A strange new chelonioid turtle from the Latest Cretaceous Phosphates of Morocco

Abstract

A new genus and species of huge marine turtle (superfamily Chelonioidea, epifamily Dermochelyoidae)
is described from the Maastrichtian Phosphates of the Oulad Abdoun Basin
of Morocco. A new type of feeding apparatus, adapted for a powerful crushing pattern,
illustrates the noteworthy diversity of fossil vertebrates of the Maastrichtian-Ypresian
Phosphates of Morocco. No other crushing cryptodire or bothremydid pleurodire has this
morphology. During the Maastrichtian, the known crushing pattern of chelonioids was different,
close to that of modern cheloniids, as illustrated in Morocco in the Maastrichtian
Ganntour Basin and the Palaeogene Oulad Abdoun Basin. This new taxon exhibits unusual
cranial characters (fusion of premaxillae associated with a backward and dorsal retraction
of the naris, horizontal stretching of the dorsal meatus quadrati), that are shared only with
another new turtle, known also from the same Maastrichtian Phosphates of Morocco.

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Author's personal copy
C.
R.
Palevol
13
(2014)
87–95
Contents
lists
available
at
ScienceDirect
Comptes
Rendus
Palevol
w
w
w.sci
encedirect.com
General
palaeontology,
systematics
and
evolution
(Vertebrate
palaeontology)
A
strange
new
chelonioid
turtle
from
the
Latest
Cretaceous
Phosphates
of
Morocco
Une
étrange
nouvelle
tortue
chélonioïde
du
Crétacé
terminal
des
Phosphates
du
Maroc
France
de
Lapparent
de
Broina,∗,
Nathalie
Bardeta,
Mbarek
Amaghzazb,
Saïd
Meslouhc
aCR2P,
UMR
7207
CNRS-MNHN-UPMC,
département
Histoire
de
la
Terre,
MNHN,
CP38,
57,
rue
Cuvier,
75231
Paris
cedex
05,
France
bOCP,
centre
minier
de
Khouribga,
Khouribga,
Morocco
cMEMEE,
Rabat,
Morocco
a
r
t
i
c
l
e
i
n
f
o
Article
history:
Received
27
February
2013
Accepted
after
revision
19
July
2013
Available
online
25
December
2013
Presented
by:
Philippe
Taquet
Keywords:
Phosphates
Morocco
Maastrichtian
Turtle
Dermochelyoidae
New
crushing
pattern
a
b
s
t
r
a
c
t
A
new
genus
and
species
of
huge
marine
turtle
(superfamily
Chelonioidea,
epifamily
Der-
mochelyoidae)
is
described
from
the
Maastrichtian
Phosphates
of
the
Oulad
Abdoun
Basin
of
Morocco.
A
new
type
of
feeding
apparatus,
adapted
for
a
powerful
crushing
pattern,
illustrates
the
noteworthy
diversity
of
fossil
vertebrates
of
the
Maastrichtian-Ypresian
Phosphates
of
Morocco.
No
other
crushing
cryptodire
or
bothremydid
pleurodire
has
this
morphology.
During
the
Maastrichtian,
the
known
crushing
pattern
of
chelonioids
was
dif-
ferent,
close
to
that
of
modern
cheloniids,
as
illustrated
in
Morocco
in
the
Maastrichtian
Ganntour
Basin
and
the
Palaeogene
Oulad
Abdoun
Basin.
This
new
taxon
exhibits
unusual
cranial
characters
(fusion
of
premaxillae
associated
with
a
backward
and
dorsal
retraction
of
the
naris,
horizontal
stretching
of
the
dorsal
meatus
quadrati),
that
are
shared
only
with
another
new
turtle,
known
also
from
the
same
Maastrichtian
Phosphates
of
Morocco.
©
2013
Académie
des
sciences.
Published
by
Elsevier
Masson
SAS.
All
rights
reserved.
Mots
clés
:
Phosphates
Maroc
Maastrichtien
Tortue
Dermochelyoidae
Nouveau
type
broyeur
r
é
s
u
m
é
Un
nouveau
genre
et
une
nouvelle
espèce
de
grande
tortue
marine
(superfamille
Chelo-
nioidea,
épifamille
Dermochelyoidae),
des
Phosphates
maastrichtiens
du
Bassin
des
Oulad
Abdoun,
Maroc,
sont
décrits.
Un
nouveau
type
d’appareil
alimentaire,
conc¸
u
pour
un
régime
broyeur
puissant,
est
représenté,
illustrant
la
remarquable
diversité
en
vertébrés
fossiles
des
Phosphates
maastrichtiens-yprésiens
du
Maroc.
Aucun
autre
cryptodire
ou
pleurodire
bothrémydidé
broyeur
n’a
cette
morphologie.
Au
Maastrichtien,
le
seul
mode
broyeur
de
chélonioïdé
connu
était
différent,
proche
de
celui
des
Cheloniidae
modernes,
tel
qu’illustré
au
Maroc
à
la
fois
dans
le
Bassin
des
Ganntour
au
Maastrichtien
et
dans
le
Bassin
des
Oulad
Abdoun
au
Paléogène.
Ce
nouveau
taxon
présente
des
caractères
crâniens
inhabituels
(fusion
des
prémaxillaires,
associée
au
recul
dorsal
de
la
narine,
étirement
à
l’horizontale
de
la
partie
dorsale
du
meatus
quadrati)
qu’il
ne
partage
qu’avec
une
autre
nouvelle
tortue,
elle
aussi
connue
dans
les
mêmes
Phosphates
maastrichtiens
du
Maroc.
©
2013
Académie
des
sciences.
Publié
par
Elsevier
Masson
SAS.
Tous
droits
réservés.
∗Corresponding
author.
E-mail
addresses:
fdelap@mnhn.fr,
france.delapparent123@orange.fr
(F.
de
Lapparent
de
Broin).
1631-0683/$
–
see
front
matter
©
2013
Académie
des
sciences.
Published
by
Elsevier
Masson
SAS.
All
rights
reserved.
http://dx.doi.org/10.1016/j.crpv.2013.07.008

Author's personal copy
88
F.
de
Lapparent
de
Broin
et
al.
/
C.
R.
Palevol
13
(2014)
87–95
1.
Introduction
The
great
diversity
of
fossil
vertebrates
in
the
Phos-
phates
of
Morocco
has
been
known
since
Arambourg
(1952)
and
has
been
enhanced
these
last
fifteen
years
by
new
palaeontological
studies
undertaken
by
French-
Moroccan
collaborations.
In
these
Maastrichtian-Ypresian
phosphatic
deposits
that
crop
out
mainly
in
the
Oulad
Abdoun
and
Ganntour
basins,
intensively
exploited
for
economic
resources
(see
Bardet
et
al.,
2010
for
details),
turtles
are
particularly
well
represented
by
both
cryptodirans
and
pleurodirans
(Bardet
et
al.,
2010,
2013,
in
press;
Gaffney
et
al.,
2006;
Gmira,
1995;
Hirayama
and
Tong,
2003;
Jalil
et
al.,
2009;
Lapparent
de
Broin,
2000;
Tong
and
Hirayama,
2002,
2004,
2008;
Tong
and
Meylan,
2013).
Most
turtle
specimens
have
been
unearthed
from
Palaeogene
phosphatic
deposits;
Maastrichtian
discover-
ies
remain
rare
up
to
now.
The
new
turtle
here
described
is
the
fourth
taxon
from
the
Maastrichtian
levels,
along
with
Euclastes
sp.
from
Ben
Guérir
(Ganntour
Basin)
(Gmira,
1995;
Jalil
et
al.,
2009),
an
indeterminate
chelonioid
from
the
Oulad
Abdoun
Basin
(Tong
and
Hirayama,
2004)
based
only
on
a
plastral
fragment
that
could
correspond
to
the
present
new
taxon
(only
known
by
a
skull
up
to
now),
and
a
new
astonishing
giant
longirostrine
taxon
from
the
Maas-
trichtian
of
Sidi
Chennane
(Oulad
Abdoun
Basin)
(Bardet
et
al.,
2013).
2.
Geological
and
stratigraphical
settings
The
phosphatic
deposits
of
Morocco
are
part
of
the
Mediterranean
Tethyan
phosphogenic
province,
which
extends
from
North
Africa
to
the
Middle-East
(Lucas
and
Prévôt-Lucas,
1996).
They
crop
out
in
four
basins,
the
most
important
and
only
economically
exploited
ones
being
the
Oulad
Abdoun
and
Ganntour
ones
(see
Bardet
et
al.,
2010
for
details)
(Fig.
1A).
Stratigraphically,
they
extend
from
the
Late
Cretaceous
(Maastrichtian)
to
the
base
of
the
Middle
Eocene
(Lutetian),
spanning
the
largest
interval
of
time
of
all
Tethyan
Phosphates
(Lucas
and
Prévôt-Lucas,
1996).
The
Maastrichtian
phosphatic
series
of
the
Oulad
Abdoun
Basin
(“Couche
III”
of
the
miners,
or
Level
III)
is
very
condensed
in
the
Grand
Daoui
zone,
northeastern
part
of
the
basin
(near
Oued
Zem
city),
being
only
about
2–5
m
thick.
It
is
composed
from
the
bottom
to
the
top
of
a
basal
grey
bone-bed,
soft
yellow
(Lower
Level
III)
and
soft
grey
(Upper
Level
III)
exploited
Phosphates,
separated
by
a
yel-
low
marly
level
(Fig.
1B).
The
type
specimen
of
the
new
taxon,
a
complete
skull
with
its
associated
mandible,
was
discovered
in
2006
by
local
people
in
the
Upper
Level
III
(Upper
Maastrichtian)
of
the
Sidi
Daoui
area
of
Grand
Daoui
zone,
northeastern
part
of
the
Oulad
Abdoun
Basin.
Institutional
Abbreviations:
CNRS,
Centre
National
de
la
Recherche
Scientifique
(France);
MEMEE,
Ministère
de
l’Énergie,
des
Mines,
de
l’Eau
et
de
l’Environnement
(Rabat,
Morocco),
MNHN,
Muséum
National
d’Histoire
Naturelle
(Paris,
France),
RA
–
AC,
Reptiles
and
Amphibians
–
Comparative
anatomy;
OCP,
Office
Chérifien
des
Phosphates,
Service
géologique
(Khouribga,
Morocco).
3.
Systematic
palaeontology
Order:
CHELONII
Latreille,
1800
Suborder:
CRYPTODIRA
Cope,
1868
Superfamily:
CHELONIOIDEA
Oppel,
1811
Epifamily:
DERMOCHELYOIDAE
Fitzinger,
1843
Genus
Alienochelys
nov.
Alienochelys
selloumi
nov.
sp.
Holotype:
OCP
DEK/GE
393,
a
complete
large
skull
with
its
associated
lower
jaw
and
the
axis
lying
in
the
orbit;
the
left
neural
arch
of
atlas.
OCP
collections,
Khouribga
(Morocco)
(Figs.
2
and
3).
No
referred
material
known
up
to
now.
Derivatio
nominis:
Genus
name
from
aliena:
stranger
in
latin,
and
chelys:
turtle
in
latin
(issued
from
greek
␹␧␭’
␷␨);
species
name
in
honor
to
Mr.
Omar
Selloumi,
OCP
technician
geologist
(now
retired)
at
the
Geological
Sur-
vey
in
Khouribga,
in
acknowledgement
of
his
constant
help
and
friendship
since
fifteen
years
during
our
fieldwork
and
stays
in
Khouribga.
Type
locality
and
horizon:
Trench
TS,
Sidi
Daoui
area,
Grand
Daoui
zone,
Northeastern
part
of
the
Oulad
Abdoun
Basin
near
the
city
of
Oued
Zem,
Khouribga
Province,
Morocco,
(Fig.
1A).
Phosphatic
deposits,
near
the
base
of
the
Upper
Level
III
(ULIII)
(Fig.
1B),
Upper
Maastrichtian
(Uppermost
Cretaceous)
(Cappetta,
1987).
Diagnosis:
For
genus
and
species,
by
monotypy.
Alienochelys
selloumi
nov.
gen.
nov.
sp.
is
a
gigantic
chelo-
nioid
turtle
(holotype
maximal
skull
length
without
lower
jaw:
41,5
cm)
belonging
to
the
epifamily
Dermochelyoidae
notably
by
full
reduction
of
the
medial
ventral
process
of
the
jugal
beneath
the
orbit,
not
contacting
the
pterygoid
and
the
maxilla,
as
well
as
by
palatines
meeting
medially
ventroposteriorly
to
the
vomer.
It
differs
from
all
other
turtles
by
its
lower
and
upper
jaw
triturating
surfaces,
considerably
developed
and
constituting
a
powerful
crus-
hing
apparatus
that
forms
a
half
circle
arch
with
rounded
extremities,
developed
as
follows:
•lower
jaw:
wide
and
dorsoventrally
flattened
fused
den-
taries,
with
surface
barely
wider
at
the
coronoid-dentary
suture
level
than
at
the
symphysis
area
one,
dorsally
barely
undulated
with
a
small
tomial
rolled
border;
•upper
jaw:
anteroposteriorly
gently
inclined,
very
widely
protruding
snout,
with
a
low
triturating
surface
poorly
concave,
also
nearly
as
wide
medially
as
laterally
and
with
a
narrowly
rounded
tomial
border;
wide
and
long
triturating
surfaces,
anterolaterally
constituted
by
both
long
and
wide
maxillae
and
fused
premaxillae
and
medi-
ally
by
the
vomer
and
palatines
that
are
linked
anteriorly
to
the
choanae;
but,
unlike
other
crushing
chelonioids,
no
secondary
palate,
the
external
naris
being
pushed
backward
so
that
it
is
superimposed
to
the
choanae
and
the
air
conduct
being
vertical
between
the
naris
and
the
choanae;

Author's personal copy
F.
de
Lapparent
de
Broin
et
al.
/
C.
R.
Palevol
13
(2014)
87–95
89
Fig.
1.
A.
Geographical
location
of
the
main
phosphatic
basins
of
Morocco.
OCP
DEK/GE
393
comes
from
the
Sidi
Daoui
area,
in
Grand
Daoui
zone,
north-
eastern
part
of
the
Oulad
Abdoun
Basin.
B.
Grand
Daoui
quarrying
area,
synthetic
stratigraphical
column.
ULIII,
Upper
Level
III
(where
Alienochelys
has
been
found);
LLIII,
Lower
Level
III.
Fig.
1.
A.
Localisation
géographique
des
principaux
bassins
à
Phosphates
du
Maroc.
OCP
DEK/GE
393
provient
de
Sidi
Daoui,
dans
la
zone
de
Grand
Daoui,
Nord-Est
du
Bassin
des
Oulad
Abdoun.
B.
Zone
d’exploitation
de
Grand
Daoui,
colonne
stratigraphique
synthétique.
ULIII,
Couche
III
supérieure
(où
Alienochelys
a
été
trouvée)
;
LLIII,
Couche
III
inférieure.
•upper
and
lower
jaw
triturating
surfaces
all
along
partic-
ularly
rough
with
short
crests
and
protuberances
unlike
other
crushing
chelonioids.
Below
orbit
and
jugal,
considerably
elongated
pos-
terolateral
maxillary
process,
extending
posteriorly
up
to
the
anterolateral
border
of
the
quadrate,
excluding
jugal
and
quadratojugal
from
the
moderate
cheek
emargina-
tion.
Foramen
posterius
canalis
carotici
interni
forming
a
ventrally
open
groove,
at
the
basioccipital-basisphenoid
and
pterygoid
contact,
and
carotid
running
into
a
closed
canalis
carotici
interni
between
pterygoid
and
basiphenoid.
No
scute
sulci.
Alienochelys
selloumi
nov.
gen.
nov.
sp.
shares
some
unusual
characters
with
only
another
new
chelonioid
from
the
coeval
Maastrichtian
Phosphates
of
Morocco
(in
this
work
called
the
“other
new
Maastrichtian
Moroccan
form”)
that
are:
anteroposteriorly
inclined
snout
with
orbits
fac-
ing
anterolaterodorsally,
fusion
of
the
premaxillae,
dorsal
and
nearly
horizontal
external
naris
pushed
back
between
the
orbits
and
superimposed
on
the
choanae,
which
are
latero-anteriorly
framed
by
the
maxillae
processes
(which
are
horizontal
instead
of
vertical),
small
lateral
prefrontals
located
lateral
to
nasals,
and
with
processes
extending
horizontally
below
the
maxillae
(before
descending
anteri-
orly).
It
also
shares
with
this
other
new
taxon
a
horizontally
stretched
meatus
quadrati
area,
facing
ventrally
in
a
half
circle
shape,
with
the
incisura
columellae
auris
ventrally
open.
The
new
turtle
differs
from
the
other
one
by
the
pres-
ence
of
anterior
horizontal
processes
of
the
nasals
framing
lateroposteriorly
the
external
naris,
frontals
not
reaching
the
orbits
and
parietals
not
contacting
the
squamosals,
and
finally,
an
unique
crushing
apparatus,
totally
dif-
ferent
from
the
suction
feeding
one
of
the
other
new
Maastrichtian
Moroccan
form
(distinguished
by
a
bony-
pipette
longirostrine
snout).
Description:
The
holotype
skull,
preserved
with
its
associated
attached
lower
jaw
and
not
completely
pre-
pared
interiorly,
exhibits
two
fractures
on
its
anterior
left
part
(Fig.
2C)
and
is
slightly
damaged
ventroposteriorly
in
some
places
(Fig.
2B).
The
skull
is
rather
low,
roughly
rectangular
(median
skull
length:
37.5
cm;
maximal
width:
35.5
cm;
full
skull
length
with
lower
jaw:
55.5
cm;
maximal
dentary
width:
33
cm,
symphysis
length:
7.8
cm)
with
a
long
and
wide
snout
(wider
than
the
skull
at
the
orbital
level)
anteri-
orly
inclined,
and
large
orbits
(12.1
×
10.6
cm).
Snout
and
orbit
portions
constitute
half
the
skull
length.
The
skull
table
is
irregularly
bulged
dorsally.
There
are
no
scute
sulci
but
marked
nutritive
foramina
and
short
canals
on
the
whole
external
skull
surface.
Some
radiating
rounded
ridges
issue
from
a
center
that
is
situated
rather
anteriorly
on
the
midline
suture
between
the
parietals
(Fig.
2A).
Much
more
pronounced
foramina
located
on
the
snout
indicate
the
emplacement
of
the
upper
rhamphotheca
(Fig.
2C)
while
the
ventral
surface
of
the
dentary
is
covered
with
small
protruding
polygons,
completing
the
dorsal
rough
surface
and
marking
the
emplacement
of
the
lower
rham-
photheca
part
(Fig.
2B).
Skull
structures
decrease
in
size
as
follows:

Author's personal copy
90
F.
de
Lapparent
de
Broin
et
al.
/
C.
R.
Palevol
13
(2014)
87–95
Fig.
2.
Alienochelys
selloumi
nov.
gen.
nov.
sp.
Oulad
Abdoun
Basin,
Morocco,
Late
Maastrichtian.
Holotype
OCP
DEK/GE
393;
photographs
of
the
skull
with
lower
jaw
in
A,
dorsal;
B,
ventral;
C,
left
lateral
slightly
dorsal
with
the
axis
lying
in
the
left
orbit,
in
right
lateral
view;
D,
dorso-posterior;
and
E,
fronto-dorsal
views.
Atlas,
F,
left
neural
arch
with
a
fragment
of
the
intercentrum,
left
lateral
view.
Chelonia
mydas,
MNHN
RA,
AC
1975-19,
G,
atlas,
left
neural
arch,
H,
axis,
left
lateral
views.
Scale
bars,
10
cm.
Fig.
2.
Alienochelys
selloumi
nov.
gen.
nov.
sp.
Bassin
des
Oulad
Abdoun,
Maroc,
Maastrichtien
supérieur.
Holotype
OCP
DEK/GE
393
;
photographies
du
crâne
avec
mandibule
en
vues
:
A,
dorsale
;
B,
ventrale
;
C,
latérale
gauche
légèrement
dorsale,
avec
l’axis
situé
dans
l’orbite
gauche
en
vue
latérale
droite
;
D,
dorso-postérieure
;
et
E,
fronto-dorsale.
Atlas
:
F,
arc
neural
gauche
avec
fragment
d’intercentre,
vue
latérale
gauche.
Chelonia
mydas,
MNHN
RA,
AC
1975-19,
G,
atlas,
arc
neural
gauche,
H,
axis,
vues
latérales
gauches.
Échelles,
10
cm.
•the
rectangularo-ovoid
orbits;
•the
suborbital
arch,
constituted
by
both
the
jugal
and
maxillary
posterior
process
situated
one
above
the
other
all
along
the
jugal
length,
from
the
mid
orbit
length
up
to
their
posterior
extremity;
•the
interorbitary
space;
•the
rectangular
external
naris.
Through
the
external
naris
opening
the
vomer
is
visible,
contacting
the
palatines
and
maxillae.
Laterally,
the
con-
tact
of
the
descending
process
of
the
prefrontal
with
the
maxilla
and
the
dorsal
border
of
the
foramen
orbitonasale
is
visible,
but
its
contact
with
the
vomer
remains
unclear.
Ventrally,
posterior
to
the
triturating
surface,
the
vomer
forms
a
vertical
short
pillar
that
is
not
hidden
by
the
tritu-
rating
part
of
the
linked
vomer
and
palatines,
the
triangular
choanae
remaining
well
visible
on
each
side.
The
posterior
process
of
the
vomer
is
long
between
palatines,
but
does
not
reach
the
pterygoid
(shortly
sutured
palatines).
The
external
pterygoid
processes
(slightly
eroded
on
the
right
side)
are
reduced,
without
any
vertical
flange
posteriorl
to
the
slight
palatine
narrowing.
This
is
due
to
the
reduction
of
the
foramen
palatinum
posterius
and
the
fully
reduced
medioventral
process
of
the
jugal.
The
pterygoids
are
wide,
not
narrowed
and
medially
flat;
they
extend
lateroposte-
riorly
up
to
the
basiphenoid-basioccipital
boundary
and
inferiorly
extend
partly
on
the
articular
quadrate
processes.
The
moderately
large
and
flat
basisphenoid
has
an
ante-
riorly
rounded
shape.
The
carotid
canal
is
not
completely
embedded
in
bone.
It
begins
posteriorly,
not
covered
ven-
trally
at
the
basioccipital
–
pterygoid
boundary,
and
then
it
runs
anteriorly,
covered
ventrally
between
the
pterygoid

Author's personal copy
F.
de
Lapparent
de
Broin
et
al.
/
C.
R.
Palevol
13
(2014)
87–95
91
Fig.
3.
Alienochelys
selloumi
nov.
gen.
nov.
sp.,
Oulad
Abdoun
Basin,
Morocco,
Late
Maastrichtian.
Holotype
OCP
DEK/GE
393;
interpretative
drawings
of
the
skull
with
lower
jaw
in:
A,
dorsal,
B,
ventral,
C,
left
lateral
slightly
dorsal,
and
D,
frontal
views
with
the
axis
lying
in
the
left
orbit,
in
dorsal
view.
cm.
Areas
with
crosses:
not
visible
sutures.
Scale
bars,
10
cm.
ang,
angular;
art,
articular;
ax,
axis;
b
roll,
tomial
border
roll;
boc,
basioccipital;
bs,
basisphenoid;
c
art
q,
condylus
articularis
quadrati;
ch,
choana;
c
mec,
canalis
meckelii;
co,
condylus
occipitalis;
cor,
coronoid;
c
soc,
crista
supraoccipitalis;
dent,
dentary;
ex
na,
external
naris;
fon,
foramen
orbitonasale;
fpci,
foramen
posterius
canalis
carotici
interni;
fr,
frontal;
fti,
fossa
temporalis
inferior;
ica,
incisura
columellae
auris;
ju,
jugal;
l
em,
lateral
emargination;
mc,
muscular
crest;
mq,
meatus
quadrati;
mx,
maxilla;
na,
nasal;
op,
opisthotic;
orb,
orbit;
pa,
parietal;
pal,
palatine;
part,
prearticular;
pfr,
prefrontal;
pmx,
premaxilla;
po,
postorbital;
post
em,
posterior
emargination;
pter,
pterygoid;
pto,
processus
trochlearis
oticus;
q,
quadrate;
qj,
quadratojugal;
sang,
surangular;
soc,
supraoccipital;
sq,
squamosal;
vo,
vomer.
Fig.
3.
Alienochelys
selloumi
nov.
gen.
nov.
sp.,
Bassin
des
Oulad
Abdoun,
Maroc,
Maastrichtien
supérieur.
Holotype
OCP
DEK/GE
393,
dessins
interprétatifs
du
crâne
avec
mandibule
en
vues
:
A,
dorsale,
B,
ventrale,
C,
latérale
gauche
légèrement
dorsale
et
D,
frontale,
avec
l’axis
situé
dans
l’orbite
gauche
en
vue
dorsale.
Zones
avec
croix
:
sutures
non
visibles.
Échelles,
10
cm.
and
the
basisphenoid;
as
a
result,
the
posterior
foramen
of
the
carotid
is
not
embedded
into
the
pterygoid
extrem-
ity.
In
the
inferior
temporal
fossa,
the
cryptodiran
processus
trochlearis
oticus
is
barely
visible
ventrally
(Figs.
1B
and
2B),
largely
constituted
by
the
quadrate:
no
prootic
participa-
tion
is
visible
at
least
ventrally
(incomplete
preparation).
It
constitutes
a
wide
facet,
anteriorly
oriented
and
slightly
laterally
protruding,
with
a
concave
anterior
border.
The
condylus
articularis
quadrati
is
wide,
rectangular,
with
rounded
angles
and
is
slightly
biconcave.
The
occipital
condyle
is
tripartite,
formed
by
the
two
exoccipitals
and
the
basioccipital,
and
is
located
behind
and
above
the
rounded
tuberculum
basioccipitale
(better
preserved
on
the
right).
The
columella
auris
groove
is
ventrally
visible
on
the
left
side,
at
the
base
of
the
quadrate
articular
process.
The
pos-
teroventral
part
of
the
squamosals
has
a
medial
longitudi-
nal
muscular
crest.
The
supraoccipital
crest
shortly
appears
posterior
to
the
condyle.
Laterally,
the
maxillae
are
strink-
ingly
so
much
posteriorly
expanded
up
to
the
quadrate
that
they
exclude
both
the
jugal
and
quadratojugal
from
the

Author's personal copy
92
F.
de
Lapparent
de
Broin
et
al.
/
C.
R.
Palevol
13
(2014)
87–95
cheek
emargination.
The
quadratojugal
and
dorsal
part
of
the
quadrate
are
horizontally
stretched,
so
that
the
meatus
quadrati
forms
a
half
circular
structure
ventrally
facing
with
the
incisura
columellae
auris.
However,
the
articular
process
of
the
quadrate
is
ventrally
not
stretched
and
well
protrud-
ing.
The
antrum
postoticum
is
moderately
developed.
The
lower
jaw
of
Alienochelys
presents
numerous
adap-
tations
for
a
strong
crushing
diet
above
described,
such
as
a
semi-circular
broadly
enlarged
shape,
global
robustness,
particularly
medially
flattening
with
anterior
thinning,
and
roughness
surfaces.
On
the
other
hand,
it
possesses
the
gen-
eral
structure
of
a
moderately
advanced
chelonioid
in
the
evolutionary
schema
presented
in
the
various
phylogenetic
analyses
such
as
those
of
Hirayama
(1995),
Kear
and
Lee
(2006)
and
Parham
and
Pyenson
(2010),
with
a
dentary
not
expanded
posteriorly
toward
the
articular
surface
and
a
moderately
anteriorly
developed
surangular.
The
coronoid
is
wide,
low
and
rounded,
and
integrated
into
the
triturat-
ing
surface
(Fig.
2A,
B).
The
area
posterior
to
the
triturating
surface
is
as
long
as
it,
and
robust
to
support
its
weight.
The
axis
is
preserved
into
the
left
orbit
(posteriorly
fac-
ing)
(Figs.
2C
and
3C)
and
a
part
of
the
atlas
(neural
arch
with
a
fragment
of
the
intercentrum)
was
preserved
in
the
matrix
just
lateral
to
the
skull
(Figs.
2F
and
3D).
They
are
similar
to
those
of
Chelonia,
being
just
slightly
rela-
tively
deeper.
The
atlas
postzygapophyses
are
slightly
more
robust.
The
axis
postzygapophyses
are
more
curved
and
the
axis
anterior
process
is
more
dorsally
protruding.
4.
Phylogenetic
Relationships
4.1.
Material
and
Methods
Alienochelys
selloumi
nov.
gen.
nov.
sp.
has
been
included
into
a
matrix
(103
characters,
22
taxa)
slightly
modified
from
that
of
Bardet
et
al.
(2013),
performed
for
the
other
new
Maastrichtian
Moroccan
form,
itself
slightly
modified
from
that
of
Kear
and
Lee
(2006)
based
on
Hirayama
(1998).
See
the
Electronic
appendix
(complementary
details,
char-
acters,
results
and
various
obtained
cladograms).
4.2.
Comparative
results
The
absence
of
postcranial
material
and
the
impossi-
bility
of
observing
the
inner
skull
characters
do
not
permit
us
to
detect
all
the
chelonioid
apomorphies
corresponding
to
the
character
analysis
(such
as
those
of
the
paddles,
dorsum
sellae,
rostrum
basisphenoidale,
position
of
inner
carotid
foramina
in
the
sella
turcica,
relative
size
of
the
inner
arterial
canals
and
of
the
processus
inferior
parietalis).
However,
the
results
(Fig.
4;
Appendix
Figs.
1
to
3)
clearly
show
that,
among
Chelonioidea,
Alienochelys
nov.
gen.
belongs
to
the
epifamily
Dermochelyoidae
(sensu
Bour
and
Dubois,
1986,
emend.
Gaffney
and
Meylan,
1988),
sharing
several
synapomorphies
of
the
superfamily
(homoplastic
here
or
there
in
few
Chelomacryptodira
and
Plesiochelyi-
dae)
including:
the
complete
ventral
reduction
of
the
jugal
below
the
orbit,
not
contacting
the
pterygoid
(10/1
and
11/1),
and
the
palatines
posteriorly
meeting
(vomer
not
contacting
the
pterygoids
(20/1))
as
in
other
taxa
of
the
epifamily
(except
Dermochelys,
but
by
reversion
as
shown
Fig.
4.
Strict
consensus
tree
obtained
from
the
matrix,
slightly
mod-
ified,
provided
in
Bardet
et
al.
(2013),
for
another
new
chelonioid
from
the
Maastrichtian
Phosphates
of
Moroccan.
Modified
from
Kear
and
Lee
(2006)
with,
as
outgroups,
an
hypothetical
taxon
and
Che-
lomacryptodira,
and
including
Alienochelys
selloumi
nov.
gen.
nov.
sp.
A.
Chelonioidea.
B.
Cheloniidae.
C.
Dermochelyoidae.
D.
Protostegidae.
L
=
222,
Ci
=
51,
Ri
=
59.
Fig.
4.
Arbre
de
Consensus
Strict
obtenu
à
partir
de
la
matrice
légère-
ment
modifiée
produite
par
Bardet
et
al.
(2013)
pour
un
autre
chélonioïdé
nouveau
des
Phosphates
maastrichtiens
du
Maroc.
Modifiée
de
Kear
et
Lee
(2006),
avec,
comme
groupes
externes,
un
taxon
hypothétique
et
les
Chelomacryptodira
et
avec
addition
d’Alienochelys
selloumi
nov.
gen.
nov.
sp.
A.
Chelonioidea.
B.
Cheloniidae.
C.
Dermochelyoidae.
D.
Protostegidae.
L
=
222,
Ci
=
51,
Ri
=
59.
by
the
visible
anterior
extraordinary
pterygoid
expansion).
The
absence
of
foramen
palatinum
posterius
(21/1)
is
also
shared
with
Dermochelyoidae
in
which
the
character
reduction
presents
two
states:
•the
most
basal
state
(here
shown),
with
the
foramen
lat-
erally
widely
open,
making
a
weak
lateral
notch
anterior
to
the
pterygoid
process,
that
is
still
slightly
protruded
as
in
Bouliachelys
Kear
and
Lee,
2006,
the
primitive
protostegids
Santanachelys
Hirayama,
1998,
Rhinochelys
pulchriceps
(Owen,
1851)
(Collins,
1970),
and
Desma-
tochelys
lowi
Williston,
1894
(Zangerl
and
Sloan,
1960)
(Hirayama,
1995)
(particularly
morphologically
close
in
that
point
to
Alienochelys
nov.
gen.
due
to
an
elongation
of
the
maxillary-palatine
gulf);
•the
pterygoid
process
and
lateral
notch
fully
disap-
pear
together
with
a
strong
pterygoid
narrowing
in
the
advanced
Protostegidae
Protostega
gigas
Cope,
1872
(Hirayama,
1995),
Archelon
Wieland,
1896
(Hay,
1908),
and
Calcarichelys
gemma
Zangerl,
1953a
(Hooks,
1998).
The
jugal-pterygoid
contact
is
also
lost
in
the
Maas-
trichtian
cheloniid
Allopleuron
hofmanni
(Gray,
1831),
the
jugal
being
ventrally
partly
reduced,
but
the
foramen
palat-
inum
posterius
is
present,
small
and
often
laterally
fissured,
and
the
pterygoid
process
is
not
reduced.
The
foramen
fully
disappears
in
all
other
Cheloniidae
with
a
secondary
palate,
and
derived
compared
to
Toxochelys
and
Ctenochelys
(Zangerl,
1953b),
but
in
which
the
jugal-pterygoid
contact
is
preserved
(Hirayama,
1995).
The
absence
of
foramina

Author's personal copy
F.
de
Lapparent
de
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et
al.
/
C.
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(2014)
87–95
93
praepalatina
(14/1)
is
shared
by
the
Dermochelyoidae
and
all
Cheloniidae
more
derived
than
Toxochelys.
However,
Alienochelys
nov.
gen.
is
not
a
cheloniid
because
of
its
pterygoid
(not
contacting
the
jugal
which
is
reduced
here
(10,
11)),
the
lack
of
any
sagittal
crest
unlike
most
cheloniids
(23/0),
its
basisphenoid
without
a
V-shaped
crest
and
posterolateral
projections
(31/0),
and
finally
because
of
the
preservation
of
other
primitive
char-
acters,
beside
the
advanced
Dermochelyoidae
characters
given
above.
The
posterior
carotid
foramen
(29/1),
located
posteriorly
but
not
fully
embedded
into
the
pterygoids,
is
primitive
in
relation
to
Cheloniidae,
slightly
less
derived
than
in
the
other
new
Maastrichtian
Moroccan
form
(29/2),
but
derived
in
relation
to
Protostegidae
(29/0).
The
absence
of
a
parietal-squamosal
contact
(7/1)
is
variable
in
Der-
mochelyoidae,
so
it
does
not
appear
to
be
decisive.
In
chelonioids,
the
deep
modification
of
the
limbs
into
rigid
paddles
reflects
adaptation
to
an
open
marine
life
–
the
fore
paddles
being
much
longer
than
the
hind
ones
–
by
a
series
of
changes
that
may
vary
according
to
the
groups
and
is
very
important
in
the
resulting
relation-
ships
of
the
taxa.
So,
if
Alienochelys
nov.
gen.
is
clearly
a
Dermochelyoidae,
its
current
lack
of
postcranial
material
means
that
its
ingroup
relationships
remain
unresolved
with
respect
to
Dermochelys
and
Protostegidae.
For
compa-
rable
reasons,
the
affinities
of
Bouliachelys
and
of
the
other
new
Maastrichtian
Moroccan
form
(Bardet
et
al.,
2013)
also
remain
unresolved.
Alienochelys
nov.
gen.
approaches
the
protostegid
con-
dition
by
an
expansion
of
the
pterygoid
on
the
quadrate
articular
process
(24/1)
that
however
remains
less
exten-
sive,
as
in
the
other
new
Maastrichtian
Moroccan
form
(24/0),
although
slightly
more
extended
in
Alienochelys
nov.
gen.
than
in
the
latter
because
it
reaches
the
condyle
(Bardet
et
al.,
2013.
See
Electronic
appendix).
Its
rela-
tive
basal
position
and
derivation
is
explained
by
the
presence
of
a
combination
of
primitive
characters,
some
of
them
being
shared
with
the
other
new
Maastrichtian
Moroccan
form
(long
nasals
and
lateral
position
of
the
prefrontals,
short
crista
supraoccipitalis,
relatively
short
posterior
emargination),
as
with
the
most
primitive
pro-
tostegids
(those
presenting
various
relative
shape
and
position
of
the
bones
surrounding
the
external
naris
and
orbits)
(Hirayama,
1995).
Both
Moroccan
forms
are
derived
in
relation
to
cheloniids
and
primitive
protostegids
by
the
absence
of
scute
sulci
(1/1)
(like
Dermochelys
and
advanced
protostegids)
and
a
more
posterior
position
of
the
carotid
foramen
(29/1)
than
in
protostegids.
This
character
is
how-
ever
less
derived
in
Alienochelys
than
in
the
other
new
Maastrichtian
Moroccan
form
in
which
the
carotid
fora-
men
is
located
as
posteriorly
as
in
Dermochelys
and
like
that
of
Cheloniidae
(29/2).
Moreover,
they
are
the
only
two
taxa
to
share
the
following
derived
characters,
not
yet
included
in
the
performed
analyses,
and
that
probably
indi-
cate
close
relationships:
fused
premaxillae,
posterodorsal
migration
of
the
external
naris
and
consecutive
modifi-
cation
of
the
surrounding
bones
and
structures,
quadrate
meatus
area
becoming
horizontally
stretched,
and
ventral
orientation
of
the
incisura
columellae
auris
notch.
Among
these,
the
random
distribution
in
Dermochelyoidae
of
some
derived
features
approaching
those
of
these
two
taxa
is
noteworthy.
For
example,
Dermochelys
has
a
maxilla
slightly
prolonged
posteriorly
to
the
orbit
(but
not
exclud-
ing
the
jugal
from
the
cheek
emargination)
(unlike
the
other
new
Maastrichtian
Moroccan
form)
and
an
incisura
facing
rather
ventrally
(like
both
Moroccan
forms,
but
with-
out
meatus
stretching).
In
Protosteginae
(Hooks,
1998;
Wieland,
1909;
Zangerl,
1953a),
the
skull
remains
poorly
described:
Archelon
ischyros
Wieland,
1896
and
Protostega
gigas
Cope,
1872
seem
to
have
an
external
naris
open-
ing
dorsally
and
partly
moved
above
the
choanae,
and
the
anterior
part
of
the
palate
of
Protostega
gigas
could
be
elon-
gated
without
hiding
the
choanae
(Hirayama,
1995,
Figs.
2i
and
3i)
whereas
Microstega
copei
(Wieland,
1909)
has
an
incisura
columellae
auris
ventrally
facing
(but
without
meatus
stretching).
This
new
chelonioid,
Alienochelys
nov.
gen.,
exhibiting
a
new
type
of
feeding
apparatus
among
the
turtles
from
the
Maastrichtian
Phosphates
of
the
Oulad
Abdoun
Basin,
is
adapted
for
a
powerful
crushing
diet
of
hard
preys.
As
in
the
other
new
Maastrichtian
Moroccan
form,
(a
suc-
tion
feeder
provided
by
an
unique
bony
elongated
tubular
snout),
whereas
the
choanae
remain
posterior,
the
elon-
gation
of
the
triturating
surfaces
is
accompanied
by
the
step
backward
of
the
external
naris
up
to
be
superimposed
on
the
choanae.
In
cheloniids,
when
the
triturating
sur-
faces
are
elongated,
the
skull
is
provided
with
a
secondary
palate,
permitting
the
conduction
of
air
to
be
prolonged
inside
the
skull
between
the
external
naris
and
the
choanae.
When
these
cheloniids
are
adapted
to
a
crushing
feed-
ing,
the
upper
jaw
fits
with
the
skull
by
an
expanded
lower
jaw
symphysis
(exception
in
“Syllomus
aegyptiacus”
in
Hasegawa
et
al.,
2005,
with
a
short
but
very
dentic-
ulated
dentary)
and
the
triturating
surfaces
are
more
or
less
crested,
directly
on
the
bony
structure
and/or
on
the
rhamphothecae,
differently
according
to
the
preys
and
the
taxa,
but
not
as
in
Alienochelys
nov.
gen.
Among
such
che-
loniids,
are
represented
all
the
members
of
the
“Euclastes
group”
(sensu
Jalil
et
al.,
2009,
i.e.
the
“Durophagous
Stem
Cheloniids”
of
Parham
and
Pyenson,
2010,
includ-
ing
Erquelinnesia,
Pacifichelys
and
Mexichelys),
as
well
as
“Argillochelys”
africana
Tong
and
Hirayama,
2008
and
Tas-
backa
(Tong
and
Hirayama,
2002).
It
should
be
noted
that
these
last
two
taxa
and
Euclastes
are
present
in
the
Phos-
phates
of
Morocco.
They
have
smooth
bony
triturating
surfaces
like
some
living
and
Tertiary
forms.
Other
extant
taxa
and
the
Maastrichtian
cheloniid
Allopleuron
have
much
more
crested
jaws.
In
conclusion,
Alienochelys
nov.
gen.
is
unique
among
chelonioids
by
the
lack
of
a
secondary
palate
(also
absent
in
other
Dermochelyoidae
and
primitive
cheloniids)
despite
of
a
crushing
diet.
Among
Dermochely-
oidae,
Dermochelys
is
a
ram
feeder
of
soft
preys
that
has
very
thin
and
smooth
“triturating”
surfaces,
contrasting
with
the
strong
ridged
and
tuberculated
triturating
sur-
faces
of
Alienochelys
nov.
gen.
However,
like
Dermochelys
and
unlike
the
other
new
Maastrichtian
Moroccan
form,
Alienochelys
bears
a
palatine
extension
up
to
the
choanae
and
a
vertical
short
vomer
pillar
that
is,
like
the
choanae,
not
hidden
ventrally.
Among
sea
turtles,
the
Alienochelys
nov.
gen.
jaw
sur-
faces,
though
exhibiting
a
different
morphology,
evoke
the
crushing
diet
also
present
in
various
bothremydid

Author's personal copy
94
F.
de
Lapparent
de
Broin
et
al.
/
C.
R.
Palevol
13
(2014)
87–95
pleurodires,
a
family
worldwidely
spread
during
the
Maastrichtian-Palaeogene.
In
bothremydids,
the
enlarge-
ment
of
the
surfaces
is
realized
around
the
palatal
medial
sulcus,
without
secondary
palate,
and
the
choanae
are
not
moved
backward.
Among
them,
the
Bothremydini
in
particular
(only
known
during
the
Palaeogene
in
Phos-
phates
of
Morocco
but
present
otherwise
in
the
Late
Cretaceous,
for
example
in
Europe;
Gaffney
et
al.,
2006),
have
wide
and
smooth
dorsal
triturating
surfaces,
later-
ally
expanded
in
width
with
posterior
characteristic
pits.
Although
the
Nigeremydinae
(sensu
“Nigeremys
group”
Broin,
1988,
Lapparent
de
Broin
and
Werner,
1998,
Nigere-
mydina
Gaffney
et
al.,
2006,
2007)
have
strong
ridged
dorsal
triturating
surfaces
on
a
widened
triangular
snout,
with
crests
that
are
parallel
to
the
labial
border
(see
origi-
nal
figures
in
Lapparent
de
Broin
and
Werner,
1998),
they
more
evoke
the
Alienochelys
pattern
by
their
strength
and
mode
of
action.
The
Nigeremydinae
are
known
during
the
Maastrichtian-Palaeocene
in
the
northern
part
of
Africa
(Egypt,
Mali,
Niger),
but
not
in
the
Moroccan
Phosphates
together
with
Alienochelys.
In
these
phosphatic
deposits,
Alienochelys
gen.
nov.
is,
up
to
now,
the
only
Maastrichtian
turtle
adapted
to
a
crushing
diet,
i.e.
occupying
such
an
ecological
niche.
It
shows
that
marine
turtles
were
much
more
diversified
than
previously
thought
in
the
Maas-
trichtian
Phosphates
of
Morocco.
Together
with
the
other
new
Maastrichtian
Moroccan
form
which
also
exhibits
a
unique
suction
feeding
apparatus
among
tetrapods
(Bardet
et
al.,
2013),
they
illustrate
the
important
taxonomic
and
ecological
diversity
of
the
fossil
vertebrates
of
the
Maastrichtian-Ypresian
Phosphates
of
Morocco.
Acknowledgments
This
work
is
part
of
a
French-Moroccan
Palaeontolog-
ical
Convention
of
Collaboration
between
MNHN/CNRS
(Paris),
OCP
(Casablanca),
MEMEE
(Rabat),
and
Cadi
Ayyad
(UCAM,
Marrakech)
and
Chouaîb
Doukkali
(UCDJ,
El
Jadida)
Universities.
We
are
grateful
to
all
OCP
staff
members
for
their
hospitality
and
logistic
support
and,
more
specially,
to
B.
Bouya,
O.
Selloum
and
M.
Bichara.
We
also
thank
N.-E.
Jalil
(UCAM)
for
his
friendship
support
during
field-
work
and
when
taking
photographs,
S.
Xerri
(Rabat)
for
informations,
R.
Vacant
for
preparation
of
the
specimen,
and
R.
Allain
(MNHN,
F,
Paleontology)
and
S.
Bailon
(MNHN,
ZA.AC,
Comparative
Anatomy
collection)
for
access
of
the
collections
under
their
care.
We
are
grateful
to
Roger
Bour,
Juliana
Sterli
and
an
anonymous
referee.
We
thank
Kevin
Padian
for
english
edits.
This
research
has
been
supported
by
funds
from
the
MNHN
(BQR
program,
Dept.
Histoire
de
la
Terre)
and
CNRS/CNRST
programs
of
collaborations
AC
no18567
and
MARPIC
no4892.
Appendix
A.
Supplementary
data
Supplementary
data
(Alienochelys
gen.
nov.
–
Phylogenetical
Analysis
elements)
associated
with
this
article
can
be
found,
in
the
online
version,
at
http://
dx.doi.org/10.1016/j.crpv.2013.07.008.
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Lapparent
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et
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/
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Palevol
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95
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