Additional hominoid material from Miocene of

Contr. Tert.
Quatern.
Geol.
figs,
9
25-39
36(1-4)
tab.
1
1999
Leiden, December
Additional hominoid material from
the Miocene of Spain
and remarks
on
into Europe
hominoid dispersals
J. van der Made
Museo Nacional de Ciencias Naturales
Madrid, Spain
and
F. Ribot
Paleontologic
Institut
Dr M. Crusafont
Sabadell, Spain
Made,
J. van der & F.
Contr. Tert.
A hominoid
(Barcelona)
Vallès-Penedès.
first
With
hominoid may
Griphopithecus
Europe.
J.
Van
The
der
9
entered
ages
of
—
Ma
Anatolia
hominoids
from the
collections
this
and such
may
later
events
(beginning
ago
are
likely
than
Dryopithecus
to
ago
Çandir
may
of
Museo
Dr M.
Nacional
Crusafont,
de
Ciencias
C. Escola
and remarks
Spain
have
Naturales, CSIC,
Industrial
23,
E-08201
from
the
the
be
were
1920s housed
first
a
(late
are
related
MN
close;
5),
Introduction
Description
p. 25
comparison
The ages of Turkish and
material
.
also
Dispersal
Summary
p.
European
p.
dryopithecines
into
28
p. 34
Europe
and conclusions
p. 35
hominoid
age,
Gutierrez
in
level
only.
an area,
Bruijn
Geologia
del Semi-
Geologia
the
in
global climate.
the
as
The
European remains of
different
a
genus.
ago
(late
MN
the
same
species. Remains
from
entered
6) it
Africa.
F.
Madrid, Spain;
Institut
Ribot,
Dryopithecus
to
in the
MLGSB.
a
At
'locality',
of
(JvdM)
us
suid
1920s, which
that
et
1992),
al.,
MN
MN
9)
one,
Can
7
record
8;
an
later
was
time, Hostalets
Neogene
to
Late
a
a
to
was
single
represent
Aragonian
Mammal Units,
Lower Vallesian
be
older
Llobateres localities
that the present
old collection
from
collec-
a
with fossils from
considered
In view of the fact
an
+
and
recog-
fossils
tooth is from
stratigraphical levels,
two
(Upper
than
(Agusti
et
the
al.,
Dryopithecus
('Hostalets oc'),
its
exact
provenance remains unknown.
Villalta Cornelia &
first
one
amongst
To date, 'Hostalets' is considered
stratigraphical
the col-
Ma
de
discussed here
are
and
coming directly
Guérin
be
to
tooth is from
in
ago,
new
Ponsic and
1984).
suids
12.5
than
specimen
(Middle Miocene,
Can
lections of the Museu i Laboratori de
Laboratori
represent
even
(MLGSB),
the
to
with
Miocene,
fossil
i
clear. Since
might
E-28006
2,
M.
by
transferred
p. 37
studying
—
dispersals.
present
tion made
Acknowledgements
and
Europe.
have been collected
changes
might represent
Barcelona
the
References
cataloguing
later
Abascal
de
When
into
Sabadell, Spain.
p. 37
Introduction
and
12.5 Ma
still assumed
Miocene hominoids
of
.
27
not
are
these occurrences
entered Europe
Jose
and not
Hostalets. The
new
Museu
tooth to
sea level
eustatic
to
5); its affinities
nised
of the
at the
dryopithecine
different species,
nari
Contents
and
dispersals
hominoid
on
1999.
well
MN
represent
14 Ma
Hadersdorf and
Klein
younger.
Ma
they
not
probably
in
Dryopithecus;
mammals,
15.5
younger,
Miocene of
Leiden, December
1 tab.
older European Dryopithecus localities, hominoid dispersals into Europe
of other
from
much
genus
one of the
dispersals
3
figs,
Dryopithecus, Hominoidea, Vallès-Penedès, stratigraphic
Made,
Paleontologic
as
to the
entered Europe
least
material
hominoid
Hostalets, recently recognised
assigned
with
have
at
are
from
is
Neudorf-Sandbergare
Key words
Additional
Hostalets
These coincided
Griphopithecus
from
molar
upper
del Seminari
well.
Ribot.
Quatern. Geol., 36(1-4): 25-39,
a
Crusafont Pairo
dryopithecine
(1941)
were
the
from the Valles-Penedes.
26
1.
Fig.
MLGSB
48486, right
posterior views,
This fossil
Villalta
(IPS
1),
Cornelia
consisted of
lower
Crusafont
teeth united
two
and
IPS
Crusafont
1827
in the
from Can Vila
font Pairo
Late
Aragonian)
Can
Mata-Ocata
authors
not
were
represent
were
stated
(=
a
age,
area
nian
Vallesian age.
Aragonian
age
that
the
(Agusti
originally
of mandible,
numbered IPS
specimen
near
1826
found
was
track
a
now
and
noted
had
that other beds
yielded
of the fact that
none
specimen
et
from
of these
is
figures
upper
-
stereo
pair
of occlusal
middle
view,
-
buccal
and
present
oldest
recorded
to
Llobateres
(=
the
et
al.,
date from the
(attributed
1990).
remains of
right
molar of
material of D.
(= type
to
In
European
hominoid
area.
D.
laietanus)
In
light
additional data
hominoid localities
into
have
to
being
been
Material from Can Ponsic
of
on
one
this,
the
and
1992a)
and Can
is younger
addition, it represents
dispersal
interest in
special
Dryopithecus
crusafonti Begun,
to
Dryopithecus.
present
is of
specimen
upper
(=
These
species
the beds
favours
work has
an
et
that
Arago-
resulted
considered
al., 1984; Begun
The
the
(Begun
of the oldest
the time
(relative)
discuss
a
seems
ages
of
model of
Europe.
Hostalets did
Six other mammal
Subsequent
in
Hipparion.
collected
4;
by
leading
be of Vindobonian
to
x
occidentalis
1949,
Mata. Villalta Cornelia & Crusa-
been
mandible;
consensus
piece
now
single locality.
have
a
Hostalets,
lingual views.
Pairo,
The
aware
the
the
by
considered it
yielded
or
and
of Hostalets,
Can
already
to
are
M3).
area
to
(1941)
from
of Dryopithecus
anterior
the type of Sivapithecus
&
lost. The isolated molars
M2)
M²
-
to
be
Abbreviations
ied;
the
to
—
Numerous collections have
indicate the
following
repositories
abbreviations
are
of
specimens
used:
in
of
al., 1990).
GML
Geological
HGSB
Hungarian Geological Survey, Budapest
Museum, Lisbon
been studexamined
27
IGGML
Institut fiir
Geowissenschaften/Geologie
der Montan-
universitat
Mesio-distal diametre
IPS
Institut
Paleontologic
IPUW
Institut
fur Palaontologie, Universitat
Dr M.
IVAU
Instituut
Wien
Bucco-lingual
Universiteit
Aardwetenschappen,
voor
11.3
Crusafont, Sabadell
width
of the
first
lobe
11.5
(= maximum width)
Utrecht
MGB
Museo
MGL
Museum
MLGSB
Museu
Bucco-lingual width between the lobes
Barcelona
Geologico,
Guimet, Lyon
of the
crown,
Maximum diagonal
MNCN
Museo Nacional
MNHN
Museum
MPZ
Museo
MTA
Maden Tetkik
NMB
Naturhistorisches Museum, Basel
Minimum diagonal of
NMM
Naturhistorisches Museum, Mainz
d2
national
Ciencias Naturales, Madrid
d'Histoire
the
Minimum diagonal of
crown,
D1
12.5
dl
11.5
Paris
naturelle,
de la
Paleontologico
9.7
posterior lobe
Barce-
lona
de
the
Bucco-lingual width of
i Laboratori de Geologia del Seminari,
10.4
Universidad
de
Maximum diagonal of the
Zaragoza
occlusal
10.2
surface,
D2
ve
Arama,
Ankara
NMW
Naturhistorisches Museum, Wien
PIMUZ
Palaontologisches
PDTFAU
Paleoantropoloii,
Institut
und
Mesio-distal length of
Museum, Universitat
Zurich
Mesio-distal
Ankara
Dil
Tarih
ve
SMNS
Staatliches Museum fiir Naturkunde, Stuttgart
UCBL
Universite
Landesmuseum
Claude
Joanneum,Graz
Table
1.
AND
COMPARISON
OF
THE
NEW
probably
a
the
present
for its
reason
this
though
morphologies.
ent
The
primate.
indicating
in
or
occur
and
suids,
includes very
term
hypocone
include
in
being
has
slightly
a
trianglular
a
given
merges
the
recognisable,
are
Aragonian
mesio-distally
a
and Valle-
reduced
or
The
with
nearly
un-
M'
or
an
M.
Meas-
mesio-lingual
transcrista,
The
crest
crest
pule,
crest
is
a
is short and adds
of the
to
longer
protocone (proto-
of the paracone forms part
which ends
abruptly
mesio-marginal
and anterior transcrista converge in
reminiscent of
the
protoconule.
one
at
crest,
point,
A small
a
the base
protocrista
small
cus-
disto-lingual
the
M²
48486, right
from Hostalets.
metaconule in this
a
crista
separates
On
obliqua.
marginal-distal
the
by
trigonal
wide
on
at
much variation in
a
wall
relatively
wide;
(Dvinska
Nova
a
an
hypocone.
assumed
of 11.6
(IPUW, cast).
wider than the
Dryopithecus
the
is much
larger
Valles-Penedes
ranges for the M2
the M
a
1
which
There is
a
no
have
deep
a
furrow
hypocone,
to
mm
NT
(Steininger,
width of
in
13.5
particular,
upper molars
Rudabanya,
(Fig. 2),
an
a
Ml of
are
Neudorf-Sandberg
be
and
larger and,
than the
and
darwini
Griphopithecus
of
It
2).
smooth. There is
upper molar from
Ves),
length
The tooth is
a
the buccal wall.
on
molars
upper
has
the
are
pliopithecids,
between the protocone and
similar furrow
The
tooth
unlike in
cingulum,
lingual
have
not
all.
cingulum along
the
and
cusp
is
transcrista
of the Carabelli cusp in the upper molars
four sides of the
The
the
near
hypocone
posterior
of D. laietanus; the tooth from Hostalets does
Carabelli
of
crest
'postmetacone'
a
The
There is
crest.
tri-
large
the distal side of the
cusp'.
accessory
The
crest.
the
directed and ends
small accessory cusp,
a
'postmetacone'
development
1967),
crest, which is
there is
to
and the
absent metacone,
outline. The tooth
trapezoidal
mesio-marginal
the mesial
protocrista.
a
bucco-distally,
or even
an
elongated,
has
crown
rounded
mesial crista of the paracone
of the anterior
of
to
in Table 1.
the formation of the
crista).
morphologi-
united
pliopithecids, Dryopith-
from Hostalets is therefore either
are
3.5
of MLGSB
(in mm)
mesio-buccally
'postero-external
lingual
surface
like the M' which have
and
pri-
differ-
specimen belongs
outline. The base of the
square outline,
The
ursids and
Griphopithecus.
rectangular
urements
1)
is
end of the
being
and
ecus
occlusal
posterior fovea
direction and
diagonal
a
hypoconc
lingual
metacone
upper molar.
an
European primates
resulting
bunodont is
confounded with that of
descriptive
trigon
that it is
is
(Fig.
The low cusps and other
sian
The
4.6
indication of
no
tooth
cal features indicate that the present
a
fovea
basin from the smaller talon. The
gonid
suid. 'Bunodont molars'
mates,
anterior
of the
length
Measurements
is in
crest
the
the
the
Bernard, Lyon
MATERIAL
that
7.2
of Dryopithecus
There is
fact
surface,
Universitesi, Ankara
Steiermarkisches
The
occlusal
Facultesi,
Cografya
SLJG
DESCRIPTION
the
(compare Fig.
Dryopithecus
but
from
into
enters
and is similar in
mm
much
morphology
the
to
2
and M
detailed
of D. laietanus
description).
that of any
of the
M
1
(see Golpe Posse,
1993 for
The size of MGSB 48486 exceeds
of
Dryopithecus
from the
Vallès2
directed from the
the protocone. These
anterior
transcrista
tinuous and
two crests
is
connects
paracone
do
the buccal
not meet
formed. The
proto- and
to
crista
metacone.
and
crest
no
obliqua
of
second
is
con-
Pencdès and
(Fig. 2).
Ribot
We
(1993;
Rudabänya,
but is within the ranges of the M
believe the tooth is
see
also
Ribot
et
an
al.,
M' of
Dryopithecus.
1996)
assigned
material from the Valles-Penedes to D. laietanus.
all
28
2.
Fig.
M¹
THE
and transverse
Bivariate plots of meso-distal diameter (DAP)
and M²
AGES
TURKISH
OF
and central
of Dryopithecus from Rudabánya
EUROPEAN
AND
diameter
control of
graphic
material is
type
material
from
and
cally
This is
ited
reasons
it is
important
the
hominoid localities.
often
have
and
incomplete,
several
close,
by precise stratigraphic (and
Before
reason.
passed through Anatolia;
areas
are
are
the
important
Fig.
3.
still
Correlation
age
many
European
scheme;
indication
in
assumed
to
dispersal
events.
units
have
left column
this
eustatic
allowed
The next
for
and
columns
African localities
central columns show suoid
that
particular lineage,
taxonomic
Taucanamo
with
and
the
lineage (Van
der
der
sea
level
corrected
faunas
two
give
the
according
samples
are
of Haq et al.
cycles
age
Miller
by
of the
ages
and their
above the
double
described
in
Parachleuastochoerus lineage (Van
der
to
8. There
explanation,
was
of the ages of
MN
to
MN
6,
European
place
the
tremendous difference
a
on
placed
in
the
of MN
boundary
of MN 5. In
content
where the reference localities of the
be
included
situated, palaeontologists
Aragonian
5, but
D
zone
in
&
(Daams
central
Europe
Freudenthal,
similar faunas
in MN4.
Van
(1987), and
al.
(Pickford,
European
are
der
Each
animals.
The
1986;
and
more
elsewhere;
der
ages
(1996).
Vallesian
(1994, 1996). The
Made, 1997a, 1998),Bunolistriodon
Made, 1996a),Conohyus lineage (Van
1997a, 1998),Tethytragus langai lineage (Fig. 5;
in
Aragonian and
et al.
lines
detail
et
of terrestrial
Krijgsman
their radiometric
1992).
are
zonation
to
of
placed
were
dispersals
bovid lineages
change. The lineages
lineage (Van
the
intercontinental
with dates of the lower limits
(1981)
in
-
column being
there
study
Mammal
For instance,
forward arguments
localities, put
MN units tend
correlations of
hominoid localities;
a
the
mm.
the estima-
Neudorf-Sandberg
in
(1986),
and France,
Spain
1988)
in the
problems
and
al.,
in
Neogene
in the estimate of the age of the lower
obviously important.
There
Hadersdorf
used
et
5. There has been discussion
hominoids
correlations between these
measurements
without
(1992),
al.
et
localities in MN
is
history
Bruijn
hominoid
united.
but may be lim-
Europe,
Bruijn
de
(MN units;
whereas Mein
palaeobiogeographic)
in
arriving
commonly
Units
Klein
reason
is
(triangles) compared to
individual localities and with
tion of the age of the
de
geographi-
are
information. The reconstruction of hominoid
another
strati-
Hominoid
frequently
subjective element,
a
good
a
for that
localities which
stratigraphically
obviously
to
the tooth from Hostalets
with
problems
MIOCENE
HOMINOIDS
For several
(DT) of
Europe (dots) and the Vallès-Penedès (crosses);
(in Ma)
latidens
each
with
thick
black
horizontal
(Daams
&
last column
other references
advanced than
of
starts
Turkish localities.
Bunolistriodon
of onset
cycle
given by
those
cycle, the second
level low,
sea
the Faunal
Van
lines
below;
a
and
Sets of
der Made,
in
is
the
MN
Pickford
1996a).
The
column indicate that
generally
lineage (Van
this is
der
reflected
Made,
in
a
1996a),
lineage (Van der Made, 1996a), Listriodon splendens
Made, 1989, 1990a, 1999b), Schizochoerus lineage (Van
Made, 1994), Tethytragus
Made, 1990a, 1999b).
which
lines indicate these
Freudenthal, 1988)
gives
Double
lockharti
a
koehlerae
lineage (Van
der
der
Made,
Made,
1994),
29
30
As
compromise,
a
al.,
et
man
D
zone
correlations with marine
and Spanish
ties and
sections
(Krijgs-
great difference be-
a
in the
strata
mammal localiUnits MN 7 and
sequences.
united into
MN 7+8
(de Bruijn
In the present paper, MN 7 and MN 8
informal
indicate
to
manner
early
on
Paratethys
western
superimposed
with
long palaeomagnetic
have been
8
1992).
an
is
the estimates of the ages of the MN units based
tween
MN
included in MN 5
was
there
Still,
1996).
used in
are
late
or
al.,
et
MN
7+8,
The
in
stratigraphy
3.
Fig.
their ages
given
are
present
events
of their lower boundaries
ages
African
in the
column. The central col-
right
selection of suoid and bovid
a
these
ual, though fluctuations in evolutionary
occurred.
in time. In both
spaced
tive level
be
can
ble lines in
the
Besides,
seen
cases
subsequent
clearly
but
ally,
evolved
more
than
aspect
Douline-
particular
fossils with
those below.
this coincides
invariably,
not
to
evenly
samples.
yielded
age, the localities above the line have
a
likely
not
are
grad-
clear difference in evolu-
a
between
is
rates are
samples
and
lineages
lineages
column indicate that for that
a
(Pick-
with
Gener-
change
a
in
Andrews
and
Pasalar
Hadersdorf
al.
et
in
MN5-6
and
hominoids from
to
(1996,
Qandir
G.
to
yielded
1996),
ment
opinion
and
This
course
species
or
is
the
both
the
impression
that
the
gap between
so
al.,
and,
Griphopithecus
1999). Engelswies
be
cannot
from
Qandir,
Andrews
et
copithecus
wished
a
(1996)
has been
one
to
a
possibility
to
be
that
taxonomical
is
present
of different
considered.
defined
a
on
stehlini’
type
include
study;
the
grouping.
four
posed
with
The
in MN
fossils
ma-
bones
Austria-
not
based
led
cerus
There
(1986)
to
Bruijn
et
de
localities have
are
localities,
studies
few
our
of various
was
above
The
(1992).
sup-
MN 6,
though
in MN
high
to
the
have
to
Protragoin
locality
MN
the
8.
locality
with
boundary
MN
mammals.
mainly large
large
a
mainly
have
placed
usually
above
been
Pasalar,
that
suggests
it
is
in
MN5.
to
though
usu-
position
much
to
The
placed
assigned
for this. The
provided
us
framework.
stratigraphic
on
study
mammals would allow
Qandir
6
based
being
the former has also been
arguments
no
the
a
though
supposed
of
(last
played
The detailed
century ago.
a
of
localities Pasalar and
has
locality,
(1992) placed
knowledge
half
lineages
of the
place
al.
yielded
these localities in
of
younger
Neudorf-Sandberg.
The best documented evidence for the relative strati-
graphic positions
of Pasalar and
Bunolistriodon latidens
distal
that
diameter of the
et
al.,
and
Pasalar and
VI
and
in the
langai
£andir
I
der
from the
in
meso-
indicates
1996a;
Made,
and
(Bovidae)
than
(Fig.
4;
size
der
Suoidea)
I is older than
(Van
der
on
the
that
IV &
Made,
1994,
Arago-
1988). Changes
lineage
indicate that Pasalar and
Pasalar
and
del Val
section of the
& Freudenthal,
in
suggest
Arroyo
Van
is in the type
(Daams
incisors
splendens
hypsodonty
older
younger than Sansan
inonii
suids
Taucanamo sansaniensis-inonuensis
laeochoeridae,
are
are
Manchones i
C2
in
lineage
Manchones
zone
these
Qandir (Van
in Listriodon
increase
Tethytragus
of
incisors
comes
increase
19%).
Similar trends
canines
Qandir
The
lineage.
Pasalar is older than
Fortelius
1996a).
different ages
was
‘Gazella
revisions of the faunas from these
recent
current
published
nian in
the
of
The presence
in MN6, close
very low
grouping
or
al.
been
subsequently
(Mein, 1986).
Mein
5. These
underlying
possibility
Neudorf-
Hadersdorf
presence
et
Bruijn
3
to
localities.
Neudorf-Sandberg
at
5)
have
Nevertheless,
place
mi-
Ma;
therefore 1.5
Klein
in the estimation of the age
these
on
lockharti
presence of the suid Bunolistriodon
been reworked
al.
gives
de
be
14
and in the top of MN6, far
(Mein, 1986)
occurrence
role
based
16 and
some
the
more
der Made,
Hadersdorf and
of
is
probably
opinion,
widely.
because
than
et
Klein
Neudorf-Sandberg, by
material
localities
8
current
material from Klein
is
MN
£andir
long
geographical
in all
species
to
obviously
morphological
seems
species
with the
proposed
as
two
in
placed
for
varied
have
or
is between
ad-
an
that from
low in MN 5, which
accepted Griphopithecus
estimates
Age
Sandberg
should
the latter date and
to
Ma older than the
ally
The
accepted.
closer
probably
(1992)
from Neudorf. Nevertheless,
darwini. This
comparative
corresponds
not
al.
in G.
assumption
that
cromammals. The age
is
and
Aragonian (Van
(1995)
the
suid Bunolistri-
stage which
locality
Ziegler's
to
corresponds
the
to
(Heizmann,
Ravensburg
in the later part of zone D,
assign-
minimise
to
Engelswies
The
1996a).
E of the
zone
are
and localities for which the
directly
with that
weinfurteri,
Hadersdorf,
on
compared
nor
the
than
primitive
the
yielded
from
one
evolutionary
an
is
MN5-6, which
terial from Klein Hadersdorf consists of
and
an
has
lower
Sweet Water Molasse and
of the end of MN5,
form, typical
Engelswies
transition of the
is in the middle unit
The
Generally, (Jandir
al.,
et
Heizmann
though
doing they
to
maintained in
in MN6, Andrews
to
and
Engelswies
(Heizmann
Although
placed
Engelswies
Pasalar
difficult basis for
localities
in
from
from
one
et
the
and Klein Hadersdorf
tooth
MN5
is that Pasalar is
assigned
a
Klein
assigned
classification is
genus.
in
They
those
Köhler
(e.g.
Engelswies
presence of
MN6.
fragmentary
a
which of
to
(1996)
papers
only
placed
sp.
Engelswies
Neudorf-Sandberg,
in
darwini,
alpani
?i
Griphopithecus
subsequent
and
Neudorf-Sandberg
Griphopithecus
Qandir
12.7) placed
table
the
Upper
localities have
loackharti.
vanced
again
taxonomy.
in
Ravensburg
Both
1992).
odon
and Arabian localities and
their localities. The evolution in
have
and Vallesian
Aragonian
the
synthesised
are
the left, while Faunal Sets
on
selected
1981),
umns
units,
MN
in columns
given
ford,
dispersal
and the estimated
zones
are
The
and
of
locality
placed
respectively.
is
Engelswies
middle unit of the
Made, 1993,
the basis of
two
(Pa-
Inonii I
1997a).
listrio-
31
Fig.
4.
Increase
in
in
length (DAP
mm)
Tethytragus langai lineage.
Manchones
and
Arroyo
The
del Val
and
hypsodonty (100 Hp/DT; Hp
localities
(IVAU),
in
are
Sari
approximate
Çay (studied
while
=
order
in
the
of the
height
from
old
DT
entoconid,
of
width)
=
the
the
M3 in
in
while
(studied
Pasalar
young:
IPS),
the
IPS).
dont
lineages.
F of
the
Sansan
The
Sansan is
A
Aragonian.
(Sen,
date
Van der
1997)
for
(Krijgsman
Made
low in MN 6 and in
placed
short
be
may
the
palaeomagnetic
in various
interpreted
of
boundary
the
zone
section
in
ways.
G1/G2
zones
1994, 1996) and the correlations by
al.,
et
(1996a)
older
slightly
ages
to
for
suggest
and
than
Pasalar and
close
12.5
to
Qandir
Ma, respec-
tively.
‘Gazella
present
in
to
from
the
citation of
prior
MN
to
described
odon
8.
are
The
remains
Bunolistriodon
as
splendens;
fossils
Protragoceros,
there
reworked
is
no
from
der
(Van
defi-
a
appeared
Neudorf-Sandberg,
to
reason
have
to
re-
rise
given
poor for
too
likely
lockharti,
der
(Van
have
belong
assume
to
Listri-
that these
Made, 1996a).
Trends in incisor width and canine size and
ogy
of MN 8;
typical
may
are
and the genus is
is
Tethytragus,
to
not
that
Neudorf-Sandberg
assignment
nite
referred
now
it is found in late MN 6 and in MN7. The
Europe
mains
stehlini’,
in Klein Hadersdorf, but is
morphol-
data from Mottl,
Made, 1996a; using
1957)
of Listriodon suggest that:
—
and
—
age
—
Klein Hadersdorf is of about the
Arroyo
St
del Val IV, i.e.
is
Stephan
as
or
Neudorf-Sandberg
but closer
5.
in
Bivariate plot of length (DAP;
mm)
of the
Dot
=
P3
of Conohyus and
Conohyus
(IPUW).
Cross
=
simorrensis
Conohyus
mm)
hyus
=
from
Göriach
ebroensis
from
from
Parachleuastochoerus
(MGL,
UCBL,
heimensis
from
Klein
simorrensis
NMB).
(DT;
do Pinheiro
Asterisks
Pasalar
C.
=
simor-
Cono-
(GML). Triangles
steinheimensis
Inverted
=
from
triangles
Steinheim (SMNS, NMB).
=
to
is
oc
oc
same
age
as
12.5 Ma;
Quirze and of the
(oc
about
=
old
the
same
collections);
same
age
as
between 12.5 and 11
(i.e.
or
Ma,
the latter date.
in
Hadersdorf
from
crosses
(SLJG, NMW).
Fonte
width
Parachleua-stochoerus.
(PDTFAU, MTA, PIMUZ). Oblique
rensis
vs
to
older than San
older than La Grive
Grive
younger than La
Fig.
close
La
Grive
P.
stein-
Following
hyus
Thenius
simorrensis
(1952),
Rabeder
(1978)
cited Cono-
from Klein Hadersdorf.
steinheimensis
However, it has been shown that Para-chleuastochoerus
steinheimensis
Conohyus
actually is no subspecies of
simorrensis,
but
(Chen Guanfang,
Conohyus
(crosses
different
a
reduced
in
Fig.
species
1984; Fortelius
premolar
5)
to
et
size
C.
Parachleuastochoerus has still
in
a
al.,
different
from
C.
ebroensis
smaller
genus
1996). Although
simorrensis
(asterisks),
premolars (trian-
32
gles).
dorf
The
indicates
Within
Conohyus.
C. simorrensis
der
Increase
these
Fortelius
from
order
MTA, PIMUZ), Klein
At
figures
that
and
these do
Taucanamo sansaniensis,
bly
the suid
evolved
Klein Hadersdorf
MN6.
The
are
rate
of
dorf.
The size
Le Fousseret
small suoids,
and
(1952) suggest
and
that author,
by
(see
and
but
possi-
Van der Made,
observation
is
correct,
castellensis
T.
(MN
sansaniensis
and
8)
lineage
late
et
older than
geographic
is
El
al.
late
in
be
in
MN7+8,
Qandir
one
between
age
1
over
the
(G.
( G.
but it is
Ma.
Qandir
al.,
MN
more
a
This
and
is
realistic than the
of
one
(19%).
is
present
Grive;
in St
Stephan
and in the old
these localities
The
Dryopithecus
are
as
old
stratigraphic
and
as
and
Gryphopithecus
The lower levels of Hostalets
(including
younger than Sant
Can
Mata)
Quirze (Agustf
et
is
was
et
Llobateres
al.
is
sequence
(range
MN
placed
and
rynochoerus
meini
aff. lissiensis
are
cise
correlation.
The
use
of the
in
of the
Propotamochoerus
(Ginsburg,
prior
Hipparion
is
locality
we
fossils
such
list
a
as
to
younger than
correlated
did
on
not
most
of
succeed
in
which the cita-
with Can
Rudabanya
of
shared
supposedly
Aceratherium
(MN6-11/12),
naui
not
Eomyops
entry
used
Litho-
the Ara-
just
but
not
present
(MN
incisivum
al.
8-9/10),
to
(1984)
MN 9-
Listriodon
Rudabanya),
useful
of Cricetulodon
et
at
catalaunicus
particularly
by Agusti
and
Ko-
Myoglis
Glirulus
for such
a
pre-
subdivide MN 9,
and the evolution
suid Parachleuastochoerus huenermanni into
crusafonti during
the
the later part of the Crictulodon
following
sequence:
Hostalets,
Can Llobateres and La Tarumba
1990a).
in
appeared
palaeochoerus
(MN5-10),
originally
units.
high
9-12), Hipparion primigenium (range
splendens (MN 6-9,
are
(PDTFAU,
locality
(1996)
Dorcatherium
11),
localities
this
the
taxa
Haders-
from
the basis of
However,
taxa.
Conohyus
Pasalar
different
as
palaeochoerus
on
The
an
Le
in MN 8, when MN 7
placed
the MN 7+8 localities. However,
the P.
and
for Klein
lineage.
treated
cited
species
and
6)
Pinheiro (GML).
locality
local
and indicates that
gives
overlapped.
be
This
tion is based.
that the
weinfurteri)
darwini).
Neudorf-Sandberg.
to
of the
in
and
the
6
Vallecas (MNCN),
still
were
palaeochoerus
1971).
de
St Gaudens
8
stratigraphically,
gonian
MN
6) suggest
MN
(early
Listriodon
on
late in
Fonte do
(MPZ),
1984).
and
Data
Puente
finding
and
Considering
possibility exists,
species
another
ranges of
assumed
Buste
7).
simorrensis-ebroensis
T.
late in MN6 and Pasalar is earlier
might
speculation,
Andrews
Dryopithecus
are
(MNHN),
Hadersdorf (Fig.
between Pasalar
position
a
molars in the Conohyus
late MN 7+8 age. A. pygmaeus
evolution,
collections from La
or
a
difference
Neudorf-Sandberg
by
1996).
grandaeuvus
If this
the
localities have
a
(MN
corroborate
Agustf
Neudorf-Sandberg
course
Fousseret
in MN 7.
or
Neudorf-Sandberg
older
increased form
(MN 8-9) (Van
Göriach (SLJG, NMW, IGGML, NMB),
Thenius
stated
as
into A.
grandaevus replaced
realistic
level
(IPUW),
two
are
evolutive
the first and second
young:
castellensis
Albanohyus
this would corroborate
in
in
Hadersdorf
Taucanamo
1996b, for systematics).
MN 6
to
of the molars from Klein
represent
represent Taucanamo pygmaeum
not
palaeochoerid
al.,
et
mm)
descriptions by
the
(MN 7)
in
old
there
Neudorf-Sandberg,
the
fossils
C. ebroensis
to
in length (DAP;
approximate
suid
molar size
Conohyus,
(MN 5-7)
from Klein Haders-
premolar
that
Made, 1989, 1998;
6.
Fig.
size of the
large
(dot)
(Fig. 7;
Can
P.
zone
Ponsic
I,
Van der Made,
33
Fig.
7.
in
Decrease
La Tarumba
The
this
Ponsic
than
younger
manni
P.
to
of
faunal
samples
Van
The
being
der Made,
transition
be
to
seems
of MN
part
known.
of
10
marked
are
important
an
that these
caused
changes
a
and
in suoid
et
di-
al., 1994).
short
period
of
evolution in Parachleuastochoerus. An alternative
explanation
is
with P.
could be
more
P.
who
and
material
fontani
assign
antiquus,
we
area
younger
D.
Can
Ponsic
think.
than
more
assign
localities
Ponsic
It is
to
and
D.
to
Sinap
like
D. laietanus
has
the
large
species
from Can
localities
1992a),
e/
a/.,
of
Llo-
laietanus and
(Ribot
Eppelsheim (Fortelius
locality
(first locality
one
older
suid
(last
of interest that
fossils
crusafonti (e.g. Begun,
it also
Middle
the
record, implying that
Can
and Can Llobateres
recognise
in
from
or
gap in the
than
crusafonti)
Dryopithecus
bateres
a
time between
huenermanni)
authors
ers
a
by important
decrease
It is
with
clearly
by important changes
der Made, 1990a, b; Fortelius
there
huener-
no
Vallesian is
versity (Van
possible
P.
abrupt,
The
and
1998)
called the 'mid-Vallesian crisis'
of the effects is
rapid
that
and the MN 9-10 transition
caused
probably
in climate. This is
one
The
Parachleuastochoerus huenermanni-crusafonti lineage.
Cricetulodon
Rudabanya suggests
at
7;
(Fig.
important changes
changes,
of the
than Can Llobateres 1 and close in
Hostalets.
and the earlier
M3
Wissberg (NMM),
and
huenermanni
crusafonti
intermediate
period
P.
is older
Can
of the
young:
El Firal
(MGB), Rudabánya (HGSB),
Can
Ponsic I,
localities
Can
are
Llobateres
in
and
(IPS).
al., 1996, fig. 5)
locality
to
age
et
mm)
order from old to
of
presence
(Agusti
in
length (DAP;
approximate
to
the
al.,
is
major.
In
Made &
replaced
1; this
Mein
1994).
al.
et
absence
and in
followed
Mein
11.
common
locality
major
to
tooth
presence
be MN 9
from
and it is
that
possibly
in
by
MN
9
based
Andrews
and
occurrence
al.,
a
on
the
which
(1983)
de-
and believed
Salmendingen
(last
Sal-
his data
of Anchitherium and
yielded
al.,
of that
genus).
possible Microstonyx
of the
1996), suggestive
10-12 elements in the collection from
There
not
are
several
clear where
(Abusch-Siewert,
assume
et
Dipoides problematicus,
et
(Fortelius
of MN
Salmendingen.
area
is
in the Turolian. Abusch-Siewert
However, the 'locality'
to
seems
(Fortelius
locality
der
(Van
major (Kostopoulos,
(1986)
Salmendingen
scribed Anchitherium from
the
10
this
Turkey
subsequently
Melchingen
placed
MN
M.
Microstonyx
MN
same moment
yielded
presence of the castorid
is
in
place
Graecopithecus
(1986),
at
1983).
fissure
the old
It
fillings
in
collections
seems more
hominoid material
the
prudent
from
comes
the
come
to
older
associations, i.e. from MN 9.
This
leaves
Europe
1996).
Dryopithecus
1996).
the
at
locality
in
while oth-
This
case
(1996),
mendingen
the
took
Moya Sola, 1989)
The oldest
1996).
Nikiti
its smaller relative
by
this
Europe
have been the
D.
Hippopotamodon
et
species
as
the
early
went
crisis and there is
as
suggested by
latest
MN
no
record
10
(La
extinct
proof
Mein
of
Dryopithecus
Tarumba,
during
the
in
II);
mid-Vallesian
that it extended
(1986)
Polinya
and Andrews
into MN
et
al.
11,
(19%).
34
THE
DISPERSAL
EUROPE
OF DRYOPITHECINES INTO
The
next
MN
Africa
times,
Aragonian
In
and
Arabia
nected and the Balkan and Anatolia formed
nent, that
tral
was
Europe
continent
faunal
in
seas,
eustatic
sea
with
occurred
East these
shallow
occurred
Steininger,
contact
exchange
Middle
the
&
(Rogl
was
connected
always
not
the
between
At
the
der
(Van
Made,
the
to
two
steps through
Europe
ing periods
al.
level low.
sea
and
ages
mals
that
Miller
Subconti-
Five of such
and
Early
from Af-
place
in
et
al.
climate.
cycles,
each
(1996)
with
cyclical
to
mam-
level
sea
lows
should be simultaneous in the different continents.
persal
events on
eustatic
sea
the continents
level
lows
correspond
given by Haq
as
intercontinental scheme of correlations
the basis of suoid and bovid
used
to
et
to
each
al.
correlate
other and
(1987).
to
This
scheme
magnetic
sections in the
al., 1994, 1996),
For
instance,
boundary
(then
et
Krijgsman
tion in the
suoid
on
new
der
correlations
Made,
of the
based
dispersals
1996a)
are
only
developed
der
on
Made,
was
level
of
cycles
Haq
corresponding
study
of the
area
very well.
D/E
MN
boundary)
palaeomagnetic
a
4/5
by
sec-
Made,
with
et
and
al.'s
the
cycles
hundreds of thou-
some
sands of
years apart and both differ
1992b)
Haq
some
3 Ma with pre-
vious estimates.
In
this
1995, 1996a, 1997b, 1999a),
change
between Africa
ago; this does
not
Set
origin
and
Ilia,
within MN
entered the
or
during
dispersals (Van
the first
Eurasia
the
major
took
have involved
(beginning
next
faunal
21
the
Engelswies
and
ex-
Ma
zone
possibility
&
at
Qiu, 1995).
this
is
The
event
boundary
a
E
wave
of
Pasalar
belong
in
cycle
this
had
southeast
Ma
possible
that
already
for
There
records
record
ous
of
Asian
some
der
(Van
of
Asian(?)
Asian
virtually
mammal
four
entered
as
Dryopithecus
(La
have
and
Grive
oc,
entered
St
entered
ori-
Anatolia
record
of
and
large
Rotem ba-
et
al.,
1987)
bovid
The
zone
primitive
Tethytragus
with
a
with
previ-
the bovid Turcoceros,
Africa,
the
and
deer
Anatolian
origin
in
to
in
western
and
Europe
Made,
been
a
number of
and
the deer
was
central
Dicroce-
reduced
disappeared
have
of
furcatus,
Euprox
der Made, 1993, 1994,
Around this time,
extinct
Crocodiles
der
boundary
6).
Albanohyus,
Europe (Van
seem
(Van
al., 1994, 1996)
to
the suid Parachleuastochoerus,
diversity
there
the
to
belongs
origin
lower
European
went
species.
and
the
the suid Hyotherium
tragulid
to one
Europe
and
1997b, 1999a).
e.g.
and
rus,
no
the end of MN
1999a),
lineages
Protrago-
of African
Fossils from the
in Anatolia,
origin,
origin
1996a, b,
the
more
Not later than
next.
time in
(TB2.6 cycle,
to
southeast
Made,
still
species
long
a
is
in Fort Ternan,
or
or
closer
are
Qandir
primate (Tchernov
Hispanomeryx
previous
their
and Ple-
age.
Set V, close
bovoid
The bovids
of African
Griphopithecus
12.5 Ma
Around
and
again
have
one
from
cycle, though they
a
G
5)
MN
Cricetodon),
the earlier part of the
be of this
lower
entered Africa. Fort Ternan and
14 Ma.
may
into
and
clear whether
include
finds
rodent
(Conohyus)
sin
and
moment,
cycle,
that
Europe
to
to
Europe.
leakeyi
A.
Chinese
(TB2.5
or
Israel
and
within
IV,
(the
mammals older than
in
ago
not
lived
between
by Afropithecus
is
cycle,
Van der
see
this
at
assess
(Sanitherium).
this
to
placed
of this level
relationship
It
Anatolia. It is
gin
to
event
Set
Gentrytragus
end of it.
this
14
Anatolia
Indian Subcontinent
and
a
typical
(Africa;
European
migrants
areas
no
with
Dabtiyah
should be
(the pliopithecids Pliopithecus
siopliopithecus),
cerus
virtually
investigation.
and
zone
of these three
the
dispersal
about
at
in Africa
origin
of
difficult
that
single
a
or
Anatolia and
hominid, Platodontopithecus,
Eurasia merits further
of African
Dionysopithecus
which is
possibility
Griphopithecus
but the
in
Ad
rec-
cycle
the basis of the presence of the suid
on
The
1996a).
in the avifauna
cycle,
entered China
cycle (Qiu
Made,
primates.
4), Dionysopithecus
may have
der
place
of TB2.3
Indian subcontinent.
Platodontopithecus
moment,
and
seem to
About 16.5 Ma ago
C,
Made,
Europe;
model of faunal
of
locality
and which is found also in Maboko
palaeo-
(Krijgsman
of the
age
ages
and the estimates based
der
(Van
An
(1987).
1999a, b) and
zone
on
area
al.
correspond
of the
the
at
Aragonian type
correlation of mammal
(Van
its
but the results
(1994),
dates of the
(Van
Aragonian type
estimates
placed
al.
sea
with
Dis-
in different continents
events
the eustatic
developed independently
was
et
dispersal
et
evolution
1992b, 1993, 1994, 1996a, 1997a, b,
to
was
a
corrected the
of terrestrial
eustatic
by
starting
cycles
Dispersals
allowed for
fluctua-
slightly
level
sea
level
There is
this age
Illb,
to
have reached
to
seems
leakyi (= Heliopithecus)
in Faunal Set Illb,
brought
sea
of
Bunolistriodon akatikubas,
one or
level.
The
Europe.
Afropithecus
represent
dispersals
mammals
Set
this
During
events.
the first hominoid
large
Middle Miocene
have taken
sea
eustatic
global
are
Indian
studied eustatic
(1987)
related
in
changes
exchange
Subcontinent).
to
named and dated the
tions,
of low
the Turkish-Balkan subcontinent dur-
of low eustatic
et
by
minus Balkan
Hominoid
expected
are
In
area.
of
southeast
little
one
previous
D,
zone
makes it difficult
Europe (the Engelswies hominoid).
record
(Europe
in the
1996a).
rica
Haq
Indian
recognised
are
cen-
separated
faunal
moments
Africa-Arabia,
Asia minus the
events
were
during periods
nent, Anatolia-Balkan and Eurasia
and
Himalayan
masses
such
and
The Indian Sub-
1983).
that became land
levels.
subconti-
a
western
boundary
possible dispersal
the anterior
con-
Asia, but apparently
through
land
to
still
were
million after the
one
This short time interval
5).
ognise
but
event was
Ma ago, lower
cycle (15.5
from
from central
important changes
1992a, 1993; Fortelius
et
well.
was
present
Gaudens,
Europe
as
around 12.5 Ma. However,
in
St
part
the
Europe prior
Stephan)
of
the
and
faunal
Aragonian
to
11
may
Ma
well
exchange
hominoid
rec-
35
ord
is
area
sibly
cal
very
in
poor
is
several
it has
other
in
Europe
for
Germany).
in
Spain
If this is the
Stratigraphy
et
very poor
the record
areas
southern
(Fig. 9).
ecological
environments
may have arrived
8.
a
Similarly,
vourable
basin,
for
partially
reasons
MN5-7.
Fig.
The
Europe.
Valles-Penedes is
richest in hominoid localities in
case,
reasons,
but
fossil record
for this
that
might
Alternatively,
in
for
is
its arrival
had
(e.g.
Styrian
fa-
Dryopithecus
seems
to
of European hominoid localities,
al., 1991), and
in
have
the later part of MN
selected African and Arabian
7+8
coincide
the
oldest
localities
an
ridae
range
poor
with
increase in
end of MN 7+8,
geologi-
in the
period
Dryopithecus
the
which pos-
Europe,
(Daams
as
&
hominoid does
12.5 Ma.
absence in
the
to
areas
ecological
by
large parts
of
Spain
Europe
as
in
Europe
with
a
1988).
If the
the
Engelswies
this
genus
faunal exchange
part of the
the
case
of
Dryopithecus,
may have been due
to
of both
hominoids
its
the lack
favourable environment,
preferences
at
the distribution of Casto-
represent Griphopithecus,
Like
western
of record in
in
Freudenthal,
not
may have entered
around
humidity
indicated
are
though
likely
have been different.
Sivapithecus localities
in
(position according
Pickford, 1986;
to
the
Indian Subcontinent (data
Van
der
from
Kappelmann
Made, 1996a).
36
Si-
The 12.5 Ma
dispersal
is also noted in the
event
Indian
der Made,
(Van
9.
Fig.
There are
differences
on the reconstruction
L
=
La
Grive, NS
The arrows
Agustf
et
al.
15.5
Ma
who
(1996),
Dryopithecus
MN
12.5
Ma
there is
have
also
5-6
for
been later
of that
persals)
of
much shorter
were
tiquus,
in
to
confirm
and
the date of the
are
this
is
dispersals
few
of
Si-
MN
geographical
9
10
tapirs
or
and
9, hyraxes
to
7+8 (but
al.,
close
have
habitat
continuous
geographical
lineages
Ankarapithecus
to
assumed
now
not
Anatolian branch
and
it
This
but
is
central
Europe,
Stephan.
Many
and
four
by
and size
are
demonstrate
a
distribution
in favour of
not
how
parallel
pendently
of
to
a
area
was
al.,
on
southeast
from
period
some
starting
time into
to
at
MN 10,
the
MN
MN
early
9-10
10. The
and
et
European
Dryopithecus
its way
may have
into this
to
but is
Dryopithecus
to
an
western
Ankara-
moved inde-
area.
The mid-Vallesian crisis occurred either
tion from MN 9
supposed
1996),
alternatively,
lineage
Dryopithecus
et
Sivapithecus (Alpagut
evolved
though
pithecus -Graecopithecus
of
to
this
in
to
St
(e.g.
clear
not
related
that
passed through
In
have fluctuated and appears
to
Sinap (MN 9)
suggestive
barriers.
distribu-
Hadersdorf,
=
geographic
events are
is
rather
dispersed
an-
their
It
be
to
and
Hippopotamodon
used
been
not
from
for
and murids and in MN
recognised
morphology
Sivapithecus (Andrews
1996).
ecological
SS
al.
if this is what occurred.
al.,
show
Klein
Quirze,
et
impact
should have evolved and maintained themselves,
early
Europe
=
their
overlapping temporal
been
dispersal
no
in
with
separation.
nearly
a
Si-
discussion in main text).
and
Graecopithecus,
even
K
Mata,
Cahuzac
have
likely
very
San
=
Their
1996).
and
in
and
with that of the hominoids.
have
ranges
et
separation
and the
of Can
Dryopithecus
geographic
Andrews
which
see
largely
long,
and Schizochoerus. The
seems
MN
and
Steininger (1983)
Gaudens, SQ
too
palaeochoerus
Hipparion
Microstonyx major
species
be related
al.
et
of
&
inclusive
Saint
=
have coincided
rather
Rögl
much
stasis,
mammals
SG
Pasalar,
dis-
Ma pe-
Agusti
=
when
still later in MN
tion of the
Dryopithecus
assumed
there
must
Sivapithecus
by
that
seen,
the entry
stasis
8, Propotamochoerus
Europe,
first
therefore
of
date of
have
we
so
15.5 Ma. The 5
is
al.
et
period.
Subsequent
MN
As
locality,
not
of
dates for
the
rather than
fact
fossils
or
old
and
P
(Kappelman
Dryopithecus
have been coeval.
stratigraphy,
Hostalets,
=
possible dispersal during
appearance
use
Ma)
independently,
Dryopithecus
the
did
boundary.
Whether
a
Ves),
outside
Steininger
first
12.5 Ma. The
morphological
Dryopithecus
there
than
age.
from
apart
who
Dryopithecus
12.5 Ma
for
(1996)
but
MN 6 and
dispersed
to
the
(16.5-15.2
MN6-7
dispersed
is
discuss
Europe,
the
Nova
after
of
dispersals
seem to
and marine
H
The
1991).
reconstruction
of continental
common ancestor
first
refer
They
not
boundary
MN 6
no
vapithecus
riod
in
that the
Sivapethecus
ago.
did
correlation
palaeogeography. Localities: Ç = Çandir,
indicate the dispersal during
and
in
opinion
Neudorf-Sandberg(Dvinska
(1996) thought
Dryopithecus
Africa
=
of the
of
al.,
vapethecus
Dispersal of Dryopithecinae into Europe; palaeogeographical
(1992).
the
et
12.5 Ma ago, the hominoid
1996a).
entered the Indian Subcontinent
vapithecus
Subcontinent with the entry of Helicoportax and Gazella
in MN 10
boundary
composition
at
or
the transiin
and
of the
a
short
lasting
bovid
37
faunas
in
marked
by
other
Spain
of
parts
changed
the
to
Old
der Made,
(Van
diversity
World
related
towards
the
to
it
1988);
but
Europe,
der
(Van
a
The
is
is
1988,
of murids
as
that
Himalayas (Van Dam,
Abusch-Siewert,
is
1997).
have
we
claims of Turolian
above,
seen
(MN
11)
upheld
and
the
dryopithecids
mainland
on
Europe
be
cannot
that
seems
they
while
crisis,
extinct
went
the mid-Vallesian
during
survived
Graecopithecus
in
southeast
Europe.
M.
tion
Suoidea tend
be
to
hominoid localities
localities
shown
from
in
8:
Fig.
and
for
of
correlating
of
stratigraphic position
of
means
the
and
Indian
is
Griphopithecus
considerable
a
selection of
a
Subcontinent
and
Dryopithecus
coexisted
probably
a
The
(Fig. 3).
Africa
time
in
central
events
these
of known
events
into
dispersals
related
are
Europe
The
chronology (Figs 8, 9).
tend
presented here,
as
published
dates from
nian type
persal
events
About
into
15.5
rise
given
related
not
and
not
to,
ago the
hominoid
arrived
later than
sea
first
in
with
the
the
Arago-
it
level
changes.
hominoid moved
have
may
been,
or
might
If this hominoid
Griphopithecus.
Griphopithecus,
to
phopithecus
dated eustatic
16.5 Ma
This
Europe.
have
and well
sections
seems
likely
that
was
Gri-
later than 14 Ma ago in Anatolia
not
12.5 Ma
ago in
entered
sal within
12.5 Ma
Europe
Europe
Europe. Dryopithecus
ago,
though
Harrison,
M.
W.
E.
its
'Bioeventos
Asian
Fahlbusch
& H.W.
Eurasian
Spain).
Mittmann
Sinap
R.L.
Sucesiones
y
Sabadell
Kappelman,
A
I.
specimen
new
Formation
of cen-
Bernor
of
In:
R.L.
faunas:
Martin,
and
V.
Bernor,
west-
168-208.
Dryopithecus crusafonti
from
Can
New
Phyletic
and
Am.
—
new
(northeastern
291-309.
diversity
hominids.
a
nov.,
sp.
Ponsic
87:
phys. Anthrop.,
primitive European
L.
Univ. Press).
(Columbia
1992b.
&
European
The evolution of
(eds).
mammal
Neogene
Am. J.
—
D.R.,
the
catarrhines.
Mioceqe hominoid species
Begun,
J.
1996.
biochronology
Miocene
1992a.
Begun, D.R.,
(Mam-
Coloquio
—
Iberico',
Fortelius,
Lindsay,
Delson,
and
southwest
York/Chichester
Bovidae
penin-
33-46.
Nature, 382: 349-351.
Distribution
in
locomotion
J.
phys.
Anthrop.,
87:311-340.
S.
Begun, D.R.,
homonoid
& M.
Moya-Sola
Evol.,
19:
1990.
Kohler,
form Can
specimens
Llobateres
and
Spain) and their geological
J. human
probably
&
dates for
consistent
—
5.
global
model of intercontinental dis-
a
T.
1988.
Continental
of Ankarapithecus metai from
—
14:
Peninsula Iberica.
la
Adrover,
Qelebi
tral Anatolia.
es-
distribu-
Iberian
cont.,
Moya-Sola,
Andrews,
H.
Temizsoy,
1996.
to
P.
B.,
Alpagut,
of
hu-
Sinopsis
Mammal
of the
margin
Paleont.
—
Terciario
el
en
Resumenes:
1988,
be younger than
to
are
palaeomagnetic
and with
or
but
previously,
long
area
Miocene.
Rafael
a
J.
—
Valles-Penedes.
1984b.
Gibert,
J.
S.
faunisticas
1984a.
Cabrera,
eastern
de
ern
those
the
the
&
Andrews, P.,
Europe.
Hominoid
in
Morales
homenaje
provides
hominoid localities
European
most
L.
in hominoid localities. Suoid
common
and bovid evolution
&
Moya-Sola
J.
&
timing
reconsidered.
radiation
&
malia) del Neogeno
Alcala, L.,
Garces
and
18: 53-81.
dynamics
sula during
conclusions
M.
Cabrera,
the pattern
Neogeno de la fosa del
Evol.,
S.
Agusti J.,
and
Fund-
143-155.
Moya-Sola
del
L.
Llobateres:
hominoid
31:
S.
tratigafica
Paleont.
Summary
der
Beriicksichtigung
Cour. Forsch.-Inst. Senckenb., 62:
Moya-Sola,
1996. Can
Evol.,
Agusti, J.,
S.
Kohler,
Pares,
Vallesian
man
it
—
(Equidae,
Anchitherien
1-401.
Agusti, J.,
J.M.
As
besonderer
unter
Sandelzhausen.
a
Unter-
CiebiBmorphologischc
eurasiatischen
an
Mammalia)
1983.
S.,
suchungen
stelle
interpreted
climate
REFERENCES
in
not
Made,
dispersal
event
seasonal
more
of the
uplift
and the
1996a)
Europe (Van Dam, 1997).
global change
al.,
et
in
the worldwide extinction of Listriodon-
1990a, b, 1991),
tinae
(Alcalä
in suoid
drop
a
New
Miocene
(Valles Penedes,
context.
paleoecological
—
255-268.
disperBruijn,
may have been diachronous.
H.
P.
A.J.
hew,
G.
Daams,
J.
Mein,
der
van
1992.
Antunes,
fossil
R.
de,
Ginsburg,
Meulen,
Report
E.
Fahlbusch,
Heizmann,
RCMNS
1990.
May-
M.
Telles
working
Newsl.
—
L.
D.F.
N. Schmidt-Kittler &
of the
mammals, Reisensburg
Acknowledgements
V.
Daxner-Hock,
Morales,
on
group
26:
Stratigr.,
65-118.
J.
Cahuzac, B.,
The late Dr L. Via and Dr S.
tor, allowed
us
to
L.
Alcala,
B.
and
study
noid fossils from the
Calzada,
publish
the present direc-
both suid and homi-
MLGSB collections. Drs J.
B.
Alpagut,
M.
Ertiirk,
Azanza,
Fortelius,
G.
Daxner-Hock,
M.
Freudenthal,
Engesser,
Q.
Ginsburg,
J. Gomez Alba, W. Graf, C. Guerin, E.
E.P.J.
Heizmann,
Kohler,
L.
Moya-Sola,
G.
M.
Kordos,
R.
Hugueney,
H.
Niederl,
Scharfe,
Lutz,
M.
P.
K.A.
Philippe,
J.
K.
B.
L.
Giileg,
Hiinermann,
Mein,
M. Telles Antunes,
Agusti,
Morales,
Rauscher,
M.
S.
G.
P.Y. Sondaar and
Pujol,
Unay
Daams,
—
R.
&
us
allowed
us
to
study
material
in
their
care
M.
A.J.
Suidae
the
and
Miocene
Freudenthal,
—
van,
of
Montenat
&
northeastern
Mediterranean
C.
At-
sea.
—
Tayassuidae (Artiodactyla,
of
184:
1988.
program
in
Steinheim
The
small
A.
a.
(Ger-
79-83.
Synopsis
the
of
the
Aragonian
Scripta Geol., Spec. Issue,
1997.
the
the
C.
the
of
279-294.
Palaeontographica,
1975-1986.
Dam,
maps
relation with
collaboration
Spanish
mammals
1:
Dutch-
type
area,
3-18.
from
the
Upper
Teruel-AIfambra region (Spain): paleobiol-
or
ogy
helped
1984.
Mammalia) from
Miocene
E.
and
Evol., 24/25:
Chen Guanfang,
many).
Lauriat-Rage,
Palaeogeograpic
lantic Neogene
Paleont.
A.
Alvinerie,
1992.
and
paleoclimatic
otherwise.
tina,
156:
1-204.
reconstructions.
—
Geol.
Ultraiec-
38
P.
Fortelius, M.,
in
provinciality
the
Abstr.:
the
1994.
Werdelin,
land
large
Eurasia.
western
of
L.
diversity,
mammals
from
and
Neogene
—
Krakow
Palaearctic,
and
turnover
1994,
J.
Van
der
and
Late
ern
Mediterranean:
Miocene
In:
cology.
R.L.
Bernor,
central
V.
and
Europe
Fahlbusch
of western
east-
paleoe-
Mittmann
Neogene mammal
Eurasian
York/Chichester
New
the
and
H.W.
&
Middle
1996.
Evolution, biogeography
Bemor,
348-377.
R.L.
Made &
Suoidea of
The evolution
faunas:
Univ.
(Columbia
1971.
Ginsburg, L.,
vindoboniens
J.Ma.
de
gidae)
los
II:
Espana).
de
bassins
78:
mammiferes
faunes de
Les
des
BRGM,
Golpe Posse,
tute
de
la
et
Loire
burdigaliens
de la
Garonne.
et
—
Los
1993.
del
del
Description
Pon-
hispanopithecos (Primates,
yacimientos
Valles-Penedes
material existente
de
Paleontologfa
J.
B.U.,
&
Sabadell.
P.R.
Insti-
el
en
Evol.,
1987.
Vail,
of
Chronology
235:
Science,
—
1156-1166.
Beitr.
Stuttg.
F.
Grosssaugetiere.
Kappelman, J.,
Raza &
Pakistan.
mates. In:
G.E.
Rossner
mammals of
the
nia, Greece;
Soc.
Greece,
Meulen,
Miocene
climate
Aragonian
type
Earth
Dam,
A.J.
record
Made,
van
J.
in
Coloquio
—
der
Made, J.
dactyla)
—
ont.,
J.
Made,
Made,
J.
and
sity
van
van
in
the
Kon.
of
of
Pri-
of 'Nikiti
species.
!', Macedo-
—
Bull.
the
A
der,
Made,
Made,
J.
&
A.J.
dating
128:
of
van
the
der
middle
of the
Agusti
R.
& L.
to late
Planet. Sci.
a
513-526.
Langereis,
J.
Suoidea
J.
Cabrera,
(pigs
142:
and
Van
1996. A
J.
peccaries).
'Bioeventos
y
Iberico',
Conohyus-lineage (Suidae, Artio-
Miocene
of
Europe.
—
Rev.
esp.
Pale-
Iberian
Suoidea.
—
Paleont. Evoi.,
1990b.
der,
Suoidea.
—
1991.
—
A
range
Paleont.
chart for
Evol., 23:
Climatical
China
Suidae
99-104.
changes
XIIIINQUA,
European
and
species diver-
1991,
Abstr.: 365.
pig.
—
classification
a
Mammalia).
Middle
and
Proc.
—
21: 75-81.
Montpellier,
from
events,
Miocene stratigra-
the
mine
open-cast
Aureliachoerus
from
Aragonian pigs
other
the
of
stratigraphy
and
Intercontinental dispersal
Styria.
—
Wien, 99A: 225-277.
Intercontinental
relationship Europe
Subcontinent. In:
Miocene
land
G.E.
mammals
Biometrical
Sciences 89:
&
S.
from
28:
paleont. It.,
trends
Rossner
of
&
-
K.
457-
Europe:
European Neogene.
evolution:
the
TetraconoSoc.
Roy.
Edin-
199-225.
1989.
Moya-Sola,
the
in
Trans.
—
Miocene
Late
Suinae
European
onwards.
Boll.
—
329-339.
of
Mein, P., 1986. Chronological succession
mate
pis.
Miocene
(F. Pfeil).
der
van
In:
J.G. Else
59-71.
&
hominoids
Ph.C.
Cambridge
Lee
the
in
Pri-
(eds).
Univ.
(Cambridge
Press).
New
G.S.
Mountain
et
global
and
level,
sea
sequences:
records.
Exxon
and
Drilling
1996.
al.,
Jersey Oligocene-Miocene
—
ice
dating
volume,
271:
Science,
1092-1095.
aus
Bericht
Osterreich,
von
iiber
St.
die
neuen
Stefan
im
Menschenaffenfunde
Lavanttal,
Karnten.
—
Carinthia, (11)67: 39-84.
1981.
Pickford, M.,
Pickford, M.,
faunas
of
Preliminary Miocene mammalian
western
Kenya.
—
J.
human
Evol.,
10:
biostrati73-97.
Geochronology of Miocene higher primate
1986.
East
Africa.
evolution:
In:
J.C. Else &
19-34.
Ph.C.
Cambridge
Lee
(eds).
Pri-
Univ.
(Cambridge
Press).
Qiu Zhanxiang
der,
and
Mus.
The
time and
19
127-139.
Vertebrates
1999a.
(Artiodactyla)
mate
1990a.
and
(Suoidea,
subfamily of pigs.
a
graphy for
21.
small
a
in
3-254,
33:
Schizochoerus
Early
der, 1999b.
Earth
burgh:
and
the Indian
Mottl, M., 1957.
367-380.
Terciario Continental
20,
J.
Miocene continental
Lett.,
van
Miller, K.G.,
Daams,
Rafael Adrover,
el
der,
and
Pa-
(Suidae, Mammalia),
(Western Styria Basin, Austria).
472. Miinchen
Made,
and
1998.
der,
XI J.
—
187-189.
distribution
Albanohyus,
Mem. Trav. EPHE Inst.
van
Africa
and
Quatern. Geol.,
1997b.
sea level
van
antelopes.
Resumenes:
Wet., 100:
der,
van
—
J.
Made,
geol.
Calatayud-Teruel basin (Central
Iberian
en
J.
eustatic
dontinae,
(F. Pfeil).
level changes and disper-
sea
Miocene
1997a. Systematics
Akad.
of the
Eustatic
Palaeochoeridae
Ned.
Europe
38: 293-303.
Taucanamo
Heissig (eds).
Order
continental deposits
the
Resumenes:
Tayassuidae.
J.
of
der,
van
Oberdorf
19-28.
van
J.
in
Mammals
78-79.
systematics
cracov.,
naturhist.
83-97.
23:
Made,
4:
Made,
europ.
1994, Abstr.:
1996b.
der,
Ann.
Formation
1999.
Daams
Sci. Lett.,
Earth
1989.
der,
from
the
homenaje
1988,
van
R.
for the middle
Sucesiones faum'sticas
Sabadell
in
Meulen,
1988.
der,
van
Acta zool.
Soc.
Garces, C.G.
Spain.
van
in
Planet.
chronology
new
Made, J.
from
occurrence
Chinji
Miinchen
about the
Langereis,
change
Neogene
Contr. Tert.
Ward,
Microstonyx major (Suidae, Artiodac-
area
M.
—
Congr.
Quaternary
1996a. Listriodontinae
der,
van
Johnson,
Heissig (eds). The Miocene
Magnetostratigraphic
Spain).
Krijgsman, W.,
N.M.
earliest
Andrews,
91-104.
remarks
C.G.
1994.
—
J.
Miocene
Barcelona
of a Middle
timing
the
Premier
—
and
leonotologi'a, Tremp 1995,
Made,
and Middle
Paleontologla,
antelope Caprotragoides
and Middle
Early
J.
128.
der, 1995.
van
of
Oberdorf
30: 341-355.
W.,
Krijgsman,
P.
Late Miocene locality
some
S.
Sheikh,
Hake,
Miocene
& K.
Europe:
1994.
Kostopoulos, D.,
tyla) from
&
Mioziine
21: 61-73.
Evol.,
Moya-Sola
K.A.
P.
1991. The
middle
1996.
Tassy,
Naturk., C39: 1-60.
Brown,
S.M.I. Shah
J. human
—
S.
M.,
land
Made, J.
change.
Krakow
palaearctic,
Lower
—
Artiodactyla and
1993:
—
Cour
—
87-97.
1994. The
der,
Turkey.
phy.
P.
Pilbeam,
B.
from the
&
Beitr.
D.
Barry,
—
1-61.
Stuttg.
Kelley,
J.
Sivapithecus
Kohler,
C33:
Duranthon
—
J.C.
Anwar,
DasTertiar in Stidwestdeutschland.
Naturk.,
Heizmann, E.P.J.,
S.M.
Made J. van
and
African
climatical
Paleont., Lyon
climate.
27-39.
153:
peccaries).
1993.
der,
Miocene
Made,
1992.
Heizmann, E.P.J.,
&
(pigs
1992, Resumenes:
space.
(Catalufia,
Paleont.
—
the Triassic.
levels since
sea
1992b.
der,
van
genera
Hardenbol
fluctuating
M.
J.
Suoidea
Senckenb.,
their evolution,
153-167.
26/27: 151-224.
Haq,
Made,
and
1992a. Migrations
der,
van
Forsch.-lnst.
sals
Press).
Mem.
J.
Made,
Made J. van
23.
Fortelius, M.,
(eds).
of
subsample
a
mammals
Quaternary
Benor &
of taxonomic
Miocene of
later
R.L.
Andrews,
Preliminary analysis
and
—
Qiu Zhuding,
of
Chinese
1995.
Chronological sequence
Neogene
mammalian
Palaeogeogr., Palaeoclimatol., Palaeoecol.,
Rabeder, G.,
1.
&
subdivision
Cicha,
1978.
J.
Die
Senes
Saugetiere
&
F.
des
Badenien.
Steininger (eds).
116:
In:
faunas.
41-70.
A.
Papp,
Chronostra-
39
und
tigraphie
1993.
Ribot, F.,
Miozan
Neostratotypcn,
480. Bratislava
Akad.
(Slowakische
M4
Revision taxonomica de los
Catalunya. Barcelona (Univ.
Badenien:
467-
Barcelona),
auton.
de
dryopithecinos
pp.
1-419
the
J. Gibert
&
T.
of
taxonomy
Catalonia (Spain).
Rogl,
F.
&
F.F.
Mediterran
85A:
Sen,
S.,
pean
Harrison,
—
J. human
Parathethys.
—
1997.
reinterpretation
of
Vallees-Penedes,
31:
Neogene
dem
7:
der
Zerfall
Ann.
naturhist.
Tethys
Mus.
zu
F.,
Neudorf
Wien,
Abh.
Villalta
mammal
chronology.
1967.
133:
Einweiterer
Miozan
the
Euro-
Palaeogeogr.,
Pa-
Zahn
von
Dryopithecus
(Mammalia, Pongidae)
Beckens.
—
Folia
P.
Ginsburg,
Tassy
&
N.F.
faunas:
Goldsmith,
Negev (Israel).
of the
1952.
an
Die
der
Saugetierfauna
March
(CSR).
7-
J.
—
1987.
Pale-
vert.
aus
Neues
Jb.
dem
Torton
Geol.
u.
von
Palaon-
96(1): 27-136.
J.F.
de
&
M.
‘Dryopithecus fontani’, Lartet,
Crusafont
en
el
de
la
55:
Die untermiozanen Kleinsaugerfaunen
aus
Valles-Penedes.
geol.
min.
1941.
Espana,
—
Bol. Inst.
Pairo,
Vindoboniense
3-15.
Ziegler, R.,
den
181-204.
1902
des Wiener
—
of
L.
mammals
Cornelia,
cuenca
calibration
mammal
284-310.
Thenius, E.,
tol.,
129-141.
Vom
Magnetostratigraphic
(Dry.) fontani darwini Abel,
7:
Evol.,
1983.
laeoclimatol., Palaeoecol.,
aus
from
Dryopitehcus
Steininger,
und
1996. A
135-163.
Steininger,
Miocene
Neogene
York/Chichester (Columbia Univ. Press).
Tchernov, E.,
ont.,
(unpubl.).
Ribot, F.,
of western Eurasian
evolution
46. New
Wiss.).
bei
1995.
Siisswasserkalken
von
Engelswies
Sigmaringen (Baden-Wiirttemberg).
Naturkunde,
B228:
Schellenfeld
und
—
Stuttg.
Beitr.
1-53.
primatol.,
243-275.
Steininger, F.F.,
Sen
&
J.
chronologic
R.L.
W.A.
Agusti,
Bernor,
Berggren,
1996.
correlations
V.
D.V.
Kent,
R.L.
Bernor,
S.
Circum-Mediterranean Neogene
of
Fahlbusch
European
&
H.W.
mammal
Mittmann
units.
(eds).
In:
The
Manuscript received
September
1999.
4
May
1999, revised
version
accepted
10