New observations on the anatomy and paleobiology of the Eocene requiem shark †Eogaleus bolcensis (Carcharhiniformes, Carcharhinidae) from Bolca Lagerstätte, Italy

Abstract

Here we provide new information about the anatomy of the extinct shark †Eogaleus bolcensis from the early Eocene Bolca Konservat-Lagerstätte. The combination of morphological and dental characters of the three known articulated specimens and a single isolated tooth collected from the Pesciara and Monte Postale sites (e.g., head bulky with rounded snout, pectoral fins of semiplesodic type, cutting-clutching dentition, dermal den-ticles shell-shaped and of drag-reducing type, caudal fin accounting for one fourth of the entire body length, 135-153 vertebrae) supports the assignment of †Eogaleus to the family Carcharhinidae and allows us to confirm definitively its distinct taxonomic status from the sympatric carcharhiniform, the triakid †Galeorhinus cuvieri. Moreover, the analysis of the dermal denticle morphology and the size and maturity age estimates of the studied specimens provide new paleobiological information about †Eogaleus, suggesting a close association of this small top predator with the tropical shallow-water marine context hypothesized for the Bolca paleobiotopes.

Full text

C.
R.
Palevol
17
(2018)
443–459
Contents
lists
available
at
ScienceDirect
Comptes
Rendus
Palevol
w
w
w.sci
encedirect.com
General
Palaeontology,
Systematics
and
Evolution
(Vertebrate
Palaeontology)
New
observations
on
the
anatomy
and
paleobiology
of
the
Eocene
requiem
shark
†Eogaleus
bolcensis
(Carcharhiniformes,
Carcharhinidae)
from
Bolca
Lagerstätte,
Italy
Nouvelles
observations
sur
l’anatomie
et
la
paléobiologie
du
requin
requiem
de
l’Éocène
†Eogaleus
bolcensis
(Carcharhiniformes,
Carcharhinidae)
du
Lagerstätte
de
Bolca,
Italie
Giuseppe
Marramàa,∗,
Giorgio
Carnevaleb,
Jürgen
Kriweta
aUniversity
of
Vienna,
Department
of
Palaeontology,
Althanstrasse
14,
1090
Vienna,
Austria
bUniversità
degli
Studi
di
Torino,
Dipartimento
di
Scienze
della
Terra,
via
Valperga
Caluso
35,
10125
Torino,
Italy
a
r
t
i
c
l
e
i
n
f
o
Article
history:
Received
3
December
2017
Accepted
after
revision
22
April
2018
Available
online
20
May
2018
Handled
by
Annalisa
Ferretti.
Keywords:
Elasmobranchii
Selachii
Ypresian
Von
Bertalanffy
equation
Paleoecology
a
b
s
t
r
a
c
t
Here
we
provide
new
information
about
the
anatomy
of
the
extinct
shark
†Eogaleus
bolcensis
from
the
early
Eocene
Bolca
Konservat-Lagerstätte.
The
combination
of
mor-
phological
and
dental
characters
of
the
three
known
articulated
specimens
and
a
single
isolated
tooth
collected
from
the
Pesciara
and
Monte
Postale
sites
(e.g.,
head
bulky
with
rounded
snout,
pectoral
fins
of
semiplesodic
type,
cutting-clutching
dentition,
dermal
den-
ticles
shell-shaped
and
of
drag-reducing
type,
caudal
fin
accounting
for
one
fourth
of
the
entire
body
length,
135–153
vertebrae)
supports
the
assignment
of
†Eogaleus
to
the
fam-
ily
Carcharhinidae
and
allows
us
to
confirm
definitively
its
distinct
taxonomic
status
from
the
sympatric
carcharhiniform,
the
triakid
†Galeorhinus
cuvieri.
Moreover,
the
analysis
of
the
dermal
denticle
morphology
and
the
size
and
maturity
age
estimates
of
the
studied
specimens
provide
new
paleobiological
information
about
†Eogaleus,
suggesting
a
close
association
of
this
small
top
predator
with
the
tropical
shallow-water
marine
context
hypothesized
for
the
Bolca
paleobiotopes.
©
2018
Acad´
emie
des
sciences.
Published
by
Elsevier
Masson
SAS.
This
is
an
open
access
article
under
the
CC
BY-NC-ND
license
(http://creativecommons.org/licenses/
by-nc-nd/4.0/).
Mots
clés
:
Elasmobranchii
Salachii
Yprésien
Équation
de
von
Bertalanffy
Paléoécologie
r
é
s
u
m
é
De
nouvelles
informations
sont
fournies
sur
l’anatomie
du
requin
disparu
†Eogaleus
bolcensis,
du
Lagerstätte
de
Bolca
de
l’Éocène
inférieur.
La
combinaison
des
caractères
mor-
phologiques
et
dentaires
des
trois
spécimens
articulés
connus
et
d’une
unique
dent
isolée
récoltée
dans
les
sites
de
Pesciara
et
de
Monte
Postale
(à
savoir,
tête
massive
avec
museau
arrondi,
nageoires
pectorales
de
type
semiplésodique,
dentition
agrippante/coupante,
den-
ticules
dermiques
en
forme
de
coquille
et
de
type
drag-reducing,
nageoire
caudale
atteignant
le
quart
de
la
longueur
totale
du
corps,
135
à
153
vertèbres)
donne
du
poids
à
l’attribution
∗Corresponding
author.
E-mail
address:
giuseppe.marrama@univie.ac.at
(G.
Marramà).
https://doi.org/10.1016/j.crpv.2018.04.005
1631-0683/©
2018
Acad´
emie
des
sciences.
Published
by
Elsevier
Masson
SAS.
This
is
an
open
access
article
under
the
CC
BY-NC-ND
license
(http://
creativecommons.org/licenses/by-nc-nd/4.0/).

444
G.
Marramà
et
al.
/
C.
R.
Palevol
17
(2018)
443–459
de
†Eogaleus
à
la
famille
des
Carcharhinidae,
et
nous
autorise
à
confirmer
définitivement
son
statut
taxonomique
distinct
de
celui
du
carcharhiniforme
sympatrique,
le
triakidé
†Galeorhinus
cuvieri.
En
outre,
l’analyse
de
la
morphologie
des
denticules
dermiques
et
l’estimation
de
la
taille
et
de
l’âge
de
maturité
des
spécimens
étudiés
fournissent
de
nou-
velles
informations
paléobiologiques
sur
†Eogaleus,
suggérant
une
association
étroite
de
ce
petit
prédateur
avec
un
environnement
tropical
marin
de
faible
profondeur,
comme
le
sont
supposés
les
paléobiotopes
de
Bolca.
©
2018
Acad´
emie
des
sciences.
Publi´
e
par
Elsevier
Masson
SAS.
Cet
article
est
publi´
e
en
Open
Access
sous
licence
CC
BY-NC-ND
(http://creativecommons.org/licenses/
by-nc-nd/4.0/).
1.
Introduction
The
family
Carcharhinidae
Jordan
&
Evermann
(1896),
whose
representatives
are
commonly
known
as
requiem
sharks,
is
a
diverse
group
of
small-
to
large-sized
selachi-
ans
(possible
maximum
body
length
of
7.4
m
attained
in
Galeocerdo
cuvier)
within
the
order
Carcharhiniformes
Compagno
(1973).
These
sharks
primarily
occur
in
tropical
and
warm-temperate
waters,
and
are
worldwide
dis-
tributed
in
coastal
and
pelagic
ecosystems,
although
some
species
can
enter
brackish
or
freshwaters
(Compagno,
1984,
1988;
Nelson
et
al.,
2016).
Although
the
phylo-
genetic
relationships
within
the
family
remain
largely
unclear
(Cappetta,
2012;
but
see
also
Naylor,
1992;
Iglésias
et
al.,
2005;
etc.),
several
authors
have
recognized
a
set
of
morphological
characters
that
are
useful
to
distinguish
requiem
sharks
from
other
carcharhiniforms,
including
well-developed
nictitating
membranes,
last
gills
open
above
the
base
of
the
pectoral
fins,
caudal
pits,
absence
of
oro-nasal
grooves,
intestine
with
scroll
valve
but
lack-
ing
a
spiral
valve,
strongly
asterospondylic
vertebrae
with
secondary
calcification
in
the
shape
of
a
Maltese
cross,
and
shell-shaped
dermal
denticles
of
drag-reducing
type
(Cappetta,
2012;
Compagno,
1988;
Nelson
et
al.,
2016;
Reif,
1985;
White,
1938).
This
family
currently
includes
about
70
species
in
12
living
and
at
least
eight
fossil
gen-
era
dating
back
to
the
Early
Cretaceous
(Cappetta,
2012;
Guinot
et
al.,
2014).
However,
the
fossil
record
of
Car-
charhinidae
is
almost
entirely
represented
by
isolated
teeth
predominantly
recovered
in
Cenozoic
deposits
in
the
North
Hemisphere
(e.g.,
Adnet
et
al.,
2010;
Cappetta,
2012;
Cappetta
and
Case,
2016;
Noubhani
and
Cappetta,
1997),
although
occurrences
have
been
recently
reported
also
in
South
America
(e.g.,
Landini
et
al.,
2017;
Staig
et
al.,
2015).
So
far,
the
only
carcharhinid
taxon
represented
by
complete
and
articulated
specimens
is
†Eogaleus
bolcensis
Cappetta
(1975)
recovered
from
the
Ypresian
(early
Eocene)
fos-
sil
fish-bearing
strata
of
the
Bolca
Lagerstätte,
one
of
the
few
Paleogene
deposits
in
which
chondrichthyans
are
exquisitely
preserved.
Although
several
recent
stud-
ies
contributed
to
our
knowledge
of
the
extraordinary
paleobiodiversity
and
evolutionary
significance
of
this
deposit,
with
more
than
230
described
teleost
species
(e.g.,
Bannikov,
2004,
2006,
2008;
Bannikov
and
Carnevale,
2009,
2010,
2016;
Blot,
1969;
Blot
and
Tyler,
1990;
Carnevale
and
Pietsch,
2009,
2010,
2011,
2012;
Carnevale
et
al.,
2014,
2017;
Marramà
and
Carnevale,
2017a,
2017b;
Marramà
et
al.,
2016a,
2016b;
Monsch,
2006;
Tyler
and
Santini,
2002),
the
cartilaginous
fishes
of
Bolca
have
received
only
little
attention
so
far
(see
Marramà
et
al.,
2017a,
2017b,
2017c).
Sharks
from
the
Bolca
Lagerstätte
are
mentioned
in
the
literature
at
least
since
the
end
of
the
XVIII
cen-
tury
(Volta,
1796)
and
are
currently
represented,
besides
†Eogaleus
bolcensis,
by
several
articulated
skeletons
of
the
triakid
†Galeorhinus
cuvieri
(Agassiz,
1835)
and
by
isolated
teeth
of
the
odontaspidid
†Brachycarcharias
lerichei
(Casier,
1946)
(Cappetta,
1975;
Fanti
et
al.,
2016;
Marramà
et
al.,
2017b,
2017c).
The
goal
of
this
paper
is
to
provide
new
information
about
the
anatomy
of
the
requiem
shark,
†Eogaleus
bol-
censis.
Such
a
revisionary
analysis
is
necessary
in
order
to
conclusively
exclude
Applegate’s
(1978)
hypothesis,
who
regarded
it
as
a
synonym
of
†G.
cuvieri
(see
“Remarks”
sec-
tion).
Moreover,
new
analyses
of
size
and
age
estimation,
as
well
as
the
study
of
dermal
denticles
provide
new
insights
into
the
paleobiology
and
paleoecology
of
this
Paleogene
requiem
shark.
2.
Geological
setting
The
examined
material
comes
from
the
early
Eocene
Bolca
Konservat-Lagerstätte,
which
is
located
in
Verona
Province,
northeastern
Italy
(Fig.
1).
The
preservational
quality
and
lithological
features
of
the
slabs
containing
specimens
MCSNV
T.311,
MCSNV
T.414/T415
and
MGP-PD
8869
C/8870
C
are
consistent
with
the
lithology
of
the
fish-
bearing
strata
of
the
Pesciara
site,
whereas
MCSNV
VII.B.94
has
been
extracted
from
the
strata
of
the
Monte
Postale
site
as
previously
suggested
by
Jaekel
(1894)
and
Cappetta
(1975).
The
fossiliferous
layers
of
the
Pesciara
site
have
been
traditionally
referred
to
the
so-called
‘Calcari
Nummulitici’,
an
informal
unit
of
Eocene
age
widely
distributed
in
northeastern
Italy
(Papazzoni
and
Trevisani,
2006).
This
succession
consists
of
a
cyclic
alternation
of
about
20
meters
of
finely
laminated
micritic
limestones
(containing
exquisitely
preserved
fishes,
plants
and
invertebrates)
and
coarse-grained
biocalcarenite/biocalcirudite
with
a
rich
benthic
fauna.
Based
on
their
larger
benthic
foraminiferan
content,
the
fish-bearing
limestones
of
the
Pesciara
site
were
referred
to
the
†Alveolina
dainelli
Zone,
corresponding
to
the
late
Cuisian
(late
Ypresian,
about
49
Ma;
Papazzoni
and
Trevisani,
2006;
Papazzoni
et
al.,
2014).
The
results
of
a
recent
quantitative
paleoecological
analysis
by
Marramà
et
al.
(2016c)
suggested
that
the
Pesciara
fish
assem-
blage
was
characterized
by
a
sharp
oligarchic
structure
dominated
by
zooplanktivorous
fishes
(mostly
clupeoids),
whereas
the
taphonomic
features
suggest
that
the

G.
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/
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R.
Palevol
17
(2018)
443–459
445
Fig.
1.
Location
and
geological
map
of
the
Bolca
area
showing
the
Pesciara
and
Monte
Postale
sites
of
Ypresian
age
where
†Eogaleus
bolcensis
Cappetta
(1975)
has
been
found.
Fig.
1.
Carte
géologique
et
de
localisation
de
la
zone
de
Bolca,
montrant
les
sites
de
Pesciara
et
de
Monte
Postale
d’âge
Yprésien
où
a
été
trouvé
†
Eogaleus
bolcensis
Cappetta
(1975).
Modifié
selon
Marramà
et
al.
(2016c).
Modified
from
Marramà
et
al.
(2016c).
fossiliferous
sediments
accumulated
in
a
shallow
intraplat-
form
basin
in
which
anoxic
conditions
and
the
develop-
ment
of
a
biofilm
at
the
bottom
promoted
the
high-quality
preservation
of
the
fossils
(see
also
Papazzoni
and
Trevisani,
2006).
The
Monte
Postale
site
is
located
about
300
m
north
of
the
better
known
Pesciara
site,
and
exhibits
similar
age
and
sedimentological
features,
mostly
comprising
finely
laminated
micritic
limestones
with
fish
and
plant
remains.
Papazzoni
et
al.
(2017)
recently
investigated
the
stratigraphical
relationships
between
the
two
fossiliferous
deposits,
suggesting
that
the
uppermost
strata
of
the
Monte
Postale
should
correlate
with
those
of
the
Pesciara
site,
although
the
fossiliferous
laminites
of
Pesciara
appear
to
be
slightly
younger.
The
Monte
Postale
succession
includes
the
Cretaceous
Scaglia
Rossa
Formation
up
to
Ypresian
fossiliferous
limestone,
containing
larger
benthic
foraminifera
of
the
genus
†Alveolina
and
also
marine
and
brackish
molluscs
in
its
uppermost
part
(Papazzoni
et
al.,
2014).
Solid
evidence
of
a
coralgal
rim,
lagoonal
deposits,
and
a
fore-reef
system
was
detected
for
the
Monte
Postale
paleobiotope
(Vescogni
et
al.,
2016);
this
interpreta-
tion
was
also
supported
by
recent
paleoecological
and
taphonomic
studies
of
the
Monte
Postale
fish
assemblage
based
on
the
abundance
of
marine
and
terrestrial
plants,
the
large
number
of
invertebrates
(including
abundant
corals),
and
reef-associated
fishes
(Marramà
et
al.,
2016c).
Disarticulation
of
fish
skeletons,
unimodal
dispersion
of
the
elements
and
bioturbation
were
the
result
of
at
least
periodic
oxic
bottom
conditions
(Marramà
et
al.,
2016c).
3.
Material
and
methods
The
present
study
is
based
on
the
three
articulated
specimens
formerly
studied
by
Cappetta
(1975)
and
on
new
material
represented
by
a
single
tooth.
Specimens
are
housed
in
the
collections
of
the
Museo
Civico
di
Storia
Naturale
di
Verona
(MCSNV)
and
Museo
di
Geologia
e
Pale-
ontologia
dell’Università
degli
Studi
di
Padova
(MGP-PD).
The
articulated
specimens
MCSNV
VII.B.94
and
MGP-PD
8869
C/8870
C
were
examined
under
UV
light
to
distinguish
preserved
soft
tissues
from
grout
or
pigments
tradition-
ally
used
in
the
ancient
restorations
of
the
fossil
fishes.
Measurements
of
the
specimens
were
taken
to
the
near-
est
0.1
cm
and
the
total
length
(TL;
from
the
anterior
tip
of
the
snout
to
the
posteriormost
tip
of
the
caudal
fin)
is
used
throughout.
The
dermal
denticles
of
MCSNV
VII.B.94
were
examined
and
photographed
with
a
Scan-
ning
Electron
Microscope
(SEM)
Jeol
6400
at
the
University
of
Vienna.
Osteological
and
tooth
terminology
follows
Cappetta
(1975,
2012),
Reif
(1985),
Compagno
(1988)
and
Herman
et
al.
(2003).
Morphometric
terminology
follows
Cappetta
(1975).

446
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/
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Palevol
17
(2018)
443–459
The
ratio
between
the
ridge
spacing
on
dermal
denti-
cles
and
their
crown
width
is
used
here
to
hypothesize
the
possible
paleoecology
of
†Eogaleus
follows
Reif
(1985).
The
von
Bertalanffy
growth
equation
is
employed
to
cal-
culate
the
estimate
age
for
the
three
specimens
following
the
methods
applied
to
living
and
fossil
chondrichthyans
(Allen,
1966;
Chen
et
al.,
1992;
Fanti
et
al.,
2016).
The
paleo-
biological
implications
of
the
age
estimates
for
†E.
bolcensis
are
not
directly
based
on
the
absolute
numerical
value;
they
rather
are
based
on
the
comparison
of
these
values
with
the
age
at
maturity
of
living
species,
in
order
to
predict
if
the
fossil
specimens
might
represent
sexually
mature
or
immature
individuals.
4.
Systematic
palaeontology
Class
Chondrichthyes
Huxley
(1880)
Subclass
Elasmobranchii
Bonaparte
(1838)
Order
Carcharhiniformes
Compagno
(1973)
Family
Carcharhinidae
Jordan
&
Evermann
(1896)
Genus
†Eogaleus
Cappetta
(1975)
4.1.
Type
species
†Eogaleus
bolcensis
Cappetta
(1975).
4.2.
Diagnosis
(Emended).
A
carcharhinid
shark
with
bulky
head
and
short,
broad
and
rounded
snout,
not
pointed
or
trian-
gular;
preoral
length
much
smaller
than
mouth
width;
first
dorsal
fin
located
in
front
of
the
mid-length
of
the
body;
its
base
equidistant
from
pectoral-
and
pelvic-fin
bases;
height
of
first
dorsal
fin
roughly
equals
the
length
of
its
base;
posterior
margin
of
first
dorsal
fin
located
well
anterior
to
pelvic-fin
insertion;
second
dorsal
fin
well-
developed
and
inserting
close
to
anal-fin
origin
and
about
2/3
as
high
as
the
first
dorsal
fin;
pectoral
fin
almost
equals
the
size
of
first
dorsal
fin
and
containing
about
18–19
radials;
postero-ventral
margin
of
caudal
fin
deeply
incised;
135–153
vertebrae
of
which
50–55
monospondy-
lous
precaudal,
25–35
diplospondylous
precaudals,
and
60–65
caudals;
gradient
monognathic
and
marked
dig-
nathic
heterodonty;
about
15
tooth
files
per
half
jaw
in
both
jaws
(total
row
count
30/30);
teeth
small,
not
exceeding
12
mm
in
total
height;
distal
cusplets
on
lower
and
upper
jaw
teeth;
lower
anterior
teeth
with
weakly
mesio-distally
expanded
crown
and
a
high
central
cusp,
slightly
bent
lingually;
labial
face
strongly
convex,
overhanging
labial
face
of
root
by
a
distinct
bulge;
distal
heels
of
lower
lat-
eral
teeth
bearing
up
to
four
cusplets;
teeth
with
marked
sigmoid
profile
and
strongly
convex
lingual
face
of
cusp;
upper
teeth
with
comparatively
higher
and
wider
crown
and
root;
distal
heel
high
and
oblique,
bearing
one
to
four
strong
cusplets;
basal
edge
of
root
broad,
not
rectilinear,
with
a
distinct
and
deep
nutritive
furrow;
dermal
denticles
on
pectoral
fins
rounded
or
subrhombic,
relatively
small
and
not
ornamented;
rhombic
placoid
scales
of
trunk
and
caudal
fin
bearing
5–6
prominent
longitudinal
ridges
with
ectodermal
pits
along
their
posterior
edge.
4.3.
Remarks
The
genus
†Eogaleus
was
created
by
Cappetta
(1975)
to
accommodate
three
articulated
specimens,
MCSNV
T.331
(holotype),
MCSNV
VII.B.94
and
MGP-PD
8869
C/8870
C,
because
of
their
evident
differences
with
the
other
car-
charhiniform
shark
from
Bolca,
the
triakid
†Galeorhinus
cuvieri,
in
body
proportions,
and
tooth
and
denticle
size
and
morphology
(Cappetta,
1987,
2012;
Fanti
et
al.,
2016).
Nevertheless,
Applegate
(1978)
regarded
†G.
cuvieri
as
a
conspecific
of
†E.
bolcensis
and
referred
both
to
†Alopiopsis.
The
taxon
†Alopiopsis
plejodon
was
created
by
Lioy
(1865)
based
on
a
single
specimen
deposited
in
Vicenza,
Italy.
The
diagnosis
and
descriptions
of
this
taxon
provided
by
Lioy
(1865)
and
later
by
de
Zigno
(1874)
included
a
total
length
of
more
than
150
cm,
probably
about
200
vertebrae,
caudal
fin
about
one
third
of
the
total
length,
and
teeth
defined
as
“completely
filled
inside”
probably
referring
to
the
absence
of
a
pulp
cavity
and
therefore
to
the
osteodont
type.
In
any
case,
this
set
of
characters
does
not
fit
with
the
morpho-
logical
features
of
the
three
specimens
of
†Eogaleus
(see
Description
and
Discussion).
Unfortunately,
the
holotype
of
†A.
plejodon
was
destroyed
during
the
Second
World
War
(Blot,
1980;
Cappetta,
1975)
and
the
original
illustrations
of
Lioy
(1865)
are
unclear
and
difficult
to
interpret,
making
its
validity
and
taxonomic
affinities
impossible
to
define
[see
Cappetta
(1975)
for
a
more
detailed
taxonomic
history
of
the
taxon
†Alopiopsis
plejodon
Lioy
(1865)].
Consequently,
this
taxon
should
be
considered
as
nomen
dubium.
4.4.
Included
species
Type
species
only.
†Eogaleus
bolcensis
Cappetta
(1975)
(Figs.
2–9)
†Alopiopsis
plejodon
Lioy
(1865):
Jaekel
(1894):
p.
171,
pl.
8
†Eogaleus
bolcensis
Cappetta
(1975):
p.
282,
figs.
2–4,
pls.
1–4
(original
occurrence
of
name
and
photographs)
†Alopiopsis
plejodon
Lioy
(1865):
Applegate
(1978):
p.
55
†Eogaleus
bolcensis
Cappetta
(1975);
Blot
(1980):
p.
343
†Eogaleus
bolcensis
Cappetta
(1975);
Cappetta
(1987):
p.
122,
fig.
104
†Eogaleus
bolcensis
Cappetta
(1975);
Frickhinger
(1991):
p.
185
†Eogaleus
bolcensis
Cappetta
(1975);
Cappetta
(2012):
p.
310,
fig.
294
†Eogaleus
bolcensis
Cappetta
(1975);
Carnevale
et
al.
(2014):
p.
41
†Eogaleus
bolcensis
Cappetta
(1975);
Fanti
et
al.
(2016):
p.
8
†Eogaleus
bolcensis
Cappetta
(1975);
Marramà
et
al.
(2017c):
p.
1
4.5.
Holotype
MCSNV
T.331,
nearly
completely
articulated
specimen,
121.6
cm
TL,
Pesciara
site
(Fig.
2A)

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447
Fig.
2.
†Eogaleus
bolcensis
Cappetta
(1975)
from
the
Eocene
Bolca
Konservat-Lagerstätte.
A.
MCSNV
T.331,
holotype.
B.
MCSNV
VII.B.94.
C.
MGP-PD
8869
C.
Scale
bars
50
mm.
Fig.
2.
†Eogaleus
bolcensis
Cappetta,
1975
du
Lagerstätte
éocène
de
Bolca.
A.
MCSNV
T.
331,
holotype.
B.
MCSNV
VII.B.94.
C.
MGP-PD
8869
C.
Barre
d’échelle
=
50
mm.
4.6.
Referred
material
MCSNV
VII.B.94,
partially
articulated
specimen,
123.5
cm
TL,
Monte
Postale
site
(Fig.
2B);
MGP-PD
8869
C/8870
C,
partially
articulated
specimen,
in
part
and
counterpart,
139.5
cm
TL,
Pesciara
site
(Fig.
2
C);
MCSNV
T.414/415,
single
isolated
tooth
in
part
and
counterpart,
lacking
the
root,
Pesciara
site
(Fig.
3).
4.7.
Type
locality
and
horizon
Pesciara
site,
Bolca
Lagerstätte,
Italy;
early
Eocene,
late
Ypresian,
middle
Cuisian,
SBZ
11,
†Alveolina
dainelli
Zone
(see
Papazzoni
et
al.,
2014).
4.8.
Diagnosis
As
for
the
genus.
4.9.
Description
4.9.1.
Overall
body
morphology
Measurements
and
counts
for
†Eogaleus
bolcensis
are
summarized
in
Tables
1
and
2
(see
also
Fig.
4).
The
three
articulated
specimens
examined
exhibit
a
similar
size
rang-
ing
from
121.6
to
139.5
cm
total
length
(TL),
thereby
suggesting
similar
ontogenetic
ages.
The
holotype
MCSNV
T.311
and
specimen
MCSNV
VII.B.94
are
dorso-ventrally
preserved
in
the
anterior
portion
of
the
body,
whereas
their
posterior
half
is
exposed
in
lateral
profile
(Fig.
2A–B).
On
the
contrary,
specimen
MGP-PD
8869
C/8870
C
appears
to
be
preserved
in
lateral
view,
although
the
arched
disposition

448
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et
al.
/
C.
R.
Palevol
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(2018)
443–459
Fig.
3.
†Eogaleus
bolcensis
Cappetta
(1975)
from
the
Eocene
Bolca
Konservat-Lagerstätte;
single
upper
lateral
tooth
in
part
and
counterpart.
A.
MCSNV
T.415.
B.
MCSNV
T.414.
C.
close
up
of
MCSNV
T.415.
Scale
bars
1
mm.
en:
enameloid;
ort:
orthodentine;
pc:
pulp
cavity.
Fig.
3.
†Eogaleus
bolcensis
Cappetta
(1975)
du
Lagerstätte
éocène
de
Bolca;
unique
dent
latérale
supérieure
en
empreinte
et
contre-empreinte.
A.
MCSNV
T.415.
B.
MCSNV
T.414.
C.
Obturation
de
MCSNV
T.415.
Barres
d’échelle
:
1
mm.
en
:
de
type
émail
;
ort
:
orthodentine
;
pc
:
cavité
pulpaire.
of
teeth
might
suggest
that
at
least
the
head
is
dorso-
ventrally
flattened
(Fig.
2C,
5A–B).
The
body
of
†E.
bolcensis
is
elongate,
gradually
tapering
posteriorly,
but
more
mas-
sive
than
that
of
the
Bolca
triakid
†G.
cuvieri.
The
head
is
broad
and
bulky,
as
long
as
broad,
about
18%
of
TL
(Fig.
5).
The
snout
is
short,
broad
and
bluntly
rounded,
not
pointed
or
triangular;
the
preoral
length
is
distinctly
smaller
than
the
mouth
width.
The
mouth
is
ventrally
positioned.
The
pectoral
fin
is
pointed,
being
almost
of
the
same
size
as
the
first
dorsal
fin.
Part
of
the
internal
skeleton
is
preserved
in
specimen
MCSNV
VII.B.94
showing
that
the
pectoral
fin
is
of
semiplesodic
type,
with
the
longest
radial
being
about
60–70%
of
the
anterior
margin
length
(Fig.
6);
therefore,
it
is
consistent
with
the
pectoral
fin-type
that
is
characteristic
of
the
family
Carcharhinidae
(Compagno,
1988).
There
are
about
18–19
pectoral
radials.
The
ante-
rior
distal
radials
are
pointed,
tapering
and
elongated,
with
the
longest
element
being
about
three
times
the
length
of
the
longest
proximal
radial.
Conversely,
the
posterior-
most
distal
radials
are
shorter,
subrectangular
in
shape,
and
higher
than
long.
The
first
dorsal
fin
originates
at
about
25%
TL
and
its
posterior
margin
is
well
anterior
to
the
pelvic-fin
insertion
(Fig.
2).
The
height
of
the
first
dorsal
fin
more
or
less
equals
its
base.
The
second
dorsal
fin
is
well-developed
and
its
origin
is
located
approximately
at
the
same
level
as
the
anal-fin
origin,
at
about
64%
TL;
the
second
dorsal
fin
is
about
2/3
as
high
as
the
first
dorsal
fin.
The
pelvic-fin
origin
is
located
at
about
mid-length
of
the
body
(ca.
51%
TL).
The
anal
fin
is
slightly
smaller
than
the
second
dorsal
fin,
with
its
height
and
base
length
being
about
0.76
and
0.57
times
those
of
the
second
dorsal
fin.
The
posterior
margin
of
the
anal
fin
seems
to
be
deeply
notched
in
MCSNV
VII.B.94.
The
caudal
fin
is
heterocercal
and
accounts
for
one
quar-
ter
of
the
entire
body
length;
its
postero-ventral
margin
is
deeply
incised
and
its
dorsal
lobe
possesses
a
small
accessory
lobe,
the
depth
of
which
is
about
5%
TL.
Undu-
lations
along
the
dorsal
margin
typical
of
carcharhinids
(Compagno,
1988)
are
difficult
to
detect.
The
ventral
lobe
of
the
caudal
fin
is
well
developed.
Although
Cappetta
(1975)
supposed
the
presence
of
a
precaudal
pit
in
MGP-
PD
8869
C/8870
C,
this
feature
cannot
be
unambiguously
detected
on
the
articulated
specimens
here
studied.
No
sex-
ual
dimorphism
can
be
inferred
based
on
the
presence
of
claspers
since
all
specimens
are
poorly
preserved
in
the
pelvic
region.
The
differences
that
can
be
seen
in
the
shape
of
some
structures
(e.g.,
fin
shape
and
size,
robustness
of
the
caudal
fin
peduncle)
among
specimens
might
be
related,
at
least
in
part,
to
artificial
historical
reconstructions
of
part
the
body
outlines,
although
we
do
not
exclude
that
taphonomic
biases
might
have
led
such
a
variation.
The
skeletal
structures
are
poorly
preserved
in
all
the
specimens
and
their
characters
are
difficult
to
define.
Due
to
the
extensive
covering
of
grout
and/or
pigment
used
to
restore
all
the
specimens,
UV
light
was
not
useful
to
distin-
guish
further
details
of
preserved
bones
or
other
tissues,
with
the
exception
of
MGP-PD
8869
C/8870
C
(Fig.
5A–B).
In
this
specimen,
the
rostrum
appears
small,
short
and
measuring
about
one
quarter
of
the
cranial
length.
As
in
extant
carcharhiniforms
(see
Compagno,
1988),
the
rostrum
is
tripartite
with
a
single
medial
and
two
lat-
eral
rostral
cartilages.
The
nasal
capsules
appear
ovoid
and
laterally
expanded,
with
their
width
reaching
about
1.5–2.0
times
their
length.
The
nasal
apertures
appear
subrectangular
in
outline
and
laterally
elongate.
No
other
structures
of
the
chondrocranium
are
preserved
in
MGP-
PD
8869
C/8870
C.
However,
this
specimen
shows
traces
of
an
eye
as
a
brown-coloured
pigmented
area
(Fig.
5A–B).
The
eyeball
is
small,
rounded
in
outline,
located
at
the
level
of
the
anterior
margin
of
the
mouth,
and
possibly
consists
of
a
dense
accumulation
of
lens
pigments
exhibit-
ing
a
pattern
very
similar
to
that
reported
in
modern
and
fossil
sharks
(Hueter
et
al.,
2004;
Vullo
et
al.,
2016;
Zigman,
1991).
The
palatoquadrate
and
Meckel’s
carti-
lage
run
nearly
parallel
to
each
other,
and
their
antimeres
meet
at
their
respective
symphyses
with
an
angle
of
about
80–90◦.

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449
Fig.
4.
Simplified
outline
of
†Eogaleus
bolcensis
Cappetta
(1975)
from
the
Eocene
Bolca
Lagerstätte
showing
the
measurements
used
in
Table
1.
The
outline
of
†E.
bolcensis
represents
the
average
body
proportions
of
the
three
fossil
specimens
and
is
compared
to
the
body
outlines
(not
to
scale)
of
extant
carcharhinid
genera:
A.
Carcharhinus.
B.
Prionace.
C.
Nasolamia.
D.
Isogomphodon.
E.
Triaenodon.
F.
Lamiopsis.
G.
Negaprion.
H.
Glyphis.
I.
Scoliodon.
J.
Rhizoprionodon.
K.
Galeocerdo.
L.
Loxodon.
Morphometric
scheme
adopted
from
Cappetta
(1975).
Body
outlines
adopted
and
modified
from
Compagno
(1984,
1988).
Fig.
4.
Contour
simplifié
de
†
Eogaleus
bolcensis
Cappetta
(1975)
du
Lagerstätte
éocène
de
Bolca,
montrant
les
mesures
utilisées
dans
le
Tableau
1.
Le
contour
de
E.
bolcensis
représente
les
proportions
moyennes
corporelles
des
trois
spécimens
fossiles
et
est
comparé
aux
contours
corporels
(pas
à
l’échelle),
des
genres
de
carcharhinidés
actuels
(A
à
L).
Schéma
morphométrique
adopté
d’après
Cappetta
(1975).
Contours
corporels
adoptés
et
modifiés
selon
Compagno
(1984,
1988).
The
vertebral
column
of
†Eogaleus
consists
of
about
135–153
vertebral
centra;
of
these,
50–55
are
monospondylous
precaudal
(35–37%
of
the
total
vertebral
count),
25–35
are
diplospondylous
precaudals
(19–23%),
and
60–65
are
caudals
(40–44%).
The
vertebrae
are
strongly
calcified,
subrectangular
in
shape
and
antero-posteriorly
compressed.
4.9.2.
Dentition
The
teeth
of
†Eogaleus
are
small,
up
to
12
mm
in
total
height.
The
dentition
is
of
cutting-clutching
type
and
gradient
monognatic.
Additionally,
a
marked
dig-
nathic
heterodonty
is
developed.
Although
most
of
the
teeth
are
strongly
displaced
in
all
examined
specimens,
it
is
possible
to
recognize
about
15
tooth
files
in
each
jaw
half
in
both
upper
and
lower
jaws
(total
row
count
30/30).
The
lower
anterior
teeth
are
small,
high
and
show
a
weakly
mesio-distally
expanded
crown
with
a
high
and
narrow
cusp,
slightly
bent
lingually
(Fig.
7A–B).
Contrary
to
†Galeorhinus
cuvieri,
the
height
of
the
anterior
teeth
is
greater
than
the
fore-aft
basal
length
and
the
crown
is

450
G.
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/
C.
R.
Palevol
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(2018)
443–459
Fig.
5.
†Eogaleus
bolcensis
Cappetta
(1975)
from
the
Eocene
Bolca
Lagerstätte;
close-up
of
the
head
of
A.
MGP-PD
8869
C
under
UV
light.
B.
Reconstruction
of
MGP-PD
8869
C.
C.
MCSNV
T.311,
holotype.
D.
MCSNV
VII.B.94.
Scale
bars
50
mm.
e:
eye;
mc:
Meckel’s
cartilage;
na:
nasal
capsule;
pa:
palatoquadrate;
ro:
rostrum;
vc:
vertebral
centra.
Fig.
5.
†Eogaleus
bolcensis
Cappetta
(1975)
du
Lagerstätte
éocène
de
Bolca;
obturation
de
la
tête
de
A.
MGP-PD
8869
C
sous
éclairage
UV.
B.
Reconstitution
de
MGP-PD
8869
C.
C.
MCSNV
T.311,
holotype.
D.
MCSNV
VII.B.
94.
Barres
d’échelle
50
mm.
e
:
oeil
;
mc
:
cartilage
de
Meckel
;
na
:
capsule
nasale
;
pa
:
palato-carré
;
r
:
rostre
;
vc
:
centres
vertébraux.
never
upturned
near
the
tip
(Fanti
et
al.,
2016).
The
labial
face
of
the
crown
is
strongly
convex
and
overhangs
the
labial
face
of
the
root
by
a
distinct
bulge.
The
root
is
low
and
very
stout
with
a
distinct
nutritive
furrow,
bearing
a
large
foramen.
Cusplets
are
absent
in
the
lower
ante-
rior
teeth.
The
teeth
of
more
lateral
files
possess
a
wider
crown
base
and
their
labial
face
shows
a
low
central
bulb
(Fig.
7C–D).
The
distal
cutting-edge
is
more
bent
creating
a
notch
at
the
distal
heel
that
is
incised
in
more
lateral
files
and
may
bear
up
to
four
distinct
cusplets.
The
cusp
is
strongly
bent
posteriorly
and
the
weak
mesial
heel
dis-
appears.
Some
of
the
lower
lateral
teeth
show
a
marked
sigmoid
profile
with
a
strongly
convex
lingual
face
of
the
cusp.
Upper
antero-lateral
teeth
have
high
and
wide
crowns
and
roots,
robust
and
oblique
cusps
that
become
more
bent,
wide
and
flat
laterally
in
the
series
(Fig.
8).
The
mesial
cutting
edges
are
less
concave
than
in
lower
lat-
eral
teeth.
There
is
no
differentiated
mesial
heel.
The
base
of
the
mesial
cutting
edge
of
some
teeth
may
bear
some
irregular
weakly
developed
serrations.
The
distal
heel
is
higher
than
in
lower
teeth,
oblique
and
bearing
up
to
four
rather
strong
cusplets.
The
basal
face
of
the
root
is
broad
and
flat,
with
a
distinct
and
deep
nutritive
furrow.

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/
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451
Fig.
6.
Pectoral
fin
of
†Eogaleus
bolcensis
Cappetta
(1975)
from
the
Eocene
Bolca
Lagerstätte.
A.
MCSNV
VII.B.94.
B.
Reconstruction
of
the
radials.
Scale
bars:
20
mm.
Fig.
6.
Nageoire
pectorale
de
†Eogaleus
bolcensis
Cappetta
(1975)
du
Lagerstätte
éocène
de
Bolca.
A.
MCSNV
VII.B.94.
B.
Reconstitution
des
rayons.
Barres
d’échelle
:
20
mm.
Most
of
these
characters
can
be
also
recognized
in
the
sin-
gle
isolated
tooth
MCSNV
T.414/415
(Fig.
3).
Although
the
specimen
is
incomplete,
lacking
the
root
and
distal
heel,
a
strongly
bent
main
cusp
with
a
broad
base
having
with
weak
serrations
at
the
base
of
the
mesial
cutting
edge
is
recognizable.
The
specimen
shows
its
internal
structure,
useful
to
detect
the
tooth
histology
of
†E.
bolcensis.
The
presence
of
a
prominent
coat
of
orthodentine
surrounding
a
wide
pulp
cavity
clearly
indicates
that
the
tooth
repre-
sents
a
functional
tooth
of
orthodont
type,
resembling
the
condition
of
most
of
the
Carcharhiniformes
(Compagno,
1988;
Herman
et
al.,
2003).
This
excludes
its
assignment
to
the
odontaspidid
†Brachycarcharias
from
Bolca
charac-
terized
by
teeth
of
osteodont
type
(Marramà
et
al.,
2017b),
whereas
the
bulky
crown
and
the
absence
of
an
upturned
crown
tip
distinguishes
from
the
sympatric
carcharhini-
form
†Galeorhinus
cuvieri.
4.9.3.
Dermal
denticles
We
sampled
three
loci
with
preserved
dermal
denticles
in
specimen
MCSNV
VII.B.94
(Fig.
9).
The
placoid
scales
of
†E.
bolcensis
are
consistent
with
the
typical
carcharhinid
morphology
being
shell-shaped,
of
drag-reducing
type
(White,
1938;
Reif,
1985).
Their
shape
and
size
vary
con-
siderably
depending
on
the
body
regions.
The
leading
edge
of
the
pectoral
fins
(Fig.
9A)
is
characterized
by
small
dermal
denticles
(250–300
␮m
in
diameter),
rounded
or
slightly
rhomboidal
in
shape,
without
cusp
and
with
a
rounded
posterior
margin.
The
crown
is
not
ornamented
and
the
ridges
are
apparently
absent.
The
dermal
denti-
cles
of
the
ventral
lobe
of
the
caudal
fin
(Fig.
9B–C)
are
closely
imbricated,
larger
(about
500
␮m
wide)
and
char-
acterized
by
a
more
complex
structure.
They
are
rhombic
in
outline
with
a
crown
usually
as
long
as
wide,
orna-
mented
with
six
prominent
longitudinal
ridges
extending
over
the
entire
length
of
the
crown.
The
ridges
are
well
spaced
from
each
other
(ca.
80–90
␮m).
The
two
central
ridges
are
slightly
stronger
and
more
pronounced
than
the
lateral
ones.
The
ectodermal
pits
are
present
along
the
pos-
terior
edge
(Fig.
9C).
The
shape
and
size
of
the
dermal
denticles
of
the
trunk
(Fig.
9D)
are
similar
to
those
of
the
caudal
fin
although
they
can
support
up
to
five
ridges,
some
of
which
bifurcate
at
least
once
and
with
their
posterior
margins
more
pointed
than
that
of
the
caudal
fin
denti-
cles.
Following
the
method
of
Reif
(1985),
the
ratio
between
ridge
spacing
(ca.
80–90
␮m)
and
denticle
diameter
(ca.
500
␮m)
of
†E.
bolcensis
falls
within
the
range
of
the
living
Carcharhinus
amblyrhincos,
C.
melanopterus,
and
C.
plumbeus
(Fig.
10),
which
are
considered
near-shore
species
characterized
by
moderate
swimming
speed
(Compagno,
1984;
Reif,
1985).

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443–459
Fig.
7.
†Eogaleus
bolcensis
Cappetta
(1975)
from
the
Eocene
Bolca
Lagerstätte.
A–B.
Lower
anterior
and
C–D.
Lower
lateral
teeth
of
MCSNV
T.311.
Scale
bars:
10
mm.
Fig.
7.
†Eogaleus
bolcensis
Cappetta
(1975)
du
Lagerstätte
éocène
de
Bolca.
A–B.
Dents
antérieures
inférieures.
C–D.
Dents
latérales
inférieures
de
MCSNV
T.311.
Barres
d’échelle
:
10
mm.
Table
1
Measurements
of
the
three
specimens
of
†Eogaleus
bolcensis
Cappetta
(1975)
from
the
Eocene
Bolca
Lagerstätte.
See
also
the
morphometric
scheme
in
Fig.
3,
which
is
adapted
from
Cappetta
(1975).
Tableau
1
Mesures
des
trois
specimens
de
†Eogaleus
bolcensis
Cappetta
(1975)
du
Lagerstätte
éocène
de
Bolca.
Voir
aussi
le
schéma
morphométrique
de
la
Fig.
3,
issu
de
Cappetta
(1975).
MCSNV
T.311
MCSNV
VII.B.94
MGP-PD
8870
Mean
%
TL
Measurements
cm
%
TL
cm
%
TL
cm
%
TL
1
121.6
100.0
123.5
100.0
139.5
100.0
100.0
2
20.0
16.4
21.1
17.1
27.6
19.8
17.8
3
70.6
58.1
71.7
58.1
78.2
56.1
57.4
4
31.5
25.9
32.1
26.0
33.6
24.1
25.3
5
31.5
25.9
24.6
19.9
42.9
30.8
25.5
6
81.5
67.0
75.6
61.2
89.2
63.9
64.1
7
60.3
49.6
63.4
51.3
74.2
53.2
51.4
8
75.9
62.4
77.2
62.5
91.4
65.5
63.5
9
91.1
74.9
92.2
74.7
105.7
75.8
75.1
10
22.0
18.1
19.1
15.5
?
?
16.78
11
?
?
5.9
4.8
6.6
4.7
4.75
12a
9.9
8.1
?
?
14.2
10.2
9.16
12b
18.1
14.9
?
?
19.2
13.8
14.32
12c
14.2
11.7
?
?
9.9
7.1
9.39
13a
6.6
5.4
10.9
8.8
7.7
5.5
6.6
13b
11.1
9.1
15.4
12.5
8.7
6.2
9.3
13c
9.4
7.7
9.1
7.4
6.2
4.4
6.5
14a
11.2
9.2
14.0
11.3
?
?
10.3
14b
17.7
14.6
20.3
16.4
14.4
10.3
13.8
14c
12.4
10.2
15.3
12.4
?
?
11.3
15a
6.2
5.1
7.6
6.2
7.0
5.0
5.4
15b
7.3
6.0
7.8
6.3
6.4
4.6
5.6
15c
5.8
4.8
6.4
5.2
4.5
3.2
4.4
16a
6.3
5.2
5.9
4.8
7.1
5.1
5.0
16b
6.0
4.9
7.3
5.9
6.4
4.6
5.1
16c
4.8
3.9
3.9
3.2
5.4
3.9
3.7

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453
Fig.
8.
†Eogaleus
bolcensis
Cappetta
(1975)
from
the
Eocene
Bolca
Lagerstätte.
A–B.
Upper
antero-lateral.
C–D.
Lateral.
E–F.
Lateralmost
teeth
of
MCSNV
VII.B.94.
Scale
bars:
5
mm.
Fig.
8.
†Eogaleus
bolcensis
Cappetta
(1975)
du
Lagerstätte
éocène
de
Bolca.
A–B.
Dents
antéro-latérales
supérieures.
C–D.
Dents
latérales.
E–F.
Dents
les
plus
latérales
de
MCSNV
VII.B.94.
Barres
d’échelle
:
5
mm.
4.9.4.
Age
and
maturity
estimation
The
three
specimens
of
†E.
bolcensis
have
a
similar
body
size
ranging
from
121.6
to
139.5
cm
total
length
(TL),
sug-
gesting
similar
ontogenetic
ages
of
the
individuals.
Based
on
literature
data
(Table
3)
the
von
Bertalanffy
growth
curves
of
some
of
the
living
carcharhinids
indicate
rather
large
age
ranges
for
the
three
specimens
of
†E.
bolcensis:
0.7–12.8
years
for
MCSNV
T.311,
0.7–15
years
for
MCSNV
VII.B.94,
and
1.2–19.2
years
for
MGP-PD
8869
C/8870
C.
The
comparison
of
the
age
at
maturity
of
living
taxa
with
the
age
of
each
fossil
specimen
would
suggest
that
they
represent
sexually
immature
individuals.
However,
some
exceptions
occur
when
†Eogaleus
body
sizes
are
compared
with
Iso-
gomphodon
oxyrhynchus,
Triaenodon
obesus,
Carcharhinus
amblyrhynchos,
C.
melanopterus,
C.
sorrah,
and
C.
plumbeus,
suggesting
that
the
fossil
specimens
(and,
in
particular,
the
comparatively
larger
MGP-PD
8869
C/8870
C)
possibly
rep-
resent
sexually
mature
individuals.
5.
Discussion
5.1.
Comparisons
The
analysis
of
the
material
referred
to
†Eogaleus
bol-
censis
has
revealed
the
presence
of
several
characters
that
unquestionably
support
the
assignment
of
this
selachian
to
the
carcharhiniform
family
Carcharhinidae,
and
con-
sequently
its
exclusion
from
the
families
to
which
other
Bolca
Lagerstätten
selachians
were
assigned.
In
partic-
ular,
the
typical
carcharhinid
characters
of
†E.
bolcensis
include
semiplesiodic
pectoral
fins
(aplesodic
in
the
tri-
akid
Galeorhinus),
dentition
of
cutting-clutching
type
with
orthodont
teeth
with
low
roots
(osteodont,
of
tearing
type
and
teeth
with
high
roots
in
the
odontaspidid
†Brachycarcharias),
dermal
denticles
shell
shaped,
of
drag-
reducing
type
(lanceolate
or
teardrop
shaped
in
†G.
cuvieri).
Other
carcharhinid
characters,
such
as
the
vertebrae
of

454
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/
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443–459
Fig.
9.
The
specimen
MCSNV
VII.B.94
showing
the
areas
where
dermal
denticles
were
sampled.
A–C.
SEM
micrographs.
D.
Image
from
Leica
MZ12
stere-
omicroscope
photographed
with
a
3D
digital
microscope
Keyence
VHX-1000D
3D.
Dermal
denticles
were
sampled
from
A.
Anterior
margin
of
pectoral
fin.
B–C.
Anterior
margin
of
the
ventral
caudal-fin
lobe.
D.
Trunk.
Scale
bars
in
A,
B,
D
are
500
␮m;
scale
bar
in
C
is
50
␮m.
Fig.
9.
Spécimen
MCSNV
VII.B.94
montrant
les
zones
où
les
denticules
dermiques
ont
été
échantillonnés.
A–C.
microphotos
au
microscope
électronique
à
balayage.
D.
image
obtenue
au
stéréomicroscope
Leica
MZ12,
photographiée
au
microscope
digital
3D
Keyence
VHX-1000
3D.
Les
denticules
dermiques
ont
été
échantillonnés
à
partir
:
A,
de
la
marge
antérieure
de
la
nageoire
pectorale
;
B–C,
de
la
marge
antérieure
du
lobe
de
la
nageoire
caudale
ventrale.
D,
du
tronc.
Barres
d’échelle
pour
A,
B,
D
=
500
␮m;
barre
d’échelle
pour
C
=
50
␮m.
asterospondylic
type
with
secondary
calcification
having
the
characteristic
shapes
of
a
Maltese
crosss
and
caudal
pits
(Compagno,
1984,
1988;
White,
1938)
are
difficult
to
detect
in
the
examined
material.
Moreover,
in
†E.
bolcensis
the
presence
of
a
pectoral
fin
having
almost
the
same
size
as
the
first
dorsal
fin,
the
caudal
fin
which
occupies
one
quarter
of
the
entire
body
length,
and
the
vertebral
num-
ber
ranging
between
135
and
153
allow
us
to
definitively
exclude
any
possible
alignment
to
the
triakid
†Galerorhinus
cuvieri
(that
on
the
contrary
has
pectoral
fins
with
a
surface
that
is
approximately
twice
that
of
the
first
dorsal
fin,
a
caudal
fin
occupying
a
third
of
TL,
and
200–213
vertebrae;
Cappetta
(1975);
Fanti
et
al.
(2016)),
thereby
providing
a
conclusive
argument
against
the
taxonomic
hypothesis
of
Applegate
(1978).
†Eogaleus
can
be
separated
from
modern
carcharhinids
by
its
unique
combination
of
meristic
features
(see
Table
2).
In
particular,
the
total
number
of
tooth
rows
(30/30)
is
use-
ful
to
distinguish
it
from
Galeocerdo,
Isogomphodon,
Lamiop-
sis,
and
Loxodon.
The
number
of
vertebrae
of
†Eogaleus
(135–153)
separates
this
extinct
genus
from
Galeocerdo,
Lamiopsis,
Nasolamia,
Negaprion,
Prionace
and
Triaenodon,
whereas
the
low
number
of
pectoral
radials
(18–19)
of
†E.
bolcensis
appears
to
be
unique
among
carcharhinids,

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Table
2
Selected
meristic
features
used
to
discriminate
†Eogaleus
bolcensis
Cappetta
(1975)
from
other
living
carcharhinid
genera.
Tableau
2
Caractéristiques
méristiques
sélectionnées,
utilisées
pour
différencier
†
Eogaleus
bolcensis
Cappetta
(1975)
des
autres
genres
de
carcharhinidés
actuels.
Les
données
sont
issues
de
Compagno
(1988).
Genus
Tooth
rows Head
width/Head
length
ratio
Vertebrae
%
monospondy-
lous
precaudal
v.
%
diplospondy-
lous
precaudal
v.
%
diplospondy-
lous
caudal
v.
Pectoral
radials
Carcharhinus
24–37/23–35
0.55–0.81
97–244
23.3–36.4
8.3–28.2
40.8–57.7
22–34
†Eogaleus
30/30
1.00
135–153
34.6–37.0
18.5–23.3
40.0–44.4
18–19
Galeocerdo
18–26/18–25
0.94
216–234
25.2–27.8
19.2–21.3
52.9–54.5
23
Glyphis
27–37/26–34
0.80
149–217
31.1–32.3
18.7–24.9
42.9–50.0
25
Isogomphodon
49–61/49–56
0.45
147–158
30.3–33.3
14.7–15.7
51.0–53.7
30
Lamiopsis
39–43/37–41
0.76
174–179
28.7–29.6
25.2–27.7
44.3–45.2
25
Loxodon
25–28/24–28 0.62
148–191 24.2–28.6
22.0–28.5
46.3–50.3
24
Nasolamia
27–30/24–28
0.52
176–182
28.1–28.9
19.2–19.3
51.7–52.4
?
Negaprion
27–33/27–33
0.91
197–227
33.0–36.1
24.3–27.6
36.8–41.8
25
Prionace
24–31/25–34
0.51
239–252
32.3–37.4
24.3–26.9
37.9–41.9
29
Rhizoprionodon
23–30/21–28
0.65
121–170
20.9–30.1
19.4–30.5
43.1–56.1
22–25
Scoliodon
25–33/24–34
0.60
148–171
27.3–33.0
31.4–38.5
31.6–36.6
22–23
Triaenodon
?
0.85
208–218
30.3–33.2
27.5–31.9
36.6–40.9
24–25
Data
from
Compagno
(1988).
Fig.
10.
Scatter
plot
showing
the
correlation
between
crown
width
and
ridge
spacing
in
dermal
denticles
of
selected
pelagic
fast
swimming
(full
line)
and
near-shore
of
moderate
speed
(dotted
line)
selachians.
Adopted
and
modified
from
Reif
(1985).
Note
that
†E.
bolcensis
falls
in
the
clouds
of
correlation
of
near-shore
moderate
speed
sharks.
The
“C.”
species
belong
to
Carcharhinus.
Fig.
10.
Diagramme
montrant
la
corrélation
entre
la
largeur
de
la
couronne
et
l’espacement
des
crêtes
dans
les
denticules
dermiques
de
sélaciens
pélagiques
nageant
vite
(ligne
pleine)
et
de
sélaciens
nageant
à
vitesse
modérée
près
des
rivages
(ligne
en
tireté).
Adopté
et
modifié
d’après
Reil
(1985).
On
notera
que
†E.
bolcensis
se
place
dans
les
champs
de
corrélation
des
requins
nageant
à
vitesse
modérée
près
des
rivages.
Les
espèces
«
C
»
appartiennent
à
Carcharhinus.
usually
possessing
a
higher
number
of
radials.
Moreover,
†Eogaleus
can
be
distinguished
from
other
early
Eocene
car-
charhinids
for
its
unique
combination
of
dental
characters.
For
example,
teeth
of
Galeocerdo,
collected
from
Ypresian
deposits
of
northern
Africa
(Cappetta,
2012;
Noubhani
and
Cappetta,
1997)
are
larger
(up
to
30
mm),
with
the
apex
strongly
bent
lingually
and
with
always
strongly
serrated
mesial
and
distal
heels;
in
addition,
the
crown
apex
is
only
slightly
bent,
there
is
no
mesial
heel
and
weak
ser-
rations
are
sometimes
present
on
mesial
cutting
edge
in
†Eogaleus.
The
teeth
of
Rhizoprionodon,
which
were
also
recognized
in
lower
Eocene
deposits
of
Morocco
(Cappetta,
2012;
Noubhani
and
Cappetta,
1997),
are
very
small
(less
than
4
mm)
and
usually
without
cusplets,
the
main
cusp
of
upper
teeth
is
narrower
and
the
basal
edge
of
the
root
is
almost
rectilinear
(Cappetta,
2012;
Compagno,
1988).
The
anterior
teeth
of
†Eogaleus
can
be
easily
distinguished
from
those
of
the
Palaeocene
to
Oligocene
†Abdounia
for
the
absence
of
large
and
tall
lateral
cusplets.
The
labial
face
is
almost
flat
in
the
teeth
of
†Abdounia
and
the
crown
does
not
overhang
the
labial
face
of
the
root
(the
labial
face
is
con-
vex
in
†Eogaleus
and
the
crown
overhangs
the
labial
face
of
root).
Furthermore,
the
distal
and
mesial
heels
of
the
upper
teeth
of
†Abdounia
are
usually
characterized
by
the
presence
of
up
to
three
strong
cusplets,
whereas
up
to
four
lower
cusplets
are
present
only
on
the
distal
heel
in
the
upper
teeth
of
†Eogaleus
(see
also
Cappetta
(1980,
2012)).
Although
the
upper
lateral
teeth
of
†Eogaleus
may
resemble
those
of
the
early
Eocene
to
middle
Miocene
†Physogaleus,
the
basal
edge
of
the
root
is
almost
rectilinear
in
the
latter
genus,
and
the
anterior
and
lower
antero-lateral
teeth
are
strongly
medio-distally
compressed
and
may
bear
a
sin-
gle
cusplet
on
the
distal
heel
(the
basal
edge
of
the
root
is
not
rectilinear
and
the
cusp
is
more
erected
in
upper
teeth,
whereas
the
cusplets
are
absent
in
lower
anterior
teeth
of
†Eogaleus;
see
also
Cappetta
(1980,
2012)).
However,
these
differences
are
very
slight,
and
teeth
of
†Eogaleus
bolcensis
might
resemble
those
of
the
Eocene
species
of
†Physogaleus
(including
the
type
species
†P.
secundus).
If
we
interpreted
such
differences
as
interspecific
variation
within
a
single
genus,
then
†Physogaleus
Cappetta
(1980)
would
be
a
junior
synonym
of
†Eogaleus
Cappetta,
1975.
When
erecting
the
genus
†Physogaleus,
Cappetta
(1980)
did
not
mention
the
genus
†Eogaleus
in
his
discussion
and
comparisons.
Fur-
thermore,
the
isolated
occurrence
of
the
skeleton-based
genus
†Eogaleus
(only
known
from
northern
Italy)
within
the
Ypresian
palaeobiogeographic
distribution
area
of
the
widespread
tooth-based
genus
†Physogaleus
(known
from

456
G.
Marramà
et
al.
/
C.
R.
Palevol
17
(2018)
443–459
Europe,
North
America,
North
Africa,
and
Near
East;
see
e.g.,
Cappetta
(2012))
might
support
such
a
hypothesis.
Isolated
teeth
of
†Eogaleus
have
been
also
reported
from
the
early
Eocene
of
Camba
Shale,
India
(Rana
et
al.,
2004)
and
from
the
late
Eocene
of
Wulagen
Formation,
north-
western
China
(Li,
1997).
However,
in
our
opinion
these
occurrences
should
be
regarded
as
doubtful.
The
teeth
fig-
ured
by
Rana
et
al.
2004,
fig.
2
are
very
small
in
size
(about
2–3
mm),
the
crown
of
the
upper
lateral
teeth
is
strongly
distally
bent
and
the
base
of
the
root
is
clearly
rectilinear.
The
lateral
teeth
referred
to
†E.
bolcensis
by
Li,
1997,
fig.
6K–L)
were
almost
not
described,
and
the
morphology
of
the
single
tooth
figured
in
the
paper,
which
is
characterized
by
tall
and
rounded
cusplets
on
the
distal
heel,
is
clearly
not
comparable
with
that
of
†Eogaleus.
5.2.
Paleobiological
remarks
Reif
(1985)
separated
modern
selachians
into
six
ecological
groups
based
on
the
morphology
of
their
squa-
mation,
presence
of
ridges
and
their
spacing.
According
to
this
classification,
dermal
denticles
characterized
by
paral-
lel
ridges
on
the
crown
with
inter-spaces
wider
than
80
␮m
are
typical
of
near-shore
hunters
characterized
by
a
moder-
ate
speed
during
swimming
(e.g.,
Galeorhinus
galeus,
most
of
the
Carcharhinus
species,
Galeocerdo
cuvier),
whereas
denticles
with
ridges
that
are
closely
arranged
with
spaces
of
40
and
80
␮m
between
the
ridges
are
typical
of
fast
pelagic
sharks
(e.g.,
Isurus
oxyrhinchus,
Lamna
nasus,
Car-
charodon
carcharias,
Sphyrna
lewini).
The
parallel
ridges
present
in
the
denticles
of
the
trunk
and
caudal
fin
of
MCSNV
VII.B.94
are
widely
spaced
from
each
other
and
the
ratio
between
ridge
spacing
and
denticle
diameter
falls
within
the
range
of
Carcharhinus
amblyrhyn-
chos,
C.
melanopterus,
and
C.
plumbeus
(Fig.
10),
which
are
considered
near-shore
species
with
a
moderate
swimming
speed
(Compagno,
1984;
Reif,
1985).
In
this
perspective,
we
can
hypothesize
that
†Eogaleus
possibly
had
a
simi-
lar
life
style.
This
is
also
suggested
by
the
presence
of
the
typical
ornamentation
of
the
ectodermal
pits
on
the
pos-
terior
edge
of
the
caudal
fin
and
trunk
dermal
denticles
in
our
samples.
Ectodermal
pits
typically
occur
along
the
pos-
terior
margin
of
placoid
scales
in
near
shore
sharks
with
moderate
swimming
speed
like
Carcharhinus
plumbeus,
Pri-
onace
glauca,
and
Galeorhinus
galeus,
whereas
they
appear
uniformly
distributed
along
the
entire
surface
of
the
den-
ticles
in
fast
swimming
pelagic
sharks
like
Sphyrna
(Reif,
1985).
These
structures,
along
with
the
particular
shell-
shaped
morphology
of
the
placoid
scales,
seem
to
have
a
drag-reducing
function
in
both
ecological
groups,
whereas
ectodermal
pits
appear
to
be
almost
totally
absent
in
deep-
sea
species,
pelagic
sharks
characterized
by
low
swimming
speed,
and
demersal
taxa
living
on
rocky,
sandy
or
muddy
substrates
(Reif,
1985).
Moreover,
considering
that
the
ratio
between
ridge
spacing
and
crown
width
of
†Eogaleus
bolcensis
falls
within
the
range
of
Carcharhinus
amblyrhynchos,
C.
melanopterus
and
C.
plumbeus
(Fig.
10),
the
size
of
the
fossil
specimens
fitted
with
the
von
Bertalanffy
curves
of
these
three
Carcharhinus
species
(see
Table
3)
suggests
that
the
individuals
of
†E.
bolcensis
from
Bolca
may
have
already
reached
the
age
of
maturity,
representing
adult
or
subadult
individuals.
Carcharhinus
amblyrhynchos,
C.
melanopterus
and
C.
plumbeus
are
actually
some
of
the
smallest
living
car-
charhiniform
species
(up
to
2.4
m)
that
are
very
common
in
coastal
inshore
tropical
waters
(mostly
in
the
Indo-Pacific)
often
associated
with
coral
reefs
and
shallow
lagoons
with
depths
of
only
a
few
metres
(Chin,
2013;
Compagno,
1984;
McAuley
et
al.,
2006).
These
selachians
are
active,
social
predators
feeding
on
reef
bony
fishes,
in
particular
small
fishes
less
than
30
cm
long,
but
also
on
squids,
octopuses,
crabs,
lobsters
and
shrimps
(Compagno,
1984).
The
range
of
body
sizes
of
the
†E.
bolcensis
individu-
als
described
herein
is
consistent
with
the
relatively
small
size
of
the
other
Bolca
selachians.
The
triakid
†Galeorhinus
cuvieri
is
known
from
several
completely
articulated
spec-
imens
ranging
from
54
to
92
cm
which
are
interpreted
as
juvenile
individuals
(Fanti
et
al.,
2016).
The
small
sand
tiger
shark
†Brachycarcharias
lerichei
is
known
from
several
isolated
teeth
belonging
to
juvenile
and
adult
individuals
whose
estimated
body
size
might
have
ranged
between
62
and
171
cm
(Marramà
et
al.,
2017b).
In
this
perspective,
the
small
size
of
†E.
bolcensis
specimens,
probably
representing
adult
or
subadult
individuals,
supports
the
hypothesis
of
Marramà
et
al.
(2017b)
who
suggested
that
the
presence
of
small
shark
species
like
†B.
lerichei
and
juvenile
indi-
viduals
of
†G.
cuvieri
in
the
Bolca
paleobiotopes
might
be
related
to
the
competitive
advantage
of
juveniles
and
small
shark
species
in
having
access
to
relatively
competitor-free
trophic
niches
and
food
resources
(see
also
Castro,
1993;
Motta,
2004;
Simpfendorfer
and
Milward,
1993),
which
were
probably
unavailable
for
larger
top
predators.
In
this
context,
sand/sea-grass
beds,
open
sea,
lagoon,
and
coral
reefs
concurred
to
create
heterogeneous
habitats
in
which
small
zooplanktivorous
fishes,
mainly
represented
by
clu-
peoids
(see
Marramà
and
Carnevale,
2015a,
2015b,
2016,
2018)
constituted
one
of
the
main
trophic
resources
for
most
predators
including
†E.
bolcensis.
Similar
paleoeco-
logical
settings
are
also
known
from
other
Eocene
tropical
shallow
contexts
like
the
middle
to
late
Eocene
Midawara
Formation
in
Egypt,
in
which
the
higher
levels
of
the
trophic
chain
were
also
occupied
by
small
cosmopolitan
shallow-
water
specialized
feeders
on
small
nectobenthic
prey,
such
as
small
triakids
and
carcharinids,
all
considered
as
gen-
eralist
feeders
focusing
on
small
active
preys
(Underwood
et
al.,
2011).
6.
Conclusions
The
new
anatomical
observations
on
the
Eocene
shark
†Eogaleus
bolcensis
presented
herein
definitively
confirm
its
attribution
to
the
family
Carcharhinidae
and
its
differ-
ent
taxonomic
status
from
the
sympatric
carcharhiniform
†Galeorhinus
cuvieri.
Moreover,
the
age
estimations
and
the
analysis
of
the
morphology
of
the
dermal
denticles
have
provided
new
insights
into
the
paleoecology
of
this
Paleogene
requiem
shark,
confirming
its
affinities
with
the
tropical
shallow
waters
of
the
Bolca
paleobiotopes.
The
Paleogene
was
marked
by
a
diversification
of
the
carcharhiniform
sharks
that
involved
members
of
the
family
Carcharhinidae
around
the
Palaeocene/Eocene
boundary
and
that
has
been
associated
with
the
decline
of

G.
Marramà
et
al.
/
C.
R.
Palevol
17
(2018)
443–459
457
Table
3
Age
estimates
for
the
three
specimens
of
†Eogaleus
bolcensis
Cappetta
(1975)
from
the
Eocene
Bolca
Lagerstätte
using
the
von
Bertalanffy
functions
of
living
carcharhinid
species.
In
some
cases,
it
was
not
possible
to
detect
an
estimate
because
the
specimens
of
†E.
bolcensis
are
larger
than
the
maximum
length
(L∞)
of
the
original
population.
Tableau
3
Estimations
d’âge
pour
les
trois
spécimens
de
†
Eogaleus
bolcensis
Cappetta
(1975)
du
Lagerstätte
éocène
de
Bolca,
utilisant
les
fonctions
de
von
Bertalanffy
des
espèces
actuelles
de
carcharhinidés.
Dans
certains
cas,
il
n’a
pas
été
possible
de
proposer
une
estimation,
car
les
spécimens
de
†
E.
bolcensis
sont
plus
grands
que
la
longueur
maximum
(L∞)
de
la
population
d’origine.
Taxon
Sex
k
L∞t0Age
at
maturity
Age
of
MCSNV
T.311
Age
of
MCSNV
VII.B.94
Age
of
MGP-PD
8870
Carcharhinus
amblyrhynchos
C
0.05
229.2
−7.51
7.3
7.6
8.0
11.3
Carcharhinus
brachyurus
C
0.0385
384.8
−3.477
16.5
6.4
6.6
8.2
Carcharhinus
brevipinna
M
0.203
257.4
−1.709
7.9
1.4
1.5
2.1
Carcharhinus
brevipinna
F
0.151
288.2
−1.988
7.8
1.6
1.7
2.4
Carcharhinus
falciformis
C
0.148
216.4
−1.76
6.0
3.8
4.0
5.2
Carcharhinus
leucas C
0.076 285
−3
16.0
4.3
4.5
5.8
Carcharhinus
limbatus
C
0.28
171
−1.5
6.0
2.9
3.1
4.5
Carcharhinus
longimanus
C
0.103
341.7
−2.7
4.5
1.6
1.7
2.4
Carcharhinus
longimanus
C
0.099
284.9
−3.4
7.0
2.2
2.3
3.4
Carcharhinus
melanopterus
C
0.251
158.5
2
5.6
7.8
8.0
10.5
Carcharhinus
plumbeus
C
0.04
239.6
−4.9
15.0
12.8
13.2
16.9
Carcharhinus
sorrah
F
0.34
123.9
−1.9
2.5
9.8
15.0
?
Carcharhinus
sorrah M
1.17 98.4
−0.6
?
?
?
?
Galeocerdo
cuvier
(Atlantic)
C
0.107
440
−2.35
10.0
0.7
0.7
1.2
Galeocerdo
cuvier
(Gulf
of
Mexico)
C
0.184
388
−1.13
7.5
0.9
1.0
1.3
Isogomphodon
oxyrhynchus
C
0.12
171.4
−2.612
6.0
7.7
8.0
11.4
Loxodon
macrorhinus
C
0.41
84.2
−2.2
1.6
?
?
?
Negaprion
brevirostris C
0.057 317.65 −2.302
12.2
6.2
6.3
7.8
Prionace
glauca
M
0.18
282
−1.35
15.0
1.8
1.9
2.4
Prionace
glauca
F
0.13
310
−1.77
15.0
2.1
2.1
2.8
Rhizoprionodon
lalandii
C
0.301
78.1
−1.463
2.6
?
?
?
Rhizoprionodon
porosus
C
0.171
112.99
−1.751
3.3
?
?
?
Rhizoprionodon
terraenovae
C
0.359
108
−0.98
3.5
?
?
?
Scoliodon
laticaudus
F
0.358
71.5
−0.59
1.7
?
?
?
Scoliodon
laticaudus M
0.4046
67.6
−0.59
1.3
?
?
?
Scoliodon
laticaudus
C
0.2731
75.5
−0.5664
?
?
?
?
Triaenodon
obesus
F
0.05
207.8
−9.8
9.0
7.8
8.2
12.5
Triaenodon
obesus
M
0.1
150.9
−6.6
9.0
9.8
10.5
19.2
Data
for
living
species
taken
from
Prabhakaran
Nair
(1976),
Compagno
(1984),
Branstetter
(1987),
Branstetter
and
Stiles
(1987),
Branstetter
et
al.
(1987),
Brown
and
Gruber
(1988),
Devadoss
(1998),
Walter
and
Ebert
(1991),
Seki
et
al.
(1998),
Castro
et
al.
(1999),
Lessa
et
al.
(1999,
2000),
Skomal
and
Natanson
(2002),
Chin
(2013),
Oshitani
et
al.
(2003),
Joung
et
al.
(2005),
McAuley
et
al.
(2006),
Robbins
(2006),
Lessa
et
al.
(2009),
and
Gutteridge
et
al.
(2013).
k:
growth
coefficient
(rate
of
change
in
length
increment);
L∞:
mean
maximum
length
for
the
population;
t0:
hypothetical
postnatal
length;
C:
male
and
female
combined;
F:
female;
M:
male.
Ages
are
in
years.
a
large
part
of
the
medium
to
large-sized,
fast-swimming
Cretaceous
top
predators
(Cappetta,
2012;
Friedman
and
Sallan,
2012;
Kriwet
and
Benton,
2004)
and
their
diversi-
fication
promoted
by
shallow
coral
reef
settings
(Sorenson
et
al.,
2014).
In
this
perspective,
†Eogaleus
might
be
con-
sidered,
along
with
†Abdounia,
Carcharhinus,
Galeocerdo,
†Physogaleus
and
Rhizoprionodon,
another
participant
to
the
opportunistic
ecological
replacement
in
marine
high
trophic
levels
experienced
by
Carcharhinidae
in
the
con-
text
of
the
adaptive
fish
radiation
in
the
aftermath
of
the
end-Cretaceous
extinction.
Acknowledgments
The
authors
thank
Roberto
Zorzin
and
Anna
Vac-
cari
(MCSNV)
and
Mariagabriella
Fornasiero
(MGP-PD)
for
access
to
the
facilities
and
material
under
their
care.
Special
thanks
are
due
to
Letizia
Del
Favero
(MGP-PD)
and
Davide
Quagliotto
(Padova)
for
their
technical
support
in
using
the
UV
light,
and
Christian
Baal
(University
of
Vienna)
in
using
the
Scanning
Electronic
Microscope
(SEM).
We
also
thank
Patrick
L.
Jambura
(University
of
Vienna)
for
the
useful
discussion
about
the
tooth
histology
in
carcharhiniforms.
The
manuscript
has
been
greatly
improved
by
the
valuable
comments
and
suggestions
of
Alberto
Collareta
(Università
di
Pisa)
and
an
anonymous
reviewer.
Financial
support
was
provided
by
the
Austrian
Science
Fund
(FWF)
[M2368-B25
to
G.M.];
the
Università
degli
Studi
di
Torino
[ex-60%
2017
grant
to
G.C.].
Open
access
funding
provided
by
Austrian
Science
Fund
(FWF).
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