**6. Impact of paleoclimate changes on Neanderthals and anatomically modern humans (AMH) in Central Europe**

The Balkan peninsula was likely the migration route of anatomically modern humans (AMH) into Europe [51], and the Danube valley which cuts the Pannonian Basin is one of the most important pathways of these population movements [52]. This region consists of vast lowlands associated with the middle Danube drainage basin and surrounded by the Carpathian Mountains, the Alps, Dinarides and the

Bohemian Massif. The Last Glacial loess provides widely extended sedimentary coverage of the area and provides valuable paleoclimate records [2–4]. Additional stratigraphic records are present in caves located in the highlands and peripheral mountain zones. The Zmajevac LPS, described above is located in this region. This geographical setting allows us to determine with considerable certainty the impact of climate change in the Late Pleistocene on Neanderthal and early modern human populations.

The Neanderthals clearly represent the autochthonous population of eastern Central Europe according to various research [53–55]. This is documented by a group of fossil finds, spread over space and time and in the various environments, ranging from the last interglacial to the temperate oscillations of the early Würmian glacial (OIS 5a–e). Some of the most important fossil finds of Neanderthals and AMH are located in the Pannonian Basin, especially in its central, western and southern parts (**Figure 1**). Here we briefly describe localities from Pannonian Basin: two in Croatia, one in Hungary and one in Slovakia.

*Krapina (Croatia):* Excavated layers 3–8 yielded more than 900 skeletal fragments of several Neanderthal individuals, especially cranial fragments, mandibles, teeth, and postcranial fragments. This makes Krapina one of the most important Neanderthal sites in Europe. ESR and U-series dating provided results between 178 and 120 ka, with average values pointing to 130 ka, i.e., to the last interglacial peak OIS 5e [56]. The Lithic industry is a variant of the Mousterian.

*Vindija, G3 layer (Croatia):* All layers in this cave are characterized by abundance of cave bear skeletal remains, especially in some of the layers. Within the sequence of the Mousterian industries, the Neanderthal fossils [54] appear in layer G3 in association with some endscrapers and possible leaf-point fragments [57, 58]. Age of neanderthal tibia fragment in G3 layer was dated and it is 38 ka B.P. [59].

*Vindija G1 layer (Croatia):* This layer yielded several human fragments of archaic morphology, which do not differ radically from the Neanderthals of the underlying layers and elsewhere in Central Europe [54]. However, the associated lithics, even if typical for the Initial Upper Paleolithic period in general, allow for somewhat contradictory interpretations. The leaf point suggests the Szeletian industry [59] and on the other hand the bone split-base point and the Mladeč type point suggest an Aurignacian [57, 60]. The Aurignacian industry that marked replacement of Neanderthals by anatomically modern humans (AMH) lasted from 43 to 26 ka years B.P. and it is characterized by worked bone or antler points with grooves cut in the bottom [61]. Their flint tools include fine blades struck from prepared cores rather than using crude flakes [61]. Mester [62] describes the problem of distinguishing these two cultural units and points to a possibility that Szeletian tools had been made by AMH. From this point of view Szeletian represents a sub variant of the Aurignacian. In this interpretation, Aurignacian bone points may have been the functional equivalents of Szeletian bifacial leaf points.

The associated bear bones were dated to 36–32 ka B.P., but dating of the human bones provided AMS radiocarbon dates of 29–28 ka B.P. [63]. Given the association of these objects in an 8–20 cm thick layer which is partially cryoturbated, we cannot exclude the possibility of some mixture of fossils and artifacts of various ages, as some researchers suggest [60]. However, since the two types of projectiles—the lithic leaf points and the polished bone-and-antler points – appear together in several other cave sites of the region (Dzeravá skala, Mamutowa Cave, Istá lloskö Cave, etc.) [64], it is rather unlikely that mechanical mixing was responsible in all cases. It seems that associations of these projectiles made from different materials and thus with different advantages and functions [65] with predefined cultures may not be as local as expected. The "Aurignacian" bone projectiles are actually being found more frequently in non-Aurignacian contexts, not only in the Central European caves,

**25**

*Pleistocene Climate Change in Central Europe DOI: http://dx.doi.org/10.5772/intechopen.93820*

100–75 ka years B.P., which fits into OIS 5 stage [67].

but also in other regions as far away as northeast Russia [57, 60]. This indicates that certain communication between separated Neanderthal groups could have existed. *Remete Felsö (Hungary):* The stratigraphy of this cave includes two glacial horizons or layers marked as: 5 and 4. The upper one, which is a layer characterized by loess containes limestone debris. Three human teeth (right I1-I2 and C) belonging to the same individual were found and analyzed. They are rather large and worn, but nothing more can be said about their specific features. The fauna, including cave bears, hyenas, lions and musk ox, suggests a tendency to cooling between the lower and upper horizon. All this faunal remains point to Szeletian in sensu latu and age is determined as OIS 3 [65]. The associated industry is characterized by typical leaf points and retouched flakes (including a Levallois flake), and has been classified generally as Szeletian, or, as a specific Transdanubian form of the late Middle Paleolithic—the Jankovichian [66]. *Šal'a (Slovakia):* Two Neanderthal cranial fragments, Šal'a 1 and the subsequently discovered Šal'a 2, were found in two different locations in the Vah river gravel deposits, but in secondary position and without precise dating. According to the correlation of the phylogenetic stratigraphic ranges of the vertebrate finds, the primary position of the Neanderthal Šaľa 1 specimen could be–with high probability – set into the terrestrial layers of the last interglacial age, approx.

Generally, eastern part of Central Europe provides solid evidence for the association of Neanderthals with the various Middle Paleolithic cultural entities of the interglacial and early glacial: the Taubachian, Mousterian, and Micoquian [68, 69]. Recent findings from the Neanderthal type locality Kleine Feldhofer Grotte site in the Neander Valley (Germany) also provide solid insights in various Middle and Upper Paleolithic cultural entities [70]. Preliminary analysis of the thousands of lithic artifacts recovered from this site has shown that two specific Paleolithic assemblages are represented: Micoquian artifacts typical of the late Middle Paleolithic and Upper Paleolithic artifacts from the Gravettian [70].

The question of the last Neanderthals and their relationship to the transitional or Initial Upper Paleolithic cultural entities of the region—the Szeletian and the Bohunician is far more susceptible to debate. Their "transitional" character is understood as a combination of archaic Middle Paleolithic patterns in technology,

The moment of AMH appearance in the Balkans and Central Europe has become better documented, since the new discovery at Pestera çu Oase 36–34 ka B.P. [71] and revisions of human fossil sites such as Mladeč which points to age of 35–34 ka B.P. In addition, the expansion of Aurignacian sites in Central Europe shows a specific time and space dynamic. While the early Aurignacian sites, dated as early as 42 ka B.P., are extremely rare and isolated (Willendorf II in Austria and Geissenklösterle in Germany) [72–74], the middle Aurignacian, dated between 34 and 29 ka years B.P., forms a kind of network of sites over large parts of the region. It also includes the emergence of Aurignacian figurative art. This findings point to interesting and probable conclusion. If the Aurignacian can be identified with AMH then the increased site density reflects their demographic growth. Also, if the art represents their higher social complexity and more advanced cognitive abilities, then the whole process may demonstrate the final "victory" of AMH over Neanderthals in Central Europe. Various authors have listed several possible reasons for the extinction of Neanderthals. Some have discussed the possibility that their extinction was stimulated by violent conflict with *Homo sapiens* [75]. Violence in early hunter-gatherer societies usually occurred as a result of resource competition following major natural disasters. Another possibility, proposed recently is the spread of pathogens or parasites carried by *Homo sapiens* into the Neanderthal population [76]. The fact of coexistence also leaves open the possibility of interbreeding which resulted

combined with the introduction of Upper Paleolithic tool-types [69].

#### *Pleistocene Climate Change in Central Europe DOI: http://dx.doi.org/10.5772/intechopen.93820*

*Pleistocene Archaeology - Migration, Technology, and Adaptation*

two in Croatia, one in Hungary and one in Slovakia.

OIS 5e [56]. The Lithic industry is a variant of the Mousterian.

the functional equivalents of Szeletian bifacial leaf points.

Bohemian Massif. The Last Glacial loess provides widely extended sedimentary coverage of the area and provides valuable paleoclimate records [2–4]. Additional stratigraphic records are present in caves located in the highlands and peripheral mountain zones. The Zmajevac LPS, described above is located in this region. This geographical setting allows us to determine with considerable certainty the impact of climate change in the Late Pleistocene on Neanderthal and early modern human populations. The Neanderthals clearly represent the autochthonous population of eastern Central Europe according to various research [53–55]. This is documented by a group of fossil finds, spread over space and time and in the various environments, ranging from the last interglacial to the temperate oscillations of the early Würmian glacial (OIS 5a–e). Some of the most important fossil finds of Neanderthals and AMH are located in the Pannonian Basin, especially in its central, western and southern parts (**Figure 1**). Here we briefly describe localities from Pannonian Basin:

*Krapina (Croatia):* Excavated layers 3–8 yielded more than 900 skeletal fragments of several Neanderthal individuals, especially cranial fragments, mandibles, teeth, and postcranial fragments. This makes Krapina one of the most important Neanderthal sites in Europe. ESR and U-series dating provided results between 178 and 120 ka, with average values pointing to 130 ka, i.e., to the last interglacial peak

*Vindija, G3 layer (Croatia):* All layers in this cave are characterized by abundance of cave bear skeletal remains, especially in some of the layers. Within the sequence of the Mousterian industries, the Neanderthal fossils [54] appear in layer G3 in association with some endscrapers and possible leaf-point fragments [57, 58]. Age of neanderthal tibia fragment in G3 layer was dated and it is

*Vindija G1 layer (Croatia):* This layer yielded several human fragments of archaic morphology, which do not differ radically from the Neanderthals of the underlying layers and elsewhere in Central Europe [54]. However, the associated lithics, even if typical for the Initial Upper Paleolithic period in general, allow for somewhat contradictory interpretations. The leaf point suggests the Szeletian industry [59] and on the other hand the bone split-base point and the Mladeč type point suggest an Aurignacian [57, 60]. The Aurignacian industry that marked replacement of Neanderthals by anatomically modern humans (AMH) lasted from 43 to 26 ka years B.P. and it is characterized by worked bone or antler points with grooves cut in the bottom [61]. Their flint tools include fine blades struck from prepared cores rather than using crude flakes [61]. Mester [62] describes the problem of distinguishing these two cultural units and points to a possibility that Szeletian tools had been made by AMH. From this point of view Szeletian represents a sub variant of the Aurignacian. In this interpretation, Aurignacian bone points may have been

The associated bear bones were dated to 36–32 ka B.P., but dating of the human bones provided AMS radiocarbon dates of 29–28 ka B.P. [63]. Given the association of these objects in an 8–20 cm thick layer which is partially cryoturbated, we cannot exclude the possibility of some mixture of fossils and artifacts of various ages, as some researchers suggest [60]. However, since the two types of projectiles—the lithic leaf points and the polished bone-and-antler points – appear together in several other cave sites of the region (Dzeravá skala, Mamutowa Cave, Istá lloskö Cave, etc.) [64], it is rather unlikely that mechanical mixing was responsible in all cases. It seems that associations of these projectiles made from different materials and thus with different advantages and functions [65] with predefined cultures may not be as local as expected. The "Aurignacian" bone projectiles are actually being found more frequently in non-Aurignacian contexts, not only in the Central European caves,

**24**

38 ka B.P. [59].

but also in other regions as far away as northeast Russia [57, 60]. This indicates that certain communication between separated Neanderthal groups could have existed.

*Remete Felsö (Hungary):* The stratigraphy of this cave includes two glacial horizons or layers marked as: 5 and 4. The upper one, which is a layer characterized by loess containes limestone debris. Three human teeth (right I1-I2 and C) belonging to the same individual were found and analyzed. They are rather large and worn, but nothing more can be said about their specific features. The fauna, including cave bears, hyenas, lions and musk ox, suggests a tendency to cooling between the lower and upper horizon. All this faunal remains point to Szeletian in sensu latu and age is determined as OIS 3 [65]. The associated industry is characterized by typical leaf points and retouched flakes (including a Levallois flake), and has been classified generally as Szeletian, or, as a specific Transdanubian form of the late Middle Paleolithic—the Jankovichian [66].

*Šal'a (Slovakia):* Two Neanderthal cranial fragments, Šal'a 1 and the subsequently discovered Šal'a 2, were found in two different locations in the Vah river gravel deposits, but in secondary position and without precise dating. According to the correlation of the phylogenetic stratigraphic ranges of the vertebrate finds, the primary position of the Neanderthal Šaľa 1 specimen could be–with high probability – set into the terrestrial layers of the last interglacial age, approx. 100–75 ka years B.P., which fits into OIS 5 stage [67].

Generally, eastern part of Central Europe provides solid evidence for the association of Neanderthals with the various Middle Paleolithic cultural entities of the interglacial and early glacial: the Taubachian, Mousterian, and Micoquian [68, 69]. Recent findings from the Neanderthal type locality Kleine Feldhofer Grotte site in the Neander Valley (Germany) also provide solid insights in various Middle and Upper Paleolithic cultural entities [70]. Preliminary analysis of the thousands of lithic artifacts recovered from this site has shown that two specific Paleolithic assemblages are represented: Micoquian artifacts typical of the late Middle Paleolithic and Upper Paleolithic artifacts from the Gravettian [70].

The question of the last Neanderthals and their relationship to the transitional or Initial Upper Paleolithic cultural entities of the region—the Szeletian and the Bohunician is far more susceptible to debate. Their "transitional" character is understood as a combination of archaic Middle Paleolithic patterns in technology, combined with the introduction of Upper Paleolithic tool-types [69].

The moment of AMH appearance in the Balkans and Central Europe has become better documented, since the new discovery at Pestera çu Oase 36–34 ka B.P. [71] and revisions of human fossil sites such as Mladeč which points to age of 35–34 ka B.P. In addition, the expansion of Aurignacian sites in Central Europe shows a specific time and space dynamic. While the early Aurignacian sites, dated as early as 42 ka B.P., are extremely rare and isolated (Willendorf II in Austria and Geissenklösterle in Germany) [72–74], the middle Aurignacian, dated between 34 and 29 ka years B.P., forms a kind of network of sites over large parts of the region. It also includes the emergence of Aurignacian figurative art. This findings point to interesting and probable conclusion. If the Aurignacian can be identified with AMH then the increased site density reflects their demographic growth. Also, if the art represents their higher social complexity and more advanced cognitive abilities, then the whole process may demonstrate the final "victory" of AMH over Neanderthals in Central Europe.

Various authors have listed several possible reasons for the extinction of Neanderthals. Some have discussed the possibility that their extinction was stimulated by violent conflict with *Homo sapiens* [75]. Violence in early hunter-gatherer societies usually occurred as a result of resource competition following major natural disasters. Another possibility, proposed recently is the spread of pathogens or parasites carried by *Homo sapiens* into the Neanderthal population [76]. The fact of coexistence also leaves open the possibility of interbreeding which resulted

with a genetic heritage left by the Neanderthals in the anatomically modern human (AMH) population of the Upper Paleolithic Europe.

Neanderthals possessed the brain that enabled them greater visual acuity than *Homo sapiens* did, but the latter had better language-processing abilities [77]. It can be stated with certainty that Neanderthal brains were more adapted to vision and spatial memory and that resulted in less available area for cognition and social interactions [77]. This difference in brain structure could also lead to extinction of Neanderthals during short period of competition with *Homo sapiens*.

A separate set of factors that are not connected to the interaction of Neanderthals and AMH are climate change and natural disasters. It is well documented and described that the general characteristic of the paleoclimate in Central Europe (particularly in Pannonian Basin) is repeated succession of oscillations with varying intensity. This climate teeter started with an expansion of dense forests during the interglacial peak (OIS 5e). In OIS 5e the climate was very similar to today's climate. It continued throughout the long transitional stage of the early glacial (OIS 5a–d) with several oscillations that shaped a dry, steppe environment [2, 4, 5, 31]. This climate change has affected the whole region, but we must not forget that geomorphology has conditioned specific micro-climatic conditions within [2, 4, 20, 31]. As discussed in this book chapter, climate change was constant during the Upper Pleistocene in Central Europe. Average summer temperature changes were in range from 9.5°C and up to 13.2°C [24, 26, 31] compared to present day temperatures which are significant changes that have certainly affected the Neanderthal population.

Climate changes during the glacial and interglacial periods were also the main cause for changes in vegetation. C3 plants have been most probably the main vegetation type during the Late Pleistocene. These include trees and cold steppe grasses [31]. Changes in plant life were reflected in herbivore, mammal population and they would have led to a corresponding decline in big, plant-eating mammals hunted by the Neanderthals [78].

From the aforementioned sites and findings within, we can assume that the Neanderthal extinction in eastern Central Europe was not the result of just adverse climatic conditions during the Lower Pleniglacial maximum (OIS 4), but rather originated from several millennia of coexistence with the emerging early modern humans during the OIS 3 [79]. Data indicate that the disappearance of Neanderthals occurred at different times in different regions of Europe and Asia. Comparing the data with results obtained from the earliest dated AMH sites in Europe allowed the quantification of the temporal overlap between the two groups. The results reveal a significant overlap of 2600–5400 years (at 95.4% probability) [78]. It is clear that the coexistence with AMH population was long enough for the transmission of cultural and symbolic behavior, as well as possible genetic exchanges (interbreeding), between the two groups [78], but it is hard to conclude that it was the main cause of Neanderthal extinction. After the interbreeding episode(s), Neanderthals and their material culture disappeared and was replaced across Europe and Asia by AMH [79]. The precise timing of this transitional period has remained difficult to identify in the absence of a reliable chronological framework [79, 80].

In the end and as the most obvious conclusion, we can say that the extinction of Neanderthals and the rise of AMH population in Central Europe is due to a combination of all the factors mentioned in above, but it is difficult to reliably determine which one prevailed.

## **7. Conclusion**

Data obtained from sedimentological and magnetic susceptibility analysis of Zmajevac LPS show a fairly good similarity with results from other LPS's in the

**27**

**Author details**

Adriano Banak1

Austria

Croatia

and Fabrizio Lirer5

\*, Oleg Mandic2

Central Europe and everywhere else in the World.

\*Address all correspondence to: abanak@hgi-cgs.hr

provided the original work is properly cited.

, Davor Pavelić3

Pannonian Basin [2, 4, 8]. Stable oxygen values measured in fossil snail shells show significant paleotemperature changes during the Upper Pleistocene in the Baranja region. Average growing season (AGS) temperature changes during that period were 13.2°C [24] or 9.5°C [26], depending on which formula is applied. The second calculated value is more plausible and in accordance with other results from Pannonian Basin. The overall climate was much cooler then present day climate. Stable carbon isotope values show that the C3 plants have been the main vegetation type of fossil snails for the entire time span during which the Zmajevac LPS was accumulated. This indicates that they lived in environment dominated by trees and cold steppe grasses. Comparison of the results from Zmajevac LPS with other LPS's from Central Europe [2, 4, 81] suggests that Upper Pleistocene climate in the Baranja region was similar to the paleoclimate in other regions in the Pannonian Basin. Certain differences in paleoclimate existed and they are probably an effect of local geomorphology and microclimate conditions.

The described climate change in the Upper Pleistocene is very likely a significant but not the only factor that influenced the extinction of the Neanderthal population which paved the way for the dominance of anatomically modern humans (AMH) in

2 Department of Geology and Paleontology, Naturhistorisches Museum Wien,

3 Faculty of Mining, Geology and Petroleum Engineering, University of Zagreb,

© 2020 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/ by/3.0), which permits unrestricted use, distribution, and reproduction in any medium,

4 Department of Geology, Faculty of Sciences, University of Zagreb, Croatia

5 Institute for Coastal Marine Environment (IAMC), INRC, Roma, Italy

1 Department of Geology, Croatian Geological Survey, Zagreb, Croatia

, Marijan Kovačić4

*Pleistocene Climate Change in Central Europe DOI: http://dx.doi.org/10.5772/intechopen.93820*

### *Pleistocene Climate Change in Central Europe DOI: http://dx.doi.org/10.5772/intechopen.93820*

*Pleistocene Archaeology - Migration, Technology, and Adaptation*

(AMH) population of the Upper Paleolithic Europe.

with a genetic heritage left by the Neanderthals in the anatomically modern human

Neanderthals possessed the brain that enabled them greater visual acuity than *Homo sapiens* did, but the latter had better language-processing abilities [77]. It can be stated with certainty that Neanderthal brains were more adapted to vision and spatial memory and that resulted in less available area for cognition and social interactions [77]. This difference in brain structure could also lead to extinction of

A separate set of factors that are not connected to the interaction of Neanderthals

Climate changes during the glacial and interglacial periods were also the main cause for changes in vegetation. C3 plants have been most probably the main vegetation type during the Late Pleistocene. These include trees and cold steppe grasses [31]. Changes in plant life were reflected in herbivore, mammal population and they would have led to a corresponding decline in big, plant-eating mammals hunted by the Neanderthals [78]. From the aforementioned sites and findings within, we can assume that the Neanderthal extinction in eastern Central Europe was not the result of just adverse climatic conditions during the Lower Pleniglacial maximum (OIS 4), but rather originated from several millennia of coexistence with the emerging early modern humans during the OIS 3 [79]. Data indicate that the disappearance of Neanderthals occurred at different times in different regions of Europe and Asia. Comparing the data with results obtained from the earliest dated AMH sites in Europe allowed the quantification of the temporal overlap between the two groups. The results reveal a significant overlap of 2600–5400 years (at 95.4% probability) [78]. It is clear that the coexistence with AMH population was long enough for the transmission of cultural and symbolic behavior, as well as possible genetic exchanges (interbreeding), between the two groups [78], but it is hard to conclude that it was the main cause of Neanderthal extinction. After the interbreeding episode(s), Neanderthals and their material culture disappeared and was replaced across Europe and Asia by AMH [79]. The precise timing of this transitional period has remained difficult to

Neanderthals during short period of competition with *Homo sapiens*.

changes that have certainly affected the Neanderthal population.

identify in the absence of a reliable chronological framework [79, 80].

In the end and as the most obvious conclusion, we can say that the extinction of Neanderthals and the rise of AMH population in Central Europe is due to a combination of all the factors mentioned in above, but it is difficult to reliably determine

Data obtained from sedimentological and magnetic susceptibility analysis of Zmajevac LPS show a fairly good similarity with results from other LPS's in the

and AMH are climate change and natural disasters. It is well documented and described that the general characteristic of the paleoclimate in Central Europe (particularly in Pannonian Basin) is repeated succession of oscillations with varying intensity. This climate teeter started with an expansion of dense forests during the interglacial peak (OIS 5e). In OIS 5e the climate was very similar to today's climate. It continued throughout the long transitional stage of the early glacial (OIS 5a–d) with several oscillations that shaped a dry, steppe environment [2, 4, 5, 31]. This climate change has affected the whole region, but we must not forget that geomorphology has conditioned specific micro-climatic conditions within [2, 4, 20, 31]. As discussed in this book chapter, climate change was constant during the Upper Pleistocene in Central Europe. Average summer temperature changes were in range from 9.5°C and up to 13.2°C [24, 26, 31] compared to present day temperatures which are significant

**26**

which one prevailed.

**7. Conclusion**

Pannonian Basin [2, 4, 8]. Stable oxygen values measured in fossil snail shells show significant paleotemperature changes during the Upper Pleistocene in the Baranja region. Average growing season (AGS) temperature changes during that period were 13.2°C [24] or 9.5°C [26], depending on which formula is applied. The second calculated value is more plausible and in accordance with other results from Pannonian Basin. The overall climate was much cooler then present day climate. Stable carbon isotope values show that the C3 plants have been the main vegetation type of fossil snails for the entire time span during which the Zmajevac LPS was accumulated. This indicates that they lived in environment dominated by trees and cold steppe grasses. Comparison of the results from Zmajevac LPS with other LPS's from Central Europe [2, 4, 81] suggests that Upper Pleistocene climate in the Baranja region was similar to the paleoclimate in other regions in the Pannonian Basin. Certain differences in paleoclimate existed and they are probably an effect of local geomorphology and microclimate conditions.

The described climate change in the Upper Pleistocene is very likely a significant but not the only factor that influenced the extinction of the Neanderthal population which paved the way for the dominance of anatomically modern humans (AMH) in Central Europe and everywhere else in the World.
