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During the Late Pleistocene, the Balkans came to be an important region with many isolated areas, enabling fauna, alongside Neanderthals, to thrive in the area. This work is focused on paleoenvironmental and paleoclimatic changes that occurred in the northern Balkan Peninsula with a special focus on fossil record from Smolućka cave aging from MIS 5 to MIS 3. Based on available data, an attempt has been made to establish a synthetic chronological context for the faunal assemblages recovered from Smolućka cave. Tentative attribution of layer 5 to MIS 5 relies on the interstadial pattern of our reconstructions, with favorable climate conditions for a large diversity of species and with mean annual temperature (MAT) reaching up to 3 °C higher than present values and abundant rainfall in the area. Cold conditions for MIS 4 are not present in Smolućka, although layer 4 (late MIS 5 or early MIS 4) shows somewhat dryer and cooler climate when compared with other layers. Layers 3z and 3 (MIS 3) are characterized by temperate and humid conditions together with a complex system of mosaic habitats with high environmental heterogeny but generally favorable conditions for a large diversity of life. Although still not fully chronologically constrained, Smolućka cave presents an interesting opportunity for future research, in the time period when only Neanderthals occupied the Balkans and thrived in local conditions. The transition between the Neanderthals and anatomically modern humans has not yet been established but can be expected in future research.

Los Batanes C4 is a cave-site in the Spanish Pyrenees, the minimum sediment calendar age was determined to be 15,234 ± 223 cal BP by radiocarbon dating. The cave opens on the northern bank of an eastern tributary of the River Gallego, at an altitude of 1025 m. The small vertebrates recorded are mainly bats, in order of abundance Rhinolophus euryale, R. ferrumequinum, Myotis sp. and Miniopterus schreibersii. The association suggests that at the time of the accumulation the climate was reasonably similar to the current climate; we situate it within a period of local retreat of the ice-cover in the Gallego Valley during the Lateglacial. This is the highest record of these species of Rhinolophus in the Iberian Quaternary, showing that the R. euryale altitudinal range was similar to its extant range during the favourable periods of the Lateglacial. This could indicate that this taxon was relatively quick in spreading into higher regions whenever climate conditions allowed it.

Grotta Maggiore di San Bernardino, located at an altitude of 135 m a.s.l. in the Berici Hills in northeastern Italy, is an archaeological site with a discontinuous sedimentary sequence dating from Marine Isotope Stage 7 (MIS 7) to MIS 3. In this paper we present for the first time a palaeoenvironmental and palaeoclimatic reconstruction of the sequence based on small-mammal (insectivore, bat and rodent) assemblages. Coupled with biochronological data and absolute dating together with previous studies on large mammals, birds and other studies on small mammals and pollen from comparable time-spans in Italy, the results enable us clearly to identify distinct climatic periods: the end of MIS 7 (7c to 7a) in units VIII-VII, MIS 5d in unit V, and probably MIS 5b in unit IV and an indeterminate MIS 3 interstadial in units III-II. Finally, the study shows that the early Middle Palaeolithic human occupation in Italy occurs during mild and temperate sub-stages of MIS 7 and that human groups with the same techno-cultural background (Mousterian) were well adapted to the changing environmental and climatic conditions of the Middle to Late Pleistocene in this part of southern Europe.

We have analyzed the palaeoenvironment and palaeoclimate of the beginning of the Holocene in the Iberian Peninsula on the basis of the small-mammal assemblages from three sites within the geography of the Iberian Peninsula: Mirador (Sierra de Atapuerca, Burgos), Colomera (Sant Esteve de la Sarga, Lleida) and Valdavara-1 (Becerrea, Lugo). These associations reveal that the palaeoenvironment was more humid than today in the sites under study, and the landscape was dominated by woodland and woodland margins in all the studied layers. Further, the climatic conditions were stable, but with winters colder than at present, above all in the Mediterranean area. Finally, our data have been compared with other environmental and climatic proxies, showing that human activities exerted little impact on the palaeoenvironmental conditions that occurred from 7000 to 3000 BP in the Iberian Peninsula.

Cova Eiros is emerging as a reference site in the northwestern Iberian Peninsula for the study of the development of the last Neanderthal populations and the first populations of Anatomically Modern Humans (AMH) in MIS 3. Cova Eiros is an archaeological site (with Middle and Upper Palaeolithic levels) located in Cancelo, Triacastela (Lugo, northwestern Iberian Peninsula), which has been systematically excavated from 2008 onwards. The small-vertebrate assemblage analysed came from the archaeo-palaeontological field seasons that took place from 2009 to 2014. At least 18 small-vertebrate taxa have been identified: 1 frog ( Rana temporaria ), 1 snake ( Vipera sp.), 4 insectivores ( Sorex minutus , Sorex sp., Talpa cf. occidentalis and Erinaceus europaeus ), 4 chiropters ( Myotis myotis/blythii , cf. Miniopterus sp., Myotis sp. and Rhinolophus ferrumequinum ) and 8 rodents ( Apodemus sylvaticus , Arvicola amphibius , Arvicola sapidus , Chionomys nivalis , Microtus (Terricola) lusitanicus , Microtus agrestis , Microtus arvalis and Microtus oeconomus ). Using the Habitat Weighting method to reconstruct the palaeoenvironment, we reconstruct a landscape for MIS 3 characterized by open woodland formations. The Mutual Ecogeographic Range (MER) method and the Bioclimatic Model (BM) used for the palaeoclimatic reconstruction show lower temperatures and higher precipitation than at present in the region. Our results from Cova Eiros are compared with the data obtained from several other sites in the Iberian Peninsula; it can be said that Neanderthals and AMH were well adapted to the territory that they occupied, as well as to the surrounding environment and the climatic conditions prevalent in the unstable context of MIS 3 in the Iberian Peninsula.

The fossil bat assemblage from the Lower Red Unit of Sima del Elefante (TELRU) in Atapuerca (Burgos, Spain) has been exhaustively analysed for the first time. Bat fossil assemblages are of particular relevance to palaeoenvironmental approaches to sites. Here we integrate our new data on the chiropteran fauna with the data provided previously by other authors on the basis of the small-vertebrate assemblages and the palynology of the site. Our results are consistent with the earlier results in that they indicate a generally warmer climate than at present in the area and stable environmental conditions throughout the major part of the TELRU sequence. However, fossil bat assemblages sometimes lead to problems when they are used in landscape reconstruction. These problems are discussed here and should be taken into account in future works.

Bats (Chiroptera, Mammalia) are a valuable and independent source of palaeoenvironmental information in Quaternary sites because bat assemblages are not necessarily produced by the same processes as those of other small mammals. Here, we report a small collection of bats (MNI 15) from a cave infill of the Aguilon P7 (AGP-7) site (Zaragoza, Spain) dated Late Pleistocene (MIS 3). This is one of the rare localities south of the Ebro River (right bank) with Neanderthal tracks; therefore, the palaeoenvironmental data provided here may help us to understand Neanderthal history. Nine bat taxa were identified, viz. Rhinolophus ferumequinum, R. hipposideros, Myotis gr. myotis/blythii, M. cf. bechsteinii, M. emarginatus, M. daubentonii, Plecotus gr. auritus/austriacus, Hipsugo savii vel. Pipistrellus kuhlii/nathusii, and Miniopterus schreibersii. The presence of a high number of yearlings of the former four species suggests the presence of breeding colonies. The surprisingly high diversity of the sample indicates a variegated vegetation cover and relatively high humidity for the MIS 3 stage in the region.

La Cansaladeta is a site located in a narrow passage dug out by the Francoli River in the Pre-littoral Catalan range (NE Iberia). The archaeological deposit lies on top of a +45–50 m fluvial terrace, and contains a rich Middle Pleistocene stratigraphic succession in which 10 archaeological layers have been identified. Here, we present the description of the sedimentary succession of the site, the available chronological framework (including TL, TT-OSL, and ESR/U series ages, as well as biostratigraphical information), the characteristics of the lithotechnical and faunal assemblages, as well as the first interpretation of the human occupations at the site. Three main occupation phases have been differentiated. The oldest archaeological layers are embedded in a palaeosol formed on a clayey layer that contains limestone fragments detached from the shelter's wall and roof. During the second phase, low energy fluvial sediments coming from lateral river floods were deposited and sealed the archaeological layers. After a transitional phase, the upper layers appear to be derived from slope sediments. Numerical dating yielded internally consistent dates of 380–400 ka for the sequence. However, the study of the fossil assemblages potentially suggest a tentative early Middle Pleistocene age for the lowermost horizons. The rich technological record is made up of several local raw materials, especially chert but also schist, quartzite, quartz and other minor rocks. Knapping sequences seem to be autochthonous, as all the elements and size-categories of the reduction sequence are present, and refits are common. Techno-typological features observed along the sequence are not very diagnostic, however the lower layers include some Acheulean forms among the large shaped tools, restricted variability among the small retouched tools (mostly denticulates) and a virtually absence of prepared cores. The faunal assemblage is scarce and appears strongly altered. Only the lower layers have provided fruitful information. Although neither combustion areas nor structures related to fire-use have been documented to date, heat damage has been repeatedly recorded on both lithic and faunal remains. This may point to systematic fire use within the archaeological sequence. The data provided by the ongoing excavations at La Cansaladeta provide new insight into the early human settlement history of northeast Iberia to be reconstructed. Combining these data with those coming from the close site of Barranc de la Boella, enable us to better reconstruct the timing, nature and context of early human occupation in the region.

No ideal method can be used alone to provide an accurate and irrefutable chronology for dating European Early Pleistocene localities. The solution lies in the combination of different, but complementary, approaches for complementing their chronological framework. In that regard, the results of the present study show that biochronology may be a useful additional tool to further refine the chronology of these localities, by providing not only relative chronological information between the sites, but also giving some important indications about a younger or older Jaramillo age. Our study presents the revised faunal list of each Early Pleistocene levels from the localities of Gran Dolina (TD) and Sima del Elefante (TE), in Atapuerca. Both localities have very likely the richest associations of small mammal species in the Quaternary of Europe. Perhaps the most striking observation is the lack of similarities between the assemblages from each site, which can only be explained by a chronological gap between them. Although the Jaramillo subchron has not been directly identified at Atapuerca, the existing chronostratigraphical framework combined with similarities of the faunal assemblages between Gran Dolina and Vallparadis indicate with some confidence that Sima del Elefante levels are older than the Jaramillo subchron. Consequently, the Early Pleistocene levels of both Atapuerca sites TE and TD are the only sequence from Spain that allows the study of a continuous stratigraphic succession of the interval around the Jaramillo subchron in terrestrial sequences. The biostratigraphy of the Early Pleistocene from Europe, based mainly in the few isolated localities of the Olduvai–Jaramillo time interval, is reinforced by the faunal succession of the Atapuerca sites.

The chronology of layer EVT7 of the Vallparadis Estacio section has been a hotly debated topic during recent years, with some authors supporting an age close to the Early-Middle Pleistocene boundary (0.83 Ma), others favouring an age older than the Jaramillo magnetostratigraphic subchron (ca. 1.2 Ma) and others proposing a chronology between 0.98 and 0.95 Ma for layer EVT7. Here, we review the biochronological data from EVT7, and review the biochronology based on the study of Mimomys savini using all the available remains of this species, which is the most abundant rodent species from this layer. Based on the morphologic and biometric comparison with remains of the same species from other similarly-aged Iberian localities (Gran Dolina TD4-6 from Atapuerca, and Barranco Leon D and Fuente Nueva 3 from the Guadix-Baza Basin), we propose an estimated age for EVT7 in the chronologic range from 0.99 to 0.94 Ma. We further perform a paleoenvironmental study based on the whole small-mammal assemblage from EVT7, which allows us to infer an open and humid environment in the vicinity of a lacustrine area or fluvial course. Recently published work also studied the rodent assemblage from EVT7. However, in this study only a part of the small mammals recovered from EVT7 was studied, and therefore its results can be partially biased.

In this paper, a biozonation of the Pleistocene continental record of the Guadix-Baza Basin is proposed. This biozonation is based on the small mammal succession, which ranges from the earliest Pleistocene (ca. 2.6 Ma) to the Middle Pleistocene (ca. 0.3 Ma). A total of 9 biozones have been recognized, all based on the range or concurrent-range of arvicolid species. Therefore, seven biozones have been defined for the Early Pleistocene: Kislangia gusii Zone, Mimomys cf. reidi Zone, Mimomys (Tcharinomy)s oswaldoreigi Zone, Allophaiomys ruffoi Zone, Allophaiomys aff. lavocati Zone, Iberomys huescarensis Zone and Terricola arvalidens Zone. In the Middle Pleistocene of the basin two biozones have been recognized: Iberomys brecciensis–Arvicola mosbachensis Zone and Iberomys brecciensis-Arvicola aff. sapidus Zone. According to different dating methods, the duration of each Early Pleistocene biozone ranges between 0.4 and 0.1 My, with increasing resolution from the Earliest Pleistocene to the late Early Pleistocene. The small mammal succession reveals a high degree of endemism and a persistent Eastern Mediterranean influence, as opposed to Central and Eastern Europe. However, correlation with other Iberian and European sites has been possible on the basis of each biozone associated fauna.

The early Pleistocene site of Barranco León (Guadix-Baza Basin, SE Spain), dated to 1.4 Ma (millions of years ago), records the oldest hominin occurrence in Western Europe, as evidenced by the discovery of one tooth and thousands of Mode 1 stone tools. In this paper a detailed analysis of the microvertebrate content of the D1 and D2 units from this site is presented. The early Pleistocene in the Guadix-Baza Basin is characterized by a sharp climatic deterioration, which possibly impeded the settlement of this region by the early hominin population from the southern Caucasus. Shortly afterwards, when the climatic conditions were again favorable, a hominin presence is suddenly evidenced at the units D1 and D2 of Barranco León. According to the microvertebrate analysis of these units, the mean annual temperature at the time of deposition was significantly higher than 13 °C, with prevalent humid conditions. However, although most of the species were inhabitants of water edges, an open landscape was present in the vicinity of the lake. The data reported here clearly support the idea that the early hominin occupation of Europe was strongly constrained by climatic and environmental conditions, rather than by physiography or cultural factors.

L'Arbreda Cave is one of the southernmost of the Reclau Caves (Serinya, Girona, northeastern Iberian Peninsula), located between the west margin of the Pla d'Usall and the Serinyadell river, at an altitude of about 211 m a.s.l. The Pla d'Usall forms part of the Banyoles-Besald Basin, which is bounded by Eocene and Neogene reliefs. It is filled with sediments of Plio-Pleistocene age. The cave is a shelter formed by travertine cascades with a degree of karstification, and contains sedimentary deposits ranging from the end of the Middle Pleistocene to the Early Holocene, thus constituting an important archaeo-palaeontological record for an understanding of the prehistoric occupation of northeastern Iberia. It is one of the most complete and best-dated terrestrial sequences in Iberia from the Mousterian to the Early Holocene period. In this paper we present a palaeoenvironmental and palaeoclimatic reconstruction of the sequence based on non-flying small mammals (insectivores and rodents) and pollen analyses. The environmental and climatic results, together with the absolute dating of the sequence and direct comparison with NorthGRIP delta O-18 and the sea-surface temperatures (SST) of the Alboran Sea, allow us to identify various climatic periods within our sequence: a cold period between interstadials 12 and 10 in Level I, probably interstadial 10 in Level H-2 the Last Glacial Maximum (LGM) in Levels C and B, the Bolling-Allered interstadial (B/A) in Level A, and the Preboreal/Boreal (Pr/B) in the Terra Rossa Level We want to thank the editor Dr. Paul Hesse and the three reviewers (Prof. Gloria Cuenca-Bescos and two anonymous) for their comments that improve the final version of the manuscript. This paper is part of the projects HAR2013-48784-C3-2-P, CGL2012-38358 and 2014-SGR900. Excavations ara funded by "Conselleria de Cultura Generalitat de Catalunya". J.M. Lopez-Garcia is a beneficiary of a Beatriu de Pinos postdoctoral fellowship (2011BP-A200001) from the Generalitat de Catalunya, a grant co-funded by the European Union through the Marie Curie Actions of the 7th Framework Program for R + D

Daura, J. et al, © 2015 Elsevier Ltd. The Cova del Rinoceront, a site in NE Iberia, contains a thick sedimentary fill preserving a faunal archive from the penultimate glacial and the the last interglacial periods. Layers I to III have been dated to between 74 and 147ka, coinciding with MIS 5a to 5e, a period poorly represented in the Mediterranean terrestrial record. The results from Cova del Rinoceront are of broader interest for the reconstruction of ecological dynamics during warm stages and the understanding of the evolution and geographical variation of several taxa. The palaeoecological evidence suggests a landscape dominated by mixed wooded vegetation with mild climatic conditions, slightly more humid than today. Several vertebrate taxa, including Haploidoceros mediterraneus, Stephanorhinus hundsheimensis and Glis glis, are documented for the first time in the early Upper Pleistocene of Europe, showing that these species persisted across the region for longer than previously thought. In addition, the recovery of a small lithic assemblage indicates human presence in the surroundings of the site. The 11m-thick stratigraphic section also provides an ideal setting in which to compare several geochronological methods. U-Th dating of the flowstones that cap the deposit, of speleothems formed along the cave walls, and of speleothems buried by the deposit at different elevations provides minimum and maximum ages of 74 and 175ka, respectively, for the accumulation. The ages obtained by luminescence, electron spin resonance (ESR), amino acid racemisation (AAR), palaeomagnetism and U-series dating of bone are in good agreement with each other and are stratigraphically consistent. This well-dated faunal succession presents a unique opportunity to assess changes in the Pleistocene fauna of the Mediterranean coast over an interval of more than 100ka., This paper is an output of the archaeological research project El Plistocé Superior a la costa central catalana: paleoambients i ocupació dels neandertals (2014/100639) sponsored by Servei d'Arqueologia i Paleontologia e Generalitat de Catalunya (G.C.) and Castelldefels City Council. Sampling and laboratory analyses were funded by Catalan (2014SGR-00108) and Spanish (HAR2011-26193, HAR2011-26193, CGL2010-18616, CGL2012-38358, CGL2012-38434-C03-03 and CGL2013-48441-P) MICINN Projects. J. Daura has been supported by a postdoctoral grant (Juan de la Cierva Subprogram JCI-2011-09543), J.M. López-García by a postdoctoral fellowship (2011BP-A00272) from the G.C. a grant co-funded by the European Union through the Marie Curie Actions of the 7th Framework Program for R.D. We are also grateful to R. Alvarez (UB) for lithics'photos. J. Mangope, Williams College, prepared and measured the ESR sample. D. Lawrence, Wadsworth Laboratories of the New York State Public Health Service provided access to the source for the ESR irradiations. Funding for the ESR spectrometer came from NSF grant ILI 9151111; other funding was provided by Williams College. L.Arnold and M. Demuro were supported by MINECO grant G2010- 16821, Australian Research Council Future Fellowship F130100195, and Marie Curie International Reintegration grant PIRG08-GA-2010-276810. Thanks to the Paleomagnetism Laboratory of Barcelona (CCiTUB-CSIC) where the paleomagnetic analyses were carried out.

For the first time, the fossil herpetofauna from the Middle and Late Pleistocene of Scladina and Sous-Saint-Paul caves (Sclayn, Belgium) is described. The amphibians and squamate reptiles are represented by one salamander (Salamandra salamandra), three anurans (Pelodytes punctatus, Bufo bufo and Rana temporaria), two lizards (Lacerta cf. agilis and Anguis fragilis) and two snakes (Zamenis longissimus and Vipera cf. berus). The occurrence of the Parsley Frog (Pelodytes punctatus) and the Aesculapian Snake (Zamenis longissimus), which are not currently represented in Belgium, is of particular interest. Scladina also represents one of the northernmost fossil mentions for the Fire Salamander (Salamandra salamandra) although it is within its current distribution in Europe. Finally, the presence of the Adder (Vipera cf. berus) is very probably attested in Scladina whereas today this snake is infrequent and classified as endangered in Belgium.

In 2007, remains of Mammuthus meridionalis and 334 lithic artefacts were recovered in level 2 of Barranc de la Boella. The presence of M. meridionalis allows us to assign an age older than 700 ka to this level, which represents one of the few sites of this age with palaeontological remains associated with lithic industry. It also enables us to shed light on questions pertaining to the chronology and the biogeography of the earliest European human settlements. In this paper, the small mammals from the Barranc de la Boella site are described for the first time, with special emphasis on the arvicolines. The presence of Victoriamys chalinei and Mimomys savini allows us to estimate the age of level 2. Level 2 of the Barranc de la Boella site ranges between the top of the Jaramillo subchron (0.99 Ma) and shortly after the Brunhes–Matuyama transition at 0.78 Ma, thus covering a time span of great importance for our knowledge of the earliest human occupations in Western Europe.

The habitat weighting method, the Simpson diversity index and the mutual climatic range method are applied to the small-mammal assemblage of Grotta del Romito (southern Italy) in order to reconstruct the environmental and climatic fluctuations that occurred during the latest Pleistocene–Holocene sequence of the cave. The analysed strata have a discontinuous chronological range from the middle-late Gravettian (ca. 24 ka uncal. BP) to the Mesolithic–Sauveterrian (still radiometrically undated). Gravettian and Epigravettian layers were excavated inside the cave, Mesolithic layers in the rockshelter outside the cave. The small-mammal assemblage shows significant differences along the sequence; a turning point is detected in strata D29-D11, coinciding with the end of Heinrich Event 1 (H1) and the beginning of the Bolling–Allerod interstadial (between ca. 14.9–14 ka cal. BP). These data are in accordance with the previously obtained environmental data for Grotta del Romito, as well as with other continental climatic and environmental changes detected at a regional scale.

Marine Isotope Stage 7 (MIS 7) is an interglacial that has not been a major topic of scientific interest, yet is interesting because it contains five warm and cold phases (MIS 7e-a) within a brief period of time (ca. 246-186 ka). In this context, there are several sites that contain Middle Pleistocene human remains in Europe, including Mollet cave, the subject of this study. Analyses of the small vertebrates from this cave, together with the numerical dating (ca. 215 ka), established the environmental and climatic conditions in which the hominins who frequented the cave lived. The results obtained situate the Middle Pleistocene hominins of Mollet cave in the MIS 7c substage, indicating that the occupation by humans took place in a transitional phase from a warm to a cold stage with forest development. © 2012 Elsevier Ltd and INQUA., This paper is part of projects CGL2012-38358 and SGR2009-324. This paper was partially financed by the ministerial projects HAR2010-22013, BOS2003-8938, CGL2006-13532-C03-01, CGL2006-4548/BTE, and the Catalan Ministry of Culture. J.M. López-García is beneficiary of a Beatriu de Pinós postdoctoral fellowship (2011BP-A00272) from Generalitat de Catalunya, a grant co-funded by the European Union through the Marie Curie Actions of the 7th Framework Program for R þ D.

J.M.L.-G. is a beneficiary of a Beatriu de Pinos postdoctoral fellowship (2011 BP-A00272) from the Generalitat de Catalunya, a grant cofunded by the European Union through the Marie Curie Actions of the 7th Framework Program for R + D. J.I.M. was supported by a predoctoral grant (FI) from AGAUR (Generalitat de Catalunya) cofunded by the European Social Fund. This paper is part of projects CGL2012-38358 and SGR2009-324

Over the course of their evolutionary history, some species of mammals have undergone variations in size, with a general trend towards increased morphometry. This effect can be seen very clearly in the fossil record of rodents because their high rate of reproduction that generates a high fossil record, which allows this phenomenon to be studied in detail. Furthermore, the rapid geographic distribution of rodents means that their evolution can be studied on a continental scale. If a relationship can be established between the size of individuals and their chronology, and the trend that governs a species' increase in size can be determined, then the chronologies of different sites can be estimated based on the size of the individuals of that species recovered at those particular sites. The correlation between morphometric data of micromammal fossils and age was already used by other authors. This article studies the rate at which the length of the first lower molar (m1) of Mimomys savini (a species of Palearctic arvicoline present in Europe between approximately 1.8 and 0.6 Ma ago) increased over the course of its evolution in Iberian Peninsula (south-western Europe). Because this increase in length occurred at a constant rate, a direct relationship can be established between average length of m1 and chronology, which allows us to set numerical dates to Pleistocene sites containing Mimomys savini remains.

Acknowledgements go first of all to the Atapuerca Foundation, the Institute of Vertebrate Paleontology and Paleoanthropology (IVPP) of the Chinese Academy of Sciences and the MOE Key Laboratory of Contemporary Anthropology, School of Life Sciences of Fudan University in collaboration with the Spanish Pavilion of the Shanghai Universal Expo 2010, which organized the Shanghai conference “First Human Settlements in Eurasia”. Over the last 20 years the Atapuerca excavation team has helped with the extraction of small mammals. The study and conservation of the fossils is supported by the University of Zaragoza, the Junta de Castilla y Leon, the MICINN-CGL2009-12703-C03-03project, the Fundacion Atapuerca, the Fundacion Duques de Soria, and Grupos Consolidados del Gobierno de Aragon. The SEM photographs were taken by us with the help of the SEM Service of the University of Zaragoza. J.R. (JCI-2010-06148) and J.M.L-G (JCI-2009-04026) have been supported by a postdoctoral grant from the Juan de la Cierva Subprogram, V.S.L (B019/09) and R.R.G (B113/2007) FPI program of the Gobierno de Aragon.

Hominin remains unearthed from level TD6-2 of Gran Dolina have been dated to slightly more than 0.8 Ma. Owing to the importance of this discovery, every effort is being made to reconstruct the environment where these hominins once lived. In this paper, the mutual climatic range and habitat weighting methods are applied to the herpetofaunal assemblages in order to estimate climatic and environmental parameters. TD6-2 is thus characterized by a warm and humid climate. Mean annual temperature is estimated at 2.1°C higher than at present in Burgos, with a greater increase in temperature during summer (+3.1°C) than during winter (+1.4°C). Rainfall was more abundant than at present (+407.2 mm), occurring as at present principally during spring and autumn. Humid meadows and riparian woody habitats were common. The association between warm climatic conditions and the not fully developed forest suggests that the several episodes of high-intensity occupation in TD6-2 may correspond to a recolonization by Homo antecessor of the high inland plateaus of the Iberian Peninsula from the Mediterranean seashore, using the Ebro valley as a savannah-type corridor after a cold glacial period. Copyright © 2013 John Wiley & Sons, Ltd.

The study of small mammals may provide important chronological and paleoenvironmental information in the archaeological context of a particular geographic region. Few specific studies have been conducted on fossil small mammals in North Africa, especially for the late Quaternary period. However, the Holocene site of the Guenfouda Cave, situated in the region of Oujda (Eastern Morocco), provides new data on the chronology and the paleoecology of this period. The small-mammal association from Guenfouda comprises at least 12 taxa: two soricomorphs (Crocidura russula and C. whitakeri), one macroscelian (Elephantulus rozeti), two bats (Myotis blythii and Rhinolophus hipposideros) and seven rodents (Mus spretus, Meriones shawii, Apodemus sylvaticus, Dipodillus campestris, Eliomys sp., Atlantoxerus getulus and Ctenodactylus sp.). The absence in our assemblage of species recently introduced to Morocco, such as Mus musculus and Rattus spp. (R. rattus and R. norvegicus), allows us to establish relative data for the...

DOI: 10.1016/j.palaeo.2017.01.019 URL: http://www.sciencedirect.com/science/article/pii/S0031018217300500 Filiació URV: SI Memòria El Mirador is a cave in the Sierra de Atapuerca (northern Iberian Peninsula) that contains 27 archaeological layers from the Lateglacial to the Late Holocene. A total of 4436 small-mammal remains have been analysed from these layers, and 19 taxa have been identified (three insectivores, seven chiropters and nine rodents). The palaeoenvironmental reconstruction based on a small-mammal analysis suggests that the entire sequence is dominated by a woodland landscape. Our climatic analysis characterises the climate in terms of an evolution from a cool and arid period in the Pleistocene layers (16,000 to 14,000 cal yr BP), probably related to Heinrich Event 1, to humid conditions and temperatures similar to nowadays in the Holocene layers in general. In MIR23 and MIR22 (7300 to 6800 cal yr BP) we detect an increase in temperature to levels above current values and an important decrease in rainfall. These climatic characteristics could be related to the end of the African Humid Period. Moreover, a short, slightly cooler event is registered in MIR11 (ca. 6300 to 5900 cal yr BP), coinciding with a change in the economic pattern in El Mirador. The results obtained on the basis of small-mammal studies are compared with multiproxy terrestrial data (pollen, charcoal, phytoliths, geochemistry, large mammals) and the Greenland ice-core record, as well as various other core records closer to the coasts of the Iberian Peninsula (MD95-2042, MD99-2331, MD95-2043 and SMP02-3). These comparisons show the agreement of the palaeoenvironmental and palaeoclimatic results with all the previous multiproxy and core-record data.

Kaldar Cave is a key archaeological site that provides evidence of the Middle to Upper Palaeolithic transition in Iran. Excavations at the site in 2014¿2015 led to the discovery of cultural remains generally associated with anatomically modern humans (AMHs) and evidence of a probable Neanderthalmade industry in the basal layers. Attempts have been made to establish a chronology for the site. These include four thermoluminescence (TL) dates for Layer 4, ranging from 23,100 ± 3300 to 29,400 ± 2300 BP, and three AMS radiocarbon dates from charcoal samples belonging to the lower part of the same layer, yielding ages of 38,650¿36,750 cal BP, 44,200¿42,350 cal BP, and 54,400¿46,050 cal BP (all at the 95.4% confidence level). Kaldar Cave is the first well-stratified Late Palaeolithic locality to be excavated in the Zagros which is one of the earliest sites with cultural materials attributed to early AMHs in western Asia. It also offers an opportunity to study the technological differences between the Mousterian and the first Upper Palaeolithic lithic technologies as well as the human behaviour in the region. In this study, we present a detailed description of the newly excavated stratigraphy, quantified results from the lithic assemblages, preliminary faunal remains analyses, geochronologic data, taphonomic aspects, and an interpretation of the regional paleoenvironment., J. van der Made contributed to the project CGL2015-65387-C3-3-P (MINECO/FEDER).

In this paper are described the arvicolines from El Chaparral site (Villaluenga del Rosario, Cádiz, Southwestern Spain). Due to their wide geographical distributions and rapid evolutionary rates, arvicolines are especially useful for biostratigraphical purposes. The stratigraphic layers of El Chaparral have yielded arvicolines remains of the genera Allophaiomys, Iberomys, Pliomys, and Terricola. The presence of these rodents together the other vertebrates recovered allows to determinate the age of the El Chaparral site and compare our data with other Iberian Pleistocene sites, such as of the Sierra de Atapuerca. El Chaparral site ranges between the Jaramillo Subchron (1.07-0.99 Ma) and shortly after the Brunhes-Matuyama transition at 0.78 Ma covering a important time span for the knowledge of the earliest humans occupations in western Europe. © 2012 Taylor & Francis Group, LLC.

In the Mediterranean area, which is climatically stressed by limited water resources and extremes of heat, climate variations are known to play a crucial role in the ecosystems and environment. Investigating how climate has changed in the past may help us to understand how it may change in the future and its consequences on temperature and water resources. The Gran Dolina sequence (north-central Spain) provides a unique long paleontological and archaeological record spanning the Mid-Brunhes (ca. 450 ka) climatic transition. A fossil amphibian- and squamate-based reconstruction of temperature and precipitation shows marked peaks that have been related to various interglacial peaks in accordance with numeric dates and paleomagnetic and biochronological data. An analysis of climate and herpetofaunal assemblage changes during the interglacial periods reveals that (1) post-Mid-Brunhes Event (MBE) interglacials were warmer than pre-MBE interglacials, (2) pre-MBE interglacials were warmer than present day, and (3) there were lower levels of rainfall in post-MBE interglacials than in pre-MBE interglacials. The climate trend in the Mediterranean area was found to be congruous with global climate changes as reconstructed from ice and sea-surface temperature records over the past million years.

According to pollen analysis and the 18 O-isotope curve, the first part of the Late Pleistocene (ca 128–30 ka) is mainly characterized by a dynamic that alternates cold phases (Heinrich Events) and temperate phases (interstadials). These rapid fluctuations provide the context for the Neanderthal occupations in the northeastern part of the Iberian Peninsula. In this paper we present the chronological, environmental and climatic data obtained by analyzing the pollen, the charcoal, the small vertebrates (amphibians, squamates and small mammals) and the large-mammal dental wear at the Neanderthal site of Teixoneres cave, Northeastern Iberia. Levels II and III from this cavity have provided Mousterian industries and other evidence of Neanderthal occupations, such as cut-marks in large-mammal bones. A multiproxy study such as this constitutes a new approach to the chronological, environmental and climatic context in which Neanderthal populations lived in southwestern Europe (Iberian Peninsula). The results allow us to establish a relative chronology for these two levels of between ca 30–90 ka and show that they are associated with different environmental and climatic conditions: temperate and humid for Level III and cold and dry for Level II. This demonstrates that the Neanderthals were well adapted to the territory that they occupied, irrespective of the climatic conditions.

The sequence from Abric Romani has provided several palaeoecological studies. This paper presents a synthesis of the results yielded by palynology, including the data from the entire non-excavated deposits, and the data obtained through the study of large mammals, micromammals, herpetofauna, and anthracology from the excavated layers. The palaeobotanical studies show that the sequence is mainly related to pine forests, and near or within these dominant forests there were smaller plant formations and/or secondary species with a Mediterranean and/or Atlantic character. Nevertheless, continuous forest did not dominate the entire region. The fauna occupying the territory indicate open spaces in which seasonal grasslands proliferated, dry assemblages in sunny areas and humid ones in mountain shadows. The non-excavated archaeological layers correspond to MIS 4, initially characterized by temperate and humid conditions, and thereafter cool and humid until HS 6 and MIS 3. Afterwards, at the beginning of the MIS 3, the Dansgaard–Oeschger cycles, ∼1500 yr rapid and abrupt climatic changes, occurred during the deposition of the archaeological levels O, N, M, L, K and J. Later, these events were interrupted by a cold and dry phase, between ca 49 and 47 ka BP, which affected levels Ja, I and H, culminating with HS 5. Finally, this cold episode was followed by the Hengelo interstadial or DO 12, characterized by a temperate and humid climate identified in levels E and D.

The transition from the Middle Palaeolithic (Mousterian) to the Upper Palaeolithic (Aurignacian) has been one of the prominent themes in the archaeology of the European Palaeolithic for more than 20 years. One of the most controversial questions concerning this period is the extinction of the Neanderthals and their replacement by modern humans. In this context, Cueva del Conde, located in the northern part of the Iberian Peninsula, is an archaeo-palaeontological site that records the Mousterian to Aurignacian transition. It has been excavated since the beginning of the 20th century, first by the Conde de la Vega del Sella and systematically by a team from the University of Oviedo since 2001. Three main zones have been identified: the External Zone, dated to approximately 39 110 ± 520 BP (level N104); the Entrance Platform, dated between 38 250 ± 390 BP and 34 730 ± 500 BP; and Gallery A with a radiocarbon date of approximately 31 540 ± 400 BP (level N2a2). The small-vertebrate assemblages recovered from the water-screening of all sediment from the excavation campaigns represent at least 21 small mammal, amphibian and squamate taxa. The small-vertebrate associations in the three zones suggest a patchy landscape, dominated by humid meadows and woodland areas with the existence of water in the vicinity of the cave. The climate shows a more continental pattern during the Mousterian, though it was milder during the Aurignacian. The small vertebrates of the Cueva del Conde Mousterian and Aurignacian levels suggest a climate that differed from modern day temperatures, between −1.1 and −4.4 °C (mean annual temperature), placing these assemblages during Interstadials 9 to 7 (Is9 to 7).

The present paper analyses the evidence provided by three sites (Sima del Elefante, Gran Dolina, and Galeria) located in the Trinchera del Ferrocarril of the Sierra de Atapuerca. These three sites are cave infillings that contain sediments deposited from approximately 1.2 Ma to 200 kyr. Pollen, herpetofauna, and small and large mammal remains are used as proxies to obtain a general picture of the environmental changes that occurred at the Sierra de Atapuerca throughout the one million-year period represented at these sites. Similarly, cultural changes are tracked analyzing the evidence of human behavior obtained from the study of several bone and lithic assemblages from these three sites. At least three periods with different cultural features, involving technology, subsistence and behavior, are determined from the available evidence. The first two periods correspond to the Mode 1 technology and Homo antecessor: the first is dated around 1.2 to 1.0 Ma and reflects opportunistic behavior both in technology and subsistence. The second period is around 800 kyr BP. Mode 1 technology is still maintained, but subsistence strategies include systematic hunting and the use of base camps. The third period is dated between 500 ka and 200 ka and corresponds to the Mode 2 technology and the acquisition of directional hunting and other organizational strategies by Homo heidelbergensis. A transition from Mode 2 to Mode 3 seems to appear at the end of this time-range, and may reflect the early phases of a fourth cultural change. With regard to the environment, our main conclusion is that there was an absence of extremely harsh conditions at Atapuerca throughout this time period. The presence of Mediterranean taxa was constant and the dominant landscape was a savannah-like open environment, probably with small forest patches. An alternation of Mediterranean and mesic species as the dominant component of the tree storey was induced by the climatic cycles, and steppes spread across the landscape during the drier periods. In any case, it is not possible to establish clear cut-off points separating entirely different environmental episodes. Our results show no evidence of any relationship between environmental change and cultural change at the Sierra de Atapuerca.

The Valdavara-1 cave, located in the westernmost part of the Iberian Peninsula, is an archaeological site that has been excavated since 2007. At least two main sedimentary units have been identified: an upper one, 30 cm-thick, that has yielded many fragments of ceramic characteristic of the Recent Prehistory (with a radiocarbon age of 4.490 ± 40 years BP), and a lower one, 90 cm-thick, separated from the upper one by an erosive discontinuity and late Pleistocene in age (radiocarbon ages between 13.770 ± 70 and 15.120 ± 70 years BP). The small-vertebrate assemblages recovered from the sieving-washing of all the sediment from the excavation campaigns include a total of at least 34 taxa (six amphibians, nine squamate reptiles, five insectivores, two chiropterans and 12 rodents). The materials from this locality constitutes the first mention in northwestern Spain for Microtus (Iberomys) cabrerae and Micromys minutus, currently absent from this area, as well as the first mention in the fossil record for Chioglossa lusitanica, Discoglossus galganoi and Chalcides striatus. In both layers, such small-vertebrate associations suggest a patchy landscape dominated by humid meadows and woodland areas, with the existence of water areas in the vicinity of the cave. The climate shows a more continental pattern during the late Pleistocene, as evidenced by the presence of Microtus arvalis, Sorex minutus and Rana temporaria, and was milder during the Recent Prehistory, with the occurrence of typically Mediterranean taxa such as Crocidura russula, Chioglossa lusitanica, Discoglossus galganoi, Rana iberica, Chalcides striatus and Timon lepidus. The amphibian and squamate associations suggest that the climate change recorded at the Valdavara-1 cave between the latest Pleistocene and Holocene levels may correspond to a difference of 0.8 °C in terms of mean annual temperature, although the difference is 2.8 °C for the mean temperature of the coldest month.

Among the Pleistocene localities of the Sierra de Atapuerca, the site called Sima del Elefante is famous for having delivered in its lower level (TE-LRU) some hominid remains currently considered the oldest in Western Europe (about 1.2 Ma). In its upper part (TE-URU), this site has fossiliferous levels pertaining to the late Middle Pleistocene (350-250 ka). Here we describe for the first time the amphibians and reptiles of these upper levels (TE18 and TE19), which have proved to be one of the richest assemblages of all the localities of the Sierra de Atapuerca. The faunal list is composed of 18 taxa made up of urodeles (Salamandra salamandra and Lissotriton helveticus), anurans (Discoglossus sp., Alytes sp. Pelobates cultripes, Pelodytes punctatus, Bufo bufo, Bufo calamita, Hyla arborea and cf. Rana sp.), a terrestrial tortoise (Testudo s.l.), lizards (Lacerta s.l., Podarcis sp. and Anguis fragilis) and snakes (Natrix natrix, Natrix maura, Coronella cf. girondica and Vipera latasti). For the first time, the genera Lissotriton, Discoglossus and Podarcis are mentioned in the Pleistocene localities of the Sierra de Atapuerca. TE19 also corresponds to the earliest mention for L. helveticus in the Iberian Peninsula. With the exception of tortoises, all these species are currently present in the province of Burgos, and thus suggests that the herpetofauna has a modern aspect since the Middle Pleistocene. This association of amphibians and reptiles suggests a slightly warmer climate than the current one, but may correspond to a “cold and dry” period if compared with other Pleistocene localities of the Sierra de Atapuerca. The landscape was probably composed of a gallery forest along a quiet water river within a Mediterranean environment alternating laterally between dry meadows, rocky or stony areas and open scrubland.

The site of El Portalon is the entrance to the Cueva Mayor karst system, located in the Sierra de Atapuerca (Burgos, Spain). This is an important Holocene archaeological site, which was excavated in the 1970s but from which little has been published. New excavations starting in 2000 have highlighted a deep stratigraphical sequence, with human occupations starting at the end of the Late Pleistocene. In this paper, we present for the first time, on the basis of the small-vertebrate assemblage, palaeoenvironmental and palaeoclimatic reconstructions of the Latest Pleistocene of the Sierra de Atapuerca, well known for its Early to Middle Pleistocene human-bearing localities. The small vertebrates of El Portalon comprise at least 25 species: 4 amphibians (Alytes obstetricans, Bufo bufo, Bufo calamita and Rana temporaria); 3 squamates (an indeterminate small-size lacertid, an indeterminate large colubrine and Vipera sp.); 6 insectivores (Sorex gr. coronatus-araneus, Sorex minutus, Neomys fodiens, Neomys anomalus, Talpa europaea and Galemys pyrenaicus); 2 bats (Myotis myotis and Myotis gr. myotis-blythi); 10 rodents (Microtus arvalis, Microtus agrestis, Microtus oeconomus, Iberomys cabrerae, Chionomys nivalis, Terricola duodecimcostatus, Arvicola sapidus, Arvicola terrestris, Apodemus sylvaticus and Eliomys quercinus). These taxa, many of which are ecothermal (sensitive to temperature) show variations in their taxonomic diversity throughout the sequence. Although, with the exception of M. oeconomus, they do not differ from the extant fauna of the Iberian Peninsula, they do so in the abundance of their taxonomic assemblage. When the small vertebrates are grouped and studied in terms of vertical trends through the sequence, it is possible to follow environmental and climatic changes. Results from the small-vertebrate associations indicate that the landscape had open habitats in the vicinity of the Atapuerca caves throughout the sequence, with wet locales in the surrounding area. Woodland and water stream meadows were more developed during “warm” periods (Is5, Is6/Is7 and Is3/Is4), whereas during “cold” periods (H3 and LGM) the environment was slightly more humid in response to winter precipitation and the opening of the landscape. These results are compared with pollen analysis and marine isotopic curves, giving a scenario for the palaeoclimatic and palaeoenvironmental changes that occurred during the Latest Pleistocene in the Sierra de Atapuerca.

The site of Guenfouda, located 30 km South of Oujda in the Jebel Metsila, which forms part of the Monts d'Oujda, was discovered in 2003. The site is of significant archaeological importance and sys...

Thousands of fossils of small vertebrates are recorded in one of the most complete stratigraphic sequences of the continental European Pleistocene, in the Sierra de Atapuerca. This sequence, with a few interruptions, spans the last 1.5 million years. The stratigraphic series is recorded in six karstic cavities in the Sierra de Atapuerca. From the oldest record to the most modern one, these are the Sima del Elefante, Gran Dolina, Galeria–Zarpazos, Sima de los Huesos, El Portalon and El Mirador. The faunal successions of continental microvertebrates have made it possible to divide the Atapuerca sequence into 7 faunal units (Atapuerca Faunal Units, ATA FUs), which are based on the first and last local appearances of taxa (LO, HO). The Spanish Quaternary small vertebrate faunas are grouped into 5 distinct biozones, from the oldest to the most recent: Allophaiomys pliocaenicus , Allophaiomys lavocati , Allophaiomys chalinei , Iberomys brecciensis and Iberomys cabrerae . These are calibrated with radiometric and palaeomagnetic dating, allowing the proposal of a continental biochronological scale for the Pleistocene of the Iberian Peninsula.

In addition to the Neanderthals, other mammal species vanished locally or totally in southern Europe during relatively a short period, between about 40 and 10 kyr. The late Upper Pleistocene was in general a time of extinction and extirpation in southern Europe, ranging from musk-oxen and reindeer to Nordic voles (extirpations) to Pliomys and Neanderthals (extinctions). The southern regions of Europe have been glacial refugia for several mammalian species (including humans), and when climatic conditions ameliorated, some spread back to the North and East European Plains (reindeer, saiga, the marsh voles and narrow-skull voles, among others), whence they originally had come. Others remained at their refugia in southern France and on the Iberian, Italian and Balkan Peninsulas, and, after periods of survival, eventually became extinct (Homo neanderthalensis, Pliomys lenki). The demise of the Neanderthals is, of course, well known, but that of the extinct vole, P. lenki, is not well known at all. Their long survival in southwestern European regions must be related to the existence of suitable climatic, vegetational and geographic conditions. Did these regions become traps when new competitors arrived?

Although ongoing, excavations at the Valdavara-1 cave (Galicia, northwestern Spain) have already yielded a rich and highly diverse terrestrial small vertebrate assemblage. Among the findings, two precaudal vertebrae from the Holocene level (with a radiocarbon age of 4490 ± 40 years BP) are attributed to the golden-striped salamander (Chioglossa lusitanica) and constitute the first fossil evidence for this species. Because of the poor intraspecific genetic diversity of living Spanish populations it has been proposed that they have a recent origin linked with a postglacial expansion from southern refuges. The palaeontological data presented in this paper thus show that the northernmost expansion of the species took place at least as early as the Chalcolithic period and permit the estimation of its dispersion speed within its potential distribution area.

Filiació URV: SI The Sierra de Atapuerca (Burgos, Spain) contains the most comprehensive archaeological and paleontological evidence of human evolution in Europe during the Quaternary. The time scale based on the microfossil content of the sedimentary infillings of the caves of Atapuerca has been used to attribute relative ages to the various paleontological and archaeological levels and correlate amongst the different sites. Microfossils are particularly significant as they are the dominant constituents of the caves from the Pleistocene to the Holocene of the Sierra de Atapuerca. The microfossils of the Atapuerca sites are mainly composed of isolated bones and teeth from small vertebrates, and the hard tissue of the vertebrate skeleton can be taxonomically classified. In revisiting the distribution and the classification of the small-mammal taxa along five of the main stratigraphic sequences from Atapuerca (Sima del Elefante, Gran Dolina, Galería-Zarpazos, Sima de los Huesos, and Portalón), we apply a new method, the "Unitary Associations" method, in order to refine the existing biozones, adding five new Faunal Units, and interpret their relative age. Finally, we correlate Atapuerca with other Pleistocene sites of Europe.

The habitat weighting method, chorotype classification and the bioclimatic model, as well as the Simpson diversity index, are applied to the small-vertebrate assemblage of Cova del Rinoceront (Castelldefels, Barcelona) in order to reconstruct the environmental and climatic fluctuations that are reflected in the MIS 5 sequence of the cave. The small-vertebrate data analysed are from Unit 1 of the sequence, which comprises three MIS 5 layers (III to I). They allow one cool and two warm episodes to be identified within the section. The warm episodes are related with open-woodland formations and temperatures higher than at present in the area, though in layer III our data suggest moist conditions, with the precipitation higher than today, which could probably be related with the beginning of MIS 5e. Layer II is associated with drier conditions, with precipitation lower than nowadays, which could probably be related with MIS 5c. By contrast, layer I is associated with open woodland formations, but with cooler and relatively humid conditions than today to judge by the temperature and precipitation data obtained. This could probably be related with MIS 5b. Our results are consistent with the available chronological, large-mammal, bird and palaeobotanical datasets for this upper part of the sequence. They are also consistent with the pollen record from the marine margins of Iberia.

The Middle to Upper Palaeolithic transition, between 50 000 and 40 000 years ago, is a period of important ecological and cultural changes. In this framework, the Rock Shelter of Uluzzo C (Apulia, southern Italy) represents an important site due to Late Mousterian and Uluzzian evidence preserved in its stratigraphic sequence. Here, we present the results of a multidisciplinary analysis performed on the materials collected between 2016 and 2018 from the Uluzzian stratigraphic units (SUs) 3, 15 and 17. The analysis involved lithic technology, use-wear, zooarchaeology, ancient DNA of sediments and palaeoproteomics, completed by quartz single-grain optically stimulated luminescence dating of the cave sediments. The lithic assemblage is characterized by a volumetric production and a debitage with no or little management of the convexities (by using the bipolar technique), with the objective to produce bladelets and flakelets. The zooarchaeological study found evidence of butchery activity and of the possible exploitation of marine resources, while drawing a picture of a patchy landscape, composed of open forests and dry open environments surrounding the shelter. Ancient mitochondrial DNA from two mammalian taxa were recovered from the sediments. Preliminary zooarchaeology by mass spectrometry results are consistent with ancient DNA and zooarchaeological taxonomic information, while further palaeoproteomics investigations are ongoing. Our new data from the re-discovery of the Uluzzo C Rock Shelter represent an important contribution to better understand the meaning of the Uluzzian in the context of the Middle/Upper Palaeolithic transition in south-eastern Italy.