Escolar Documentos
Profissional Documentos
Cultura Documentos
The MN
System and Continental Stage/Age Concepts Discussed
Author(s): J. A. Van Dam, L. Alcal, A. Alonso Zarza, J. P. Calvo, M. Garcs and W.
Krijgsman
Source: Journal of Vertebrate Paleontology, Vol. 21, No. 2 (Jul. 20, 2001), pp. 367-385
Published by: Taylor & Francis, Ltd. on behalf of The Society of Vertebrate Paleontology
Stable URL: http://www.jstor.org/stable/20061959
Accessed: 26-06-2016 14:22 UTC
Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at
http://about.jstor.org/terms
JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted
digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about
JSTOR, please contact support@jstor.org.
Taylor & Francis, Ltd., The Society of Vertebrate Paleontology are collaborating with JSTOR to
digitize, preserve and extend access to Journal of Vertebrate Paleontology
This content downloaded from 163.118.172.206 on Sun, 26 Jun 2016 14:22:20 UTC
All use subject to http://about.jstor.org/terms
institute of Earth Sciences Jaume Alera, CSIC, Mart? Franqu?s s/n, 08028 Barcelona, Spain;
5Paleomagnetic Laboratory, Ford Hoofddijk, Faculty of Earth Sciences, Utrecht University, P.O. 80021, 3508 TA,
Utrecht, the Netherlands
ABSTRACT?An extended and revised mammal succession of 99 fossil localities from the Upper Miocene sediments
of the Teruel-Alfambra region (NE Spain) is presented. An updated biozonation is proposed. The biostratigraphic
justification for the correlation of the magnetic polarity patterns of the La Gloria, El Bunker, Masada Ruea, Masada
del Valle and Mas?a de la Roma sections to the Geomagnetic Polarity Time Scale (GPTS) is discussed.
A comparison with Late Miocene faunas from elsewhere in Europe demonstrates that faunal resemblance across the
continent is very low. As illustrated by an analysis of the "Progonomys event," local appearances of genera may be
strongly diachronous and even species should not a priori be assumed to be isochronous at resolutions higher than
several hundred thousands of years. These observations have implications for European continental stratigraphy and
chronology: (1) The usefulness of the European mammal-based Stages/Ages can be doubted because their biostrati
graphic significance is mainly local, and because more and more direct calibrations of mammal faunas to the numerical
time scale are becoming established; (2) The European Mammal Neogene (MN) system, currently defined as a series
of 16 time-ordered faunas, should not be divided into sub-units, because this weakens its power for cross-continental
faunal correlation. In addition, the use of MN "boundaries" is erroneous and misleading, both from a philosophic and
technical point of view.
INTRODUCTION
namics studies (van Dam, 1997; van Dam and Weltje, 1999).
human) bones from Concud were taxonomically described.
A shift towards paleoenvironmental reconstructions has also
From the 1940s onwards, the Spanish paleontologists Crusa characterized the work on the macromammals. For instance Al
operation. Among these are the monograph on Hipparion by should be regarded as a half-graben, with the NNE-SSW ori
ented master fault running along the eastern border of the basin
(Fig. 1).
367
This content downloaded from 163.118.172.206 on Sun, 26 Jun 2016 14:22:20 UTC
All use subject to http://about.jstor.org/terms
purposes, for instance between the middle part of the Mas?a del
Barbo section and the upper part of the Puente Minero section,
and between the La Gloria and El Bunker sections (Fig. 3), The
carbonates may alternate with greyish marls and clays or black
lignites, all of which may contain mammal remains. The unit
reaches a maximum thickness of more than 60 m.
(NE Spain).
The studied fossiliferous organic-rich marls belong to the the degree of resolution for those assemblages is not better than
(1976). The carbonates associated with these facies units dis of isolated teeth, is exceptionally dense. The total number of
teeth exceeds 20,000. Rodent data for MBA-B, PER4, PERA
play features typical of sedimentation in shallow lake environ
ments and include: (1) carbonate paleosols showing distinct D, ALF (partly), TOA-C, VIP (a small fraction), MDV2-7, TO,
VB, VB2 (a small fraction), CC, CCB, CCL, CC2-3, TO, LM,
maturation stages (Machette, 1985); (2) carbonate pond se
quences (Sanz et al., 1995); (3) palustrine carbonates, i.e., root and VDC3-4 are from van de Weerd (1976). Species identifi
ed micrites and biomicrites (Platt, 1989; Alonso-Zarza et al., cations of Desmaninae (Talpidae) are from R?mke (1985). Des
1992), mottled carbonates and marls, and nodular carbonates;
manella (Talpidae) from CC3 and LM was described by R?mke
(4) tufa deposits (Pedley, 1990); (5) carbonate channel fills,
(1974). Lagomorphs from MBB, VIP, CC3 and MDV2, 4 and
locally rich in gastropod opercula; and (6) tabular, gastropod 6 were described by L?pez Mart?nez (1989). The remaining
rich carbonate beds. Superposition of carbonates displaying ex
tensive rooting and other palustrine features on primary car
bonate lithofacies, such as tufas and carbonate channel fills, is
micromammal data from ALJB and a part of the data from VIP
This content downloaded from 163.118.172.206 on Sun, 26 Jun 2016 14:22:20 UTC
All use subject to http://about.jstor.org/terms
369
LEGEND
Quaternary,
including Villanyian
* fossil locality/
localities
~vl Neogene
1 i ' i ' I Neogene
Neogene
red/orange conglomerates,
|^\^N| Mesozoic
FIGURE 2. Schematic geological map of the northern part of the Teruel basin (Teruel-Alfambra region) with positions of sections and mammal
localities. See Figures 7-8 for locality names. Note that GLO = La Gloria 6, 14A and B, GL = La Gloria 4 and 5, and GLO/AG = other La
Gloria and Los Aguanaces localities. MNT = Montalvos.
LOCAL BIOZONATION
R2, ALF, VIP, and VB2/2C are extended. For a review of older
Macromammal identifications are from Alcal? et al. (1986), omys ramblensis Zone. Here we propose a second three-parti
for PM from Alcal? et al. (1991), for VDC5 from Adrover et tion subdividing the Progonomys hispanicus Zone.
al. (1986), for BUN and VDC3 from Alberdi et Alcal? (1990).
For the sake of convenience, we adopt and extend the letter
The data for AG were compiled by Alcal? and Montoya (1990),
system (A, B up to I) introduced by Daams and Freudenthal
and those from ALJB by Alcal? (1987). The rest of the macro
(1981) for the Early to early Late Miocene sediments of the
(1994). Besides new data, this work includes the updating and
completion of faunal lists from Alcal? et al. (1994), Alcal? and
This content downloaded from 163.118.172.206 on Sun, 26 Jun 2016 14:22:20 UTC
All use subject to http://about.jstor.org/terms
T~2
?==3 CC(MN12-L)
?==3 CCB(MN12-L)
LAS CASIONES
SECTION
claystones
palustrine carbonates
(including palustrine paleosols
and tufa)
lithostratigraphic correlations:
- ascertained
-probable
- -?- - possible
MASADA RUEA
SECTION
no diagnostic faci?s
FIGURE 3. Stratigraphie columns for the southern part of the Teruel-Alfambra region.
Zone I?After Daams and Freudenthal (1981). Diagnosis: Hispanomys peralensis is mainly restricted to this zone. Sciur
The lower boundary is defined by the entry of the cricetid Cri ids (Heteroxerus) are rare. Many species have their last occur
cetulodon and the glirid Ramys multicrestatus. The base of the rences: the cricetids Cricetulodon and Democricetodon, the
next-higher zone is not defined by Daams and Freudenthal, but glirids Tempestia, Muscardinus hartenbergeri, and Myomimus
is taken here at the first regular occurrence of Progonomys his dehmi, and the soricids Miosorex, Crusafontina and genus and
species 1. Crusafontina shows its greatest abundance. Another
panicus.
soricid, Paenelimnoecus has its first occurrence.
Additional micromammal data: Cricetulodon and Hispano
Additional macromammal data: Macromammals are scarce
mys are important. Megacricetodon debruijni is present, M. ib
ericus is absent. Among the Gliridae Myomimus dehmi, Tem and poorly studied. The presence of the robust equid Hipparion
pestia hartenbergeri, Muscardinus hispanicus and Ramys mul primigenium and rhinocerotids is characteristic. The latter fam
ticrestatus are common. Some rare eomyids (Leptodontomys ily is represented by three genera, of which Alicomops alf?m
brense is restricted to this zone. The giraffid Decennatherium
catalaunicus) and murids (Progonomys cf. hispanicus) occur in
pachecoi and the suid Microstonyx are common. Bovids are
this zone. Miosorex is a common soricid, together with an as
yet unnamed genus (gen. et sp. 1). Characteristic macromam represented by Tragoportax gaudryi (Boselaphini), a species
which is also present in zones K and L, and by Aragoral mu
Zone J?Progonomys hispanicus Zone of van de Weerd dejar, a form considered to be ancestral to the caprines. No
(1976). Diagnosis: Progonomys hispanicus-Parapodemus lug cervids are documented although Micromeryx (Moschidae) is
This content downloaded from 163.118.172.206 on Sun, 26 Jun 2016 14:22:20 UTC
All use subject to http://about.jstor.org/terms
371
a
?^3 BUN(MN13-I\
A?A
?A?
A?A
?A?
A?A
A?A
7\?7\
N^^3 PM3(MN11-K)
?^3PM(MN11-K)
- -i
?-^-l 6
A', i i,
r
, J&MB2B(MN10-J2)
?^3 MB2A(MN10-J1 )
5AG6 (MN10-J3) - -r
3AG4(MN10-J3)
7T^T
Hi
3GL011 (MN10-J3)
?AG5B (MN10-J3)
AG5A (MN10-J3)
^3\PM13(MN10-J3)
?
?T^3 PM10(MN10-J2)
3 ;
^^3 PM8(MN10-J2)
?^=3 PM2(MN10-J2)\
<^^> PM1 (MN10-J1) ^
PUENTE MINERO
SECTION
EL BUNKER
LA GLORIA
SECTION
SECTION
A A
A
A A
A
A? A
A A
AAA A
A|
A A Al
MAS?A DEL
BARBO SECTION
LOS MANSUETOS
SECTION
FIGURE 3. Continued.
gacricetodon is absent and Cricetulodon is rare. Among the Additional micromammal data: Progonomys cathalai and P.
glirids, Muscardinus, Tempestia, and particularly Myomimus hisp?nicas are the only murids. Cricetulodon and glirids are
dehmi are common. The soricid Miosorex is a common insec rare. Crusafontina is a common insectivore. Desmanella is rare.
tivore. The talpid Desmanella is absent.
This content downloaded from 163.118.172.206 on Sun, 26 Jun 2016 14:22:20 UTC
All use subject to http://about.jstor.org/terms
^^ &
mas?a de la roma
SECTION
alcalai and Castromys inflatus enter the area at the base of this
zone. Castromys inflatus is present only in the lower part of the
FIGURE 4. Stratigraphie columns for the northern part of the Teruel zone and Paraethomys miocaenicus in the uppermost part.
Sciurids are very rare. The insectivore association resembles
Alfambra region (the Peralejos area).
of Parapodemus lugdunensis.
occurrence of Hystrix.
cuta eximia and Paramachairodus orientalis. The giraffid De sia interval zone, from the entry of Paraethomys miocaenicus
cennatherium is replaced by Birgerbohlinia schaubi. Dorcath to the entry of Celadensia.
This content downloaded from 163.118.172.206 on Sun, 26 Jun 2016 14:22:20 UTC
All use subject to http://about.jstor.org/terms
FIGURE 5. The Mas?a de la Roma (ROM) area. All mammal localities (indicated by numbers) belong to carbonate faci?s unit M3 of Hernandez
et al. (1983). The local top of this unit can be easily recognized by the extensive white limestone bed, which is situated about 8 m above locality
ROM11. The limestones are followed by some 25 m of predominantly reddish mudstones, which, in turn, are succeeded by limestones constituting
the top of the cuesta. The presence of a small down-glided block containing the locality ROM1 precludes the exact bed-to-bed correlation of ROM2
3 to the main section containing ROM 4-11. Unfortunately, most of the original fossil localities were destroyed by recent road construction works.
NUMERICAL AGES
sediments in the area usually provide reliable paleomagnetic lation of quantified evolutionary stages of Progonomys and Oc
results. The whitish limestones appear to have much lower citanomys (Muridae, Rodentia). For paleomagnetic details the
magnetic intensities, and their demagnetization diagrams are reader is referred to Krijgsman (1996), Krijgsman et al. (1996),
generally more difficult to interpret. Our preferred correlations and Garces et al. (1999). The correlation to the GPTS of the
of the polarity sequences of the La Gloria, El Bunker de Val middle part of the La Gloria section is different from that pro
decebro, Masada Ruea, Masada del Valle, and Mas?a de la La posed by Garces et al. (1999), due to a re-interpretation of a
FIGURE 6. The La Gloria-Los Aguanaces area. Numbers refer to mammal localities. Mammal localities are indicated by dots. AGI and 3, of
which the stratigraphie positions are indicated by arrows, are situated 400 m south-west of the other AG localities. Three main carbonate units
can be seen: the lower one contains AG4-7, the middle one AGI and 3 and the upper one the lower Pliocene locality GL04 (not included in
this study). The carbonate units alternate with detrital sediments units which mainly consist of reddish mudstones.
This content downloaded from 163.118.172.206 on Sun, 26 Jun 2016 14:22:20 UTC
All use subject to http://about.jstor.org/terms
374
TABLE 1. The sequence of Late Miocene localities of the Teruel-Alfambra region: micromammals. Estimated ages according to the time scale
of Cande and Kent (1995). Use of Muridae and Cricctidae according to Chaline et al. (1977). Symbols: cf with a lowercase first letter of a species
(e.g., cf h) refers to the species part of the name only (Progonomys cf. hispanicus). cf with an uppercase first letter of a genus (cf P) refers to
the total name (cf. Progonomys hispanicus).
"5 P
II
II
?1 Arquillo 4
El Arquillo 1*
La Olor?a 5
ARQ4
ARQ1
Bunker de Valdeecbro
GL05
ML
KSS
KS
VB1
MOD
CAP3
CAP2
REG5
BUN4/5
VDC6
VDC3
MDV7
BUN
GLOl
MDV6
VDC5
Milagros
Las Casioncs superior
Las Casioncs
Villalba Baja 1
Modorras
El Capon 3
El Capon 2
Regajo 5
Valdeecbro 6
Valdeecbro 3
Masada del Valle 7
La Gloria 6
La Gloria 1
Coneud
Concud 3
Los Mansuetos*
Aljezar B
Concud barranco
Concud 2
Tortajada
Regajo 4
Regajo 3
Tortajada D
Tortajada C
Las Casioncs 2
Concud B
Masada Ruca 4
Masada Ruca 3
Tortajada B
Valdeecbro 4
Vivero de Pinos
Tortajada A
Los Aguanaces 3
Los Aguanaces
Regajo 2
Alfambra
Puente Minero 3
La Gloria 10
Los Aguanaces 1
Masada Ruca 2
Los Aguanaces 7
Peralejos D
La Gloria 11
La Cantera
Los Aguanaces 6
Los Aguanaces 5B
Los Aguanaces 5A
Peralejos C
Puente Minero 13
Masada Ruca/M.R.2A
Los Aguanaces 4
La Roma 2
Peralejos B
La Roma 1
La Cantera 2
La Gloria 14A/B
Puente Minero 10
Masia de la Roma 11
Puente Minero 8
Mas?a del Barbo B*
Puente Minero 2
Peralejos A
La Saue
Masia de la Roma 1
Peralejos 4
Masia de la Roma 9
Masia de la Roma 8
Puente Minero 1
Mas?a del Barbo A
Mas?a de la Roma 7
Masia de la Roma 6
Masia de la Roma 5
Mas?a de la Roma 4C
Masia de la Roma 4B
Mas?a de la Roma 3
Peralejos 3_
GL06
CC
CC3
LM
ALTO
CCL
CC2
TO
VB2/2C
REG4
REG3
MDV5
MDV4
TOD
TOC
KS2
MDV3
CCB
MRU4
MRIT3
MDV2
LM2
TOB
BITN6/7
VDC4
VIP
TOA
AG3
AG
REG2
ALF
PM3
GLO10
AG?
MRU2
PM5A/B
PM
AG7
PERD
GLOll
CAT
AG6
AG5B
AG5A
PERC
PM13
MRU
AG4
R2
Rl
CAT2
GL14A/B
PM10
ROM11
PM8
MBB
PM2
PERA
SAL
ROM1
PER4
ROM9
ROM8
PM1
MBA
ROM7
ROM6
ROM5
ROM4C
ROM4B
6.2
6.2
6.3
6.7
6.7
6.7
?fP
6.9
6.9
6.9
7.0
7.0
7.1
7.1
7.1
7.1
7.1
7.3
7.3
7.5
7.9
7.9
8.1
8.1
8.2
8.2
8.2
8.3
8.3
8.3
8.3
8.6
8.7
efH ?fH
8.9
8.9
8.9
8.9
9.0
9.0
9.2
9.2
9.2
9.3
9.3
9.3
9.3
9.3
9.4
9.4
9.4
9.4
9.4
9.4
9.5
9.5
9.6
ROM3 9.7
PER5 9.7
This content downloaded from 163.118.172.206 on Sun, 26 Jun 2016 14:22:20 UTC
All use subject to http://about.jstor.org/terms
I3*
375
ill
III
ARQ4
ARQ1
111
Talptdae
* *
111111
L-L
GL05
ML
KSS
KS
VB1
MOD
CAP3
CAP2
REG5
BUN4/5
VDC6
VDC3
MDV7
BUN
GL06
GLOl
MDV6
VDC5
CC
CC3
LM
ALJB
CCL
CC2
TO
VB2/2C
REG4
REG3
MDV5
MDV4
TOD
TOC
KS2
MDV3
CCB
MRU4
MRU3
MDV2
LM2
TOB
BUN6/7
VDC4
VIP
TOA
AGO
AG
REG2
ALF
PM3
GLO10
AGI
MRU2
PM5A/B
PM
AG7
PERD
GLOll
CAT
AG6
AG5B
AG5A
PERC
PM13
MRU
AG4
R2
PERB
Rl
CAT2
GL14A/B
PM10
R0M11
PM8
MBB
PM2
PERA
X X X X
SAL
ROM1
PER4
ROMS?
ROM8
PMI
MBA
ROM7
ROM6
ROM5
ROM4C
ROM4B
ROM3
PER5
Lagomorpha
X X X X
This content downloaded from 163.118.172.206 on Sun, 26 Jun 2016 14:22:20 UTC
All use subject to http://about.jstor.org/terms
376
TABLE 2. The sequence of Late Miocene localities of the Teruel-Alfambra region: macromammals. cf refers to the species part of the name
only (e.g., Hipparion cf. primigenium)
9 10 10 10 10 11 11 11 11 12 12 12 12 12 12 12 12 12 13 13 13 13 13
I Jl J2 J3 J3 K K K K L L L L L L L L L Ml Ml M2 M2 M2
local Zone
U %
I S
218I8~
tf> 1 U JS -g ?5 .I
S 11 I | i ?
Carn?vora
Canidae
Ursidae
C<j?/'j cipio
Indarctos atticus
Mustelidae
Agriotherium roblesi
Simocyon primigenias
x x
x x
Plesiogulo sp.
Plesiogulo monspessulanus
Mustela sp.
Baranogale adroveri
Martes cf. paleosinensis
x cf
Martes basilii
Sivaonyx lluecai
XXX
Ptioviverrops guerini
Ictitheriinae indet.
XXX
Mustelidae indet.
Hyaenidae
cf cf
Felidae
Thalassictis hipparionum
Thalassictis sp.
Lycyaena chaeretis
Percrocuta gigantea
Adcrocuta eximia
Metailurus major
Metailurus parvulus
cf
Paramachairodus orientalis
Machairodus sp.
Amphimachairodus giganteus
Felidae indet._
XXX
Tragoportax sp.
Boselaphini indet. 1
Boselaphini indet. 2
Palaeoryx pallasi
Protoryx carolinae
Oazella deperdita
Aragoral mudejar
Bovidae indet.
x x
cf x
Turiacemas concudensis
X X
X X
Pliocervus turolensis
_Cervidae indet._
Perissodactyla Equidae
Hipparion
Hipparion
Hipparion
Hipparion
primigenium
concudense
gromovae
periqfricanum
x cf cf
x cf
XXX
Hipparion sp.
Rhinocerotidae Alicornops alf?mbrense
Aceratherium incisivum
Lartetotherium schleiermacheri
Rhinocerotidae indet._
This content downloaded from 163.118.172.206 on Sun, 26 Jun 2016 14:22:20 UTC
All use subject to http://about.jstor.org/terms
x cf
X X X X
X X
311
EL BUNKER
?*>BUN4-5 (M1)
?%?BUN <M1)
LA GLORIA
FIGURE 7. Correlation of key sections to the Cande and Kent (1995) Geomagnetic Polarity Time Scale (Krijgsman, 1996; Krijgsman et al.,
1996; Garces et al., 1999). See text for ages of local zone boundaries. Ages of MN reference localities: MN9 = Can Llobateres: 9.7 Ma (Garc?s
et al., 1996; Agust? et al., 1997); MN10 = Mas?a del Barbo B: 9.3 Ma (Garc?s et al., 1999; this paper); MN11 = Crevillente 2: age is tentatively
set at 8.1 Ma, and corresponds to the midpoint of zone K, which contains faunas that correlate to this reference locality; MN12 = Los Mansuetos:
interpolated age of 6.9 Ma (van Dam, 1997; this paper); MN13 = El Arquillo 1: 6.2 Ma, corresponds to midpoint of zone M2 (this paper). Ages
of Serravallian-Tortonian boundary after Berggren et al. (1996), Tortonian-Messinian boundary after Krijgsman et al. (1994), and Messinian
Zanclean boundary after Lourens et al. (1996).
sample in the middle part of the La Gloria section, which was The relative thicknesses of the zones, and hence the sedimen
previously assumed to be reversed. This re-interpretation does
tation rates, are not constant from section to section and they
do not show a linear relation with relative durations in the cor
not have consequences for the ages of fossil localities. Detailed
information on the estimation of the ages of individual localities responding part of the GPTS (Fig. 7).
thermore, we assume the long reversed interval identified in the able for magnetostratigraphy because they are associated with
top of the La Gloria and the lower part of the El Bunker parallel carbonate lithofacies (e.g., results for Mas?a de la Roma show
sections to be C4r. This chron contains zone-K faunas, implying
low paleomagnetic intensities in a large part of the section,
a minimum age for zone-J faunas of ?8.7 Ma. The interpreta
Krijgsman, 1996). Figure 4 shows that in this area zones Jl, J2
tion of the intermediate polarity patterns (which would corre and J3 are all approximately equally thick. We use the assump
tion of constant sedimentation rates in this area as an additional
spond to the intermediate chron C4A) is not straightforward.
This content downloaded from 163.118.172.206 on Sun, 26 Jun 2016 14:22:20 UTC
All use subject to http://about.jstor.org/terms
estimation not too close or older than the next older zone
boundary (K-L, 7.5 Ma) is the correlation to C3An.ln. A min
imum age of 6.7 Ma for the L-Ml boundary is consistent with
The I?Jl boundary (=base of the Progonomys hispanicus than 6.3 Ma. It is set at 6.3 Ma, because of the slightly younger
zone), is situated between ROM3 and ROM4B in the Mas?a de estimate of 6.1 Ma for the M2-M3 boundary.
la Roma area and is estimated at 9.6 Ma (Fig. 5). The J1-J2
boundary is positioned between MBA and MBB in the Mas?a
FAUNAL CORRELATIONS ACROSS EUROPE
del Barbo-Masada Ruea sections (Fig. 3) and is estimated at
9.3 Ma (Garc?s et al., 1999). The J2-J3 boundary is estimated
Magnetostratigraphic control for zones L and M is poor. Al lian chronology, and discuss both philosophical and technical
aspects of the two commonly used systems: the MN "zone"
TABLE 3. Faunal resemblance between four more or less contemporaneous mammals faunas (?8.5-7.5 Ma) from Spain, Germany and Greece.
The faunas contain both micro- and macromammals. Crevillente 2 is MN11 reference locality. MN11 correlations for Puente Minero and Dorn
D?rkheim according to de Bruijn et al. (1992) and MN11/12 assignment for Pikermi/Chomateri according to Bernor et al. (1996). sp, species;
gen, genus; R, faunal resemblance index of Simpson (1960); N, number of taxa in fauna; C, number of faunas that two faunas have in common.
The smallest N is used as the denominator in the calculation of R.
Puente Minero
28
30, Nee
Nm
Puente Minero
(Teruel basin, Spain; MN11)
Crevillente 2
Nm 29, N?,
Cgf
Crevillente 2
(Alicante, Spain; MN11)
Csp
14
48
Dorn-D?rkheim
(Rhine basin, Germany; MN11)
Csp
RSP
20
RSP
Pikermi/Chomateri
(Rafina basin, Greece; MN11/12)
Rm
20
R,n
K?
64, N?
57
14
50
28
Dorn-D?rkheim
Nm 65, Ngen = 63
Cgen - 6
Rgen = 21
25
Csp
c
1
24
28
Pikermi/Chomateri
cet
21
29
14
C
^sp
Rm
13
This content downloaded from 163.118.172.206 on Sun, 26 Jun 2016 14:22:20 UTC
All use subject to http://about.jstor.org/terms
h-1 00
05 ?1
"IS
en
" S ^ or
la 3^
11 I
is?
^ ^ K
11 lia
IB
Q
<
5 s
*>
S
^? p
I mil
? ii ? ? s ?1
II ? ?
O tu a.
o?
Sa ?
o"
II
U?CJft,
II
a,
o>?
ft
kj
kj
?l
H oo
? s .5 S
Il ?
il
PQ ^
PQ .
il
22
'S S
si
Il .^ ?o ?
?o
(N
-S?
& ^ si
*?I
Il ?
Il ?
Il K5 ^?00
?
Il
O?,
(N
fi
Cm
J3 ? c
c?
o ^
"3 c
(N
00
O ^ Il
?> CO "
"
'S ^
r-H Cd .
u X? O
*t3 f_h ^h
^ Oh
00 <? Il
?^ Il
<$ -o K
Z d) <
-CL
.
a
00
S g II
e5>^
2 '^ s
~T?2
o? ? ?
5 S?
:?uJ
5 o
?2 s
O . ^D ,2 ^ *-H
sS
es
? s*?
S? a, O S ft,
<D C^ ^
This content downloaded from 163.118.172.206 on Sun, 26 Jun 2016 14:22:20 UTC
All use subject to http://about.jstor.org/terms
379
.1
co ,?j
SQ S&J
Sk ^3
"t*O
s8SB
cd cd
te
son (1960)
?^?
R = (C/min(A0) * 100,
with: R, faunal resemblance (the symbol is ours), C, number of
taxa that two faunas have in common, and min(A0, number of
taxa in the smaller fauna.
R varies between 0 and 100. The denominator, min(A0, is the
number of taxa in the smallest of the two faunas; use of the the
total number of taxa in both faunas as the denominator instead
of the minimum number of taxa will result in bias if the num
Ph o
? Il
?j cd
cd t?
Xi t?
both the species and genus levels (cf. designations have been
.a 5
03 -Cl
5-?
f ?s
p2?
o
?=
? ?0-?
25S
?.i *-i
ti
f ^ ti
*> -S
5
B ??
SP
? i
Oh
00
!? ? *>
O
K
O
Jk Q
cd ^
rj <
cd ^
? o
p VO
*2
?3 y
(D
cd fi
S 73
<D
cd
73 cd
.1
21
S M
O
D
? ^
s? 2
cd
o a s
s ^s
II*
Oft;
o
O
nas.
fi
O
?2
si
IS
?.I
>> cd
N <*
00
? ?3
fi
(U O
w
<? .Si cd
del Barbo (MBB) from the Teruel basin (Table 1), the combined
faunas of Trinxera Nort Autopista and Trinxera Sud Autopista
2 (we will refer to these two stratigraphically equivalent levels
? O fi
> D .S
Bat?
,-H ^ 0 -~
? Mn fi
g
?-1
<J->
ft cd cd
Oh
>
On On'
> ?
(N
ON
Ph
>
t?.HTl
^ C ?5
cd
fi
Ph S
Cu
fi
CO
Ses
pq g
o ?
HUH
<?
'1
, o
(N
U
(N
fi
^ O
?&
?t? +H ^t w W
B PQ
X) (D
O 73
? S
<D cl
II
H <
p< fi o
oQ?
q_i cd cd
D 4) 1)
Cd O? t/3
the species level is 19, i.e., only 19% of the number of species
present in the smallest fauna is present in the larger fauna to
which it is compared. Three comparisons between different ar
This content downloaded from 163.118.172.206 on Sun, 26 Jun 2016 14:22:20 UTC
All use subject to http://about.jstor.org/terms
381
TABLE 6. Age ranges for the entries of various Progonomys species into Spanish and Greek basins. Rodent assemblages with less than 100
specimens are not considered as evidence for species absence.
entry
references
Calatayud-Daroca/Teruel
basins (NE Central
Spain)
magneto stratigraphy :
Progonomys hispanicus
Progonomys woelferi
(1997)
gueras 2C (10.0-9.6
4B (9.6 Ma)
(1996)
Progonomys cathalai
Spain)
Greece)
(1996)
(9.6 Ma)
absent
absent
absent
Ma)
(9.2 Ma)
of the three pairwise comparisons there is only one species over large distances, "ecological resistance" to dispersal is to
shared (Rsp = 1/6 * 100 = 17). Trinxera Autopista (NE Spain) be expected as a result of interspecific competition. Very rapid
perior, resemble each other reasonably well (Rsp and Rgen are 43
and 57, respectively), but the younger assemblages have not ble 6). In the broader context of the Old World, the term "Pro
more than one and two species in common, respectively, in gonomys event" only makes sense at a very coarse level of
inland areas may have had a different fauna (e.g., the Vall?s
cies is involved, genus "events" (e.g., Progonomys, Hipparion pudia 9 is followed by a 23% presence in Torremormoj?n 1
event) will always take more time than species events. For in
(from Alvarez Sierra, 1983). Such a two-step expansion is not
stance speciation from one congeneric species to another has to recorded in the Vall?s-Pened?s basin, where P. hispanicus nev
be assumed to occur within the time of the event. The repro
er reaches high percentages (e.g., 10% in Trinxera Nort Auto
ductive isolation associated with allopatric speciation typically
pista, Agust?, 1990). Obviously, isochrony of the first occur
This content downloaded from 163.118.172.206 on Sun, 26 Jun 2016 14:22:20 UTC
All use subject to http://about.jstor.org/terms
chronous at 9.6 Ma in the Vall?s-Pened?s basin in Spain and related to an MN unit. This means that it has to be decided
on Crete.
which reference fauna is most similar to the new fauna. In some
In conclusion, the dispersal of each of the three Progonomys
therefore not be used for MN allocations. The MN scale is an European Mammal-based Stages/Ages: Are They Still
Necessary?
expressions such as "late MN9," "early MN10" are therefore nologic units independently from the marine-based Global
not correct. Last but not least, so-called MN "boundaries" (as Chronological Scale, because there were no reliable correlations
This content downloaded from 163.118.172.206 on Sun, 26 Jun 2016 14:22:20 UTC
All use subject to http://about.jstor.org/terms
edition (Salvador, 1994), which explicitly required the defini Mieres, Andr? Nieuwenhuizen, Fred Rogl, Ed van Slobbe, Ka
tion of boundaries by Boundary Stratotypes. A Vallesian-Tur
rel Steensma, Agnes and Paul Tijman op Smeijers, Marleen
olian Boundary Stratotype could in principle be defined in one Vriends, Maaike Dekkers, Caspar Geraedts, Kees Hordijk, Wil
lem Renema, Lena Sel?nne, Wilma Wessels, and the late Con
of the Teruel sections. It could for instance correspond to the
local J-K boundary, which is characterized by the simultaneous
stantin Theocharopoulos. We thank the facilities found in the
entry of the rodent taxa Parapodemus and Eozapus (8.7 Ma). Museo Paleontol?gico de la Universidad de Zaragoza, the Mu
As a chronostratigraphical unit, the Vallesian-Turolian bound seo Nacional de Ciencias Naturales (CSIC) in Madrid. We
ary would then, at least in theory, exist world wide. However, thank also Pascual Ca?ada and Joaqu?n Mel?ndez for their help
biostratigraphically, this boundary has a very local significance.
storing materials at the Gabinete Geol?gico de la Diputaci?n
For example, in the Rhone Basin, SE France (the closest area Provincial de Teruel, and the owners of the lands where fossil
with a well-documented contemporaneous record across a po localities are excavated. We are grateful to Gijs van Dam, Hans
tential Vallesian-Turolian boundary), the patterns of co-occur
Brinkerink, Juliette Richter, Willem Renema, Joost Smets, and
rence of species are quite different (Mein, 1999): Parapodemus
Frank Geerts for their help in sorting the residues for small
(not present before 8.7 Ma in Teruel, i.e., this would be "Tur
mammals. Hans de Bruijn, Albert van der Meulen, Johan Meu
olian") is consistently associated with Progonomys (not present lenkamp and two anonymous reviewers are thanked for their
after 8.7 Ma in Teruel, i.e., "Vallesian"). The different species
critical reading of the manuscript. We are indebted to Jaap Lu
compositions in the two areas can easily be explained by dif teijn, Wil den Hartog, Paul van Oudenallen, Izaak Santoe, Jaco
ferent environmental conditions, as indicated by the presence
van Bergenhenegouwen and Fred Trappenburg for their help in
of many forest-dwelling rodents in the Rhone basin, and ab preparing the figures and to Marnella van der Toi for her help
sence of those forms in Spain. The relevant question in this in preparing the manuscript.
case is not whether the French faunas have to be called Valle
Jan van Dam and Wout Krijgsman were supported by the
sian or Turolian, but to which stratigraphie positions in the Earth and Life Sciences Foundation (ALW) and Geosciences
Spanish sections these faunas correlate best.
Foundation (GOA) with financial aid from the Netherlands Or
The pertinent question is therefore whether or not European ganization for Scientific Research (NWO). Jan van Dam and
Neogene continental Stages are useful. Knowledge has in Miguel Garc?s thank the Faculty of Earth Sciences of Utrecht
creased enormously since Thaler (1966) proposed his "Echelle University for financial support. Field campaigns were partly
des zones de Mammif?res du Tertiaire d'Europe" in an effort carried out within the framework of the projects "Paleoecologia
to free mammal paleontologists from marine stratigraphic/chro
nological terminology. In terms of faunal correlation across Eu
rope, however, these continental units (Stages/Ages) have not
ACKNOWLEDGEMENTS
ruel 67:7-21.
We dedicate this paper to the late Remmert Daams who has -, and P. Mein. 1996. Nuevo Ruscinomys (Rodentia, Mammalia)
en el Mioceno Superior de la region de Teruel (Espa?a). Estudios
contributed enormously to the development of mammal pale
geol?gicos 52:261-365.
ontology in Spain, and who helped us a lot, both in and outside
-, -, and E. Moissenet. 1993. Roedores de la transici?n
the field. We are very grateful to Albert, Joke and Michiel van
Mio-Plioceno
de la regi?n de Teruel. Paleontolog?a i Evoluci? 26
der Meulen, Pablo Pelaez Campomanes, Marian Alvarez Sierra,
27:47-84.
and Martine Bestebreurtje for their structural help with the col
paigns. Many people joined the campaigns for shorter and lon
ger time and digged, carried, and washed sediments from the
This content downloaded from 163.118.172.206 on Sun, 26 Jun 2016 14:22:20 UTC
All use subject to http://about.jstor.org/terms
?rica 22:239-264.
gaceta 3:61-63.
schappen, B 68:121-126.
Daams, R., L. Alcal?, M. Alvarez Sierra, B. Azanza, J. A. van Dam,
243-258.
Volume 54.
This content downloaded from 163.118.172.206 on Sun, 26 Jun 2016 14:22:20 UTC
All use subject to http://about.jstor.org/terms
Zarza. 1999. Late Miocene alluvial sediments from the Teruel area:
magnetostratigraphy, magnetic susceptibility, and facies organisa
665-684.
pen, B 79:256-270.
-, F. J. Hilgen, C. G. Langereis, and W. J. Zachariasse. 1994. The
- 1985. A review of fossil and recent Desmaninae (Talpidae, In
age of the Tortonian/Messinian boundary. Earth and Planetary Sci
3:1-343.
superior del sector central del la Cuenca del Duero. Studia Geo
l?gica Salmanticensia 22:191-212.
Lourens, L. J., F. J. Hilgen, W. J. Zachariasse, A. A. M. van Hoof, A.
Antonarakou, and C. Vergnaud-Grazzini. 1996. Evaluation of the
Pliocene to early Pleistocene astronomical time scale. Paleocean
ography 11:391-413.
53:321-334.
Geol?gicos 17:209-305.
367-387.
-, E. Moissenet, and R. Adrover, 1990. Biostratigraphie du N?o Weerd, A. van de. 1976. Rodent faunas of the Mio-Pliocene continental
g?ne sup?rieur de Teruel. Paleontolog?a i Evoluci? 23:121-139.
sediments of the Teruel-Alfambra region, Spain. Utrecht Micropa
Moissenet, E. 1983. Aspectos de la neotect?nica en la Fosa de Teruel;
leontological Bulletin, Special Publication 2:1-217.
pp. 427-446 in Geolog?a de Espa?a, Libro Jubilar J. M. R?os, Vol. Woodburne, M. O. 1996. Precision and resolution in mammalian chron
2. Instituto Geol?gico y Minero de Espa?a, Madrid.
This content downloaded from 163.118.172.206 on Sun, 26 Jun 2016 14:22:20 UTC
All use subject to http://about.jstor.org/terms