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. Ade­ mas 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 Euro­ pa. 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 infe­ rior 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 most famous localities in Western Europe after the discovery of The Atapuerca sites are part of a complex karst system at Sierra Homo antecessor. de At apuerca. This Sierra is a Mesozoic-core hill, related with The longest stratig raph ical sequence at Atapuerca is the th e Iberian range, 14 km to the East of Burgos (Fig. 1). Trinchera Dofina (TO ) site , a 18 m cave fill ing divided in 11 The Sierra de Atapuerca has two main cave systems: Cueva stratigraphical levels (Fig. 1). Almost all of them (TD3-TD1 1) are Mayor and Trinchera del Ferrocarril. Th e first comprises th e rich in fauna and artefacts (Carbonell ef alii, 1999). ESR dating Sima de los Huesos, Galerla del Silex, G aleria Baja, Galerla del and U·series analysis allocate th e Trinche ra Oolina fossil iferous Silo, Galeria de las Estatuas and Portal6n sites , being the 8ima levels between about 200 ka and 800 ka (Falgueres ef alii, de los Huesos one of the most import ant collections of Middle 1999). The paleomagnetic Matuyama·Brunhes boundary was Pleistocene age fossil hu man remains. detected at level TD7 in dicating there an age of 780 ka (Pares The Trinchera del Ferrocarril is an ancient railway cut that & Perez Gonzalez 1995, 1999) thus the levels placed below exposed several fossiliferous as well as fossil cave fillings that TD7 (TD6·TD3/4), are older than 780 ka. Direct dates on the constitutes th e Gran Oolina, Trinchera Pen al, Galerfa-Tres T06 fossils us ing ESR and U/Th methods are consisten t, Simas and Elefante sites (Figure 2). The red , conspicuous cave ranging from 780 to 886 ka (Falgueres ef alii, 1999), a time sediments from the railway trench attracted the attention of span that could correspond with oxygen isotope stages 19, 20 archaeologists si nce th e seco nd half of the last century, or 21 . Next to th e top of T06 a stratum named Aurora yielded nevertheless, is not until th e 1960s th at th e en terprise of lithic artefacts, abundant faunal remains and nearly 80 human archaeological studies began by Clark, Strauss, Apellaniz fossils attributed to a new species, Homo antecessor (Bermudez (Ortega, 1999). The first palaeontological study of the Sierra de Castro ef alii, 1997). TD8a is dated between 563 ± 84 ka was made by Torres in 1976, in his investigation on the Iberian and 653 ± 98 ka and TO 1 0- 11 from 400 to 300 ka. (Falgueres fossil ursids (Torres 1987). Emiliano Aguirre in 1978 began the ef alii, 1999). Atapuerca Project that we inherit today (Aguirre, 1995, 2001, We wi ll give here a brief description of the main lithological Carbonell ef alii, 1999). Gran Dolina is probably one of the 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 • Area de Paleonlologia. Oplo. C,encias de la Tierra, F.Cienclas. U. Zaragoza. Trinchera Dolina section is divided in 11 li thostratigraphic units, E,50009 Zaragoza. Spam. [email protected] from TD1 , at the bottom, to TD11 , at the top of the section. TO 1 ,. Dpto. de Paleontologla, Facullad de CienclSS Geol6glcas. Universidad Complutense. E- 28040 Madrid, Spain, [email protected] and T02 are sediments of interior facies and T03 to TD11 are "'Museo Nacional de Ciencias Naluralcs. c. Jose Gullerrez Abascal. 2. exterior deposits with some breakdowns or sediments derived E-28006 Madrid. Spain. [email protected] from the erosion or weathering of the host limestone. Units T03 142 Mlscclanea en homenaJe a Emiliano Aguirre. Pafeontologia

FRANCE

• TP Legend []I] Palaeozoic SPAIN MId~." ...... eomp,e$Or Mesozoic .. EI 100m TRINCHERA c=J Cenozoic DEL FERROCARRIL { ATA : Sierra de Atapuerca • Main City Cueva ptlluda

BUREBA • Logroflo CORRIDOR CullYa dol Silo BURGOS • '" ATrA -. Ebro Basin

Duero Basin

''''.

Fig. 1. Geographic situation of the Sierra de Alapuerca (Burgos, Spain). Note the Fig. 2. The Trinchera del FerroCBrril (railway trench) and Cueva Mayor karst strategic pasillon of the Sterra at the cenlre of the Ebro·Quero corridor (Bureba Systems of Atapuerca (Burgos. Spain). Labels indicate the main cave 10cali!lolj; corridor) In the pass from Ihe Medite rranean Sea to the Atlantic Ocean. TP Penal, TD=Dotina, TZ. TG, TN-Tres Simas Complex (Zarpazos. Gahlll'l, Atapuerca conshtutes a fossil relieve in the Neogene sediments of the Bureba Norce). TE Sima del Elefante, SH=Sima de los Huesos. The' points to Ihe sitos corridor thai communicates the Ebro with the Duero basins (The Mediterranean with fossil human remains (Redrawn Irom Martin Menno el alii: 1981). 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 mainly by sandy clays and mud levels. At the top of the vertebrate remains, actually it only contains root-concretions sequence, the terra rosa fills the cracks and joints the limestone and bat guano at the top. Levels TD3 and TD4 are 2 m thick and at the top of the site. 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 thick and Biostratigraphy of Gran Dolina is more clastic with very little clay matrix than the previous ones. The mammal assemblage of levels 3/4 to 8a in Sierra de The Aurora Stratum containing the human remains is a 20 cm Atapuerca, includes the carnivores: Ursus sp., Crocuta thick massive yellowish red lutite with limestone clasts (some of crocuta, Mustefa paferminea, Canis mosbachensis, Vufpes them 22cm large). Level T07 is a 1.5m calcarenite layer. The praegJaciafis, Lynx sp., Homotherium fatidens and Panthern paleomagnetic reversal Matuyama/Brunhes (and therefore the gombaszoegensis; the large hervibores Mammuthus sp., Early/Middle Pleistocene boundary) took placed upon the level Stephanorhinus etruscus, Equus altidens, Dama 'nestii' T07. The T08 unit is a 2.5m thick unit formed by heterometric val/onnetensis. Eucfadoceros giulii, Cervus e/aphus clasts flows, poor in matrix. Unit T09 is a O.35m thick red clay acoronatus, Bison cf. voigtstedtensis, Sus scrofa, and bat guano layer without fossil contents. Unit TO lOis a 2m Hippopotamus amp/:libius, Pra eovibos sp.; and the rodents thick matrix consisting on clastic flows containing large Allophaiomys chafinei, Stenocranius grega/aides, Terricola limestone clasts (as large as 1.5m) that seems an opening of arvalidens, Pliomys episcopafis, Mimomys savini, fberomys the Gran Oolina cave. Unit TO 11 with a 3 .5m thickness, huescarensis, Microtus seseae, Microtus aft. oeconomus, and contains coarse clasts or boulders (up to 1m) and is composed Microtus agrestis. These taxa are characteristic of the G. CUENCA BESCOS, N. GAR CIA & J. VAN OER MADE I Fossil mammals 01 the Lower to Middle PleIstocene sito. of Trinchera Dolina, Atapuerca 143

Ca ll1ivorcs Small herbivores Large herbivores

g "0" .c .-..J-

I

. k. H.a • • • • • I I

fig. 3, Faunal distribution of th e mammals at Gran Dolina Site (Atapuerca, Burgos, SpaIn). The dolled line in T08 (wi th a lateral arrow) represents a strallgraphic hiatus (there IS no foss~ record within a timespan of sevcralthousands of years) : 10 th is chart this change on th e 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, The carnivores of Gran Dolina Garcia & Arsuaga, 1 999 & Made, 1999). The Canids Some of the species coming from levels 3·8a (i,e. Vulpes Vulpes praeglacialis is present in Dolina levels 4·6. This species praeglacialis, Homotherium latiden s, Panlhera gombasz­ seems to be the ancestor of the arctic fox, Alopex /agopus, so oegensis, Dama 'nestii' vallonnetensis, Stephanorhinus its presence suggest s cool conditions, H owever, our c t fliSCUS, Bison cf. voigtstedtensis, Mimomys savini, interpretation is that, given that none of the other taxa (rodent Stenocranius grega/oides and Terricola arvalidens) survive and un gulates) as well as the palinological data show any beyond the Early to Middle Pleistocene boundary and can also evidence of cold conditions, it is likely that Vulpes praeglacialis be found in early Middle Pleistocene localities in central and had not incorporated the typical 'arctic traits' of a cold adapted North Europe. species that the arc tic fox has, This same line of argumentation The fau nal assemblage of levels 10 and 11 include Ursus ct. is proposed for the Praeovibos·Ovibos evolutive line (Kahlke, nrctos, cf. Cuan a/pinus, Pan thera leo, Vu lpes vulpes, 1999). Vulpes vulpes makes its appearance in the Atapuerca Homotherium la tidens, Meles meles, Mustela sp. sites after the Matuyama·Sruhnes reversal, in TO 10 and TO 11, (erminealn;valis), Hemitragus bonnali, Equus cabal/us, Tri nchera Galerfa and Sima de los Huesos, being all post­ Bos/Bison, Stephanorhinus cf, hemitoechus, Sus scrofa, Cromerian sites form the Sierra. Dama dama clactoniana, Cervus elaphus ct, priscus, Arvicola The presence of cf. Cuon a/pinus at T011 layer is pointed out cf. sapidus, Pliomys lenk;, Microtus agrestis, Microtus arvalis, due to an isolated fragmentary lower fourth premolar, which Jberomys brecciensis and Terricola atapuerquensis. The faunal shows a secondary posterior cusplet. This feature is never distribution is shown in Figure 3, 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 presence of a proboscidean in T06 is confirmed by a fragm ont one that appears in the Oolina layers. of another milk tooth in situ. The small -sized wolf Canis mosbachensis is found in Trinchera-Oohna levels 4-8a. The features that identify the C. Stephanorhinus etruscus mosbachensis line are (among others), a poorly-developed Fossils from TOW4, TOW5, T06 and T08a have been assign/tll hypoconulid basin of M and a second posterior cusplet of Plow to S. elruscus (Van der Made, 1998, 1999). Guerin (1980) and separated from the posterium cingulum. Both traits are recognized the lineage: Dieerorhinus etruscus etruscus - () observed in the Oolina specimens from T06-Aurora stratum. etruseus brachycephafus - D. hemitoeehus. Fortelius at nf/l (1993) placed these rhinos in Stephanorhinus, assigned man I The Hyaenids from Gran Defina site of Guerin's D. e. brachyeephalus to S. hundsheimensis and dkl The genus Crocuta is always present from the T04 Early not consider these forms, and a small S. ct. hundsheimensis, I Pleistocene level up to the T08a Middle Pleistocene level, thus be a sin91e lineage. Van der Made (2000) largely foliowod a coexistence with the T06 level ancient hominids is confirmed. Fortelius et alii (1993), but considered the three forms to form II The top level, T011 which can represent the last-Middle sequence in time. In several cases, late Early Pleistocene alld Pleistocene (around oxygen stage 6), has been already early Middle Pleistocene rhinos have (because of the supposed excavated in a large extension (about 800 square feet) and age?) been assigned to D. e. brachycephalus and later this WOH none hyaena remains were recovered. translated into S. hundsheimensis. The fossils of the time period There are other two european sites with possible Crocuta should be revised. T08a is possibly the youngest, or at least onO remains corresponding to an early chronology (Selva Vecchia of the youngest localities with S. etruscus. Voigtstedl wllh and Belfia V) but bolh cases are ambiguous (Garcia et S. hundsheimensis is probably only slightly younger. Arsuaga, 1999). Th e Crocuta crocuta remains from the lowermost levels of TO (T04-5) can be considered as the Stephanerhinus ef. hemitoeehus ol dest occurrence of undoubted stratigraphical and Poor remains from TOl 0-11 have been assigned to S. ct, chronological position in Europe. hemiloechus (Van der Made, 1998). This species replaced S. hundsheimensis, which is still present in Mauer (Von The Large Felids Koenigswald & Heinrich, 1999). It is more hypsodont and has u Homotherium latidens cat is present in level T05E of Gran smaller second premolar. This does not exclude that one evolved Oolina. This large cat is known throughout Europe an Asia from into the other, whether this is really the case is the question, and around 3 Ma, having its last occurrence in Western Europe 458 the possibility exists that S. hemitoechus arrived in Europe by ka ago in Fontana Ranuccio (Gliozzi et afii, 1997). H. lat/dens dispersal and is first recorded in localities such as Arago was one of the longest-lived of the larger carnivores. occurring (Moigne el alii, 2000) and Bilzingsleben (Van der Made, 2000). throughout the Villafranchian and into the Middle Pleistocene. The oldest currently record for Panthera leo is at Isern ia, placed EqutJs in the Middle Pleistocene, between 500 ka and 783 ka Remains from TDW4, T06 and T08a were assigned to W£quus (Kolfschoten, 1996; Garcia & Arsuaga, 1999). Homolherium sp. stenonid type" or to Equus ct. altidens and remains from survived in Europe until the end of the Cromer, coinciding with T010 and T011 were assigned to "Equus sp. cabalioid type" the first occurrences of the African lion (with a short co­ (Van der Made, 1998, 1999). In a general sense, stenonid existence of both species at some late Cromerian sites as horses are more abundant in the Early and early Middle Mauer, Westbury, Mosbach or Vertesszolos). Their size similarity Pleistocene, and cabaUoid horses more in the younger localities. and resource competition (Hutchinson, 1959) could explain the extinction of Homotherium.ln the Sierra de Atapuerca, Panthera Sus serofa leo is first recorded in the uppermost levels otTO (T01 0-T011), A single premolar from T06 was assigned to Sus serefa (Van placed between ois. 9 and 11 , when Homotherium was already der Made, 1999). A 1hird cuneiform from TOE5 and half a molar extinct. It seems that Panthera leo had not arrived in Europe prior from TO 10 belong very probably to the same species. Two to Cromerian III-IV interglacials or was very scarce; species of suids are recognized in the Pleistocene of Europe: Homotherium was still the large felid predator in the ecosystem. Sus strozzii and Sus scrofa (Faure & Guerin, 1984). Sus strozzii has extremely wide premolars and European S. scrofa The herbivores at Gran Dolina site tends to have particularly narrow premolars. Mammuthus sp. Part of the fossils from T06 were recovered in 1985 from Hippopotamus amphibius blocks that were falien down. They include a much worn 04, An upper incisor from T08a was assigned to this species (Van that either belongs to an evolved Mammuthus meridionalis or der Made, 1998). Pleistocene European hippos have either one of the first typical M. Irogonlherii (A9uirre, 1999). The 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 to have fluctuated. This means that the metapodials became (Faure, 1985; Mazza, 1991 ). The incisor does not permit a relatively larger. If this model is correct, the following localities precise identification. are in order from old to young: Venta Micena, Unlermassfeld, Atapuerca T04 and Apollonia-1 . II is now clear that some very Duma large and very worn teeth assigned to Cervidae indet. (Van der Remains from TOW4, T06 and T08a have been assigned to Made, 1999) represent this species.

Dnma IInestii" vallonnetensis (Van der Made, 1998, 1999). ' ~ece ntly excavated material from TOE5 belongs to the same Mega/oceros giganteus form. Remains from T01 0 have been assigned to Dama dama Van der Made (1998) assigned a molar fragment from TOI 0 to IIff. clactoniana (Van der Made, 1998). There are various "Megaloceros giganteus?". Several years of excavation of the opini ons on the Dama-like deer. Azzaroli (1992) ptaced the upper levels of Gran Oolina failed to confirm the presence of oarl ier forms in a different genus Pseudodama and assumed this species. this genus to be not closely related to Dama. Van der Made (1996, 1998, 1999) considered these early forms and recent Praeovibos/Ovibos Dama a single lineage and a single genus. Pfeifer (1997, 1999) Two hind limbs of a single individual from T07 represent either Included Pseudodama as a subgenus in Dama . Dama Ovibos or Praeovibos (Van der Made, 1998). Praeovibos sp. Inc reased antler complexity and finally evolved palma ted with slender metapodials is known from Venta Micena (Maya­ on tiers. Size increased much from the earliest Pleistocene till Sola, 1987). This form was probably not yet adapted to arctic the time of Bilzingsleben and Atapuerca TGl 0-11 for later or glacial environments, but to arid environments; later forms decreasing in size (Van der Made, 1998, 1999a, 1999b). The aquired more robust metapodials and were probably more cold Dama from Atapuerca T04-6 and T08a is a little larger than adapted (Sher, 1992; Van der Made & Rodriguez, 1999). The Dama from Vallonnet and Untermassfeld. individual from T07 has stlll relatively gracile proportions. Dama from TOI 0 is a little smaller than from TG 1 0-1 ,. and was assigned to Dama dama clactoniana (Van der Made, Hemitragus bonali 1999b). The lack of distal parts of the antlers in T010 is a Some fossils from T01 0 have been assigned to Caprini indet. problem, since at this time palmation started to evolve. (Van der Made, 199 8). A liUle more material is now known, also from T011 , and permits an assignation to Hemitragus bonali. Cervus elaphus This form is first known from Vallonnet (Moulle, 1998) and is Material from TOW4, T06, T08a, TOI 0 and TOIl was replaced during the late Middle Pleistocene by H. cedrensis assigned to Cervus e/aphus (Van der Made, 1998, 1999). (Faure & Guerin, 1994). European Pleistocene Cervus was considered to belong to two species C. acoronatus and C. ela phus or two (chrono) Bison ct. voigtstedtensis subspecies. Recently, Oi Stefano & Petronio (1993) Material from TOW4, T06 andT08 info was assigned tocl. "Bison recognized many SUbspecies. It seems however, that there are voigtstedtensis" (Van der Made, 1998). There is new material from four chrono·subspecies: C. e. acoronatus (large, antler without TOE5. A skull from the old excavations was assigned to Bison crown), c. e. priscus (small, antlers with crown), C. e. spe/aeus schoetensackicf. voigtstedtensis (Soto, 1 987). It is now clear that (large, with c rown) and C. e. e/aphus (small, c rown, short there are two lineages or groups: the B. degiulii - schoetensacki metapodials) (Van der Made, in prep.). The size changes are lineage and the B. menneri . voigtstedtensis group or lineage (Van important and occurred in the fashion of the ~ punctuated der Made, 1998, 1999a). B. menneri is a large form with slender equilibria ". The remains from TDl 0 -11 belong probably to C. e. metapodials (Sher, 1997); B. voigtstedtensis is a little smaller. priscus and those from the lower levels to C. e. acoronatus, but Remains of a large bovine from TOl 0 and TOll were assigned to the distal parts of the antlers are lacking in all levels. Bas/Bison (Van der Made, 1998). The size is more or less comparable to B. schoetensacki. TG10 and TGll have very Eucladoceros giulii similar faunas, but differ in the bovine, which is very small there Material from TOW4 and T06 was assigned to E. giulii (Van der (Van der Made, 1999b). Made, 1999) and material from T08a with some reservation; alternatively it might represent M. solilhacus (Van der Made, The rodents at Gran Dolina site 1998). New material from TOE5 belongs to E. giulii. This Mimomys savini and Arvicola species was only recently recognised (Kahlke, 1997). It now The MimomyslArvicola lineage is well known and widespread in seems the dominant large deer from the later part of the Early the European Pleistocene. Moreover the MIA transition marks Pleistocene. The form is characterize by large and slender the BiharianfToringian boundary in the European chronology. The metapodials, whereas those of M. solilhacus are robust. species M. savini is the largest vole of the rodent assemblage in Metapedial size increased with time, though general size seems 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 gregaloides and T. arvalidens are present together at numerous in young individuals and Mimomys fold. The proportion of localities in central and southern Europe dated to Cromerian 5.1. individuals with enamel islet, (in early ontogenetic stages), and (Sutcliffe and Kowalski, 1976, Rekovets & Nadachowski, with closed linea sinuosa or presence of roots, (late ontogenetic 1995). The primitive appearance of S. gregaloides of TD3-TD6 stages), are successively reduced with time. In Arvicola, this could indicate an early radiation of this species at the end of the character is absent or present vestigial at such sites as Cullar de Early Pleistocene. Baza. In young specimens, without roots, the enamel islet is present in 14% of the assemblage, and the typical Mimomys Microtus seseae enamel differentiation is not evident. The enamel islet is less First described by Gil (1997), this is a species restricted to the frequent in modern populations (Huescar 14%, West Runton G ran Oolina levels T0 3-T0 7. Our preliminary work allows us to 7%, Voigtstedt and Prezletice 0 - 1%) than in older populations propose that M. seseae is closely related to Ihe Middle of Mimomys savini (Chiscau 1 and TD6 14%). Pleistocene and extant Microtus species as M. nivaloides, M. malei, M. nivalinus and Tyrrhenicola and it may represent a Iberomys aft. huescarensis and Iberomys breccien5is second radiation of the Microtus group at the end of the Lower The extant Iberomys cabrerae is an endemic vole in the Iberian Pleistocene. Peninsula tougth fossil species of this genus (I. aft. huescarensis, I. brecciensis) are found in l ower to Middle Aflophaiomys chalinei Pleistocene Mediterranean localities in Spain, France and Ityaly The species A. chalinei is the second largest vole of the (Ventura et alii, 1998, Cuenca Besc6s ef alil~ 1999, ). This Gran Oolina section. Its size and morphology split it from the lineage probably split from the Microtus s.l. group as the rest of the microtines. The species is typical of l ower "Iberomys clade" by the end of the lower Pleistocene. The Pleistocene localities of Spain and Italy, such as Cueva Iberomys origin may be related with Allophaiomys nufiensis Victoria, Bagur 2, Casablanca 3(=Almenara 3), Castelldefells, from les Valerots and Monte Peglia. We also fo und primitive l as Cabezas, Fuentenueva 3 and Pietrafitta, The last morphotypes of Iberomys in the species Microtus hintoni and appearance of Allophaiomys chalinei in Ool ina is at the M. theniifrom Untermaf3feld and Neu leiningen 5, 15 (l ap lana ef midd le of level 6 (Cuenca Besc6s et alii, 1999, Lapiana, alii, 1999). The study of the relationships among these species 1999). Level TD 6 records the lasl app earance of the is still in progress. The autapomorphy of the lineage may be species also in Europe. defined as the asymmetry of the bucco-lingual salient angles: the ratio Lalli is always lower than th e same ratio in other Microtus aff. oeconomus lineages of Microtus (Cuenca Besc6s et alii, 1995, 1999). The This species is found exclusively in level T0 8a. Several spec;ies I, huescarensis, with advanced "nutiensis" species with the ratticepoid morphology that characterises morphology, is the first representative of the Iberomys lineage the Pallasiinus li neage are cited in several localities of late and it is found as f. aft. huescarensis in Atapuerca, in Trinchera Early Pleistocene to early Middle Pleistocene age as Kozi del Elefante, and Gran Dolina levels 3-8a (Lapiana & Cuenca Grzbiel, Hohensiilzen, West Runton, Voiglstedt, Belfia 7, Besc6s, 2000). The middle Pleistocene species I. brecciensis Holsteijn, Shamin, Colle Curti. All of the contain Mimomys appears in Oolina 8b, 10 and 11, and it is characterised by the savini faunas. advanced stage of th e trai t asymmet ry. The species I. brecciensis is also present in the Trinchera Galeria. The species Terricola atapuerquensis I. huescarensis (and relatives) is of late Early Pleistocene age The species is present in levels T08b till TO 11. It is a large and I. brecciensis characterises the early Middle Pleistocene in Terricola defined by Gil (1996). It is characterised by its large Spain. size and an terior complex. It is found also in Trinchera G aleria (Cuenca Besc6s et alii, 1999b). Stenocranius gregaloides and Terricola arvalidens Both species are more frequent in the lower levels, T03-T05 Microtus agrestis jansoni than in T06, being T. arvalidens somewhat more abundant and This species is found in T01 0, 11 and also in Trinchera Galeria one of the scarce faunas of the level T07. The differences (Cuenca Besc6s et alii, 1999). It is a species of Microtus with between S. gregaloides and T. arvalidens from Gran OoHna and agrestis or agrestoide morphology, probably related with the its represe nt ants from Middle Pleistocene localities can point to extant M. agrestis. The species M. janson; was first described different chronospecies. The holotype of S. grega/oides from by Chaline (1972) as a subspecies of M. agrestis in the early West Runton, is somewhat different from the S. gregaloides Middle Pleistocene site of La Grotte de l'Escale a Saint Esteve from T D3-TD6 in the deeper LRA5 in the West Runton Janson. The author distinguish the species from M. agrestis association. All increases from older to younger assemblages aubinensis from the late Pleistocene of Pointe du Bois a as Rekovets & Nadachowski (1995) show. The species S. 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 southeastern Europe (Kolfschoten & Turner, 1996; Rekovets & The rooted vole Pliomys is represented in Gran Oolina by two Nadachowski, 1995; Cuenca 8esc6s, this volume). species: P episcopalis in the levels T03-T06 and P.lenki at the Sus scrota and Cervus efaphus, seem to have entered T01 0-1 1. Both species are found together in several localities Europe during the late Early Pleistocene while Eucladoceros of late Early-early Middle Pleistocene age (Bartolomei et alii, giufii was already present. 1975). Its taxonomic status needs further revision. Panthera leo does not enter into Euro pe from Africa until the Early Middle Pleistocene, being first present at Isernia, placed between 783 and 500 ka (Garcia et Arsuaga, 1999). This large Conclusions size cat could have been the substitute (in ecological terms) of The MimomyslArvicola transition marks the BiharianfTo ringian the large sabretooth Homotherium latidens, previously present boundary and it was earlier coincident with the Lower/Middle in the Sierra (in level TD5E). At the Atapuerca sites, Panthera Pleistocene boundary in the continental Ouaternary biostratig­ leo remains have only been recovered from layers dated raphy. The updated chronology of the Ouaternary uses the between 500 and 200 ka. paleomagnetic reversal Matuyama/Brunhes boundary as the The Crocuta crocuta record is continuous from the lowermost Lower/Middle Pleistocene boundary and M. savini crosses this Oolina levels up to layer TOSa (just above the MatuyamalBruhnes limit up to the early Middle Pleistocene (West Runton, Voigtstedt boundary). Here is the first undubious oldest record of this taxon and Cromer faunas in general). Thus, the Late B iharian, in Europe. The coexistence of Homo antecessor and this Mimomys savini Rodent Zone includes the paleomagnetic specialized hunter-scavenger could have resulted on resource boundary Matuyama/Brunhes in Europe, such as Stranska competition and final displacement of the hyaenid from the Sierra. Skala, Grace, Shamin, Mahlis, Karlich, Atapuerca TD7 (Cuenca From the stratigraphic hiatus at TOSb, no Crocuta remains nor Besc6s et alii, 1999). TD6 is below the TD7 level which shows coprolites have been found, at the TOl l (already excavated) level the paleomagnetic Matuyama/Brunhes boundary in the Gran nor at TD l 0 (that is still being excavated). Dolina Section (Pan,s & Perez-Gonzalez, 1995, 1999). The The mammal association and the magnetochronology and Matuyama/Brunhes boundary is fixed in the late Biharian combined U'series/ESR dating methods of Pares & Perez­ (Microtus-Mimomys rodent Superzone). In this biozone, the Gonzalez (1999), Falgueres et alii (1999, van der Made, 1999, species Mimomys savini presents advanced characters as the Garcia & Arsuaga, 1999) provides confirmation of an age range loosening of roots and lack of the UMimomys islet" and this zone between 780 and 850 for t he Dolina levels 3-6. The is characterized by the absence of small Mimomys species as M. Matuyama/Bru nhes boundary at the level T07 gives an age of pusilfus or M. blanci. The assemblage and characteristics of the 780ky for this level and the dates for TD8 are between roughly species M. savini, T arvalidens, S. gregaloides indicates that 500 and 700 ky (Falgueres et alii, 1999). TD3-TD61evels are older than West Runton (type Cromerian). In Two main associations chronologically very different can be the G ran Oolina Section we can calibrate, for the first time, the detected in the Sierra de Atapuerca sites: one of a late Early evolutionary rank of those important biochronological markers Pleistocene age and another placed on the Middle Pleistocene. with magnetostratigraphy. We propose that a radiation of We can establish the relative evolutionary stage of the taxa Microtus s.I., with the appearance of the fi rst, primitive S. coming from different layers and tentatively correlate the gregaloides, T arvalidens and Iberomys took place just before Atapuerca sites among them and relate them with other the Matuyama/ B runhes boundary. Very close to this first Pleistocene Western European sites. radiation of Microtus a second took place short before the The macro- and micromammals association together with the paleomagnetic event, leading to the appearance of new species magnetochronology and combined U-series/ESR dating as M. seseae in Gran Oolina and other species of the Microtus methods, provides confirmation of an age range between 7S0 and group in Italy (i.e. M. arvalis, M. multiplex-subterraneus of Selva 850 ka for the Dolina levels 3-6 (Cuenca Besc6s et alii, 1999; Vecchia, Fontignano, Sardelia et alii, 1998) or the latest Early Falgueres et alii, 1999; Garcia et Arsuaga, 1999; Van der Made Pleistocene faunas of Eastern Europe with the first appearance 1999; Pares et Perez-Gonzalez, 1999). The dates for TD8a are of the M. gr. oeconomus (Markova, 1998). between roughly 500 and 700 ka (Falgueres et alii, 1999). The last appearance of Allophaiomys chalineiin Oolina is at the The lower levels of G ran Oolina (T03 to TOSa) may represent middle of level 6. This species is typical of other Early Pleistocene the climatically unstable period of the end of the Lower localities of Spain and Italy, such as Cueva Victoria, Bagur 2, Pleistocene. Arid, open country mammals as Marmota and a Casablanca 3 (=Almenara 3) , Castelidefeli s, Las Cabezas, high diversity of arvicolids are founded in T05. The warmer and Fuentenueva 3 and Pietrafitta. Their age range between about relatively wooded country may be represented in T04 and T06 1.6 (Bagur 2) to 0.8 Ma (Gran Dolina 6) (Lapiana, 2000, Cuenca with a diversified soricidae fauna as well as the presence of Besc6s this volume). The species M. savini survive beyond the Hystrix and Castor. In the level TD8a, the presence of Hystflx Early to Middle Pleistocene boundary and can also be found in as well as Hippopotamus and the low diversity of voles may early Middle Pleistocene localities in central, north and indicate a relatively warm period. Levels TOSb and TO 10, TO 11 1' 111 MI II( IllunWI lin IlIhlUfilOIIl II [,"thana AgUirro. Paleontologia

- j LAPLANA CONESA, c.; CANUOO, J.I. & A RSUAGA, J.l. (1997): ·Small mammal:. may represent a longer and complex period with alternating from Sima de los Huesos·. Journal of Human evo/ullOnt 33: 175-190. climate between warmer and cooler conditions (Van der Made, -: CANUOO, J.1. Y LAPLANA, C. (19 9 9) : "A nalisis bioestratigr8lico de lot: 1998, Cuenca Besc6s et alii, 1999). This is coherent with the roedores del Plelstoceno Med io del yacimienl o de G alerla (S ierra do O xigen Isotope Stages (O IS): levels T01 0, 11 are correlated Atapuerca, B u r gos)~. Memorias. Arque% gfa en Castilla y Leon, 7, 1B9·2 1O . with OI S 9 to 11 , T06 is correlated with the OIS 19 or 21 -; LAPLANA, C. & CANUDO. J.1. (1999): "B iochronological implications of tho Arvicolidae (Rodentia, Mammaha) from tho Lower Pleistocene hominid (Falgueres et alii, 1999) and T05 may be correlated with the bearing level of Gran Dolina 6 (TD6, Atapuerca, Spain)". Journal of HUn)IIf) OIS 22 and TD4 with the OIS 23. Evolulion, 37(3·4): 353·373.

CI1AlINE. J. (1972): ~ Les rongueurs du Pleistocene moyen et superteUf du France". Ca h. Paleont. 41 Opp.

Aknowledgements 0 1 STEFANO, G . & PETRONIO, C. (1993): ~A new Cervus e/apllus subspeclOll The Atapuerca crew (represented by its directors Juan Luis Arsuaga, Jose Maria of Middle Pleistocene age". Neues Jahfbllch fUr Geologie und Paliionlologie, Bermudez de Castro and Eudald Carbonell) helped us in diUerent ways Wi th the Abhandlungen, 190: 1-18. ext raction of the fossil materiats during the Alap uerca field season every. FALGUERES, C.; B AHAIN. J.J.; YOKOYAMA, Y.; A RSU .... G ... , J.l.: B ERMUOEZ OE Research has partially supported by Consejeria de EducaCl6n y Gultura 01 Junta CASTRO. J. M .; CARBONELL. E.: B ISCHOFf. j ,l. & DOLO , J. M . (1999)' de C astilla y Leon, Direcci6n General de Investi9aci6n Cienllfica y Tecnica "Eafliest human s in Europe: the age of TD6 Gran Oolina, Atapuerca, Spain", (P B93·0066·C03, PB96-00 ·COI , PB 9 S'1 0 2 S-C0 3·02, PB96-00-C 03), J. Hum Evol .. 37{3 ·4): 343·52. Ministerio de Educaci6n (M EC, HA-97-22) and "Unidad Asociada" program. Th e FAURE, M . (19 83): Les Hippopotamidae (Mammalia. Artiodactyla) d'Europu follOWing students helped us in different ways: Cesar Lapiana. Jose Ignacio occidentale. TMse, 3' cycle Paleontologie. Univ. Claude Bernard . Lyon " Canudo, Beatriz Romero. SergiO B ajo, Raquel L6pez, Carlos GarCia, Toni Canals, 1367: 233 pp. M. Dolores Garcia-Ant6n and Josetina Barreifos. & C. G UERIN , ( 19B4): ·Sus strozzi; at Sus scro!a. d eux mammifcrOfl artiod acty las marq uers des p aleoenvjfonnaments". Pa laeogeography, Bibl10g raphy Palaeoclimatology, Palaeocecology, 4H: 21 5-228.

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