Fossil and subfossil herpetofauna from Cadet 2 Cave (Marie-Galante, Guadeloupe Islands, F. W. I.): Evolution of an insular herpetofauna since the Late Pleistocene

Abstract

This study deals with the herpetofaunal fossil and subfossil remains from the Cadet 2 site (Marie-Galante, Guadeloupean Archipelago). This study provides new data concerning the herpetofaunal community since the Late Pleistocene by revealing the early local occurrence of some taxa (Eleutherodactylus sp., cf. Sphaerodactylus sp., Ameiva sp., cf. Capitellum maria-galantae, Anolis cf. ferreus, cf. Antillotyphlops sp., cf. Alsophis sp. and Colubroidea sp. 1) and possible Pleistocene extinctions (Boa sp. and Colubroidea sp. 2). Moreover, the first metric data for fossil Marie-Galante anoles show clear size stability throughout time. As regards the evolution of the island herpetofaunal biodiversity, our work points to the long-term stability of the fauna before human colonization and subsequently to the marked impact of human-caused environmental disturbances during colonial but also Pre-Columbian periods.

Full text

Please
cite
this
article
in
press
as:Bochaton,
C.,
et
al.,
Fossil
and
subfossil
herpetofauna
from
Cadet
2
Cave
(Marie-
Galante,
Guadeloupe
Islands,
F.
W.
I.):
Evolution
of
an
insular
herpetofauna
since
the
Late
Pleistocene.
C.
R.
Palevol
(2015),
http://dx.doi.org/10.1016/j.crpv.2014.10.005
ARTICLE IN PRESS
G Model
PALEVO-803;
No.
of
Pages
10
C.
R.
Palevol
xxx
(2015)
xxx–xxx
Contents
lists
available
at
ScienceDirect
Comptes
Rendus
Palevol
w
w
w.sci
encedirect.com
General
palaeontology,
systematics
and
evolution
(Vertebrate
palaeontology)
Fossil
and
subfossil
herpetofauna
from
Cadet
2
Cave
(Marie-Galante,
Guadeloupe
Islands,
F.
W.
I.):
Evolution
of
an
insular
herpetofauna
since
the
Late
Pleistocene
L’herpétofaune
fossile
et
subfossile
de
la
grotte
Cadet
2
(Marie-Galante,
Archipel
de
la
Guadeloupe,
P.-A.
F.)
:
évolution
d’une
herpétofaune
insulaire
depuis
le
Pléistocène
supérieur
Corentin
Bochatona,b,∗,
Sandrine
Grouarda,
Raphaël
Cornetteb,
Ivan
Ineichb,
Arnaud
Lenoblec,
Anne
Tresseta,
Salvador
Bailona
aMuséum
national
d’Histoire
naturelle
(MNHN),
UMR
7209
«
Archéozoologie
et
archéobotanique
:
sociétés,
pratiques
et
environnements
»
(CNRS),
CP
56,
55,
rue
Buffon,
75005
Paris,
France
bMuséum
national
d’Histoire
naturelle
(MNHN),
ISYEB
(UMR
7205
CNRS,
EPHE,
UPMC),
CP
30,
57,
rue
Cuvier,
75251
Paris,
France
cUniversité
de
Bordeaux,
UMR
5199
PACEA,
avenue
des
Facultés,
33405
Talence,
France
a
r
t
i
c
l
e
i
n
f
o
Article
history:
Received
1st September
2014
Accepted
after
revision
26
October
2014
Available
online
xxx
Handled
by
Michel
Laurin
Keywords:
West
Indies
Herpetofauna
Paleontology
Biodiversity
Extinction
Guadeloupe
islands
Faunal
turnover
a
b
s
t
r
a
c
t
This
study
deals
with
the
herpetofaunal
fossil
and
subfossil
remains
from
the
Cadet
2
site
(Marie-Galante,
Guadeloupean
Archipelago).
This
study
provides
new
data
concerning
the
herpetofaunal
community
since
the
Late
Pleistocene
by
revealing
the
early
local
occurrence
of
some
taxa
(Eleutherodactylus
sp.,
cf.
Sphaerodactylus
sp.,
Ameiva
sp.,
cf.
Capitellum
maria-
galantae,
Anolis
cf.
ferreus,
cf.
Antillotyphlops
sp.,
cf.
Alsophis
sp.
and
Colubroidea
sp.
1)
and
possible
Pleistocene
extinctions
(Boa
sp.
and
Colubroidea
sp.
2).
Moreover,
the
first
metric
data
for
fossil
Marie-Galante
anoles
show
clear
size
stability
throughout
time.
As
regards
the
evolution
of
the
island
herpetofaunal
biodiversity,
our
work
points
to
the
long-term
stability
of
the
fauna
before
human
colonization
and
subsequently
to
the
marked
impact
of
human-
caused
environmental
disturbances
during
colonial
but
also
Pre-Columbian
periods.
©
2014
Académie
des
sciences.
Published
by
Elsevier
Masson
SAS.
All
rights
reserved.
Mots
clés
:
Antilles
Herpétofaune
Paléontologie
Biodiversité
Extinction
Guadeloupe
Remplacement
de
faune
r
é
s
u
m
é
Nous
étudions
ici
le
matériel
herpétofaunistique
provenant
du
site
de
Cadet
2,
situé
sur
l’île
de
Marie-Galante,
dans
l’archipel
de
la
Guadeloupe.
L’analyse
de
ces
restes
apporte
des
données
inédites
concernant
le
peuplement
herpétologique
de
l’île
depuis
le
Pléistocène
supérieur.
Elle
met
en
évidence
la
présence
ancienne
de
certains
taxons
sur
l’île
(Eleuthero-
dactylus
sp.,
cf.
Sphaerodactylus
sp.,
Ameiva
sp.,
cf.
Capitellum
mariagalantae,
Anolis
cf.
ferreus,
cf.
Antillotyphlops
sp.,
cf.
Alsophis
sp.
et
Colubroidea
sp.
1).
Elle
montre
aussi
de
possibles
extinctions
datant
du
Pléistocène
(Boa
sp.
et
Colubroidea
sp.
2).
Nos
premiers
résultats
∗Corresponding
author
at:
Laboratoire
“Reptiles
et
Amphibiens”,
57,
rue
Cuvier,
CP
30,
75005
Paris,
France.
E-mail
address:
corentin.bochaton@mnhn.fr
(C.
Bochaton).
http://dx.doi.org/10.1016/j.crpv.2014.10.005
1631-0683/©
2014
Académie
des
sciences.
Published
by
Elsevier
Masson
SAS.
All
rights
reserved.

Please
cite
this
article
in
press
as:Bochaton,
C.,
et
al.,
Fossil
and
subfossil
herpetofauna
from
Cadet
2
Cave
(Marie-
Galante,
Guadeloupe
Islands,
F.
W.
I.):
Evolution
of
an
insular
herpetofauna
since
the
Late
Pleistocene.
C.
R.
Palevol
(2015),
http://dx.doi.org/10.1016/j.crpv.2014.10.005
ARTICLE IN PRESS
G Model
PALEVO-803;
No.
of
Pages
10
2
C.
Bochaton
et
al.
/
C.
R.
Palevol
xxx
(2015)
xxx–xxx
obtenus
sur
les
faibles
variations
de
la
taille
des
anolis
fossiles
de
Marie-Galante
démontrent
la
grande
stabilité
des
peuplements
au
cours
du
temps.
Ce
travail
met
en
avant
la
stabilité
de
l’herpétofaune
de
l’île
durant
les
périodes
précédant
l’arrivée
de
l’homme,
puis
les
mod-
ifications
considérables
qui
suivent
son
installation
pendant
les
périodes
précolombiennes,
puis
modernes.
©
2014
Académie
des
sciences.
Publié
par
Elsevier
Masson
SAS.
Tous
droits
réservés.
1.
Introduction
The
biogeography
of
the
Greater
and
Lesser
Antilles
herpetofauna
has
been
the
subject
of
several
publications
during
the
past
decades
(Hedges,
1996,
2006;
Lescure,
1987;
Lescure
et
al.,
1991).
Thus,
the
present-day
com-
position
and
distribution
of
the
herpetofauna
on
the
Caribbean
Islands
is
well
known
(Henderson
and
Powell,
2009;
Powell
and
Henderson,
2012).
In
the
same
way,
the
modern
herpetofauna
on
the
Guadeloupe
Islands
is
well
documented
and
has
recently
been
the
subject
of
a
monograph
(Breuil,
2002).
In
contrast,
the
fossil
and
subfossil
herpetofauna
is
poorly
known,
in
particular
in
the
Lesser
Antilles
(Pregill
and
Olson,
1981;
Pregill
et
al.,
1988,
1994).
Many
questions
remain
unresolved
as
regards
the
past
Guadeloupean
herpetofaunal
community
and
in
spite
of
several
studies
focusing
on
archaeological
fauna
(Grouard,
2001,
2003,
2007,
2010),
pre-anthropic
fossil
data
for
reptiles
and
amphibians
have
remained
scant
since
the
pioneering
work
of
Pregill
et
al.
(1994).
In
addition,
the
impact
of
human
populations
on
her-
petological
communities
during
Amerindian
and
colonial
periods
is
sparsely
documented
although
some
works
have
attempted
to
tackle
this
question
(Pregill,
1986;
Steadman
et
al.,
1984).
The
recent
exploration
and
excavation
of
fossil-bearing
deposits
in
Guadeloupe
and
Marie-Galante
considerably
improve
our
knowledge
of
past
Guadeloupean
biodiversity
(Grouard
et
al.,
2014;
Lenoble
et
al.,
2009;
Stouvenot
et
al.,
2014).
The
present
study
focuses
on
reptile
and
amphibian
remains
collected
from
one
of
these
sites,
the
Cadet
2
Cave,
which
lies
on
the
southeastern
coast
of
Marie-Galante,
and
documents
the
composition
and
evolution
of
the
Marie-
Galante
herpetofauna
over
the
past
thirty
thousand
years.
Marie-Galante
island
and
site
description
Marie-Galante
(61.223358
W;
15.893134
N)
(Fig.
1)
is
a
small,
low-lying
limestone
island
of
approximately
158
km2(maximum
altitude
=
202
m).
It
probably
emerged
during
the
Late
Calabrian
about
800,000
years
ago
(Münch
et
al.,
2013).
Six
squamate
species
occur
nowadays
on
the
island:
three
of
them
are
considered
to
be
native
species
(Anolis
ferreus,
Sphaerodactylus
fantasticus
anidrotus
and
Thecadactylus
rapicauda)
and
three
others
as
recently
introduced
species
(Iguana
iguana,
Gymnophtalmus
under-
woodi
and
Hemidactylus
mabouia).
There
are
also
three
species
of
frogs,
one
native
(Eleutherodactylus
martinicen-
sis)
and
two
allochthonous
(Eleutherodactylus
johnstonei
and
Scinax
cf.
x-signatus).
Several
other
currently
extinct
taxa
have
been
mentioned
by
naturalists
over
the
past
centuries,
including
three
squamates,
the
scincid
Capitel-
lum
mariagalantae
and
two
snakes
(Alsophis
antillensis
and
Liophis
juliae)
(Breuil,
2002).
In
addition,
the
genus
Iguana
have
been
mention
in
Marie-Galante
archaeologi-
cal
deposits
dated
from
200
to
1000
AD
(Grouard,
2001).
The
Cadet
3
fossil
deposit
(Sierpe,
2011;
Stouvenot
et
al.,
2014)
containing
layers
dated
from
13,800
BC
to
mod-
ern
time
provides
evidence
of
three
additional
extinct
taxa,
hitherto
never
mentioned
on
the
island
(Boa
sp.,
Ameiva
sp.
and
Leiocephalus
cuneus).
Ameiva
was
previously
thought
to
have
been
present
on
Marie-Galante
during
the
past
by
Breuil
(2002),
but
clear
evidence
was
lack-
ing.
Cadet
2
is
a
flank
margin
cave
(Lenoble
et
al.,
2009)
opening
onto
the
cliff
of
the
Capesterre
terrace
near
the
Cadet
3
site,
about
250
m
from
the
coast
(Fig.
1).
The
site
was
first
excavated
by
P.
Courtaud
from
2004
to
2007
in
order
to
record
Amerindian
burials
(Courtaud,
2011;
Courtaud
et
al.,
2005).
Due
to
the
paleontological
poten-
tial
of
the
site
(Lenoble
et
al.,
2009)
another
excavation
was
subsequently
directed
by
S.
Grouard
in
2010
to
inves-
tigate
the
pre-anthropic
layers
and
to
collect
the
vertebrate
remains.
This
last
excavation
focused
on
two
loci
described
below.
The
first
locus
(Fig.
1)
is
located
in
the
deepest
part
of
the
cave.
It
measures
one
square
meter
and
reaches
a
depth
of
1.4
meters.
This
locus
contains
a
large
part
of
the
cave
infill-
ing.
Above
a
basal
accumulation
of
sands
rich
in
seashells
and
land
crab
fragments
(U5-D),
the
deposits
correspond
to
a
succession
of
bedded
silt,
divided
into
three
subunits
following
the
sedimentary
structure
(U5-C
to
U5-A).
The
organic
silt
fraction
was
radiocarbon
dated
at
the
Centre
de
datation
par
le
radiocarbone,
Lyon,
(France).
The
dates
range
between
34,229–31,888
cal.
B.C.
and
11,909–11,530
cal.
B.C.
(Ref.:
Ly
8496–8492)
(Fig.
1).
Above
these
levels,
lies
layer
U4-U4,
corresponding
to
an
undated
non-excavated
archaeological
level,
probably
of
Holocene
age.
An
archae-
ological
pit
was
identified
in
layer
U4
(ST
500
see
Fig.
1).
This
structure
is
contemporaneous
with
the
Amerindian
occupation
of
the
cavity,
estimated
to
date
from
the
14th
century
AD
(Courtaud,
2011;
Courtaud
et
al.,
2005).
The
second
locus
was
excavated
over
half
a
square
meter
and
divided
into
six
levels.
The
radiocarbon
dating
of
a
char-
coal
from
level
U5-b
at
the
Erlangen
laboratory
(Germany)
yielded
ages
of
28,413–27,425
cal.
B.C.
(Ref.:
Erl
14,011).
On
the
basis
of
the
characteristics
of
the
sediment
facies,
the
stratigraphic
units
of
this
locus
were
correlated
with
layers
U5-C
and
U5-B
from
the
first
locus,
and
are
thus
considered
to
be
of
Pleistocene
age.
The
whole
assemblage
is
made
up
of
131,571
osteologi-
cal
remains,
including
57,468
(48%)
reptile
and
amphibian
remains
collected
in
all
the
layers
except
U5-D,
which
con-
tained
only
few
very
fragmented
bones.

Please
cite
this
article
in
press
as:Bochaton,
C.,
et
al.,
Fossil
and
subfossil
herpetofauna
from
Cadet
2
Cave
(Marie-
Galante,
Guadeloupe
Islands,
F.
W.
I.):
Evolution
of
an
insular
herpetofauna
since
the
Late
Pleistocene.
C.
R.
Palevol
(2015),
http://dx.doi.org/10.1016/j.crpv.2014.10.005
ARTICLE IN PRESS
G Model
PALEVO-803;
No.
of
Pages
10
C.
Bochaton
et
al.
/
C.
R.
Palevol
xxx
(2015)
xxx–xxx
3
Fig.
1.
Site
location
and
stratigraphy
of
the
first
locus
with
radiocarbon
dates.
Fig.
1.
Localisation
du
site
et
stratigraphie
du
premier
locus
avec
datations
radiocarbone.
2.
Material
and
methods
The
sediment
samples
from
the
site
were
washed
with
water
using
a
0.5
mm
sieve
mesh.
The
osteologi-
cal
remains
were
collected
with
a
binocular
scope
(Nikon
SMZ1000).
The
osteological
collections
used
for
the
identifi-
cation
of
the
remains
(see
Appendix
1)
come
from
the
Muséum
national
d’Histoire
naturelle
(MNHN,
Paris,
France)
(Comparative
Anatomy—MNHN-ZA-AC—,
Reptile
and
Amphibians—MNHN-RA—and
UMR
7209
“Archéozoologie
et
archéobotanique”
“Caraïbes”—MNHN-
UMR7209—collections)
and
UMR
5199
CNRS
“PACEA”
(PACEA,
Bordeaux,
France).
We
also
referred
to
several
published
osteological
characteristics.
In
this
study,
we
only
describe
the
main
osteological
features
used
for
the
identification
of
each
taxon.
The
classifications
employed
here
derive
from
several
works
concerning
amphibians
(Frost
et
al.,
2006;
Hedges
et
al.,
2008)
and
squamates
(Conrad,
2008;
Townsend
et
al.,
2011),
including
some
on
snakes
(Lee
and
Scanlon,
2002;
Vidal
et
al.,
2010).
The
osteological
features
for
frog
identification
are
based
on
the
synopsis
by
Hedges
et
al.
(2008)
and
Lynch
(1971).
The
characteristics
used
to
identify
squamate
cranial
remains
are
those
cited
by
Conrad
(2008),
Estes
et
al.
(1988)
and
Evans
(2008).
Post-cranial
identification
is
based
on
the
works
of
Etheridge
(1967),
Hoffstetter
and
Gasc
(1969)
and
Lécuru
(1968,
1969).
Several
other
works
were
used
to
specifically
identify
the
Dactyloidae
(Etheridge,
1959;
Etheridge
and
De
Queiroz,
1988;
Frost
and
Etheridge,
1989;
Nicholson
et
al.,
2012;
Poe,
1998,
2004),
the
Scin-
cidae
(Greer,
1970;
Rao
and
Ramaswami,
1952)
and
the
Teidae
(Kosma,
2004;
Pujos
et
al.,
2009;
Tedesco
et
al.,
1999).
Snake
vertebra
identification
is
based
on
the
works
of
Albino
and
Carlini
(2008),
Albino
(2011),
Lee
and
Scanlon
(2002)
and
Rage
(1984).
3.
Results
Systematic
account
Anura
Duméril,
1806

Please
cite
this
article
in
press
as:Bochaton,
C.,
et
al.,
Fossil
and
subfossil
herpetofauna
from
Cadet
2
Cave
(Marie-
Galante,
Guadeloupe
Islands,
F.
W.
I.):
Evolution
of
an
insular
herpetofauna
since
the
Late
Pleistocene.
C.
R.
Palevol
(2015),
http://dx.doi.org/10.1016/j.crpv.2014.10.005
ARTICLE IN PRESS
G Model
PALEVO-803;
No.
of
Pages
10
4
C.
Bochaton
et
al.
/
C.
R.
Palevol
xxx
(2015)
xxx–xxx
Eleutherodactylidae
Lutz,
1954
Eleutherodactylus
sp.
Duméril
and
Bibron,
1841—48,330
remains
representing
most
anatomical
parts.
Minimal
Number
of
Individuals
(MNI)
=
4,405
based
on
the
ilia.
The
eleutherodactylid
frog
attribution
is
based
on
the
combined
occurrence
of
the
following
features:
elon-
gated
maxillary
bearing
a
high
number
of
teeth
(t.),
without
lateral
dermal
ornamentation
but
with
a
thick
maxillary
shelf
(M.
s.)
and
short
posterior
process
(M.
p.
pr)
(Fig.
2A–B);
fused
sphenethmoids;
unfused
fron-
toparietals;
fused
prootic
and
frontoparietal;
type
1
atlas
articulation
(sensu
Lynch,
1971);
atlas
and
second
vertebra
neural
arches
not
fully
ossified;
sacral
vertebra
with
dis-
tally
weakly
expanded
and
slightly
posteriorly
orientated
sacral
processes;
elongated
scapula
with
well
individual-
ized
acromial
and
glenoid
processes;
ilium
bearing
a
weak,
medially
incurved
dorsal
crest
with
a
globular
and
lat-
erally
placed
superior
tuberosity
and
distally
fused
tibial
and
fibular.
In
addition,
the
occurrence
of
vomerine
teeth
is
a
feature
encountered
in
the
Eleutherodactylus
genus
(Hedges
et
al.,
2008)
(Fig.
2C).
These
osteological
features,
combined
with
present
geo-
graphical
distribution
data,
allow
us
to
attribute
this
frog
to
the
genus
Eleutherodactylus,
and
perhaps
to
a
member
of
the
martinicensis
series
(sensu
Hedges
et
al.,
2008).
The
only
extant
members
of
this
genus
occur
in
the
Lesser
Antilles.
These
remains
are
present
in
all
the
Cadet
2
layers.
Squamata
Oppel,
1811
Dactyloidae
Fitzinger,
1843
Anolis
cf.
ferreus
(Cope,
1864)—8638
remains
corre-
sponding
to
most
skeletal
parts.
MNI
=
215
based
on
the
dentaries.
Many
of
the
observed
features
point
towards
an
attribu-
tion
of
these
bones
to
an
iguanid
whereas
others
are
typical
of
polychrotine
iguanids
(sensu
Evans,
2008
=
Dactyloidae
sensu
Townsend
et
al.,
2011):
maxillary
bearing
a
trian-
gular
facial
process
forming
an
anterodorsal
plate
(ad.
p.)
grooved
by
a
furrow
(canthus)
(c.)
(Fig.
2D);
fused
frontals
without
dorsal
keel
or
pineal
foramen
and
medially
fused
ventral
cranial
crests
(c.
cr.)
(Fig.
2E);
parietal
bearing
a
Y-
shaped
crest
and
a
pineal
foramen
on
its
anterior
margin;
jugal
of
angular
type
morphology;
no
pterygoidian
teeth.
In
addition,
the
blunt
extremity
of
the
dentary
posterior
process
(p.
pr.)
(Fig.
2F)
is
characteristic
of
the
Anolis
bimac-
ulatus
series
(Poe,
2004),
which
includes,
among
others,
the
North
Lesser
Antillean
anoles.
The
maxillary
and
dentary
tooth
morphology
is
also
characteristic
of
anoles;
teeth
are
pleurodont
and
slightly
labio-lingually
flattened.
The
pos-
terior
teeth
are
tricuspid
with
a
wider
central
cusp
and
two
reduced
anterior
and
posterior
cusps.
The
anterior
teeth
tend
to
become
more
and
more
monocuspid.
The
observed
osteological
features
on
all
skeletal
parts
show
that
this
taxon
corresponds
to
one
or
more
species
of
Anolis
of
the
bimaculatus
series
(Poe,
2004).
These
remains
are
abundant
in
every
Cadet
2
layer.
We
estimated
the
fossil
anole
SVL
size
using
the
dental
row
length
of
each
complete
dentary
by
comparison
with
modern
A.
ferreus
of
known
size.
According
to
our
estima-
tions,
fossil
size
ranges
from
53
to
123
mm
SVL,
a
range
consistent
with
the
present-day
size
of
the
Marie-Galante
anole,
A.
ferreus
(see
Breuil,
2002).
No
significant
size
differences
emerge
between
the
Pleistocene
and
Holocene
Cadet
2
layers
(Wilcoxon-Mann-Whitney
test;
P.
val
>
0.05)
(Fig.
3A).
In
addition,
the
clearly
bimodal
distribution
of
fossil
size
corresponds
perfectly
to
female
(73
mm
SVL
maximum)
and
male
(123
mm
SVL
maximum)
A.
ferreus
sizes
(Fig.
3B).
Sphaerodactylidae
Underwood,
1954
cf.
Sphaerodacty-
lus
sp.
Wagler,
1830
–
1
femur.
MNI
=
1
This
single
femur
has
been
identified
on
the
basis
of
its
morphological
proximity
with
S.
fantasticus
and
its
reduced
size
(3
mm
length),
compared
to
all
the
other
taxa
described
here.
This
taxon
is
only
represented
in
the
locus
2
Pleis-
tocene
layer.
Scincidae
Gray,
1825
Mabuya
(s.
l.)
sp.
Fitzinger,
1826
(=
cf.
Capitellum
maria-
galantae
sensu
Hedges
and
Conn
(2012))
–
2
maxillaries,
1
frontal,
2
parietals,
2
dentaries,
1
coronoid,
3
articular-
surangulars,
10
dorsal
vertebrae,
1
sacral
vertebra,
5
caudal
vertebrae,
3
pelvic
girdles,
1
humerus,
3
femurs,
3
ulnas.
MNI
=
2
The
following
osteological
characteristics
allow
us
to
identify
this
taxon
as
a
Scincidae:
maxillary
bearing
a
bifid
posterior
process;
fused
frontals
with
weak
intra-
orbital
constriction
bearing
slight
ornamentation
(or.)
on
its
dorsal
part
and
moderately
strong,
medially
uncon-
nected
ventral
cranial
crests
(c.
cr.)
(Fig.
4A–B);
fused
parietals
bearing
slight
dermal
ornamentation,
a
central
pineal
foramen,
long
posterior
processes
but
shorter
than
the
parietal
table
and
a
deep
posteromedial
notch;
dentary
with
a
straight
ventral
margin,
well
individualized
pos-
teroventral
(pv.
Pr.)
and
posterodorsal
(pd.
Pr.)
processes
and
Meckel
groove
(M.
g.)
completely
surrounded
by
bone
on
its
anterior
half
(Fig.
4C–D);
articular,
prearticular
(pa.)
and
surangular
(sa.)
fused
and
retroarticular
process
(ra.
pr.)
rounded,
well-developed
and
slightly
inclined
medi-
ally
(Fig.
4E);
pleurodont,
cylindrical
and
monocuspid
teeth
with
rounded
apex
on
maxillary
and
dentary
bones.
In
addition,
the
small
size
of
these
remains
and
their
morphological
similarity
with
Mabuya
(sensu
lato)
lead
us
to
attribute
them
to
that
genus,
which
is
the
only
scin-
cid
genus
traditionally
recognized
in
the
Lesser
Antilles.
Following
Hedges
and
Conn
(2012),
Mabuya
sensu
lato
is
currently
divided
into
sixteen
genera
but
no
osteo-
logical
features
have
yet
been
proposed
to
differentiate
them.
Thus,
relying
exclusively
on
geographical
criteria,
we
suggest
an
attribution
to
the
Marie-Galante
skink,
Capitellum
mariagalantae
sensu
Hedges
and
Conn
(2012),
(formerly
Mabuya
mabuya),
mentioned
for
the
last
time
by
Dunn
(1935).
These
fossil
remains
occur
from
the
old-
est
to
the
youngest
Cadet
2
layers
but
are
absent
from
U5-B.
Teiidae
Gray,
1829
Ameiva
sp.
(Meyer,
1795)
–
2
premaxillaries,
8
maxillar-
ies,
2
prefrontals,
4
frontals,
2
postorbitofrontals,
1
jugal,
2
pterygoids,
1
quadrate,
2
dentaries,
4
coronoids,
1
articu-
lar,
1
axis,
17
dorsal
vertebrae,
3
second
sacral
vertebrae,
45
caudal
vertebrae,
6
scapulocoracoids,
1
ischium,
3
humeri,
1
tibia
and
2
femurs.
MNI
=
4
The
main
elements
clearly
show
teiid
squamate
fea-
tures:
maxillary
with
long
and
high
facial
process
(f.
p.)
and
without
dermal
ornamentation
(Fig.
4F);
fused

Please
cite
this
article
in
press
as:Bochaton,
C.,
et
al.,
Fossil
and
subfossil
herpetofauna
from
Cadet
2
Cave
(Marie-
Galante,
Guadeloupe
Islands,
F.
W.
I.):
Evolution
of
an
insular
herpetofauna
since
the
Late
Pleistocene.
C.
R.
Palevol
(2015),
http://dx.doi.org/10.1016/j.crpv.2014.10.005
ARTICLE IN PRESS
G Model
PALEVO-803;
No.
of
Pages
10
C.
Bochaton
et
al.
/
C.
R.
Palevol
xxx
(2015)
xxx–xxx
5
Fig.
2.
Eleutherodactylus
sp.
A
and
B.
Maxillary,
labial
and
lingual
views.
C.
Vomer,
dorsal
view.
Anolis
cf.
ferreus.
D.
Maxillary,
lingual
view.
E.
Frontal,
ventral
view.
F.
Dentary,
lingual
view.
Fig.
2.
Eleutherodactylus
sp.
A
et
B.
Maxillaire,
vue
labiale
et
linguale.
C.
Vomer,
vue
dorsale.
Anolis
cf.
ferreus.
D.
Maxillaire,
vue
linguale.
E.
Frontal,
vue
ventrale.
F.
Dentaire,
vue
linguale.
frontals
with
deep
dorsolateral
nasal
bone
impressions
(N.
i.),
a
clearly
visible
dorsal
ornamentation
(or.)
and
low
ventral
cranial
crests
(c.
cr.)
without
anteroventral
descendant
process
(Fig.
4G–H);
dentary
Meckel
groove
(M.
g.)
fully
open
and
incurved
dentary
ventral
margin
(Fig.
4I);
articular
angular
process
ventrally
orientated;
pleurodont
bicuspid
or
monocuspid
teeth
surrounded
at
their
bases
by
dental
cement
and
basally
pierced
by
a
subcircular
replacement
pit;
procoelous
vertebrae
with
well-developed
zygosphene
that
lacks
an
anterior
notch.
In
addition,
the
absence
of
pterygoidian
teeth
and
the
Meckel
groove
opening
only
visible
in
lingual
view
are
characteris-
tic
of
Ameiva
and
the
occurrence
of
monocuspid
posterior
teeth
with
a
rounded
apex
is
a
feature
occurring
in
the
Lesser
Antillean
Ameiva
taxa
(A.
fuscata,
A.
griswoldi,
A.
plei
and
A.
pluvianotata)
(Kosma,
2004;
Pregill,
1984).
These
features
allow
us
to
identify
this
taxon
as
a
member
of
the
genus
Ameiva,
possibly
comprised
in
the
Lesser
Antillean
Ameiva
clade
on
the
basis
of
its
dental
morphology.
We
estimate
the
size
of
this
squamate
at
around
150–200
mm
SVL,
which
is
similar
to
A.
cineracea
(from
Basse-Terre),
but
smaller
than
A.
major
(from
Petite-Terre),
the
two
previously
described
Guadelou-
pean
ameivas
considered
to
have
become
extinct
recently
(Breuil,
2002)
in
Guadeloupe
islands.
A
comparison
with
these
taxa
is
impossible
as
their
skeletal
morphology
remains
unknown.
The
ameiva
remains
are
present
from
the
oldest
to
the
youngest
Cadet
2
layers
but
are
absent
from
U5-B.
Serpentes
Linnaeus,
1758
Scolecophidia
Cope,
1864
Cf.
Antillotyphlops
Richmond,
1966–169
dorsal
and
cau-
dal
vertebrae.
These
vertebrae
(Fig.
5A–B)
bear
typical
scolecophid-
ian
features.
They
are
small
with
a
centrum
length
ranging
from
0.9
to
1.65
mm.
They
bear
a
zygosphene/zygantrum
articulation
(zs.)
but
neither
neural
spine
nor
hemal
keel.
The
neural
arch
posterior
margin
is
weakly
notched.
The
prezygapophyseal
processes
(pz.
pr.)
are
long,
slender
and
Fig.
3.
A.
Distribution
of
the
Snout-Ventral
length
(SVL)
estimated
from
the
dentaries
dental
length
of
Anolis
cf.
ferreus
in
the
Holocene
and
Pleistocene
Cadet
2
layers.
B.
Histogram
containing
all
the
estimated
SVL
of
the
fossils
and
Gaussian
distributions
resulting
of
a
Gaussian
mixture
analysis.
Fig.
3.
A.
Distribution
des
longueurs
museau-cloaque
(SVL)
estimées
à
partir
des
longueurs
des
rangées
dentaires
d’Anolis
cf.
ferreus
dans
les
niveaux
d’âge
Holocène
et
Pléistocène
de
Cadet
2.
B.
Histogramme
de
toutes
les
tailles
estimées
sur
les
fossiles
et
distributions
gaussiennes
résultants
d’une
analyse
de
mélanges
gaussiens.

Please
cite
this
article
in
press
as:Bochaton,
C.,
et
al.,
Fossil
and
subfossil
herpetofauna
from
Cadet
2
Cave
(Marie-
Galante,
Guadeloupe
Islands,
F.
W.
I.):
Evolution
of
an
insular
herpetofauna
since
the
Late
Pleistocene.
C.
R.
Palevol
(2015),
http://dx.doi.org/10.1016/j.crpv.2014.10.005
ARTICLE IN PRESS
G Model
PALEVO-803;
No.
of
Pages
10
6
C.
Bochaton
et
al.
/
C.
R.
Palevol
xxx
(2015)
xxx–xxx
Fig.
4.
Cf.
Capitellum
mariagalantae.
A
and
B.
Frontal,
dorsal
and
ventral
views.
C
and
D.
Dentary,
labial
and
lingual
views.
E.
Articular-prearticular,
dorsal
view.
Ameiva
sp.
F.
Maxillary,
lingual
view.
G
and
H.
Frontal,
ventral
and
dorsal
views.
I.
Dentary,
lingual
view.
Fig.
4.
Cf.
Capitellum
mariagalantae.
A
et
B.
Frontal,
vues
dorsale
et
ventrale.
C
et
D.
Dentaire,
vues
labiale
et
linguale.
E.
Articulaire-préarticulaire,
vue
dorsale.
Ameiva
sp.
F.
Maxillaire,
vue
linguale.
G
et
H.
Frontal,
vues
ventrale
et
dorsale.
I.
Dentaire,
vue
linguale.
anteriorly
oriented.
The
cotyle
is
dorsoventrally
flattened
and
there
are
no
paracotylar
foramina.
Synapophyses
are
simple
and
rounded.
According
to
these
features,
these
bone
remains
can
be
attributed
to
a
scolecophidian
snake.
They
could
be
related
to
a
member
of
the
Antillotyphlops
genus,
the
only
genus
occurring
today
in
the
North
of
the
Lesser
Antilles
(Hedges
et
al.,
2014).
These
typical
vertebrae
occur
from
Pleistocene
to
Holocene
Cadet
2
layers
but
are
absent
from
U5-C
and
U5-B.
Alethinophidia
Nopcsa,
1923
Boidae
Gray,
1825
Boa
sp.
Linnaeus,
1758–16
dorsal
vertebrae.
These
vertebrae
(Fig.
5C–E)
bear
all
the
Boa
features:
they
are
strongly
built,
are
wider
than
they
are
long
and
have
a
high
neural
arch
and
neural
spine
(ns.).
The
neural
arch
posterior
margin
is
strongly
notched,
the
prezy-
gapophyseal
facets
(prz.
f.)
are
long
and
laterally
oriented
and
the
prezygapophyseal
processes
(pz.
pr.)
are
present
but
short.
The
zygosphene
(zs.)
is
thick
from
an
anterior
view
and
its
anterior
margin
is
more
or
less
concave
from
a
dorsal
view.
From
a
ventral
view,
there
is
a
strong
pre-
condylar
constriction
and
the
hemal
keel
(h.
k.)
is
thin
with
a
blunt
ventral
margin.
A
paracotylar
foramen
(p.
f.)
is
present
on
each
cotyle
side.
These
features
allow
us
to
attribute
these
vertebrae
to
a
member
of
the
genus
Boa.
The
small
size
of
the
vertebrae
(centrum
length
ranging
from
2.8
to
4.2
mm)
suggests
very
small
specimens
with
a
SVL
size
of
around
80
cm.
This
raises
the
question
of
the
juvenile
character
of
these
vertebrae,
a
hypothesis
also
supported
by
the
fact
that
vertebral
cotyles
and
condyles
are
wider
than
high
(Albino,
2011).
Another
possibility
could
be
membership
in
a
dwarf
species,
but
the
material
is
too
sparse
to
provide
a
clear
answer
to
this
question.
This
taxon
is
only
present
in
the
Pleistocene
U5-C
and
Locus
2
layers
of
Cadet
2.
Colubroidea
Oppel,
1811
Cf.
Alsophis
sp.
Fitzinger,
1843
-
155
dorsal
and
caudal
vertebrae.
These
vertebrae
(Fig.
5F–H)
are
weakly
built
and
are
rel-
atively
more
elongated
than
those
of
non-colubroid
Cadet
2
snakes.
The
centrum
is
longer
than
wide
and
has
length
ranges
from
0.8
to
3.5
mm.
In
dorsal
view,
the
zygosphene
(zs.)
anterior
margin
is
slightly
lobed.
The
neural
spine
(n.
s.)
is
longer
than
it
is
high.
The
synapophysis
(sy.)
is
well
individualized
with
a
diapophysis
slightly
bigger
than
the
parapophysis.
The
prezygapophyseal
processes
(pz.
pr.)
are
conical,
thick,
blunt,
laterally
oriented
and
their
lengths
correspond
to
half
of
the
prezygapophyseal
facet
(prz.
f.)
length.
Paracotylar
foramina
(p.
f.)
are
present
on
each
side
of
the
condyle.
From
a
lateral
view,
the
hemal
keel
(h.
k.)
is
well
marked.
On
the
basis
of
these
features,
these
vertebrae
can
be
ascribed
to
a
colubroid
snake.
In
addition,
the
prezygapoph-
ysis
morphology,
thicker
and
more
laterally
oriented
than
those
of
the
other
Cadet
2
colubroid
vertebrae,
allows
us
to
advocate
their
attribution
to
the
genus
Alsophis.
These
vertebrae
occur
in
all
the
Cadet
2
layers.
Colubroidea
sp.
1–13
dorsal
vertebrae.
These
vertebrae
(Fig.
5I)
are
similar
to
those
previ-
ously
described
for
cf.
Alsophis
sp.,
except
that
they
are
smaller
(maximal
centrum
length
=
1.2
mm)
and
they
bear

Please
cite
this
article
in
press
as:Bochaton,
C.,
et
al.,
Fossil
and
subfossil
herpetofauna
from
Cadet
2
Cave
(Marie-
Galante,
Guadeloupe
Islands,
F.
W.
I.):
Evolution
of
an
insular
herpetofauna
since
the
Late
Pleistocene.
C.
R.
Palevol
(2015),
http://dx.doi.org/10.1016/j.crpv.2014.10.005
ARTICLE IN PRESS
G Model
PALEVO-803;
No.
of
Pages
10
C.
Bochaton
et
al.
/
C.
R.
Palevol
xxx
(2015)
xxx–xxx
7
Fig.
5.
Cf.
Antillotyphlops.
A–B.
Dorsal
vertebra,
dorsal
and
ventral
views;
Boa
sp.
C–E.
Dorsal
vertebra,
dorsal,
lateral
and
anterior
views.
Cf.
Alsophis
sp.
F–H.
Dorsal
vertebra,
dorsal
lateral
and
anterior
views.
Colubroidae
sp.
1.
I.
Dorsal
vertebra,
dorsal
view.
Colubroidae
sp.
2.
J–M.
Dorsal
vertebra,
dorsal,
ventral,
lateral
and
anterior
views.
Fig.
5.
Cf.
Antillotyphlops.
A
et
B.
Vertèbre
dorsale,
vues
dorsale
et
ventrale
;
Boa
sp.
C–E.
Vertèbre
dorsale,
vues
dorsale,
latérale
et
antérieure.
Cf.
Alsophis
sp.
F–H.
Vertèbre
dorsale,
vues
dorsale,
latérale
et
antérieure.
Colubroidae
sp.
1.
I.
Vertèbre
dorsale,
vue
dorsale.
Colubroidae
sp.
2.
J–M.
Vertèbre
dorsale,
vues
dorsale,
ventrale,
latérale
et
antérieure.
longer,
sharper
and
anteriorly
oriented
prezygapophyseal
processes
(pz.
pr.).
These
vertebrae
could
possibly
belong
to
a
member
of
the
genus
Liophis
(possibly
Erythrolamprus
following
Curcio
et
al.,
2009),
which
is
with
Alsophis
the
only
genus
of
colu-
broid
snake
ever
observed
on
Marie-Galante
(Breuil,
2002),
but
the
vertebral
morphology
of
the
latter
is
still
unde-
scribed
and
remains
unknown
to
us
for
the
time
being.
These
vertebrae
occur
in
all
the
Cadet
2
layers.
Colubroidea
sp.
2–1
dorsal
vertebra.
This
vertebra
(Fig.
5J–M)
is
similar
to
those
previously
described
for
cf.
Alsophis
but
it
presents
some
significant
differences.
It
bears
a
more
marked
neural
arch
median
constriction
than
the
other
vertebrae
in
dorsal
view.
The
neural
spine
(n.
s.)
is
also
different;
it
is
high
and
thin
with
anterior
and
posterior
borders
slightly
overhanging.
The
prezygapophyseal
processes
(pz.
pr.)
are
shorter
and
thicker
and
the
prezygapophysis
articular
facets
(prz.
f.)
have
a
more
elongated
shape.
For
the
time
being,
we
cannot
suggest
a
specific
attribu-
tion
for
this
vertebra.
It
could
be
from
an
unknown
taxon
from
the
island
fauna.
This
single
vertebra
was
recovered
in
the
Locus
2
Pleistocene
layer.
4.
Discussion
We
identified
at
least
ten
taxa
in
the
different
sampled
layers
of
Cadet
2,
all
of
which
occur
in
the
Pleistocene
layers,
which
are
currently
the
oldest
known
Marie-Galante
fossil
levels.
Consequently,
our
study
provides
the
first
informa-
tion
on
Marie-Galante
Pleistocene
fauna
prior
to
14,000
B.C.
(Stouvenot
et
al.,
2014).
Three
taxa
are
only
present
in
the
Pleistocene
layers
at
Cadet
2:
Boa
sp.,
Colubroidea
sp.
2
and
cf.
Sphaerodactylus
sp.
Boa
sp.
was
previously
reported
at
Cadet
3
(Stouvenot
et
al.,
2014),
where,
like
at
Cadet
2,
this
taxon
only
occurs
in
Pleistocene
levels.
Colubroidea
sp.
2
had
not
been
iden-
tified
up
until
now
but
this
attribution
will
require
further
investigation.
Sphaerodactylus
is
a
very
small
sphaero-
dactylid
gecko
still
occurring
on
Marie-Galante
(Breuil,
2002;
Powell
and
Henderson,
2012).
This
is
the
first
fos-
sil
record
of
this
taxon
on
the
island
and
it
shows
that
it
was
already
present
there
during
the
Late
Pleistocene.
This
is
not
surprising
given
the
existence
of
a
Marie-Galante
endemic
sub-species
(Sphaerodactylus
fantasticus
adrino-
tus)
(see
Breuil,
2002),
which
was
the
first
evidence
of
its
local
evolution
on
the
island.

Please
cite
this
article
in
press
as:Bochaton,
C.,
et
al.,
Fossil
and
subfossil
herpetofauna
from
Cadet
2
Cave
(Marie-
Galante,
Guadeloupe
Islands,
F.
W.
I.):
Evolution
of
an
insular
herpetofauna
since
the
Late
Pleistocene.
C.
R.
Palevol
(2015),
http://dx.doi.org/10.1016/j.crpv.2014.10.005
ARTICLE IN PRESS
G Model
PALEVO-803;
No.
of
Pages
10
8
C.
Bochaton
et
al.
/
C.
R.
Palevol
xxx
(2015)
xxx–xxx
Fig.
6.
Table
containing
the
distribution
of
the
taxa
in
the
Cadet
2
and
3
(Sierpe,
2011;
Stouvenot
et
al.,
2014)
layers
along
with
their
historical
mentions
(see
Breuil,
2002),
present
occurrence
(Powell
and
Henderson,
2012)
and
minimal
period
of
occurrence
on
Marie-Galante
(in
grey).
Fig.
6.
Tableau
de
distribution
des
taxons
dans
les
niveaux
de
Cadet
2
et
3
(Sierpe,
2011
;
Stouvenot
et
al.,
2014)
avec
leurs
mentions
historiques
(voir
Breuil,
2002),
leur
présence
actuelle
(Powell
et
Henderson,
2012)
et
leur
période
minimale
de
présence
à
Marie-Galante
(en
gris).
The
other
identified
taxa
have
been
previously
observed
alive
on
the
island
and
occur
in
all
the
Cadet
2
layers.
This
is
the
case
for
the
Eleutherodactylus
frogs,
currently
rep-
resented
on
the
island
by
two
species:
E.
marticensis
and
E.
johnstonei
(Breuil,
2002;
Powell
and
Henderson,
2012).
The
Marie-Galante
skink
(cf.
Capitellum
mariagalantae)
was
observed
on
the
island
for
the
last
time
in
1830
and
is
since
considered
to
be
extinct
(Hedges
and
Conn,
2012).
This
taxon
is
also
present
in
the
Cadet
3
site
(Stouvenot
et
al.,
2014).
Ameivas
were
reported
for
the
last
time
in
the
17th
century
(see
Breuil,
2002).
This
taxon
is
also
present
in
several
other
Marie-Galante
sites
(Grouard,
2001,
2003;
Stouvenot
et
al.,
2014).
Anoles
(Anolis
cf.
fer-
reus)
are
still
widely
present
on
the
island
(Breuil,
2002)
and
fossil
anole
remains
have
already
been
mentioned
at
other
sites
(Grouard,
2001,
2003;
Stouvenot
et
al.,
2014).
As
for
the
snakes,
Antillotyphlops
is
considered
to
have
become
extinct
very
recently
on
Marie-Galante
(Breuil,
2002),
Alsophis
disappeared
from
Marie-Galante
during
the
20th
century
(Breuil,
2002)
and
Liophis
(Colubroidea
sp.
1?)
even
earlier
at
the
end
of
the
19th
century
(Henderson,
1992).
Bones
from
these
three
snakes
were
also
reported
at
Cadet
3
(Stouvenot
et
al.,
2014).
Evolution
of
the
species
composition
Our
data
shed
light
on
the
herpetological
biodiver-
sity
stability
during
the
Pleistocene
and
Holocene
periods
(Fig.
6).
Accordingly,
only
two
taxa
(Boa
sp.
and
Colubroidea
sp.
2)
seem
to
have
disappeared
between
Pleistocene
and
Holocene
periods.
At
the
present
time,
this
fact
is
difficult
to
explain
due
to
the
lack
of
Pleistocene
environmental
data
for
the
island.
The
case
of
the
boa,
still
represented
in
the
Lesser
Antilles
on
Dominica
(Boa
nebulosa)
and
Saint
Lucia
(B.
orophias)
by
endemic
taxa,
is
very
interesting
but
the
study
of
larger
series
is
indispensable
to
enhance
our
knowledge
of
the
taxonomic
status,
biological
features
and
extinction
causes
of
this
snake.
This
apparent
stability
no
longer
subsists
for
the
extant
island
fauna.
Among
the
seven
taxa
that
appear
to
be
present
since
the
Late
Pleistocene
in
Cadet
2,
only
three
of
them
still
occur
on
the
island
today
(Eleutherodactylus
sp.,
cf.