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. cuenca@posta.unizal.es

,. Dpto. de Paleontologla, Facullad de CienclSS Geol6glcas. Universidad Complutense. E- 28040 Madrid, Spain, nurlag@eucmax.sim.ucm.es

"'Museo Nacional de Ciencias Naluralcs. c. Jose Gullerrez Abascal. 2. E-28006 Madrid. Spain. mcnJv538@mncn.csic.es

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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