The MNSystem and Continental Stage-Age Concepts Discussed PDF

The Upper Miocene Mammal Record from the Teruel-Alfambra Region (Spain).
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
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Journal of Vertebrate Paleontology 21(2):367-385, June 2001

? 2001 by the Society of Vertebrate Paleontology
THE UPPER MIOCENE MAMMAL RECORD FROM THE TERUEL-ALFAMBRA REGION

(SPAIN). THE MN SYSTEM AND CONTINENTAL STAGE/AGE CONCEPTS DISCUSSED
J. A. VAN DAM1, L. ALCAL?2, A. ALONSO ZARZA3, J. P. CALVO3, M. GARC?S4, and W. KRIJGSMAN5
faculty of Earth Sciences, Utrecht University, P.O. 80021, 3508 TA, Utrecht, the Netherlands; E-mail: jdam@geo.uu.nl;
2National Museum of Natural History, Gutierrez Abascal 2, 28006 Madrid, Spain;
3Department of Petrology and Geochemistry, Faculty of Geology, Complutense University, Ciudad Universitaria,
28040 Madrid, Spain;
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
During the nineties pal?ontologie research took various new

directions. The number of micromammal localities was consid
The Teruel-Alfambra region in NE Spain (Figs. 1 and 2) is

erably increased by renewed intensive sampling, and fossilif
a classical area for mammal paleontology. Documentation of
erous sections were sampled paleomagnetically (Krijgsman et
fossil bones dates back to the first part of the eighteenth cen
al., 1996; Opdyke et al., 1997; Garces et al., 1999). The "high
tury, when Feijoo (1736) mentioned the occurrence of "varios
huesos de el cuerpo humano, y otras, que representan huesos
resolution" magnetobiostratigraphic approach (sensu Lindsay,
see Woodburne, 1996) not only improved the chronology of
de bestias" near the village of Concud (see Alcal?, 1994, 1997,

mammal localities and zones, but also yielded the temporal
for a historical review). During the nineteenth and first part of
framework for paleoecologic, paleoclimatologic, and faunal dy
the twentieth century an increasing amount of (obviously non
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
font, Truyols and Villalta started to explore the region more

systematically, and discovered various other important macro
mammal sites. One of the results of these field campaigns was
the introduction of the term "Turolian" to indicate "les niveaux
cal? (1994) combined detailed taphonomic analyses with paleo

ecological studies of body-size distribution and ecological di
versity spectra.
de Teruel" (Crusafont, 1965). A formal definition of the Tur
GEOLOGICAL SETTING AND LITHOLOGY
olian as a continental Stage was given somewhat later (Marks,
1971a; Aguirre et al., 1975).

Many new, particularly micromammal localities became
The Teruel-Alfambra region comprises half of the northern
part of the Teruel Basin, a NNE-SSW oriented basin in the

northeastern part of the Iberian Peninsula (Fig. 1). The basin
known from the sixties onwards, when Adrover started his ex

tensive prospecting of the area around Teruel. His contribution covers an area approximately 100 km in length and 15 km in
to the mammal paleontology of the area is enormous and is con width. It is filled with a fairly complete middle to late Neogene
tained in many publications (see bibliography in Adrover, 1986). succession exceeding 500 m in thickness (Moissenet, 1983,
Some other important studies resulted from Dutch-Spanish co
1989). According to Anad?n and Moissenet (1996) the basin
operation. Among these are the monograph on Hipparion by should be regarded as a half-graben, with the NNE-SSW ori
Sondaar (1961) and the work of van de Weerd (1976) on rodents.

The latter study was the first palaeontologic study of the area
based on modern bio- and chronostratigraphic concepts, coupled
with sampling of sections with series of superposed micromam
mal localities. Van de Weerd's study resulted in a firmly-estab
lished biozonation based on Murinae. Latest Miocene to Pliocene
ented master fault running along the eastern border of the basin
(Fig. 1).
The Neogene succession of the Teruel-Alfambra region (Fig.

2) is chiefly bounded by Triassic clastic and evaporite forma
tions and Jurassic carbonates. The oldest Neogene deposits are
situated along the eastern margin of the basin and are of early
gaps in the zonation were bridged by the Spanish-French team Miocene age (Fig. 2: area around mammal locality Montai vos
of Adrover, Mein and Moissenet (Mein et al., 1990).
(MNT), which is currently under study). Most of the basin fill
367
368 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 21, NO. 2, 2001

Roma, Mas?a de la Roma and Peralejos (Fig. 4). Carbonate
channel fills are predominant in the lower part of this facies
and palustrine carbonates and paleosols in the upper part. The
carbonate beds alternate with organic-rich marls which contain
mammal and abundant gastropod remains. A maximum thick

ness of almost 50 m is observed in the Mas?a de la Roma
section.
Facies unit M4m is well exposed in the area close to Teruel,
where it typically forms the top of escarpments (e.g., Fig. 6) and
cuestas. The characteristic thick tabular and indurated capping
limestone beds (type 6 above) have been used for correlation
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.
The (reddish) claystone-dominated intervals in the studied
FIGURE 1. Schematic map of the Teruel basin and surrounding areas
(NE Spain).
sections belong to facies unit M2-P12 of Hern?ndez et al. (1983).

This unit contains the major part of the Miocene and Pliocene
clastic sediments (claystones, sandstones, conglomerates) of the
basin and is mainly developed along the basin margin. It is
more or less equivalent to the Peral Formation of van de Weerd

(1976) and may locally reach thicknesses of 200 m.
Both the carbonate and clay stone facies may laterally grade
into evaporites, e.g., in Los Mansuetos (LM) (Fig. 3) and at

sequence that can be observed in the central part (the river Los Aljezares (ALJ, see Fig. 2; Calvo et al., 1999). More to
Alfambra valley), however, consists of sediments of Late Mio the north and northwest, this evaporitic facies reaches thick
cene and Early Pliocene age. Overall, these sediments were nesses of 60 m (Tortajada and Concud sections, TO and CC,
deposited in distal alluvial fan and shallow lake environments.
Fig. 4). A maximum thickness of about 150 m is estimated
The main mammal-containing sections are schematically rep
resented in Figures 3 and 4. Figure 3 shows eleven sections in
the southern part of the area, close to the town of Teruel. Figure
4 shows three sections in the northern part, which will be re
ferred to as the Peralejos area (Fig. 2). Lithostratigraphic cor
relations between sections are based either on the positions of
individual marker beds, or on the positions of (boundaries be
tween) lithostratigraphic units described in the literature (van
de Weerd, 1976; Hernandez et al., 1983).
Most fossil mammal remains are found in organic-rich marl

stones and clay stones, that are associated with a carbonate lith
close to the village of Tortajada (Hernandez et al., 1983). These
latter, more northern gypsum exposures constitute van de

Weerd's (1976) Tortajada Formation. This formation contains
massive, meter-thick gypsum beds with a fabric consisting of
strongly bioturbated (by chironomids) lenticular gypsum, which
accumulated extensively in moderately saline lake waters (Rod
riguez-Aranda and Calvo, 1998).
THE SEQUENCE OF MAMMAL LOCALITIES
The composite upper Miocene mammal succession of the

ofacies, although some localities are associated with a distal area (Tables 1-2) consists of 99 assemblages. All except one
contain micromammals and 23 contain both micro- and macro
alluvial facies (e.g., MBA) or gypsum facies (e.g., LM and
LM2). The relation between lithofacies and the accumulation mammals. The stratigraphie order of the localities is based on
of mammal remains has been tentatively studied by Albesa et
al. (1997) but a comprehensive approach to the factors con
trolling the formation of the mammal sites is still in progress.
the interpretation of litho-, bio- and magnetostratigraphic data.
The positions of isolated, poorly documented assemblages in

the sequence of localities are only approximate, implying that
The studied fossiliferous organic-rich marls belong to the the degree of resolution for those assemblages is not better than
carbonate facies units M3 and M4m as described by Hernandez

et al. (1983), and to the Alf ambra Formation of van de Weerd
that of the biozone.
The small mammal record, which almost exclusively consists
(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
and R2, as well as the micromammal data from SAL, REG2

5, AG, BUN, VDC6, GL05-6, and AQ1 and 4 are from Ad
common. This suggests that the shallow lakes underwent epi rover (1986), Adrover and Mein (1996), Adrover et al. (1982a,
sodic lake level oscillations (Alonso-Zarza and Calvo, 2000).
b, 1984, 1993) and Mein et al. (1990). The data for PM are
Facies M3 is exposed in the Peralejos area, where it forms from Alcal? et al. (1991). All other micromammal data are new.
the lowest part of the hills bordering the river Alfambra. Its top The new localities are: PER5; ROM1, 3, 4B, 4C, 5, 6, 7, 8, 9,
11; PM1, 2, 3, 5A, 5B, 8, 10, 13; GLO10, 11, 14A, 14B; CAT2;
can easily be recognized in the field (half-way the hill slope in
Fig. 5) and is used for correlation among the sections of La Rl; MRU, MRU2, 2A, 3, 4; AGI, 3, 4, 5A, 5B, 6, 7; BUN4,
VAN DAM ET AL.?UPPER MIOCENE MAMMALS FROM SPAIN
369
LEGEND
Quaternary,
including Villanyian
* fossil locality/
localities
~vl Neogene
1 i ' i ' I Neogene
limestones and marls
Neogene
red/orange conglomerates,
sandstones and clays
|^\^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.
5, 6, 7; KS, KS2, KSS; CAP2, and MOD. The samples from
LOCAL BIOZONATION
R2, ALF, VIP, and VB2/2C are extended. For a review of older
The biozonation is a modification of the Miocene part of the

(9.6-8.0 Ma) Murinae see van Dam (1997). Up to now, most zonation of van de Weerd (1976), who defined range zones for
rodents have been identified at the species level, most insecti
the area on the basis of Murinae (Rodentia). A first refinement
vores and lagomorphs at the genus level, and part of the bats
of van de Weerd's scheme was published by Mein et al. (1990),
at the family level.
who proposed a three-partition of the upper Turolian Stephan
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
mammal associations (ROM1, MBB, R2, VIP, MDV2, LP,
neighbouring Calatayud-Daroca Basin. We use this system next
(1994). Besides new data, this work includes the updating and
completion of faunal lists from Alcal? et al. (1994), Alcal? and
introduce the local zones J to M, with J (Progonomys hispan
CCL, LM, CC, KS, ML and ARQ1) were described by Alcal?
Morales (1997), Azanza et al. (1997), Fraille et al. (1997) and

van der (Made 1997).
to the zone names of van de Weerd, which are retained. We
icus Zone) and M (Stephanomys ramblensis Zone) subdivided

into subzones Jl-3 and Ml?3, respectively (see also Daams et
al., 1998). The local zones are described below.
JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 21, NO. 2, 2001
T~2
?==3 CC(MN12-L)
?==3 CCB(MN12-L)
LAS CASIONES
SECTION
sandstones and conglomerates
claystones
palustrine carbonates
(including palustrine paleosols
and tufa)
carbonate channel fills

marlstones (generally organic-rich)
paleosols developed on alluvial fades

gypsum (massive and bioturbated)
lithostratigraphic correlations:
- ascertained
-probable
- -?- - possible
carbonates (tabular and indurated) :
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
mals have not been found in the Teruel-Alfambra region.
dunensis interval zone, from the first regular occurrence of P.

hispanicus to the entry of Parapodemus lugdunensis.
common. The carnivore record is poor.
Subzone Jl?Diagnosis: P. hispanicus-P. cathalai interval

Additional micromammal data: The murid Progonomys his zone, from the entry of P. hispanicus to the entry of P. cathalai.
panicus is the dominant rodent; the range of this species, how Additional micromammal data: Progonomys hispanicus is the
ever, is not confined exactly to the nominate zone. The cricetid dominant rodent and the only murid present. The cricetid Me
371
?-l-J f? ?^r3 BUN4-5 (MN13-M1)
a
?^3 BUN(MN13-I\
A?A
?A?
A?A
?A?
A?A
?~^3 BUN6-7 = VDC4

(MN11-K)
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.
Subzone J2?Diagnosis: Progonomys cathalai range zone,
characterized by the total range zone of Progonomys cathalai.
Subzone J3?Diagnosis: Huerzelerimys minor?Parapode
mus lugdunensis interval zone, from the entry of Huerzelerimys

minor to the entry of Parapodemus lugdunensis. Additional mi

erium and Micromeryx have their last occurrence. Typical for the
zone is the cervid Lucentia. Among the rest of the artiodactyls,
Micro stonyx and Tragoportax continue their presence. In PM,
Hipparion primigenium is associated with H. gromovae, a small
species otherwise known from zone M. The diversification of
Hipparion contrasts with a decline in rhinocerotid diversity.
^^ &
Zone L?Parapodemus barbarae Zone of van de Weerd
(1976). Diagnosis: Parapodemus barbarae-Stephanomys ram
blensis interval zone, from the entry of Parapodemus barbarae

to the entry of Stephanomys ramblensis.
Additional micromammal data: The range of Parapodemus

barbarae is not confined exactly to the nominate zone. The
zone comprises the total ranges of Occitanomys adroveri and
Huerzelerimys turoliensis. It includes the last occurrence of P.

gaudryi. Ruscinomys starts its range in this zone. Eliomys truci
and the sciurids Spermophilinus turoliensis and Atlantoxerus
adroveri are common. The latter is entirely restricted to this

zone. Insectivore diversity is much higher than in zone K. Ar
chaeodesmana turolense is common.
Additional macromammal data: The first known occurrence
of canines (Canis cipio) is characteristic. Carnivora, especially

hyaenids and felid, are diverse. Zone L is the last zone with
Microstonyx. The giraffid Birgerbohlinia continues to be pre
sent. Zone L is characterized by a marked proliferation of bo
vids, which are, apart from Boselaphini, represented by hip

potragines (Palaeoryx pallasi and Protoryx carolinae) and an
tilopines (Gazella deperdita and Hispanodorcas torrubiae).
Cervids start their range with Turiacemas concudensis. Hip
parion concudense is the common equid in this zone. Zone L
is the only zone with more than one proboscidean genus.
Zone M?Stephanomys ramblensis Zone of van de Weerd

(1976). Diagnosis (based on Mein et al., 1990): Stephanomys
ramblensis-Celadensia interval zone, from the entry of Ste
phanomys ramblensis to the entry of the cricetid Celadensia.
Additional micromammal data: The zone roughly corre

sponds to the range zone of Apodemus gudrunae and the cri
cetids Blancomys sanzi and Apocricetus alberti. Occitanomys
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).
that of zone L, except that Crusafontina is now replaced by

Amblycoptus, and Archaeodesmana turolense and A. adroveri
by A. luteyni and A. major. Among the lagomorphs, Prolagus
crusafonti is replaced by P. michauxi.
cromammal data: Huerzelerimys minor and Progonomys his

Additional macromammal data: The medium-sized hyaenid
panicus are the only murids present. Myomimus sp. A is re Thalassictis hipparionum and large-sized sabertooth tiger Am
stricted to this subzone. Myomimus dehmi, Tempestia and Mus
cardinus are absent. Among the insectivores, Desmanella is

common.
Zone K?Parapodemus lugdunensis Zone of van de Weerd
phimachairodus giganteus are abundant. Among the mustelids
a large-sized form (Plesiogulo monspessulanus) is present.

Within the group of ursids, Indarctos is replaced by Agrioth
(1976). Diagnosis: Range zone, characterized by the total range
erium. The most characteristic artiodactyls are Hexaprotodon

crusafonti (the oldest hippopotamid known from Eurasia) and
Additional micromammal data: This range corresponds more

or less to the ranges of the murids Occitanomys sondaari and
only. Tragoportax and antilopines continue to be present, and

the medium-sized suid Propotamochoerus appears for the first
of Parapodemus lugdunensis.
Huerzelerimys vireti. Parapodemus gaudryi and the cricetids
Hispanomys freudenthali and Kowalskia occidentalis start their

ranges at about the base of the zone. The range of the zapodid
Eozapus intermedius is restricted to this zone. Sciurids (Heter
oxerus) are very rare. The castorid Dipoides, the soricid Blari
nella and the talpids Archaeodesmana and Talpa appear for the
first time with very low numbers. Insectivore diversity is low,
two species of the cervid Pliocervus, known from this zone
time. Various species of Hipparion co-occur, among which is
Hipparion periafricanum, the smallest Hipparion known. Rhin

ocerotids are reduced to one species, the large-sized Lartetoth
erium schleiermacheri.
Zone Ml?Diagnosis: Range zone, characterized by the total
range of Castromys inflatus.
Zone M2?Diagnosis: Castromys inflatus-Paraethomys mio

and only three species belonging to Galerix, Desmanella and caenicus interval zone, from the exit of Castromys inflatus to
Paenelimnoecus are common. The only Rhinolophidae (Chirop the entry of Paraethomys miocaenicus.
Additional micromammal data: this zone includes the first
tera) known from the studied interval thus far occur in this zone.
Additional macromammal data: Plioviverrops guerini occurs
for the first time, and it coexists with Indarctos atticus, Adcro
occurrence of Hystrix.
Zone M3?Diagnosis: Paraethomys miocaenicus-Celaden
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.
VAN DAM ET AL.?UPPER MIOCENE MAMMALS FROM SPAIN 373
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
Roma sections are shown in Figure 7. Ages for most of the

localities (Table 1) were estimated by linear interpolation of
The most suitable sections have been subjected to magneto

sediment thicknesses within polarity zones and sections, litho
stratigraphic studies in order to provide numerical age con
straints for the fossil localities. Especially the reddish colored
stratigraphic correlations between sections, and linear interpo
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.
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
Masada del Valle 6_

Valdeecbro 5
GLOl
MDV6
VDC5
Milagros
Las Casioncs superior
Las Casioncs
Villalba Baja 1
Modorras
El Capon 3
El Capon 2
Regajo 5
Bunker de Valdeecbro 4/5
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
V?lalba Baja 2/2C
Regajo 4
Regajo 3
Masada del Valle 5

Masada del Valle 4
Tortajada D
Tortajada C
Las Casioncs 2
Masada del Valle 3
Concud B
Masada Ruca 4
Masada Ruca 3
Masada del Valle 2

Los Mansuetos 2
Tortajada B
Bunker de Valdeecbro 6/7
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
Puente Minero 5A/B

Puente Minero
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
I3*
375

TABLE 1. (Continued)
Chiroptera
Eomyidae Zapodidac Castoridac Hystricidae
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
Vespcttilionid. Rhinolophid. Ochotonidae Leporidae
X X X X
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
correlation to MN reference fauna
local Zone
U %
I S
218I8~
locality (* = MN reference fauna)
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.
cf. Sabadellictis sp.
Baranogale adroveri
Martes cf. paleosinensis
x cf
Martes basilii
Sivaonyx lluecai
XXX
Ptioviverrops guerini
Ictitheriinae indet.
XXX
Mustelidae indet.
Hyaenidae
cf cf
Ictitherium aff. pannonicum
Felidae
Thalassictis hipparionum
Thalassictis sp.
Lycyaena chaeretis
Percrocuta gigantea
Adcrocuta eximia
Felis IF. attica (a)
Metailurus major
Metailurus parvulus
cf
Paramachairodus orientalis
Machairodus sp.
Amphimachairodus giganteus
Felidae indet._
Suidae Microstonyx erymanthius

Microstonyx major
Suidae indet.
Hippopotarnidae Hexaprotodon crusafonti
Tragulidae Dorcatherium naui

Giraffidae Decennatherium pachecoi
Birgerbohlinia schaubi
Bovidae Tragoportax gaudryi
XXX
Tragoportax sp.
Boselaphini indet. 1
Boselaphini indet. 2
Palaeoryx pallasi
Protoryx carolinae
Oazella deperdita
Hispanodorcas / H. torrubiae (t)
Aragoral mudejar
Bovidae indet.
Moschidae Micromeryx /M. flourensianus (f)
x x
Cervidae Lucentia aff. pierensis
cf x
Turiacemas concudensis
Pliocervus aff. matheroni
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._
Hyracoidea Pliohyracidae Pliohyrax cf. graecus

Proboscidea Deinotheriidae Deinotherium giganteum
Gompotheriidae Tetralophodon longirostris
cf. Anancus arvernensis
Mastodon?dae Zygolophodon turicensis

Proboscidea indet.
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).
and zonal boundaries is given by van Dam (1997).

The Peralejos, La Roma and Mas?a de la Roma parallel sec
The identification of the long normal chron C5n in the La tions in the Peralejos area (Fig. 4) contain the four successive
Gloria and Masada Ruea sections implies a maximum age of biostratigraphic (sub)zone boundaries I-Jl, J1-J2, J2-J3, and
less than 10 Ma for the zone-J faunas in these sections. Fur J3-K. Unfortunately, the majority of the sediments are less suit
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.

zones we have to refer to magnetostratigraphic results from
other Spanish basins in order to calibrate our biorecord. For the
age of the K-L boundary we use 7.5 Ma, based on a magne

tostratigraphic reinterpretation by Krijgsman et al. (1996) of
Opdyke et al.'s (1990) results for the Cabriel Basin (eastern

Spain). The lower part of the Cabriel record contains a small
fauna (Balneario) with Parapodemus lugdunensis, which cor
relates to our zone K, and which is situated ?10 m below a
fauna with Parapodemus gaudryi and Occitanomys adroveri
(Fuente Podrida), which correlates to our zone L. This corre
lation implies that a gap, corresponding to a large part of chron
C4n, is present in the Masada del Valle and Alfambra parallel

sections (Fig. 7).
Another stratigraphie gap, which probably includes the entire
zone L, is present in the El Bunker section (Fig. 7), the top of
which contains Ml faunas. A minimum age of 6.7 Ma is as
sumed for the L-Ml boundary on the basis of an estimated age

FIGURE 8. Positions of the basins/areas and localities referred to in
of 6.1 Ma for the entry of our M3 marker (Paraethomys) in the
this study. (1) Calatayud-Daroca basin: Nombrevilla and Pedregueras
2C; Teruel basin: for localities see Fig. 2 and Tables 7 and 8; (2) Cabriel Fortuna basin (SE Spain; Garc?s et al., 1998). Assuming a con
basin: Balneario, Fuente Podrida, Venta del Moro; (3) Alicante: Crev stant sedimentation rate in the top part of the El Bunker section
(Fig. 7), the only magnetostratigraphic correlation for the up
illente 1-5; (4) Duero: Ampudia 9, Torremormoj?n 1, 1A and 3; (5)
Vall?s-Pened?s basin; (6) Languedoc Central: Montredon; (7) Rhone

basin; (8) Rhine basin: Dorn-D?rkheim; (9) Strimon basin: Lefkon,
Maramena; (10) Raima basin: Pikermi/Chomateri.
permost small normal interval at the top, resulting in an age
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
constraint to interpret the polarity patterns in the other sections

near Teruel. According to our preferred interpretation, each of
the Jl, J2 and J3 subzones has a duration of ?0.3 m.y.
an age of 6.8 Ma for the oldest locality in the Fortuna Basin

containing typical zone-M rodents (Garc?s et al., 1998). When
the uppermost normal interval in the El Bunker section corre
sponds to C3An.ln, the position of the Ml faunas BUN4 and
5 implies that the age of the M1-M2 boundary cannot be older
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
In this section, we will compare various faunas from the Te

at 9.0 Ma on the basis of linear interpolation of sediment thick
ruel basin with other, approximately time-equivalent faunas
nesses in the three parallel sections in the Peralejos area (Fig.
from elsewhere in Europe (Fig. 8). We will then focus on the
4) and magnetobiostratigraphic correlation to the La Gloria sec
Progonomys "event," which is one of the few events in the
tion (Fig. 7). The J3-K boundary is estimated at 8.7 Ma in the
La Gloria section (Krijgsman et al., 1996, and above). The age time interval studied where isochrony/diachrony can be tested
estimations for the zone-J localities (Table 1) are consistent at a relatively high level of resolution (a few hundred thousands
of years). Finally, we will put the faunal patterns observed in
with interpolations resulting from the quantification of evolu
the Teruel area in the wider perspective of European mamma
tionary stages in Progonomys hispanicus (van Dam, 1997).
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"
though the assumption of isochronicity of bioevents between

basins will not always be valid (see next section), for these two
system and the continental Stage/Age system.
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
Nsp = 65, Ngen = 63
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
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
S ? & <N .S?
? ii ? ? s ?1
II ? ?
O tu a.
kjTc3 ?? ?t? ? ? kj^ 03 ^ i ? ? o> ?
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^ ^
379

Low Faunal Resemblance
.1
co ,?j
Tables 3-5 show faunal resemblance indices for pairs of

more or less contemporaneous faunas across Europe. Resem
SQ S&J
Sk ^3
"t*O
s8SB
cd cd
te
blance is quantified by the following index proposed by Simp
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
bers of taxa are distinctly unequal or if one of the two faunas

is less complete than the other. Indices have been calculated for
?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
Four well-documented mammal faunas of similar age (?8.5

7.5 Ma) containing both micro- and macromammals are com
*> -S
5
B ??
SP
? i
Oh
00
treated as if they were definite). Tables 4 and 5 relate to rodents

only, because other groups in these localities were either absent,
rare or less well studied.
!? ? *>
pared in Table 3: Puente Minero (Teruel, this paper), Crevillente
O
K
O
Jk Q
2 (Alicante, SE Spain, Montoya, 1994), Dorn-D?rkheim (Rhine

basin, Germany, Franzen and Storch, 1999, Storch, pers.
comm., 1995), and Pikermi/Chomateri (Greece, faunal list
based on de Bruijn, 1976; R?mke, 1976; Bernor et al., 1996;
de Bruijn et al., 1999). Crevillente 2 is the MN11 reference
locality, and both Puente Minero and Dorn-D?rkheim have
been correlated to this unit (e.g., see de Bruijn et al., 1992).
cd ^
rj <
cd ^
? o
p VO
*2
Pikermi/Chomateri is correlative to the MN12 reference locality
Los Mansuetos according to de Bruijn et al. (1992), and to

MN11/12 according to Bernor et al. (1996).
?3 y
(D
Table 3 shows that faunal resemblances are rather low. Com
cd fi
S 73
parisons among Spain, Germany and Greece show R values

between 13 and 28 for species and 14 and 29 for genera. The
<D
two Spanish faunas have about half of their number of species
cd
73 cd
in common (Rsp = 48; Rgen = 50). In this case, species and

genus results are almost the same, which is due to the near
absence of congeneric species not only in but also across fau
.1
21
S M
O
D
? ^
s? 2
cd
o a s
s ^s
The faunal resemblance values in the terminal Miocene are

even lower. El Arquillo 1 (Teruel basin, this paper) with 31
II*
Oft;
o
O
nas.
species and Maramena (Strimon Basin, Greece, Schmidt-Kittler
et al., 1995) with 54 species share only one species: Gazella
fi
O
?2
si
IS
?.I
deperdita, i.e., Rsp = 3. (This number might be slightly under
>> cd
8) considered to be of more or less similar age (MN10). Three

Spanish faunas are included: the MN10 reference fauna Mas?a
N <*
00
estimated because the two Hipparion species from Maramena

could not be identified at the species level, but may be the
same). Generic resemblance is 27, i.e., as high as in Table 3.
Table 4 shows the R values for five rodent assemblages (Fig.
? ?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
as Trinxera Autopista, TAP) from the Vall?s-Pened?s basin

(Agusti, 1990; Aldana Carrasco, 1992; Agust? et al., 1997) and
73 ?d
fi ^-N ^ ^"N ^~s
Torremormoj?n 1 (TOI) from the Duero basin (Alvarez Sierra,
cd vo S 00 Q\
a On -*-? 00 Os
1983; L?pez Mart?nez et al., 1986). Included are furthermore
SOn u On On
the
French locality Montredon (MNR) from the central Lan
cd .
Q Id pi Id Id guedoc region (Aguilar, 1982) and the Greek locality Lefkon
O t? vu
<D cl
II
H <
p< fi o
oQ?
q_i cd cd
D 4) 1)
Cd O? t/3
(LEF) from the Strimon basin (de Bruijn, 1989).
Faunal resemblance at the species level is again very low

(numbers below the diagonal in Table 4). The average Rsp at
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
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
Vall?s-Pened?s basin (NE
fauna: van Dam (1997);
fauna and magneto stratigra fauna: Alvarez Sierra
Spain)
magneto stratigraphy :
Krijgsman et al. (1996),

Garc?s et al. (1999)
Progonomys hispanicus
Progonomys woelferi
phy: Garc?s et al.

(1996), Agust? et al.
(1997)
gueras 2C (10.0-9.6
4B (9.6 Ma)
between MBA (9.4 Ma)
and PER4 (9.3 Ma)

absent
Duero basin (NW Spain)
(1983), Garcia Moreno

(1987); magnetostratigra
phy: Krijgsman et al.
(1996)
very rare cf. P. h. in Pedre between Can Llobateres 1
Ma); first regular pres

ence P.h. between Mas?a
de la Roma 3 (9.7? Ma)
and Mas?a de la Roma
Progonomys cathalai
Spain)
(9.7 Ma) and Rubi Ter

rassa (9.7 Ma)
Kastellios Hill (Crete,
Greece)
fauna: de Bruijn and Za

chariasse (1979), van
Dam (1997); magneto

stratigraphy: Sen et al.
(1986), Woodburne et al.
(1996)
very rare P.h. in Ampudia 9between KAI (9.6 Ma) and

(?10.3 Ma); first regular KA2A (9.5 Ma)
presence between Torre
mormoj?n 3 (10.0 Ma)

and Torremormoj?n 1
(9.6 Ma)
absent
absent
between KAI (9.6 Ma) and
between Can Llobateres 1

(9.7 Ma) and Trinxera
Autopista (TNA/TSA2)
absent
below KAI (older than 9.6
KA2A (9.5 Ma)
Ma)
(9.2 Ma)
requires the formation of some barrier, a process which in most

cases will take a significant portion of geological time. But even
Spanish faunas (MBB, TAP, TOI) are very different: in each in those instances where species have been observed to migrate
eas yield Rsp values of zero (TOl-MNR, TOl-LEF, TAP-LEF),

i.e., they have not a single species in common. Even the three
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
and Montredon (SW France), however, show a value larger than

50 (Rsp = 63), while Rgen (numbers above the diagonal in Table
4) is 100, i.e., all genera known from the smallest fauna, Trinx
era Autopista, are present in Montredon. However, the average
Rgen for all comparisons is 42, i.e., less than half of the genera
in the smaller faunas occur in the larger faunas to which they
are compared. In five out of the ten comparisons the smallest
faunas have only one or two genera in common.
dispersal (less than 0.01 m.y.) across an entire continent is only

to be expected under special circumstances, e.g., when a taxon
displays some innovative key adaptation (Rosenzweig and
McCord, 1991), or when a large ecosystem crash allows pioneer

species to distribute widely.
The Progonomys "Event"
Table 5 shows the comparison between rodent assemblages

Species of the primitive murine rodent genus Progonomys
from the adjacent Daroca-Calatayud and Teruel basins with are among the rare examples of widely distributed species dur
contemporaneous (11-9 Ma) assemblages from the Vall?s-Pe ing the Late Miocene of Europe (Table 4). Recent magneto
ned?s basin, which is situated 300 km in ENE direction. The stratigraphic work has resulted in better constraints for the tim
oldest two rodent assemblages, Nombre villa and Hostalets su
ing of the so-called Progonomys event, which is a multiple
perior, resemble each other reasonably well (Rsp and Rgen are 43
event corresponding to the immigration of different species (Ta
spite of their close distance.

The main reason for the low overall faunal similarity is with
out doubt related to the paleogeography of the European conti
nent. A complex Mediterranean coastline with various peninsu
las, an extensive inland lake system (Paratethys), and mountain
ranges (pre-Alps, pre-Pyrenees) probably all have their effect on
the lowering of faunal interchange. The likely presence of dif
ferent climatic regimes (from subtropical-dry to temperate hu
mid, and from maritime to continental) will further tend to en
hance faunal differences. In addition, coastal and more elevated
resolution (2 m.y.), given the spread in entry data (e.g., 12 Ma
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
in Pakistan (Jacobs and Downs, 1994), 10.5 Ma in Anatolia

(Lunkka et al., 1999), and ~10 Ma in Spain).
In Europe there are three wide-spread species of Progono
mys: P. hispanicus, P. cathalai and P. woelferi. Sections that
are correlated to the GPTS and contain the first local occurrence
of Progonomys are exposed in Spain (the Daroca-Calatayud/

Teruel, Vall?s-Pened?s, and Duero basins) and Greece (Kastel
lios Hill on the island of Crete). Bio- and magnetostratigraphic
references are indicated in Table 6.
inland areas may have had a different fauna (e.g., the Vall?s
P. hispanicus is recorded in all four sections. Its immigration

in the Calatayud-Daroca/Teruel basins consists of two steps (Ta
The above shows that is not realistic to rely on mammal ble 6). The first step is an occurrence with very low numbers
species alone for time correlations with high resolutions (lower (0.1% of cf. P. hispanicus in Pedregueras 2C). The second step
than 0.1-0.2 m.y.) across Europe. Using genera instead of spe is its first regular occurrence which is first documented in Mas?a
cies artificially increases faunal overlap but does not solve the
de la Roma 4B, where it represents ?60% of the rodent fauna
problem of time correlation, because genera are not the units
(van Dam, 1997). A comparable pattern is observed in the Due
that migrate: species are. When more than one congeneric spe
ro basin, where a very rare occurrence of P. hispanicus in Am
Pened?s basin versus the Calatayud-Daroca/Teruel basins).
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

rence of P. hispanicus can not be tested because of the rareness
The recognition of only 16 values on the scale (the reference
of the first representatives. For the time being it is true, how faunas) is in line with the very purpose of the MN system,
ever, that the first regular occurrence of P. hispanicus is iso
which is long-distance correlation. As illustrated by the exam
chronous in Spain and Greece at ?9.6 Ma (Table 6).
A second Progonomys species with a wide distribution is P.
cathalai. Its first occurrence in the Teruel basin is at 9.4-9.3

Ma, whereas its first occurrence on Crete occurs earlier (9.6
9.5 Ma). Furthermore, the absence of the species in both the
Vall?s-Pened?s and Duero basins means that P. cathalai is of
less interest as a precise time marker across SW Europe. Pre
ples above, more detailed correlations do usually not make

sense because of low faunal resemblances and potential diach
ronicity. There have been efforts to define true (i.e., biostrati
graphic) zones for larger areas ("MN zones," Agusti and
Moy?-Sol?, 1991; zone-like units of Aguilar, 1982), but it is

evident that equivalent faunas at the species level can at best
be successfully recognized within certain parts of Europe (e.g.,

sumably due to ecological reasons (competition in a different
Western Europe in the latter cases). Most genus "events" will
environment?) the species was not able to enter particular areas. probably be diachronous at levels of 0.5-1 m.y. or more across
P. woelferi is the oldest Progonomys species in the Kastellios Europe. This does not imply that a European zonation based
Hill section, where it occurs in the lowest fossiliferous level

KA1 (9.6 Ma). In Spain (like P. cathalai), it is documented in
one basin only (Vall?s-Pened?s), with a first occurrence in
Trinxera Nort Autopista/Sud Autopista 2 at 9.2 Ma. The next
on the appearances of wide-spread taxa is useless, but zone

boundaries can not assumed to be isochronous, and because
abundant, wide-spread taxa are rare, the zones will have long
durations. We therefore reject recent recommendations (Stein

inger, 1999) to abolish the reference-fauna interpretation and
Llobateres, Garc?s et al., 1996) and the absence between 9.7 redefine all MN zones in a strictly biostratigraphic way.
and 9.2 Ma might be an effect of sample size. Because of its
In addition to these more philosophical arguments, there is
uneven distribution in Spain P. woelferi is probably not a very another, more technical argument against the use of MN
reliable time marker, although in theory its entry might be iso
"boundaries." Suppose a newly identified fauna has to be cor
older, relatively rich localities have an age of 9.7 Ma (e.g., Can
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
species across Europe seems to be isochronous at the scale of

resolution of 0.5 m.y. At the scale of 0.1-0.2 m.y., the immi
gration of P. cathalai seems to be diachronous, whereas more
data are needed to demonstrate diachrony in the case of P.
hispanicus and P. woelferi. The use of of P. woelferi (and P.
cathalai) as a precise time marker is suspect because of its
absence in various Spanish basins.
cases it will be obvious with which reference fauna the studied
fauna correlates, but in other cases this will be not so clear.

Because the total fauna is the only criterion, all faunal aspects
have in fact to be considered, i.e., presence/absence, abundanc
es, size and morphology of all taxa. Next, it has to be decided
how all this faunal information should be weighed. In principle,
a multivariate quantitative approach could be used, but there
will be multiple solutions depending on the specific technique

or program options used. Consequently, there will be multiple
The Fiction of MN "Boundaries"
solutions for the perfect "intermediate" fauna ("boundary").
The low faunal resemblance and potential diachrony of spe
Alternatively, one could be satisfied with a more qualitative
cies across Europe imply that long-distance correlations based
and/or intuitive correlation, and not bother too much about the
on mammal faunas alone cannot be expected to be precise. This problem that different workers may correlate to different MN
holds for European Neogene mammal faunas in general. It im
units. With the purpose of the MN system in mind (coarse-level
plies that European mammal chronological systems such as the
correlation), such a different opinion may not always be prob
MN "zone" system have to be interpreted in a specific and

crude way to be useful. Particularly, the introduction of MN
boundaries is erroneous and misleading as will be explained
below. Similarly, the use of boundaries between continental
Stages/Ages is problematic. Both systems will be discussed
lematic. Obviously, boundaries make no sense in this case ei
ther. The placement of faunas in MN units in the standard tables

of Mein (1990) and de Bruijn et al. (1992) has occurred in such
a qualitative manner.
The general conclusion is therefore that the MN scale is only
against the background of "high-resolution" mammal data.
useful for crude correlation (?0.5-1 m.y. resolution, i.e., in the
In the most recent major treatment of the Mammal Neogene
(MN) system (de Bruijn et al., 1992) MN "zones" are a set of order of the average time spacing between the reference faunas)
and for correlations between different parts of Europe ("prov
16 time-ordered reference faunas. They are certainly not zones,
inces"). So-called "MN boundaries" do not exist and their age
which by definition are stratigraphie units, i.e., three-dimen
estimates make no sense. The same reasoning automatically ap
sional bodies of rock (International Stratigraphie Guide; Sal

vador, 1994). In our opinion then, a MN unit (=reference fau plies to the "boundaries" of European continental "Stages/
Ages" erected during the 1975 Munich symposium on mam
na), unlike a zone, only exists in the reference locality itself.
malian
stratigraphy (Fahlbusch, 1976), which are not Stages/
This implies that non-reference localities do not "belong" to
Ages sensu the International Stratigraphie Guide (i.e., defined on

the basis of Stratotypes), but just aggregates of two or three
ing to de Bruijn et al. "allocation" (correlation) should occur reference
faunas.
by comparing faunal contents only. Other correlation criteria,
including direct calibrations to the numerical time scale, should
an MN unit, but have to be correlated to one of them. Accord
therefore not be used for MN allocations. The MN scale is an European Mammal-based Stages/Ages: Are They Still
ordinal scale. In our opinion, strictly spoken, intermediate po

sitions between the reference faunas are therefore not defined.
A fauna is either MN9 or MN10 and not something in between;
Necessary?
During the 60s and 70s European mammal paleontologists
felt the need to define continental chronostratigraphic/geochro
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
they are currently used and expressed in millions of years) have

no meaning because there is nothing but "emptiness" between
successive reference faunas. In addition, there is not only time,
between marine and continental biozones. As a part of this pro
cess, the Vallesian and Turolian (Upper/Late Miocene) were

given official status as Stages/Ages (Marks, 1971a, b). Unlike
but also space involved in the voids between the reference lo the "MN aggregates" referred to above (which have unfortu
calities, because reference localities are situated in different nately been given "ian" name endings as well) these Stages/
Ages are based on true type sections.
geological basins or parts of Europe.

don, Cor Langereis, Lars van den Hoek Ostende, Ton van Hoof,
Although the definitions of these officially defined Stages
Fran?ois L?v?que, Filip Levering, Bob Martin, Carmen Martin,
conformed to the previous edition of the International Strati
graphic Guide (Hedberg, 1976), they do not conform to the new Pierre and Marie-Th?r?se Mein, Johan Meulenkamp, Francisco
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
y paleoclimatolog?a del Ne?geno continental de la fosa Cala

tayud-Teruel" (PB92-0013) and "Evoluci?n paleoambiental de
sucesiones con alta resoluci?n bioestratigr?fica en el Ne?geno
brought us very far, and, in our opinion, we can do without
continental: area de Daroca-Teruel (Arag?n-Espa?a)" (PB 95
them. The basic problem is correlation, and because more and
more local calibrations of mammal faunas to the numerical time 0114), financed by the Direcci?n General de Investigaci?n y
T?cnica, Spain. Teruel-Alfambra fossil mammals have been re
scale are now established (e.g., see Krijgsman et al., 1996;

covered thanks to Diputaci?n General de Aragon permission
Steininger et al., 1996; Lindsay, 1997) isolated, non-calibrated
Exp. 10/91, 2/92, 40/93, 64/94, 28/96, 22/97, 118/98, and 226/
faunas can and should directly be correlated to these calibrated
99 from the Departamento de Educaci?n y Cultura.
faunal sequences, and their age should be directly estimated in
terms of millions of years. Appropriate quantitative interpola
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The Upper Miocene Mammal Record from the Teruel-Alfambra Region (Spain).

Journal of Vertebrate Paleontology 21(2):367-385, June 2001

THE UPPER MIOCENE MAMMAL RECORD FROM THE TERUEL-ALFAMBRA REGION

28040 Madrid, Spain;

During the nineties pal?ontologie research took various new

The Teruel-Alfambra region in NE Spain (Figs. 1 and 2) is

huesos de el cuerpo humano, y otras, que representan huesos

resolution" magnetobiostratigraphic approach (sensu Lindsay,

see Woodburne, 1996) not only improved the chronology of

de bestias" near the village of Concud (see Alcal?, 1994, 1997,

font, Truyols and Villalta started to explore the region more

cal? (1994) combined detailed taphonomic analyses with paleo

de Teruel" (Crusafont, 1965). A formal definition of the Tur

GEOLOGICAL SETTING AND LITHOLOGY

olian as a continental Stage was given somewhat later (Marks,

1971a; Aguirre et al., 1975).

The Teruel-Alfambra region comprises half of the northern

part of the Teruel Basin, a NNE-SSW oriented basin in the

known from the sixties onwards, when Adrover started his ex

Sondaar (1961) and the work of van de Weerd (1976) on rodents.

The Neogene succession of the Teruel-Alfambra region (Fig.

368 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 21, NO. 2, 2001

mammal and abundant gastropod remains. A maximum thick

limestone beds (type 6 above) have been used for correlation

The (reddish) claystone-dominated intervals in the studied

FIGURE 1. Schematic map of the Teruel basin and surrounding areas

sections belong to facies unit M2-P12 of Hern?ndez et al. (1983).

basin and is mainly developed along the basin margin. It is

more or less equivalent to the Peral Formation of van de Weerd

into evaporites, e.g., in Los Mansuetos (LM) (Fig. 3) and at

de Weerd, 1976; Hernandez et al., 1983).

Most fossil mammal remains are found in organic-rich marl

close to the village of Tortajada (Hernandez et al., 1983). These

latter, more northern gypsum exposures constitute van de

riguez-Aranda and Calvo, 1998).

THE SEQUENCE OF MAMMAL LOCALITIES

The composite upper Miocene mammal succession of the

al. (1997) but a comprehensive approach to the factors con

trolling the formation of the mammal sites is still in progress.

the interpretation of litho-, bio- and magnetostratigraphic data.

The positions of isolated, poorly documented assemblages in

carbonate facies units M3 and M4m as described by Hernandez

that of the biozone.

The small mammal record, which almost exclusively consists

and R2, as well as the micromammal data from SAL, REG2

VAN DAM ET AL.?UPPER MIOCENE MAMMALS FROM SPAIN

limestones and marls

sandstones and clays

5, 6, 7; KS, KS2, KSS; CAP2, and MOD. The samples from

The biozonation is a modification of the Miocene part of the

mammal associations (ROM1, MBB, R2, VIP, MDV2, LP,

neighbouring Calatayud-Daroca Basin. We use this system next

introduce the local zones J to M, with J (Progonomys hispan

CCL, LM, CC, KS, ML and ARQ1) were described by Alcal?

Morales (1997), Azanza et al. (1997), Fraille et al. (1997) and

to the zone names of van de Weerd, which are retained. We

icus Zone) and M (Stephanomys ramblensis Zone) subdivided

JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 21, NO. 2, 2001

sandstones and conglomerates

carbonate channel fills

paleosols developed on alluvial fades

carbonates (tabular and indurated) :

mals have not been found in the Teruel-Alfambra region.