Resumen EI yacimiento de Trinchera Oolina es uno de los mas importantes de la Sierra de Atapuerca par su contenido en restos humanos, Homo antecessor en el nivel de Trinchera Dolina 6. Ademas de esto la enorme riqueza en restos arqueol6gicos y paleontol6gicos hace de Trinchera Dolina un yacimiento unico, de referencia obligada para el Pleistoceno y Paleolitico de Europa. Bioestratigraficamente, el yacimiento de Trinchera Oolina (TO) puede dividirse en tres grandes unidades: la que comprende los niveles TD3 a T06; la de los niveles T07 a TOS inferior y I. de TO 8 superior. T011.
Palabras clave: Mamiferos, Pleistoceno.
Abstract Gran Dolina is one of the Pleistocene sites located at the Sierra de Atapuerca (Spain). The Gran Dolina deposits belong to different chronological periods of the Early and Middle Pleistocene. The uppermost levels of Gran Dolina (TDII, TDIO and TD8b) contain Middle Pleistocene (post-Cromerian) macro- and micromammal assemblages. The excavation works have overpassed level TDII and have not concluded yet at TDIO: TDII is poor in macromammal remains (carnivores and herbivores) but rich in rodents. The macromammals fossil material from TDtt is very scarce and this enables (for the macromammals) definite conclusions about the chronology and type of community of these levels. The lowermost levels of Gran Dolina (TD3/4, TD5, TD6 and TD8a) contain a different mammal assemblage with typical late Early Pleistocene-Cromerian species. This radical substitution of taxa is placed at TD8 layer probably due to a stratigraphic gap in this level. The level TD6 of the Gran Dolina site contains the earliest fossil human remains of Europe, Homo antecessor, and it has also a rich and diverse micromammal assemblage. Rodents, insectivores, bats, rabbits as well as birds, lizards and amphibians are well represented: Mimomys savini, Microtus seseae, Stenocranius gregaloides, Terricola arvalidens, Iberomys huescarensis, Allophaiomys chalinei, Pliomys episcopalis, Allocricetus sp., Eliomys sp., Micromys minutus, Apodemus aft. flavicollis, Castor fiber, Marmota sp., Hystrix refossa, Beremendia fissidens, Soricidae spp., Crocidura sp., Talpidae spp., Erinaceus sp., Miniopterus schreibersii, Myotis spp., Rhinolophus spp., Oryctolagus lacosti and Lepus terraerubrae. The large mammals include, Homo antecessor, as well as a diverse large mammal assemblage: Among the herbivorous there are Mammuthus sp., Stephanorhinus etruscus, Equus altidens, Sus scrofa, Dama 'nestii' vallonnetensis, Cervus elaphus acoronatus, Eucladoceros gium, Bison cf. vOigtstedtensis, and the carnivores include Ursus sp., Crocuta crocuta, Mustela palerminea, Lynx sp., Canis mosbachensis and Vulpes praeglacialis. Paleoenvironmental reconstruction based upon the relative proportion and stratigraphical distribution of the mammalian fauna of the Gran Dolina section shows that there are represented several arid, open country phases as well as wetter, warmer and more wooded phases in the sequence of this site.
Key words: Atapuerca, Gran Dolina, Carnivores, Rodents, Herbivores, Middle and Early Pleistocene, BIostratigraphy.
1 4 1
Fossil mammals of the Lower to Middle Pleistocene site of Trinchera Dolina, Atapuerca (Burgos, Spain)
Gloria Cuenca Besc6s', Nuria Garcia" & Jan van der Made'"
Introduction
The Atapuerca sites are part of a complex karst system at Sierra de Atapuerca. This Sierra is a Mesozoic-core hill, related with the Iberian range, 14 km to the East of Burgos (Fig. 1).
The Sierra de Atapuerca has two main cave systems: Cueva Mayor and Trinchera del Ferrocarril. The first comprises the
Sima de los Huesos, Galerla del Silex, G aleria Baja, Galerla del
Silo, Galeria de las Estatuas and Portal6n sites, being the 8ima de los Huesos one of the most important collections of Middle Pleistocene age fossil human remains.
The Trinchera del Ferrocarril is an ancient railway cut that exposed several fossiliferous as well as fossil cave fillings that constitutes the Gran Oolina, Trinchera Penal, Galerfa-Tres Simas and Elefante sites (Figure 2). The red, conspicuous cave sediments from the railway trench attracted the attention of archaeologists si nce the seco nd half of the last century, nevertheless, is not until the 1960s that the enterprise of archaeological studies began by Clark, Strauss, Apellan iz
(Ortega, 1999). The first palaeontological study of the Sierra
was made by Torres in 1976, in his investigation on the Iberian fossil ursids (Torres 1987) . Emiliano Aguirre in 1978 began the
Atapuerca Project that we inherit today (Aguirre, 1995, 200 1, Carbonell ef alii, 1999). Gran Dolina is probably one of the
• Area de Paleonlologia. Oplo. C,encias de la Tierra, F.Cienclas. U. Zaragoza. E, 50009 Zaragoza. Spam. [email protected]
,. Dpto. de Paleontologla, Facullad de CienclSS Geol6glcas. Universidad Complutense. E- 28040 Madrid, Spain, [email protected]
"'Museo Nacional de Ciencias Naluralcs. c. Jose Gullerrez Abascal. 2. E-28006 Madrid. Spain. [email protected]
most famous localities in Western Europe after the discovery of Homo antecessor.
The longest stratigraphical sequence at Atapuerca is the Trinchera Dofina (TO) site , a 18 m cave fill ing divided in 11 stratigraphical levels (Fig. 1). Almost all of them (TD3-TD1 1) are
rich in fauna and artefacts (Carbonell ef alii, 1999). ESR dating
and U·series analysis allocate the Trinchera Oolina fossil iferous levels between about 200 ka and 800 ka (Falgueres ef alii, 1999). The paleomagnetic Matuyama·Brunhes boundary was detected at level TD7 ind icating there an age of 780 ka (Pares
& Perez Gonzalez 1995, 1999) thus the levels placed below
TD7 (TD6·TD3/4), are older than 780 ka. Direct dates on the
T06 fossils using ESR and U/Th methods are consistent, ranging from 780 to 886 ka (Falgueres ef alii, 1999), a time
span that could correspond with oxygen isotope stages 19, 20 or 21 . Next to the top of T06 a stratum named Aurora yielded lithic artefacts, abundant faunal remains and nearly 80 human fossils attributed to a new species, Homo antecessor (Bermudez de Castro ef alii, 1997). TD8a is dated between 563 ± 84 ka
and 653 ± 98 ka and TO 1 0- 11 from 400 to 300 ka. (Falgueres
ef alii, 1999).
We wi ll give here a brief description of the main lithological units described in the section (for more detail see Pares & Perez Gonzalez, 1995, 1999). The red sands and lutites with
heterometric limestone fragments dominate the lithology. The Trinchera Dolina section is divided in 11 lithostratigraphic units, from TD 1 , at the bottom, to TD11 , at the top of the section. TO 1
and T02 are sediments of interior facies and T03 to TD11 are exterior deposits with some breakdowns or sediments derived from the erosion or weathering of the host limestone. Units T03
142 Mlscclanea en homenaJe a Emiliano Aguirre. Pafeontologia
Legend
[]I] Palaeozoic
EI Mesozoic
c=J Cenozoic
{ ATA: Sierra de Atapuerca
• Main City
BUREBA CORRIDOR
BURGOS • '" A TrA -.
Duero Basin
''''.
•
SPAIN
FRANCE
MId~." ...... ..
• Logroflo
Ebro Basin
Fig. 1. Geographic si tuation of the Sierra de Alapuerca (Burgos, Spain). Note the
strategic pasillon of the Sterra at the cenl re of the Ebro·Quero corridor (Bureba
corridor) In the pass from Ihe Medite rranean Sea to the Atlantic Ocean.
Atapuerca conshtutes a fossil relieve in the Neogene sediments of the Bureba
corridor thai communicates the Ebro with the Duero basins (The Mediterranean and Ine Atlantic realms). The small hili is of the outmost importance to Ihe faunal
migrations between bolh areas (including 10 our ancestors, Homo antecessor and Homo heidelbergensis. Arsuaga e/alii. 1997, 1999, Bermudez de Castro et
afii, 1997. Carbonell el alii, 1995, 1999). Even up to today. Burgos IS a landmark
in Ihe Saint Jacques Pilgrimage roule).
to T011 contain fossil remains except level T09 that lacks
vertebrate remains, actually it only contains root-concretions
and bat guano at the top. Levels TD3 and TD4 are 2 m thick and they are constituted by sandy lutite with limestone debris. Unit
TD5 is 2.5m thick and is composed mainly by mud with debris horizons with angular pebbles. Unit T06 is also 2.5m th ick and
is more clastic with very little clay matrix than the previous ones.
The Aurora Stratum containing the human remains is a 20 cm
thick massive yellowish red lutite with limestone clasts (some of
them 22cm large). Level T07 is a 1.5m calcarenite layer. The
paleomagnetic reversal Matuyama/Brunhes (and therefore the
Early/Middle Pleistocene boundary) took placed upon the level T07. The T08 unit is a 2.5m thick unit formed by heterometric
clasts flows, poor in matrix. Unit T09 is a O.35m thick red clay
and bat guano layer without fossil contents. Unit TO lOis a 2m
thick matrix consisting on clastic flows contain ing large
limestone clasts (as large as 1.5m) that seems an opening of
the Gran Ool ina cave. Unit TO 11 with a 3 .5m thickness,
contains coarse clasts or boulders (up to 1 m) and is composed
TP
eomp,e$Or
TRINCHERA DEL
FERROCARRIL
Cueva ptlluda
CullYa dol Silo
100m
Fig. 2. The Trinchera del FerroCBrril (railway trench) and Cueva Mayor karst
Systems of Atapuerca (Burgos. Spain). Labels indicate the main cave 10cali!lolj;
TP Penal, TD=Dotina, TZ. TG , TN-Tres Simas Complex (Zarpazos. Gahlll'l ,
Norce). TE Sima del Elefante, SH=Sima de los Huesos. The' points to Ihe sitos with fossil human remains (Redrawn Irom Martin Menno el alii: 198 1).
mainly by sandy clays and mud levels. At the top of the sequence, the terra rosa fills the cracks and joints the limestone
at the top of the site.
Biostratigraphy of Gran Dolina The mammal assemblage of levels 3/4 to 8a in Sierra de
Atapuerca, includes the carnivores: Ursus sp., Crocuta
crocuta, Mustefa paferminea, Canis mosbachensis, Vufpes
praegJaciafis, Lynx sp., Homotherium fatidens and Panthern
gombaszoegensis; the large hervibores Mammuthus sp.,
Stephanorhinus etruscus, Equus altidens, Dama 'nestii'
val/onnetensis. Euc fadoceros giulii, Cervus e/aphus
acoronatus, Bison cf. voigtstedtensis, Sus scrofa,
Hippopotamus amp/:libius, Praeovibos sp.; and the rodents
Allophaiomys chafinei, Stenocranius grega/aides, Terricola
arvalidens, Pliomys episcopafis, Mimomys savini, fberomys
huescarensis, Microtus seseae, Microtus aft. oeconomus, and
Microtus agrestis. The se taxa are characteristic of the
G. CUENCA BESCOS, N. GARCIA & J. VAN OER MADE I Fossil mammals 01 the Lower to Middle PleIstocene sito. of Trinchera Dolina, Atapuerca 143
. k.
H.a
g " "0 .c -.-..J
• •
Call1ivorcs
I
•
I I
Small herbivores Large herbivores
• • •
fig. 3, Faunal distribution of the mammals at Gran Dolina Site (Atapuerca, Burgos, SpaIn). The dolled line in T08 (with a lateral arrow) represents a strallgraphic hiatus (there IS no foss~ record within a timespan of sevcralthousands of years) : 10 this chart this change on the faunal aSSOCiation eM be clearly observed. H.a.: Homo anleceSSOf, The species on grey are also preseot at the Trinchera Galerla Middle Pleistocene site. close to the Dolina si te; (Sources: Van der MAde et alii. 2003; Cuenca Besc6s 01 alii, , 999).
European late Early Pleistocene (Cuenca Besc6s et alii, 1999 , Garcia & Arsuaga, 1 999 & Made, 1999).
Some of the species coming from levels 3·8a (i,e. Vulpes praeglacialis, Homotherium latidens, Panlhera gombaszoegensis, Dama 'nes tii' vallonnetensis, Stephanorhinus c t fliSCUS, Bison cf. voig tstedt ensis, Mimomys savini, Stenocranius grega/oides and Terricola arvalidens) survive
beyond the Early to Middle Pleistocene boundary and can also
be found in early Middle Pleistocene localities in central and
North Europe.
The faunal assemblage of levels 10 and 11 include Ursus ct. nrctos, cf. Cuan a/pinus, Pan thera leo , Vu lpes vulpes, Homotherium la tidens, Meles meles, Mustela sp.
(erminealn;valis), Hemitragus bonnali, Equus cabal/us, Bos/Bison, Stephanorhinus cf, hemitoechus, Sus scrofa, Dama dama clactoniana, Cervus elaphus ct, priscus, Arvicola cf. sapidus, Pliomys lenk;, Microtus agrestis, Microtus arvalis, Jberomys brecciensis and Terricola atapuerquensis. The faunal
distribution is shown in Figure 3,
The carnivores of Gran Dolina
The Canids Vulpes praeglacialis is present in Dolina levels 4·6. This species
seems to be the ancestor of the arctic fox, Alopex /agopus, so
its presence suggest s cool conditio ns, H owever, our
interpretation is that , given that none of the other taxa (rodent
and ungulates) as well as the palinological data show any
evidence of cold cond itions, it is likely that Vulpes praeglacialis had not incorporated the typical 'arctic traits' of a cold adapted
species that the arc tic fox has, This same line of argumentation
is proposed for the Praeovibos·Ovibos evolut ive line (Kahlke,
1999). Vulpes vulpes makes its appearance in the Atapuerca
sites after the Matuyama·Sruhnes reversal, in TO 10 and TO 11,
Trinchera Galerfa and Sima de los Huesos, being all post
Cromerian sites form the Sierra.
The presence of cf. Cuon a/pinus at T011 layer is pointed out
due to an isolated fragmentary lower fourth premolar, which
shows a secondary posterior cusplet. Th is feature is never
present in Canis lupus, Furthermore, the size of this premolar is
1411 MI8COIf\JlOa Of) hornenaje a Emlhano Aguirre. Paleontologia
too large to belong to Canis mosbachensis, at least to the small
one that appears in the Oolina layers. The small -sized wolf Canis mosbachensis is found in
Trinchera-Oohna levels 4-8a. The features that identify the C. mosbachensis line are (among others), a poorly-developed
hypoconulid basin of M and a second posterior cusplet of Plow
and separated from the posterium cingulum. Both traits are
observed in the Oolina specimens from T06-Aurora stratum.
The Hyaenids from Gran Defina site The genus Crocuta is always present from the T04 Early
Pleistocene level up to the T08a Middle Pleistocene level, thus
a coexistence with the T06 level ancient hominids is confirmed.
The top level, T011 which can represent the last-Middle
Plei stocene (around oxygen stage 6) , has been already
excavated in a large extension (about 800 square feet) and
none hyaena remains were recovered.
There are other two european sites with possible Crocuta remains corresponding to an early chronology (Selva Vecchia and Belfia V) but bolh cases are ambiguous (Garcia et
Arsuaga, 1999). The Crocuta crocuta remains from the
lowermost levels of TO (T04-5) can be considered as the
ol dest occurrence of undoubted stratigraphical and
chronological position in Europe.
The Large Felids
Homotherium latidens cat is present in level T05E of Gran
Oolina. This large cat is known throughout Europe an Asia from around 3 Ma, having its last occurrence in Western Europe 458
ka ago in Fontana Ranuccio (Gliozzi et afii, 1997). H. lat/dens was one of the longest-lived of the larger carnivores. occurring
throughout the Villafranchian and into the Middle Pleistocene.
The oldest currently record for Panthera leo is at Isern ia, placed
in the Middle Pleistocene , between 500 ka and 783 ka
(Kolfschoten, 1996; Garcia & Arsuaga, 1999). Homolherium survived in Europe until the end of the Cromer, coinciding with
the first occurrences of the African lion (with a short co
existence of both species at some late Cromerian sites as
Mauer, Westbury, Mosbach or Vertesszolos) . Their size similarity
and resource competition (Hutchinson, 1959) could explain the
extinction of Homotherium.ln the Sierra de Atapuerca, Panthera leo is first recorded in the uppermost levels otTO (T01 0-T011),
placed between ois. 9 and 11 , when Homotherium was already
extinct. It seems that Panthera leo had not arrived in Europe prior
to Cromerian III-IV interglacials or was ve ry scarce;
Homotherium was still the large felid predator in the ecosystem.
The herbivores at Gran Dolina site
Mammuthus sp. Part of the fossils from T06 were recovered in 1985 from
blocks that were falien down. They include a much worn 04,
that either belongs to an evolved Mammuthus meridionalis or
one of the first typical M. Irogonlherii (A9uirre, 1999). The
presence of a proboscidean in T06 is confirmed by a fragmont
of another milk tooth in situ.
Stephanorhinus etruscus Fossils from TOW4, TOW5, T06 and T08a have been assign/tll
to S. elruscus (Van der Made, 1998, 1999). Guerin (1980)
recognized the lineage: Dieerorhinus etruscus etruscus - () etruseus brachycephafus - D. hemitoeehus. Fortelius at nf/l (1993) placed these rhinos in Stephanorhinus, assigned man I
of Guerin's D. e. brachyeephalus to S. hundsheimensis and dkl
not consider these forms, and a small S. ct. hundsheimensis, I be a sin91e lineage. Van der Made (2000) largely foliowod
Fortelius et alii (1993), but considered the three forms to form II
sequence in time. In several cases, late Early Pleistocene alld early Middle Pleistocene rhinos have (because of the supposed
age?) been assigned to D. e. brachycephalus and later this WOH
translated into S. hundsheimensis. The fossils of the time period
should be revised. T08a is possibly the youngest, or at least onO of the youngest localities with S. etruscus. Voigtstedl w llh
S. hundsheimensis is probably only slightly younger.
Stephanerhinus ef. hemitoeehus Poor remains from TOl 0-11 have been assigned to S. ct, hemiloechus (Van der Made, 1998). This species replaced S. hundsheimensis, which is still present in Mauer (Von
Koenigswald & Heinrich, 1999). It is more hypsodont and has u
smaller second premolar. This does not exclude that one evolved
into the other, whether this is really the case is the question, and
the possibility exists that S. hemitoechus arrived in Europe by
dispersal and is first recorded in localities such as Arago
(Moigne el alii, 2000) and Bilzingsleben (Van der Made, 2000).
EqutJs Remains from TDW4, T06 and T08a were assigned to W£quus sp. stenonid type" or to Equus ct. altidens and remains from
T010 and T011 were assigned to "Equus sp. cabalioid type"
(Van der Made, 1998, 1999). In a general sense, stenonid
horses are more abundant in the Early and early Middle
Pleistocene, and cabaUoid horses more in the younger localities.
Sus serofa A single premolar from T06 was assigned to Sus serefa (Van
der Made, 1999). A 1hird cuneiform from TOE5 and half a molar
from TO 10 belong very probably to the same species. Two
species of suids are recognized in the Pleistocene of Europe:
Sus strozzii and Sus scrofa (Faure & Guerin, 1984). Sus strozzii has extremely wide premolars and European S. scrofa tends to have particularly narrow premolars.
Hippopotamus amphibius An upper incisor from T08a was assigned to this species (Van
der Made, 1998). Pleistocene European hippos have either
been seen as various chrono-subspecies of H. amphibius
G. CUE NCA BESCOS. N. GARCiA & 1. VAN OER MADE I Fossil mammals of the Lowor 10 Middle Pleistocene slle of Trinchera Dohna. Atapuerca 145
(Kahlke, 1987) or as a more complex group of different species
(Faure, 1985; Mazza, 1991 ). The incisor does not permit a
precise identification.
Duma Remains from TOW4, T06 and T08a have been assigned to
Dnma IInestii" vallonnetensis (Van der Made, 1998, 1999).
'~ece ntly excavated material from TOE5 belongs to the same
form. Remains from T01 0 have been assigned to Dama dama IIff. clactoniana (Van der Made, 1998). There are various
opinions on the Dama-like deer. Azzaroli (1992) ptaced the
oarl ier forms in a different genus Pseudodama and assumed
this genus to be not closely related to Dama. Van der Made
(1996, 1998, 1999) considered these early forms and recent
Dama a single lineage and a single genus. Pfeifer (1997, 1999)
Included Pseudodama as a su bgenu s in Dama . Dama Inc reased antle r complexity and finally evo lved palma ted
on tiers. Size increased much from the earliest Pleistocene till
the time of Bilzingsleben and Atapuerca TGl 0-11 for later
decreasing in size (Van der Made, 1998, 1999a, 1999b). The
Dama from Atapuerca T04-6 and T08a is a little larger than
Dama from Vallonnet and Untermassfeld.
Dama from TOI 0 is a little smaller than from TG 1 0-1 ,. and
was assigned to Dama dama clactoniana (Van der Made,
1999b). The lack of distal parts of the antlers in T010 is a
problem, since at this time palmation started to evolve.
Cervus elaphus Material from TOW4, T06, T08a, TOI 0 and TOIl was
assigned to Cervus e/aphus (Van der Made, 1998, 1999).
European Pleistocene Cervus was considered to belong to two
species C. acoronatus and C. elaphus or two (chrono)
subspec ie s. Rec en tly, O i Stefano & Petronio (1993)
recognized many SUbspecies. It seems however, that there are
four chrono·subspecies: C. e. acoronatus (large, antler without
crown) , c. e. priscus (small, antlers w ith crown), C. e. spe/aeus (large, with c rown) and C. e. e/aphus (small, c rown, short
metapodials) (Van der Made, in prep.). The size changes are
important and occurred in the fashion of the ~ punctuated
equilibria". The remains from TDl 0 -11 belong probably to C. e.
priscus and those from the lower levels to C. e. acoronatus, but
the distal parts of the antlers are lacking in all levels.
Eucladoceros giulii Material from TOW4 and T06 was assigned to E. giulii (Van der
Made, 1999) and material from T08a with some reservation;
alternatively it might represent M. solilhacus (Van der Made,
1998). New material from TOE5 belongs to E. giulii. This
species was only recently recognised (Kahlke, 1997). It now
seems the dominant large deer from the later part of the Early
Pleistocene. The form is characterize by large and slender
metapodials , whereas those of M. solilhacus are robust.
Metapedial size increased with time, though general size seems
to have fluctuated. This means that the metapodials became
relatively larger. If this model is correct, the following localities
are in order from old to young: Venta Micena, Unlermassfeld,
Atapuerca T04 and Apollonia-1 . II is now clear that some very
large and very worn teeth assigned to Cervidae indet. (Van der
Made, 1999) represent th is species.
Mega/oceros giganteus Van der Made (1998) assigned a molar fragment from TOI 0 to
"Megaloceros giganteus?". Several years of excavation of the
upper levels of Gran Oolina failed to confirm the presence of
this species.
Praeovibos/Ovibos Two hind limbs of a single individual from T07 represent either
Ovibos or Praeovibos (Van der Made, 1998). Praeovibos sp.
with slender metapodials is known from Venta Micena (Maya
Sola, 1987). This form was probably not yet adapted to arctic
or glacial environments, but to arid envi ronments; later forms
aquired more robust metapodials and were probably more cold
adapted (Sher, 1992; Van der Made & Rodriguez, 1999). The
individual from T07 has stlll relatively gracile proportions.
Hemitragus bonali Some fossils from T01 0 have been assigned to Caprini indet.
(Van der Made, 199 8). A liUle more material is now known, also
from T011 , and permits an assignation to Hemitragus bonali. This form is first known from Vallonnet (Moulle, 1998) and is
replaced during the late Middle Pleistocene by H. cedrensis (Faure & Guerin, 1994).
Bison ct. voigtstedtensis Material from TOW4, T06 andT08 info was assigned tocl. "Bison voigtstedtensis" (Van der Made, 1998). There is new material from
TOE5. A skull from the old excavations was assigned to Bison schoetensackicf. voigtstedtensis (Soto, 1 987). It is now clear that
there are two lineages or groups: the B. degiulii - schoetensacki lineage and the B. menneri . voigtstedtensis group or lineage (Van
der Made, 1998, 1999a). B. menneri is a large form with slender
metapodials (Sher, 1997); B. voigtstedtensis is a little smaller.
Remains of a large bovine from TOl 0 and TOll were assigned to
Bas/Bison (Van der Made, 1998). The size is more or less
comparable to B. schoetensacki. TG10 and TGll have very
similar faunas, but differ in the bovine, which is very small there
(Van der Made, 1999b).
The rodents at Gran Dol ina site
Mimomys savini and Arvicola The MimomyslArvicola lineage is well known and widespread in
the European Pleistocene. Moreover the MIA transition marks
the BiharianfToringian boundary in the European chronology. The
species M. savini is the largest vole of the rodent assemblage in
Gran Oolina levels T03-T08a. It has the Mimomys enamel
146 Miscelanea en homenaje a Emil ana Aguirre. Paleontologia
differenciation, roots, cement in the reentrant angles, enamel islet
in young individuals and Mimomys fold. The proportion of
individuals with enamel islet, (in early ontogenetic stages), and
with closed linea sinuosa or presence of roots, (late ontogenetic
stages), are successively reduced with time. In Arvicola, this
character is absent or present vestigial at such sites as Cullar de
Baza. In young specimens, without roots, the enamel islet is
present in 14% of the assemblage, and the typical Mimomys
enamel differentiation is not evident. The enamel islet is less
frequent in modern populations (Huescar 14%, West Runton
7%, Voigtstedt and Prezletice 0 - 1 %) than in older populations
of Mimomys savini (Chiscau 1 and TD6 14%).
Iberomys aft. huescarensis and Iberomys breccien5is
The extant Iberomys cabrerae is an endemic vole in the Iberian
Peninsula tougth fossil species of this genus (I. aft.
huescarensis, I. brecciensis) are found in l ower to Middle
Pleistocene Mediterranean localities in Spain, France and Ityaly
(Ventura et alii, 1998, Cuenca Besc6s ef alil~ 1999, ). This
lineage probably split from the Microtus s.l. group as the
"Iberomys clade" by the end of the lower Pleistocene. The
Iberomys origin may be related with Allophaiomys nufiensis
from les Valerots and Monte Peglia. We also found prim itive
morphotypes of Iberomys in the species Microtus hintoni and
M. theniifrom Untermaf3feld and Neuleiningen 5, 15 (l ap lana ef
alii, 1999). The study of the relationships among these species
is still in progress. The autapomorphy of the lineage may be
defined as the asymmetry of the bucco-lingual salient angles:
the ratio Lalli is always lower than the same ratio in other
lineages of Microtus (Cuenca Besc6s et alii, 1995, 1999). The
spec;ies I , huescarensis, with advanced "nutiensis"
morphology, is the first representative of the Iberomys lineage
and it is found as f. aft. huescarensis in Atapuerca, in Trinchera
del Elefante, and Gran Dolina levels 3-8a (Lapiana & Cuenca
Besc6s, 2000). The middle Pleistocene species I. brecciensis
appears in Oolina 8b, 10 and 11, and it is characterised by the
advanced stage of the t rai t asym met ry. The species I. brecciensis is also present in the Trinchera Galeria. The species
I. huescarensis (and relatives) is of late Early Pleistocene age
and I. brecciensis characterises the early Midd le Pleistocene in
Spain.
Stenocranius gregaloides and Terricola arvalidens
Both species are more frequent in the lower levels, T03-T05
than in T06, being T. arvalidens somewhat more abundant and
one of the scarce faunas of the level T07. The differences
between S. gregaloides and T. arvalidens from Gran OoHna and
its represent ants from Middle Pleistocene localities can point to
different chronospecies. The holotype of S. grega/oides from
West Runton, is somewhat different from the S. gregaloides
from TD3-TD6 in the deeper LRA5 in the West Runton
association. Al l increases from older to younger assemblages
as Rekovets & Nadachowski (1995) show. The species S.
gregaloides and T. arvalidens are present together at numerous
localities in central and southern Europe dated to Cromerian 5.1. (Sutcliffe and Kowalski, 1976, Rekovets & Nadachowski,
1995). The primitive appearance of S. gregaloides of TD3-TD6
could indicate an early radiation of this species at the end of the
Early Pleistocene.
Microtus seseae
First described by Gil (1997), this is a species restricted to the
G ran Oolina levels T0 3-T0 7. Our preliminary work allows us to
propose that M. seseae is closely related to Ihe Middle
Pleistocene and extant Microtus species as M. nivaloides, M.
malei, M. nivalinus and Tyrrhenicola and it may represent a
second radiation of the Microtus group at the end of the Lower
Pleistocene.
Aflophaiomys chalinei
The species A. chalinei is the second largest vole of the
Gran Oolina section. Its size and morphology split it from the
rest of the microt ines. The species is typical of l ower
Pleistoce ne localities of Spain and Italy, such as Cueva
Victoria, Bagur 2, Casablanca 3(=Almenara 3), Castelldefells,
l as Cabezas, Fuentenueva 3 and Pi et rafi tt a, The last
appearance of Allophaiomys chalinei in Ool ina is at the
midd le of level 6 (Cuenca Besc6s et alii, 1999, Lapiana,
1999). Leve l TD 6 records the lasl ap p earance of the
species also in Europe.
Microtus aff. oeconomus
This species is found exclusively in level T0 8a. Several
species with the ratticepoid morphology that characterises
the Pallasiinus lineage are cited in several localities of late
Early Pleistocene to early Middle Pleistocene age as Kozi
Grzbiel, Hohensiilzen, West Runton, Voiglstedt, Belfia 7,
Holsteijn, Shamin, Colle Curti. All of the contain Mimomys
savini faunas.
Terricola atapuerquensis
The species is present in levels T08b till TO 11. It is a large
Terricola defined by Gil (1996). It is characterised by its large
size and anterior complex. It is found also in Trinchera Galeria
(Cuenca Besc6s et alii, 1999b).
Microtus agrestis jansoni This species is found in T01 0, 11 and also in Trinchera Galeria
(Cuenca Besc6s et alii, 1999). It is a species of Microtus with
agrestis or agrestoide morphology, probably related with the
extant M. agrestis. The species M. janson; was first described
by Chaline (1972) as a subspecies of M. agrestis in the early
Middle Pleistocene site of La Grotte de l'Escale a Saint Esteve
Janson. The author distinguish the species from M. agrestis
aubinensis from the late Pleistocene of Pointe du Bois a
Santenay by its larger size.
G. CUENCA BESCOS, N. GARCiA & J. VAN DER MADE / Fossil mammals of the l ower to Middle Pleistocene site of Trinchera Dolina, Atapuerca 147
,
Pliomys episcopalis and Pliomys lenki
The rooted vole Pliomys is represented in Gran Oolina by two
species: P episcopalis in the levels T03-T06 and P.lenki at the
T01 0-11. Both species are found together in several localities
of late Early-early Middle Pleistocene age (Bartolomei et alii,
1975). Its taxonomic status needs further revision.
Conclusions
The MimomyslArvicola transition marks the BiharianfToringian
boundary and it was earlier coincident with the Lower/Middle
Pleistocene boundary in the continental Ouaternary biostratig
raphy. The updated chronology of the Ouaternary uses the
paleomagnetic reversal Matuyama/Brunhes boundary as the
Lower/M iddle Pleistocene boundary and M. savini crosses this
limit up to the early Middle Pleistocene (West Runton, Voigtstedt
and Cromer faunas in general). Thus, the Late B iharian,
Mimomys savini Roden t Zone includes the paleomagnetic
boundary Matuyama/ Brunhes in Europe, such as Stranska
Skala, Grace, Shamin, Mahlis, Karlich, Atapuerca TD7 (Cuenca
Besc6s et alii, 1999). TD6 is below the TD7 level which shows
the paleomagnetic Matuyama/Brunhes boundary in the Gran
Dol ina Section (Pan,s & Perez-Gonzalez, 1995, 1999). The
Matuyama/Brunhes boundary is fixed in the late Biharian
(Microtus-Mimomys rodent Superzone). In th is biozone, the
species Mimomys savini presents advanced characters as the
loosening of roots and lack of the UMimomys islet" and this zone
is characterized by the absence of small Mimomys species as M.
pusilfus or M. blanci. The assemblage and characteristics of the
species M. savini, T arvalidens, S. gregaloides ind icates that
TD3-TD61evels are older than West Runton (type Cromerian). In
the Gran Oolina Section we can calibrate, for the first time, the
evolutionary rank of those important biochronological markers
with magnetost ratig raphy. We p ropose that a radiation of
Microtus s.I., with the appearance of the fi rst, p rimitive S.
gregaloides, T arvalidens and Iberomys took place just before
the Matuyama/ B runhes boundary. Very close to this first
radiation of Microtus a second took place short before the
paleomagnetic event, leading to the appearance of new species
as M. seseae in Gran Oolina and other species of the Microtus
group in Italy (i.e. M. arvalis, M. multiplex-subterraneus of Selva
Vecchia, Fontignano, Sardelia et alii, 1998) or the latest Early
Pleistocene faunas of Eastern Europe w ith the first appearance
of the M. gr. oeconomus (Markova, 1998).
The last appearance of Allophaiomys chalineii n Oolina is at the
middle of level 6. This species is typical of other Early Pleistocene
localities of Spain and Italy, such as Cueva Victoria, Bagur 2,
Casablanca 3 (=Almenara 3) , Castelidefelis, Las Cabezas,
Fuentenueva 3 and Pietrafitta. Their age range between about
1.6 (Bagur 2) to 0.8 Ma (Gran Dolina 6) (Lapiana, 2000, Cuenca
Besc6s this volume). The species M. savini survive beyond the
Early to Middle Pleistocene boundary and can also be found in
early Middle Pleistocene localities in central, north and
southeastern Europe (Kolfschoten & Turner, 1996; Rekovets &
Nadachowski, 1995; Cuenca 8esc6s, this volume).
Sus scrota and Cervus efaphus, seem to have entered
Europe during the late Early Pleistocene while Eucladoceros
giufii was already present.
Panthera leo does not enter into Europe from Africa until the
Early Middle Pleistocene, being first present at Isernia, placed
between 783 and 500 ka (Garcia et Arsuaga, 1999). This large
size cat could have been the substitute (in ecological terms) of
the large sabretooth Homotherium latidens, previously present
in the Sierra (in level TD5E). At the Atapuerca sites, Panthera
leo remains have only been recovered from layers dated
between 500 and 200 ka.
The Crocuta crocuta record is continuous from the lowermost
Oolina levels up to layer TOSa (just above the MatuyamalBruhnes
boundary). Here is the first undubious oldest record of this taxon
in Europe. The coexistence of Homo antecessor and this
specialized hunter-scavenger could have resulted on resource
competition and final displacement of the hyaenid from the Sierra.
From the stratigraphic hiatus at TOSb, no Crocuta remains nor
coprolites have been found, at the TOl l (already excavated) level
nor at TD l 0 (that is still being excavated).
The mammal association and the magnetochronology and
combined U'series/ESR dating methods of Pares & Perez
Gonzalez (1999), Falgueres et alii (1999, van der Made, 1999,
Garcia & Arsuaga, 1999) provides confi rmation of an age range
between 780 and 850 fo r t he Dolina levels 3-6. The
Matuyama/Bru nhes boundary at the level T07 gives an age of
780ky for this level and the dates for TD8 are between roughly
500 and 700 ky (Falgueres et alii, 1999).
Two main associations chronologically very different can be
detected in the Sierra de Atapuerca sites: one of a late Early
Pleistocene age and another placed on the Middle Pleistocene.
We can establish the relative evolutionary stage of the taxa
coming from different layers and ten tat ively correlate the
Atapuerca si tes among them and relate them with other
Pleistocene Western European sites.
The macro- and micromammals association together with the
magnetochronology and combined U-series/ESR dat ing
methods, provides confirmation of an age range between 7S0 and
850 ka for the Dolina levels 3-6 (Cuenca Besc6s et alii, 1999;
Falgueres et alii, 1999; Garcia et Arsuaga, 1999; Van der Made
1999; Pares et Perez-Gonzalez, 1999). The dates for TD8a are
between roughly 500 and 700 ka (Falgueres et alii, 1999).
The lower levels of Gran Oolina (T03 to TOSa) may represent
the climatically unstable period of the end of the Lower
Pleistocene. Arid , open country mammals as Marmota and a
high diversity of arvicolids are founded in T05. The warmer and
relatively wooded country may be represented in T04 and T06
with a diversified soricidae fauna as well as the presence of
Hystrix and Castor. In the level TD8a, the presence of Hystflx
as well as Hippopotamus and the low diversity of voles may
indicate a relatively warm period. Levels TOSb and TO 10, TO 11
1' 111 MI II( I llunWI l in IlIhlUfilOIIl II [,"thana AgUirro. Paleontologia
may represent a longer and complex period with alternating
climate between warmer and cooler conditions (Van der Made,
1998, Cuenca Besc6s et alii, 1999). This is coherent with the
Oxigen Isotope Stages (O IS): levels T01 0, 11 are correlated
with OI S 9 to 11 , T06 is correlated with the OIS 19 or 21
(Falgueres et alii, 1999) and T05 may be correlated with the
OIS 22 and TD4 with the OIS 23.
Aknowledgements
The Atapuerca crew (represented by its directors Juan Luis Arsuaga, Jose Maria Bermudez de Castro and Eudald Carbonell) helped us in diUerent ways Wi th the
extraction of the fossil materiats during the Al ap uerca field season every.
Research has partially supported by Consejeria de EducaCl6n y Gultura 01 Junta
de C astilla y Leon, Direcci6n General de Investi9aci6n Cienllfica y Tecnica
(PB93·0066·C03, PB96 -0 0 ·CO I , PB 9 S'1 0 2 S-C03·02, PB96-00-C 03),
Ministerio de Educaci6n (M EC, HA-97-22) and "Unidad Asociada" program. The
follOWing students helped us in different ways : Cesar Lapiana. Jose Ignacio
Canudo, Beatriz Romero. SergiO B ajo, Raquel L6pez, Carlos GarCia, Toni Canals,
M. Dolores Garcia-Ant6n and Josetina Barreifos.
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