Cover Page The handle http://hdl.handle.net/1887/49751 holds various files of this Leiden University dissertation Author: Welker, F. Title: The palaeoproteomic identification of pleistocene hominin skeletal remains Issue Date: 2017-05-18 Chapter 1: Introduction 1. Perspectives on the Middle to Upper Palaeolithic Transition The Middle to Upper Palaeolithic Transition (MUPT) in western Europe concerns a chronological period during which local Neanderthal populations are replaced by Anatomically Modern Human populations (AMHs). At the end of the transitional period AMHs are present in large portions of the territory formerly occupied by Neanderthal populations, while these themselves have gone extinct. The replacement is a permeable one from a genetic perspective with a modest proportion of Neanderthal ancestry present in non-African modern humans. This is commonly interpreted as the occurrence of gene flow between the two populations (Green et al., 2010). Because of its chronological position and association to the spread of AMHs out of southwestern Asia, the MUPT takes a central role in discussions regarding the development and spread of behavioural modernity, including aspects of cognition and its archaeological manifestations (McBrearty and Brooks, 2000), the shaping of genetic variation in present day modern human populations (Green et al., 2010; Vernot et al., 2016) and the timing and causes of Neanderthal extinction (Higham et al., 2014). The exact processes facilitating this transition have sparked considerable controversy, especially in regards to several so-called “transitional” assemblages. These technocomplexes are placed chronologically between assemblages commonly attributed to Neanderthals and those attributed to AMHs and are present across Europe (Figure 1.1). They are “transitional” in that they seem to combine various Middle Palaeolithic (MP) or Mousterian and Upper Palaeolithic (UP) or Aurignacian elements into a single behavioural system. Specifically, the Chȃtelperronian (extending from central France to northwestern Spain), the Uluzzian (in Italy and Greece), the Szeletian (in the Czech Republic and Hungary) and the Lincombian-Ranisian-Jerzmanowician (LRJ, present across northern Europe) are termed “transitional” based on their chronostratigraphic placement and cultural attributes. These transitional technocomplexes occur at various times in some areas of western Eurasia, generally exhibiting the expected chronological east-to-west pattern (Hublin, 2015). However, they are not present in some other regions of Europe such as the majority of the Iberian peninsula (Figure 1.1; but see comments the “transitional Aurignacian” in Iberia (Bernaldo de Quirós and Maíllo-Fernández, 2009)). Furthermore, the presence of these transitional technocomplexes does not preclude the co-presence of late Middle Paleolithic assemblages in the same regions (Bobak et al., 2013; Higham et al., 2014; Jaubert et al., 2011; Slimak, 2007). Most importantly, despite some insights into their behavioural characteristics (see §1.1) and chronological occurrence (see §1.2), little direct evidence is available for the hominin population(s) responsible for the transitional technocomplexes (see §1.3). To complicate matters further, assemblages such as the Emirian (in the Levant), and the Bohunician and the Bachokirian (in central Europe) have been grouped together as Initial 1 Upper Palaeolithic (IUP) technocomplexes. Again, they follow a local MP or transitional technocomplex from a chronostratigraphic perspective but are suggested to be more fully rooted in UP technology compared to the transitional technocomplexes. The Szeletian in particular has been described as an MP acculturation in reaction to the arrival or development of the Bohunician (Nigst, 2012; Tostevin, 2007). The Szeletian and the Bohunician have significant geographic and chronological overlap (Škrdla, 2016), suggested to support such a scenario for the emergence of the Szeletian. This is in contrast to theoretical models advanced for the development of the Châtelperronian (Roussel et al., 2016). As with most transitional industries, these IUP industries are sparsely associated to hominin specimens. When they are present, such as at Üçağızlı Cave, they indicate an affiliation with AMHs (Kuhn et al., 2009). The transitional technocomplexes are preceded by local MP industries such as the MTA, Keilmessergruppen and the Micoquian that are attributed to Neanderthals (Benazzi et al., 2014a; Gabori-Csank, 1983; Schmitz et al., 2002; Toussaint et al., 2010), including hominin specimens that are directly dated and genetically studied (Krings et al., 1997). The precise chronology of the transitional and IUP technocomplexes remains elusive (see §1.2), as does their biological attribution (see §1.3). Both the transitional assemblages and the IUP are replaced by Proto-Aurignacian or Early Aurignacian populations that are invariably linked to AMHs, supported by chronometric and genetic data (Benazzi et al., 2015; Nigst et al., 2014; Verna et al., 2012). These are sometimes described as Early Upper Palaeolithic assemblages (EUP) instead. It must be noted that comparatively little is known about the “transitional” period in central and eastern Eurasia. Indeed, the existence of formal and well-defined analogous cultural technocomplexes has recently been re-addressed for these regions (Derevianko, 2011; Hoffecker, 2011; Zwyns et al., 2014). Examples include the identification of an IUP- related assemblage in central Mongolia (Zwyns et al., 2012). In part, this might be due to a low density of excavated sites and poor chronological control. As for western Europe, there are only a few hominin specimens associated with these assemblages, while the Ust’Ishim femur has been related to the IUP based on its direct AMS radiocarbon date (Fu et al., 2014). The recent report of AMHs in China by 80,000 BP (Liu et al., 2015) demonstrates that much remains to be learned about the cultural, biological and chronological arrival of AMHs in Eastern and Southern Eurasia and the local replacement (if any) of “archaic” hominins (Blinkhorn et al., 2013; Dennell, 2008; Groucutt et al., 2015; Timmermann and Friedrich, 2016). For western Eurasia, the long history of research into the transitional period has resulted in a body of data large enough for some authors to propose explicit models on the behavioural origins of the transitional assemblages and their chronological relationship to Aurignacian assemblages. These models are dominated by inferences based on the Chȃtelperronian due to a prolonged research history into this technocomplex. Therefore, in the following a majority of archaeological, chronological and biological observations are based on the Châtelperronian. Generally, the proposed hypotheses can be divided into i) those that propose that Neanderthals are associated to a transitional industry and that they independently developed its behavioural characteristics, ii) those that propose that Neanderthals are associated to a transitional industry but acquired its behavioural characteristics directly or indirectly by contact with AMHs, or iii) the association of Neanderthals to a transitional industry and its behavioural characteristics is the result of taphonomic factors causing movement of artefacts or hominin fossils between Mousterian, transitional and/or Aurignacian layers. In the third (iii) case, the transitional industries would 2 be made by AMHs. The first two models state that the transitional technocomplexes were made by Neanderthals based on the presence of Neanderthal fossil specimens or indirectly through the interpretation of technological characteristics. If this would be disproven by direct biological evidence and supported by taphonomic control, then the transitional assemblages simply become a proxy for an earlier presence of AMHs. More complex scenarios such as dual authorship or “hybrid”-origin can be formulated based on genetic evidence for gene flow between Neanderthal and AMH populations (as tentatively hypothesized by Ruebens et al. (2015) and d’Errico and Banks (2015)). They should not be excluded a priori, although they seem difficult to directly falsify based on the currently available data. In addition, the idea that all transitional technocomplexes would be related to either AMHs or Neanderthals does not need to be true (Hublin, 2015). The third hypothesis, however, proposes that taphonomic mixing of hominin specimens or cultural artifacts is the main cause of suggested attributions of Neanderthals to the transitional assemblages. This hypothesis needs to be refuted if one is to argue for either of the other two hypotheses. These hypotheses were largely formulated after the biological assignment of a skeleton located in the Chȃtelperronian layers at Saint-Césaire to Neanderthals (Lévêque and Vandermeersch, 1980). This provides an interesting perspective on the importance of biological specimens in transitional contexts, as the Chȃtelperronian had previously been associated with AMHs through its technological and behavioural characteristics (Ashton, 1983; Bordes, 1972; Breuil, 1909; Peyrony, 1948). Following the publication of the Saint- Césaire hominin and the confirmation of a Neanderthal-Chȃtelperronian association by analysis of the inner-ear morphology of a Neanderthal temporal at the Grotte du Renne, Arcy-sur-Cure (Hublin et al., 1996), there was a need to explain this association in formal behavioural or cognitive models. The implications of these models regarding the