within and around the
Iguanodon Sinkhole at Bernissart. Initially, the aim of this drilling program
was to evaluate the chances of nding more fossils, to understand the gen-
esis of the Iguanodon Sinkhole, and to test a seismic geophysical technique
for ground imaging (Tshibangu et al., a, b). In October ,
the drilling program started with a completely cored well (named  )
using the PQ wireline technique.   reached .m of Thanetian,
Late Cretaceous, Early Cretaceous, and Westphalian sediments (Yans et
al., b; Yans, ). During these operations, various parameters were
recorded: rate of penetration, core recovery, and brief core descriptions
(Tshibangu et al., a, b).   provided exceptional material to
improve our knowledge of the iguanodon-bearing Wealden facies, with
multidisciplinary research funded by - ( no. ....F).
Another borehole ( ) was also cut into the Wealden facies (Spagna
and Van Itterbeeck, ).
The formation processes of the Iguanodon Sinkhole were documented
by sedimentological studies of the lacustrine Wealden facies (including
clay mineralogy, granulometry, and magnetic susceptibility; Spagna et al.,
New Boreholes within the
Iguanodon Sinkhole
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Godefroit et al.
18
; Spagna et al., Chapter  in this book) and by characterization of the
organic matter with Rock-eval, palynofacies, soluble alkane content, and
carbon isotope and structural analyses (Schnyder et al., ). Schnyder
et al. () suggested two steps in the life of the lacustrine Wealden facies
of Bernissart: a rst step with a large supply of plant debris, and a second
step with active algal/bacterial activity with amorphous organic matter,
which followed the lake’s level variations. The paleontological content
was studied using paleohistology (de Ricqlès and Yans, ; de Ricqlès et
al., Chapter  in this book) and diagenesis of the bone fragments (Leduc,
Chapter  in this book), characterization of amber, and preparations for
diatom and ostracod analyses, which were unfortunately barren (C. Cornet,
pers. comm.; B. Andreu, pers. comm.). A late Late Barremian to earliest
Aptian age was estimated for the iguanodon-bearing sediments by both
palynology and chemostratigraphy (Yans et al., a, , ; Dejax
et al., a; Yans et al., Chapter  in this book), which permitted a better
knowledge of the initial steps of the subsidence in the Mons Basin (Spagna
et al., ). Moreover, Wealden facies samples from the  collection
(historical searches of –) and other localities in the Mons Basin
(Hautrage, Thieu, Baudour) were also investigated. Rare dinosaur fossils
are described from the Baudour Clays Formation (Godefroit et al., Chapter
 in this book). Palynology and determination of wood and plant mesofos-
sil fragments provide further information about the paleoenvironment of
the Mons Basin during the Early Cretaceous (Gerards et al., , ;
Dejax et al., b, ; Gomez et al., ; Gomez et al., Chapter  in
this book). In Thieu, the occurrence of dinoagellate cysts suggests marine
inuences in the Wealden facies of the Eastern part of the Mons Basin
(Yans et al., ). These data were integrated into the Early Cretaceous
geological context of Northwest Europe (Thiry et al., ; Quinif et al.,
). Studies are still in progress ...
Alvin, K. L. . Three abietaceous
cones from the Wealden of Belgium.
Mémoires de l’Institut Royal des Sci-
ences naturelles de Belgique : –.
. . On the two cones Pseudo-
araucaria heeri (Coemans) n. comb.
and Pityostrobus villerotensis n. sp. from
the Wealden of Belgium. Mémoires de
l’Institut Royal des Sciences naturelles
de Belgique : –.
. . Further conifers of the Pina-
ceae from the Wealden formation of
Belgium. Mémoires de l’Institut Royal
des Sciences naturelles de Belgique :
–.
. . Weichselia reticulata (Stokes
et Webb) Fontaine from the Wealden of
Belgium. Mémoires de l’Institut Royal
des Sciences naturelles de Belgique :
–.
Benton, M. J. . On the trail of the
dinosaurs. Crescent Books, New York,
 pp.
Bertrand, C. E. . Les coprolithes de
Bernissart. . Les coprolithes qui ont été
attribués aux iguanodons. Mémoires
du Musée royal d’Histoire naturelle de
Belgique : –.
Buffetaut, E. . Sur l’anatomie et la
position systématique de Bernissartia
fagesii Dollo, L. , crocodilien du
Wealdien de Bernissart, Belgique. Bul-
letin de l’Institut Royal des Sciences
Naturelles de Belgique, Sciences de la
Terre  (): –.
Bultynck, P. . Bernissart et les Iguan-
odons. French version by F.Martin
and P. Bultynck. Edition Institut royal
des Sciences naturelles de Belgique,
Brussels,  pp.
. . An assesssment of posture
and gait in Iguanodon bernissartensis
Boulenger, . Bulletin de l’Institut
Royal des Sciences Naturelles de
Belgique, Sciences de la Terre :
–.
References
1-BD Ch1 (1-20).indd 18 2/14/12 8:51 AM
IU Press uncorrected proof. Copyrighted material. For proofing purposes only.
Bernissart and the Iguanodons 19
Dejax, J., D. Pons, and J. Yans. a. Paly-
nology of the dinosaur-bearing Wealden
facies sediments in the natural pit of Ber-
nissart (Belgium). Review of Palaeobotany
and Palynology : –.
Dejax, J., E. Dumax, F. Damblon, and
J.Yans. b. Palynology of Baudour
Clays Formation (Mons Basin, Bel-
gium): correlation within the “strato-
typic” Wealden. Notebooks on Geology
e-Journal. http: //paleopolis.rediris.es/cg/
CG_M/CG_M.pdf, –.
Dejax, J., D. Pons, and J. Yans. .
Palynology of the Wealden facies from
Hautrage quarry (Mons Basin, Belgium).
Memoirs of the Geological Survey :
–.
Delguste, B. . L’exploitation du char-
bon dans le Blaton de la première moitié
du ème siècle. Mercuriale, Mélanges IX,
Cercle d’Histoire et d’Archéologie Louis
Sarot : –.
. . Les origines de la maison
“Canivez.” Mercuriale, Mélanges XV,
Cercle d’Histoire et d’Archéologie Louis
Sarot : –.
De Pauw, L. F. . Notes sur les fouilles
du charbonnage de Bernissart. Décou-
verte, solidication et montage des Iguan-
odons. Imprimerie photo-litho, J.H. and
P. Jumpertz, Bruxelles, Belgium.
Dollo, L. a. Première note sur les dino-
sauriens de Bernissart. Bulletin du Musée
royal d’Histoire naturelle de Belgique :
–.
. b. Deuxième note sur les dino-
sauriens de Bernissart. Bulletin du Musée
royal d’Histoire naturelle de Belgique :
–.
. a. Note sur la présence chez
les oiseaux du “troisième trochanter” des
dinosauriens et sur la fonction de celui-
ci. Bulletin du Musée royal d’Histoire
naturelle de Belgique : –.
. b. Troisième note sur les dino-
sauriens de Bernissart. Bulletin du Musée
royal d’Histoire naturelle de Belgique :
–.
. c. Quatrième note sur les dino-
sauriens de Bernissart. Bulletin du Musée
royal d’Histoire naturelle de Belgique :
–.
. . Cinquième note sur les dino-
sauriens de Bernissart. Bulletin du Musée
royal d’Histoire naturelle de Belgique :
–.
. a. L’appareil sternal de
l’Iguanodon. Revue des Questions
Scientiques : –.
. b. Les Iguanodons de Bernis-
sart. Revue des Questions Scientiques
: –.
. . Iguanodontidae et Camp-
tonotidae. Comptes Rendus hebdoma-
daires de l’Académie des Sciences, Paris
: –.
. . Les allures des Iguanodons,
d’après les empreintes des pieds et de la
queue. Bulletin Scientique de la France
et de la Belgique : –.
. . Le centenaire des Iguanodons
(–). Philosophical Transactions of
the Royal Society of London, Series B :
–.
Forster, C. A. . Iguanodontidae; pp.
– in P.J. Currie and K. Padian
(eds.), Encyclopedia of dinosaurs.
Academic Press, San Diego.
Gerards, T., J. Yans, and P. Gerrienne. .
Growth rings of Lower Cretaceous Soft-
woods: some plaeoclimatic implications.
Notebooks on Geology e-Journal. http://
paleopolis.rediris.es/cg/CG_M/
CG_M.pdf, –.
Gerards, T., J. Yans, P. Spagna, and
P.Gerrienne. . Wood remains and
sporomorphs from the Wealden facies
of Hautrage (Mons Basin, Belgium):
paleoclimatic and paleoenvironmental
implications. Memoirs of the Geological
Survey : –.
Gomez, B., T. Gillot, V. Daviero-Gomez,
P.Spagna, and J. Yans. . Paleoora
from the Wealden facies strata of Bel-
gium: mega- and meso-fossils of Hautrage
(Mons Basin). Memoirs of the Geological
Survey : –.
Gosselin, R. . –, ème anni-
versaire. Histoire d’une gigantesque
découverte, l’Iguanodon de Bernissart.
Administration communale de Bernissart,
 pp.
Hooley, R. W. . On the skeleton of
Iguanodon athereldensis sp. nov., from
the Wealden shales of Athereld (Isle
of Wight). Quarterly Journal of the
Geological Society of London : –.
Leidy, J. . Hadrosaurus and its discovery.
Proceedings of the Academy of Natural
Sciences, Philadelphia : –.
Mantell, G. A. . Notice on the Iguan-
odon, a newly discovered fossil reptile,
from the sandstone of Tilgate forest, in
Sussex. Philosophical Transactions of the
Royal Society of London : –.
Norell, M. A., and J. M. Clark. . A
reanalysis of Bernissartia fagesii, with
comments on its phylogenetic posi-
tion and its bearing on the origin and
diagnosis of the Eusuchia. Bulletin de
l’Institut Royal des Sciences Naturelles de
Belgique, Sciences de la Terre : –.
Norman, D. B. . On the ornithischian
dinosaur Iguanodon bernissartensis from
Belgium. Mémoires de l’Institut Royal
des Sciences naturelles de Belgique :
–.
. . The illustrated encyclopedia
of dinosaurs. Salamander Books, London,
 pp.
. . On the anatomy of Iguanodon
athereldensis (Ornithischia: Ornithop-
oda). Bulletin de l’Institut Royal des Sci-
ences Naturelles de Belgique, Sciences de
la Terre : –.
. . On the discovery of fossils at
Bernissart (–) Belgium. Archives
of Natural History : –.
. . Gideon Mantell’s Mantel-
piece: the earliest well-preserved ornith-
ischian dinosaur. Modern Geology :
–.
. . Dinosaurs. A very short intro-
duction. Oxford University Press, Oxford,
 pp.
Owen, R. . Report on British fossil
reptiles, part II. Report of the Eleventh
Meeting of the British Association for
the Advancement of Science, held at
Plymouth, July , –. London.
Poinar, G., Jr., and A. J. Boucot. . Evi-
dence of intestinal parasites of dinosaurs.
Parasitology : –.
Quinif, Y., H. Meon, and J. Yans. .
Nature and dating of karstic lling in the
Hainaut province (Belgium). Karstic,
geodynamic and paleogeographic
implications. Geodinamica Acta :
–.
Ricqlès, A. de, and J. Yans. . Bernissart’s
iguanodons: the case for “fresh” versus
“old” dinosaur bone. Journal of Vertebrate
Paleontology  (supplement to ): .
Schnyder, J., J. Dejax, E. Keppens,
T.Nguyen Tu, P. Spagna, S.Bou-
lila, B.Galbrun, A. Riboulleau, J.-P.
Tshibangu, and J. Yans. . An Early
Cretaceous lacustrine record: organic
matter and organic carbon isotopes
at Bernissart (Mons Basin, Belgium).
Palaeogeography, Palaeoclimatology,
Palaeoecology : –.
Seward, A. C. . La ore wealdienne
de Bernissart. Mémoires du Musée royal
d’Histoire naturelle de Belgique : –.
Sohier, A. . Note sur la réparation
du cuvelage du puits n° , Ste-Barbe,
du charbonnage de Bernissart. Revue
universelle des Mines (ser. ) : –.
Spagna, P., and J. Van Itterbeeck. .
Lithological description and granulomet-
ric study of the Wealden facies in two
borehole core drilled in the “Cran aux
Iguanodons de Bernissart” (N-W of the
Mons Basin, Belgium). Geologica Belgica
Meeting, Liège, abstract book, .
1-BD Ch1 (1-20).indd 19 2/14/12 8:51 AM
IU Press uncorrected proof. Copyrighted material. For proofing purposes only.
Godefroit et al.
20
Spagna, P., S. Vandycke, J. Yans, and
C.Dupuis. . Hydraulic and brittle
extensional faulting in the Wealden facies
of Hautrage (Mons Basin, Belgium).
Geologica Belgica : –.
Spagna, P., C. Dupuis, and J. Yans. .
Sedimentology of the Wealden Clays in
the Hautrage Quarry (Mons Basin, Bel-
gium). Memoirs of the Geological Survey
: –.
Stockmans, F. . Guide de la salle des
végétaux fossiles. Initiation à la paléobo-
tanique stratigraphique de la Belgique et
notions connexes. Les Naturalistes belges
et Patrimoine de de l’Institut royal des
Sciences naturelles de Belgique, Brussels,
pp.
Taverne, L. . Ostéologie et position
systématique d’Aethalionopsis robustus
(Pisces, Teleostei) du Crétacé inférieur de
Bernissart (Belgique) et considérations sur
les afnités des Gonorhynchiformes. Aca-
démie Royale de Belgique, Bulletin de la
Classe des Sciences : –.
. . Sur Pattersonella formosa
(Traquair, ) et Nybelinoides brevis
(Traquair, ), deux téléostéens sal-
moniformes argentoïdes du Wealdien
inférieur de Bernissart (Belgique) précé-
demment décrits dans le genre Leptolepis
Agassiz, . Bulletin de l’Institut Royal
des Sciences Naturelles de Belgique,
Sciences de la Terre  (): –.
. . Ostéologie et position systé-
matique d’ Arratiaelops vectensis, gen.
nov., Téléostéen élopiforme du Wealdien
(Crétacé inférieur) d’Angleterre et de
Belgique. Bulletin de l’Institut Royal
des Sciences Naturelles de Belgique,
Sciences de la Terre : –.
Thiry, M., F. Quesnel, J. Yans, R. Wyns,
A.Vergari, H. Thévenaut, R.Simon-
Coinçon, C. Ricordel, C., M.-G. Moreau,
D. Giot, C. Dupuis, L.Bruxelles, J.Bar-
barand, and J.-M. Baele. . Continen-
tal France and Belgium during the Early
Cretaceous: paleoweatherings and paleo-
landforms. Bulletin Société Géologique
de France : –.
Traquair, R. H. . Les poissons wealdiens
de Bernissart. Mémoires du Musée royal
d’Histoire naturelle de Belgique : –.
Tshibangu, J.-P., F. Dagrain, B.Deschamps,
and H. Legrain. a. Nouvelles recher-
ches dans le Cran aux Iguanodons de Ber-
nissart. Bulletin de l’Académie Royale de
Belgique, Classe des Sciences : –.
Tshibangu, J.-P., F. Dagrain, H. Legrain,
and B. Deschamps. b. Coring perfor-
mance to characterise the geology in the
“Cran aux Iguanodons” of Bernissart (Bel-
gium). Lecture Notes in Earth Sciences
: –.
Van Beneden, P.-J. . Découverte de
reptiles gigantesques dans le charbonnage
de Bernissart, près de Péruwelz. Bulletin
de l’Académie Royale de Belgique (ser. )
: .
. . Sur l’arc pelvien chez les
dinosauriens de Bernissart. Bulletin de
l’Académie Royale Belge : –.
Yans, J. . Lithostratigraphie, minéralo-
gie et diagenèse des sédiments à faciès
wealdien du Bassin de Mons (Belgique).
Mémoire de la Classe des Sciences, Aca-
démie Royale de Belgique (ser. ) :
–.
Yans, J., J. Dejax, D. Pons, C. Dupuis,
and P. Taquet. a. Paleontologic and
geodynamic implications of the palyno-
logical dating of the Bernissart Wealden
facies sediments (Mons Basin, Belgium).
Comptes Rendus Palevol : –.
Yans, J., P. Spagna, C. Vanneste, M.Hen-
nebert, S. Vandycke, J.-M. Baele, J.-P.
Tshibangu, P.Bultynck, M. Streel, and
C. Dupuis. b. Description et impli-
cations géologiques préliminaires d’un
forage carotté dans le “Cran aux Iguan-
odons” de Bernissart. Geologica Belgica
: –.
Yans, J., J. Dejax, D. Pons, L. Taverne,
and P. Bultynck. . The iguanodons
of Bernissart are middle Barremian to
earliest Aptian in age. Bulletin Institut
Sciences Naturelles Belgique : –.
Yans, J., E. Masure, J. Dejax, D. Pons, and
F. Amedro. . Inuences boréales
dans le bassin de Mons (Belgique) à
l’Albien; in L.G. Bulot, S. Ferry, and
D. Groshény (eds.), Relations entre les
marges septentrionale et méridionale
de la Téthys au Crétacé. Notebooks on
Geology, Mémoire /, Résumé 
(CG_M/).
Yans, J., T. Gerards, P. Gerrienne,
P.Spagna, J. Dejax, J. Schnyder, J.-Y.
Storme, and E. Keppens. . Carbon-
isotope of fossil wood and dispersed
organic matter from the terrestrial
Wealden facies of Hautrage (Mons
Basin, Belgium). Palaeogeography,
Palaeoclimatology, Palaeoecology :
–.
1-BD Ch1 (1-20).indd 20 2/14/12 8:51 AM
IU Press uncorrected proof. Copyrighted material. For proofing purposes only.
... Similarly, osteological fossils of the putative trackmakers are also common and well-reported in Lower Cretaceous sediments in Europe (e.g., Norman, 1980Norman, , 1986Norman, , 2010Norman, , 2012Norman, , 2015Carpenter and Ishida, 2010;Godefroit et al., 2012;Gasulla et al., 2014Gasulla et al., , 2015Gasulla et al., , 2022Verdú et al., 2015Verdú et al., , 2017Verdú et al., , 2018Verdú et al., , 2019Verdú et al., , 2021Fuentes-Vidarte et al., 2016;Lockwood et al., 2021;Santos-Cubedo et al., 2021;Figueiredo et al., 2015Figueiredo et al., , 2023Bonsor et al., 2023;Medrano-Aguado et al., 2023;Santos-Cubedo, 2023). However, the diversity of these dinosaurs has been under scrutiny in recent decades (Norman, 2004(Norman, , 2012(Norman, , 2014Paul, 2008;Carpenter and Ishida, 2010;McDonald et al., 2012;Lockwood et al., 2021). ...
... In 1878, the Sainte-Barbe mine shaft started to produce one of the greatest palaeontological discoveries of all times: more than 20 complete articulated skeletons and several incomplete specimens of the dinosaur Iguanodon along with numerous remains of plants, crocodiles, turtles, rare amphibians, insects and about 3000 actinopterygians. Godefroit et al. (2012) documented the historical background of this discovery. In a detailed monograph, the actinopterygians were studied by Traquair (1911), who identified 11 genera. ...
Article
Lepidotes bernissartensis is a species of holostean ray-finned fish from the Barremian-Aptian of Bernissart, Belgium, described by Traquair in 1911. We provide here a revision of its anatomy, which led us to include this species in the genus Scheenstia, and to consider L. brevifulcratus and L. arcuatus, both from the same site, synonymous with S. bernissartensis. We performed two cladistic analyses in order to assess the phylogenetic position of S. bernissartensis and to do an updated appraisal of the evolutionary history of the ginglymodians. Scheenstia is included in the Lepidotidae, and placed in a pectinated position between the basal genus Lepidotes and the more derived members of the family (other species of Scheenstia, Isanichthys and Camerichthys). The nodes within the lepidotids are weakly supported. Although S. bernissartensis is not directly related to S. mantelli from the Wealden of Europe, the two species have similar palaeoenvironments and stratigraphical ranges. Taken as a whole, the ginglymodians experienced several episodes of diversification that are spatially and temporally restricted. The oldest episode involved basal ginglymodians and occurred in the Middle Triassic, in marine environments along the northern margin of the Tethys. A second episode affected the Semionotidae and occurred in freshwater environments of North America and Europe in the Late Triassic and Early Jurassic. The remaining Semionotiformes, Macrosemiidae and Callipurbeckidae, ranged from the Triassic to the Early Cretaceous and were mostly marine. Among the Lepisosteiformes, two clades, the Lepidotidae and the Lepisosteoidi, show episodes of diversification, first in marine and then in freshwater environments.
... The Early Cretaceous (late Barremian-earliest Aptian) styracosternan Iguanodon bernissartensis Boulenger in Van Beneden 1881 is represented by an exceptional series of subcomplete specimens from a single locality. In 1878, miners discovered a large number of skeletons in a coal mine in Bernissart (southern Belgium; Godefroit et al., 2012); Norman (1986) estimated that at least 37e38 specimens of I. bernissartensis were unearthed from this 'Iguanodon Sinkhole'. Most of them are approximately the same size and can be considered 'adults', while three smaller specimens may represent 'subadults'. ...
Article
At the end of nineteenth century, approximately thirty nearly complete skeletons of the ornithopod dinosaur Iguanodon bernissartensis were discovered in Lower Cretaceous deposits in Bernissart (Belgium). Because most of the specimens are approximately the same size, they offer a unique opportunity for studying individual variation (i.e., differences among individuals of the same or roughly the same age) in iguanodontians. Here, we report the results of a study combining both morphometric and visual analyses of the postcranial skeleton of the Bernissart specimens. We found significant individual variation in the morphology of the axis, sacrum, caudal vertebrae, scapula, humerus, pollex, ilium, ischium, femur, and tibia. No definitive evidence of sexual dimorphism was identified in I. bernissartensis. Individual variation in I. bernissartensis has significant implications for phylogenetic and ontogenetic studies as well as for evaluating diagnostic characters in Iguanodon and other iguanodontians. For example, the status of the basal styracosternan ‘Delapparentia turolensis’ from the lower Barremian of Spain is discussed; given the individual variation in the postcranial skeleton of I. bernissartensis, it is in fact impossible to distinguish ‘Delapparentia’ from Iguanodon species based on the available material.
... From this first concentration of fossil vertebrates, the gallery was extended for approximately 50 m in an east-southeast direction across the cran. The second goniopholidid specimen (IRSNB R290) was found on October 22, 1879, approximately 38 m from the entrance of the cran (see Godefroit et al., 2012). ...
Article
Full-text available
Two specimens originally referred by Louis Dollo to Goniopholis simus from the Lower Cretaceous of Bernissart, Belgium, are described. They consist of fully articulated skeletons, one missing the skull and mandible. Comparison of these specimens with recently revised specimens from the Wealden of England allows confirmation that the Belgian specimens are referable to the goniopholidid Anteophthalmosuchus hooleyi. The Belgian specimens are the most completely known representatives not only of this species but also of any Goniopholididae. Study of the postcranial skeletons from Bernissart reveals that the appendicular skeleton closely resembles that of derived neosuchians. The dorsal and ventral shields present a morphology directly comparable to other goniopholidids and pholidosaurids. Such observations stress the necessity to gather an osteological database of postcranial elements to test relationships of the various neosuchian lineages. Goniopholididae were relatively diverse during the Early Cretaceous of Europe, and depending on taxonomic opinion, three to five genera are recognized: Anteophthalmosuchus, Goniopholis, Hulkepholis, and possibly Vectisuchus and Nannosuchus. SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP Citation for this article: Martin, J. E., M. Delfino, and T. Smith. 2016. Osteology and affinities of Dollo's goniopholidid (Mesoeucrocodylia) from the Early Cretaceous of Bernissart, Belgium. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2016.1222534.
Article
The Lower Cretaceous plant assemblage of Bernissart (Mons Basin, Belgium) was studied in detail during the 1900s, but no recent revisions have been performed. The taxonomy of plant taxa recovered from this site is updated, which includes one undetermined “aquatic plant” taxon, nine fern taxa (Cladophlebis sp., Hausmannia dichotoma, Phlebopteris dunkeri, Matoniaceae indet., Ruffordia goeppertii, Onychiopsis psilotoides, Coniopteris sp., Korallipteris sp., and Weichselia reticulata), aff. genus Taeniopteris (of unknown affinity), and five undetermined conifer organs (one stem, two types of seed, one cone, and a dispersed bract). Two lithologies are identified, both consisting of grey clays, one with a smooth surface while the other is more irregular. A large number of available specimens has permitted the study of the species richness and relative abundance of the locality and both lithologies. The taphonomical analysis of the specimens including the preservation of the remains, fragment size, and associations between taxa, together with the diversity analyses, results in four assemblages: “algae” in the water column of the lake; a vegetation composed of Weichselia and Phlebopteris closest to the lake margin; Hausmannia, Onychiopsis and the other ferns further away from the margin; and Matoniaceae indet., conifers and aff. Taeniopteris even further away from the depositional site. In general, the plant assemblage at Bernissart consists of open vegetation, which probably belonged to an early successional stage that was burnt frequently by wildfires.
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Since the description of Isisfordia duncani, a number of new extinct species and revisions of previously described species have prompted a variety of contradicting phylogenetic hypotheses on the topology of Neosuchia. As a consequence, a consensus on the rooting of Eusuchia in relation to other neosuchian clades has not been reached and the origin of the group remains unsettled. Exemplifying this, Bernissartia fagesii, from the Early Cretaceous of Belgium, has long been considered a key taxon for understanding the origin of Eusuchia, but more recent hypotheses found support for a more basal position, as an ally to goniopholidids, paralligatorids or atoposaurids. Because many details of the anatomy of the type specimen are hidden by glue and the sediment adhering to the fossils, a number of characters are pending confirmation. Based on computed tomography data, we extract bones of the cranium and mandibles, describe new characters and re-evaluate anatomical details in the lectotype specimen. Our phylogenetic analysis confirms that B. fagesii is a derived neosuchian, unrelated to atoposaurids, goniopholidids and paralligatorids. We recover B. fagesii and Koumpiodontosuchus aprosdokiti in a basal position within Eusuchia, together with Susisuchidae, a group of gondwanan neosuchians containing Susisuchus and Isisfordia, which here form a polytomy with Hylaeochampsidae. The presence/absence of pterygoid-bound internal choanae cannot be used to fully resolve relationships at the neosuchian–eusuchian transition because of the variability of this character even at the familial level, as recently reported within susisuchids and bernissartiids. There is no doubt that true eusuchians were present in Laurasia as early as the Early Cretaceous, the hylaeochampsid Hylaeochampsa vectiana being the oldest (Barremian) undoubted representative. But whether the Eusuchia were also present in southern landmasses depends on solving the phylogenetic position of susisuchids and other less known gondwanan forms within or outside Eusuchia.
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The Stavelot-Venn inlier (also called as the Stavelot-Venn Massif) exposes a thick series of sedimentary rocks that have been intensely deformed and fractured by two succeeding orogenies (Caledonian and Hercynian). In addition, the Massif has been metamorphised into greenschists (at least at its southern border), subsequently cut by a large collapse structure (the Graben of Malmedy) and is still subjected to vertical tectonic movements. The geology of the Massif is quite distinct from that of the surrounding Devonian-Carboniferous strata. The rocks that built it, as well as their complex geological histories, created a particular landscape that is of great biological and geological interest; this has led to the creation of the High Fens Natural Park. Its subcrop encloses many peculiarities of regional, international or even global importance: e.g. manganese and molybdenum mineral deposits, slightly mineralised and ferruginous waters and sharpening stones made of ‘coticule’. The building stones extracted from the Massif (slates, quartzites) and from its borders (arkoses, sandstones) provide a characteristic look to the built environment and to other smaller heritage objects. Some relicts of the former mining activities have been used for touristic purposes (e.g. the slate quarry of Recht, the Museum of the Coticule); others have evolved into natural sanctuaries for rare or even exceptional faunas and floras, while some others have been deliberately abandoned to evolve freely. The conservation of the extensive peat bogs of the High Fens receives international attention and support. Clusters of ‘viviers’ (ramparted and closed remnants of Pleistocene lithalsas) within this peat bog landscape are unique witnesses of subarctic conditions that ruled here during the youngest ice age. The cities of Spa and Chevron are true cities of water: the former is internationally renowned, having given its name to the famous and popular spa. The natural and geological heritage assets of the Stavelot-Venn Massif are thus multiple and interrelated: stunning landscapes, interesting former mining activities, exceptional ecosystems, excellent mineral waters and attractive local building stones. However, a holistic project integrating all of the latter components has still to be launched and is waiting for regional, national and international support. The Massif is well positioned between Belgium and Germany and covers different administrative units, offering excellent opportunities for creating a cross-border Geopark. © 2015 The European Association for Conservation of the Geological Heritage
Article
The discovery in 1834 of a partial skeleton of Iguanodon in a quarry near Maidstone in Kent came at the perfect moment. During the 1820s and early 1830s numerous discoveries of the fragmentary remains of large fossil reptiles had been made in England. The block of Kentish Rag, upon which were preserved the scattered remains of the dinosaur Iguanodon (Mantell's "Mantel-piece') provided the early investigators of the anatomy and relationships of these reptiles with a partial, associated skeleton in which it is possible to identify many of the key osteological characteristics of one of the three founder members of the Dinosauria (Owen, 1942). In addition to some notes on the historical context into which the Maidstone speciment fell before, and after, its descovery, a belated description of the Maidstone Iguanodon is provided in this article and a reconstruction of this dinosaur has been attempted on the basis of these preserved remains. -from Author
Article
The osteology of Arratiaelops vectensis gen. nov. from the Wealden (Early Cretaceous) of England and Belgium is studied in detail. The comparison with Pachythrissops laevis from the English Purbeckian and Pachythrissops propterus from the Tithonian of Bavaria shows that the Wealden species does not belong to the genus Pachythrissops as previously thought. Its oral dentition with large bands of numerous very small teeth, its retroarticular fused with the angular and forming a part of the joint surface for the quadrate, its autogenous articular, the posterior opening for the mandibular sensory canal on the medial surface of the angular, its parasphenoid denticulated but devoid of basipterygoid process, its very large supratemporal, and its large, roofed and posteriorly placed temporal fossa in which the exoccipital is participating are all characters which, when joined together, situate Arratiaelops vectensis within the Elopiformes. Some cranial apomorphies connect it more particularly with the Megalopidae. Comparisons with the other fossil and modern Elopiformes and especially with Flindersichthys denmeadi from the Albian of Australia are made. They attest that Arratiaelops vectensis really belongs to a new genus.