The last (?) Neanderthals in Iberia; implications of the finds

Neanderthals survived at least 3,000 years longer than we thought in Southern Iberia — what is now Spain — long after they had died out everywhere else, research suggests.


The authors of the study, an international team from research institutions, say their findings suggest that the process of modern human populations absorbing Neanderthal populations through interbreeding was not a regular, gradual wave-of-advance but a “stop-and-go, punctuated, geographically uneven history.”

Over more than ten years of fieldwork, the researchers excavated three new sites in southern Spain, where they discovered evidence of distinctly Neanderthal materials dating until 37,000 years ago.


“Technology from the Middle Paleolithic in Europe is exclusively associated with the Neanderthals,” said Dr. João Zilhão, from the University of Barcelona and lead author of the study. “In three new excavation sites, we found Neanderthal artefacts dated to thousands of years later than anywhere else in Western Europe. Even in the adjacent regions of northern Spain and southern France the latest Neanderthal sites are all significantly older.”

The Middle Paleolithic was a part of the Stone Age, and it spanned from 300,000 to 30,000 years ago. It is widely acknowledged that during this time, anatomically modern humans started to move out of Africa and assimilate coeval Eurasian populations, including Neanderthals, through interbreeding.

According to the new research, this process was not a straightforward, smooth one — instead, it seems to have been punctuated, with different evolutionary patterns in different geographical regions.

In 2010, the team published evidence from the site of Cueva Antón in Spain that provided unambiguous evidence for symbolism among Neanderthals. Putting that evidence in context and using the latest radiometric techniques to date the site, the researchers show Cueva Antón is the most recent known Neanderthal site.


“We believe that the stop-and-go, punctuated, uneven mechanism we propose must have been the rule in human evolution, which helps explaining why Paleolithic material culture tends to form patterns of geographically extensive similarity while Paleolithic genomes tend to show complex ancestry patchworks,” commented Dr. Zilhão.

The key to understanding this pattern, says Dr. Zilhão, lies in discovering and analyzing new sites, not in revisiting old ones. Although finding and excavating new sites with the latest techniques is time-consuming, he believes it is the approach that pays off.

“There is still a lot we do not know about human evolution and, especially, about the Neanderthals,” said Dr. Zilhão. “Our textbook ideas about Neanderthals and modern humans have been mostly derived from finds in France, Germany and Central Europe, but during the Ice Ages these were peripheral areas: probably as much as half of the Paleolithic people who ever lived in Europe were Iberians. Ongoing research has begun to bear fruit, and I have no doubt that there is more to come.”



Abstract The late persistence in Southern Iberia of a Neandertal-associated Middle Paleolithic is supported by the archeological stratigraphy and the radiocarbon and luminescence dating of three newly excavated localities in the Mula basin of Murcia (Spain). At Cueva Antón, Mousterian layer I-k can be no more than 37,100 years-old. At La Boja, the basal Aurignacian can be no less than 36,500 years-old. The regional Middle-to-Upper Paleolithic transition process is thereby bounded to the first half of the 37th millennium Before Present, in agreement with evidence from Andalusia, Gibraltar and Portugal. This chronology represents a lag of minimally 3000 years with the rest of Europe, where that transition and the associated process of Neandertal/modern human admixture took place between 40,000 and 42,000 years ago. The lag implies the presence of an effective barrier to migration and diffusion across the Ebro river depression, which, based on available paleoenvironmental indicators, would at that time have represented a major biogeographical divide. In addition, (a) the Phlegraean Fields caldera explosion, which occurred 39,850 years ago, would have stalled the Neandertal/modern human admixture front because of the population sink it generated in Central and Eastern Europe, and (b) the long period of ameliorated climate that came soon after (Greenland Interstadial 8, during which forests underwent a marked expansion in Iberian regions south of 40°N) would have enhanced the “Ebro Frontier” effect. These findings have two broader paleoanthropological implications: firstly, that, below the Ebro, the archeological record made prior to 37,000 years ago must be attributed, in all its aspects and components, to the Neandertals (or their ancestors); secondly, that modern human emergence is best seen as an uneven, punctuated process during which long-lasting barriers to gene flow and cultural diffusion could have existed across rather short distances, with attendant consequences for ancient genetics and models of human population history.


Introduction In the Aquitaine basin and the Pyrenees, the Middle Paleolithic (MP) Mousterian culture is followed, in succession, by the Châtelperronian, the Protoaurignacian and the Aurignacian I (a.k.a. Early Aurignacian). In Iberia, these initial phases of the Upper Paleolithic (UP) are represented in the Cantabrian strip and in Catalonia but remain unknown to the South of the Ebro basin. Based on these observations, the “Ebro Frontier” model hypothesizes that (a) in Valencia, Murcia, Andalusia, Gibraltar, the Mesetan hinterland, and Portugal, the corresponding chronostratigraphic slot is occupied by a late-persisting Mousterian and (b) the pattern is explained by the major biogeographical divide that the Ebro basin would have been at that time (Zilhão, 1993; Zilhão, 2000; Zilhão, 2006a; Zilhão, 2009).

The paleontological and ancient DNA (aDNA) evidence indicates that, in Europe, extensive admixture occurred at the time of contact between aboriginal Neandertals and in-dispersing groups of modern humans, resulting in the former’s eventual assimilation (Smith et al., 2005; Trinkaus, 2007; Pääbo, 2015). The authorship of the Châtelperronian, the Protoaurignacian, and the other so-called “transitional” industries from this time remains debated (Higham et al., 2010; Caron et al., 2011; Hublin et al., 2012; Trinkaus and Zilhão, 2013; Zilhão, 2013; Zilhão et al., 2015; Welker et al., 2016). In Western Eurasia, however, the Mousterian is exclusively associated with the Neandertals, while the Aurignacian I and the succeeding Aurignacian II (a.k.a. Evolved Aurignacian), which extend from Asturias in the West to northern Israel in the East, are associated with modern humans only (Verna et al., 2012). In this context, the broader paleoanthropological significance of the “Ebro Frontier” model resides in the implication that Neandertals persisted in Southern and Western Iberia longer than everywhere else.

It has been argued that no Aurignacian exists in Southern and Western Iberia, their Upper Paleolithic beginning with the Gravettian (de la Peña, 2013). Such views imply that (a) the Mousterian persisted even longer (Finlayson et al., 2006; Finlayson et al., 2008), or (b) after a Neandertal extinction event, Southern and Western Iberia remained uninhabited until modern human reoccupation (Bradtmöller et al., 2012; Galván et al., 2014). In these scenarios, the role of biogeographical divide played by the Ebro basin under certain climatic and environmental conditions would not have contributed to observed patterns in any significant manner.

Re-dating and critical examination of old sites and collections (Kehl et al., 2013; Wood et al., 2013) have advanced these debates. The scope of the many empirical issues involved, however, requires the excavation of new sites with the potential to settle the key points of contention. Here, we report on the progress made in that direction resulting from a decade of fieldwork in Murcia, Southeast Spain.


Results We excavated three localities <2 km apart within the Mula basin (Angelucci et al., 2017). The Supplementary Information (SI) Appendix provides a succinct geographical description of the area, as well as extensive monographic presentations of the sites’ stratigraphic sequences, dating, human occupation features, and stone tool assemblages. The sites are: Cueva Antón (CA; 38°03′51.84″N, 01°29′47.20″W), Finca Doña Martina (FDM; 38°04′43.21″N, 01°29′25.13″W), and Abrigo de La Boja (ADB; 38°04′43.37″N, 1°29′23.17″W).

The mutually exclusive presence/absence of diagnostic technologies in the Mula basin sites stands despite differences in assemblage size of up to two orders of magnitude, and is consistently seen across time. In this regard, the Late Mousterian in layer I-k of Cueva Antón is no different from the Middle Paleolithic assemblage of MIS 5 age found in the site’s layer II-l. Likewise, the equivalently small size of the Early Gravettian assemblages in OH13-OH14 of La Boja is no impediment for their fully Upper Paleolithic nature to manifest itself through such diagnostics as bladelets extracted from both prismatic and “burin” core-types, the “burins” themselves, and even the technocomplex’s index fossil (a microgravette point). Much the same applies to La Boja’s Aurignacian assemblages. At Finca Doña Martina, the lower resolution of the stratigraphic sequence means that each unit samples, and averages out, much longer time intervals. Yet, it remains that (a) Levallois and Discoid cores and blanks, sidescrapers, and denticulates are found together in this site’s basal layer 9 but not in overlying layers 8, 7b and 6/7, while (b) the reverse is true of prismatic, carinated/nosed “scraper” and “burin” core-types, endscrapers, or bladelet tools.

For the Rambla Perea rock-shelters, lateral variation between two adjacent archeological sites that, in the living past, must have functioned as a single, spatially extensive locus of human activity, suffices to explain the contrasts between coeval lithic assemblages. Through time, across the regional MP-UP transition, the use-wear evidence shows that the differences are primarily of a techno-typological nature. Hide-working, wood-working, defleshing and the use of projectiles are documented in both the Mousterian and the Aurignacian. However, (a) hides were processed with sidescrapers in the Mousterian but with endscrapers in the Aurignacian, and (b) projectiles were armed with single, axially-mounted points in the Mousterian but with multiple, laterally-mounted microlithic elements in the Aurignacian. In short, synchronic functional variability cannot explain the differences in lithic technology upon which we have assigned the stone tool assemblages of the Mula basin sites to either the Middle or the Upper Paleolithic.


Discussion The latest Middle Paleolithic south of the Ebro

The dating work carried out at the site of Sima de las Palomas, on the coast of Murcia, ca.60 km to the Southeast of Cueva Antón, provides further support for the late persistence of the Middle Paleolithic in the region — in this case, with diagnostic Neandertal remains found stratigraphically together with the lithics (Walker et al., 2008; Trinkaus and Walker, 2017). Correct understanding of the significance of the dates obtained at this key site is hindered by the samples’ provenience notations referring to arbitrary horizontal spits that do not reflect the stratigraphic layout of the sequence — something misunderstood by Wood et al. (2013) and Santamaría and de la Rasilla (2013), although explicitly stated in Walker et al. (2008). When the actual stratigraphy is considered, the dating results— obtained by radiocarbon on burnt bone treated with the ABA protocol, U-series on bone using Diffusion/Adsorption (D/A) assumptions, and multi-grain quartz OSL on sediments — are mutually consistent.

Taken together, the OSL and U-series dates are in turn consistent with the two radiocarbon dates from samples retrieved at the same stratigraphic elevation or higher up in units A-B and E. The uppermost radiocarbon result (OxA-10666) is from a faunal fragment cemented to a diagnostic Neandertal mandible that was (a) found half-way through the unit A deposit and (b) overlain by ca.50 cm of sediment containing nothing but diagnostic Middle Paleolithic stone tools and diagnostic Neandertal remains. As OxA-10666 translates into a calibrated age within the 38.6–42.0 ka interval, the Sima de las Palomas evidence strongly indicates, in line with the Cueva Antón pattern, that the Middle Paleolithic persisted in the region well beyond 40–42 ka. In addition, it shows that such a late-persisting Mousterian is indeed a Neandertal-associated technocomplex. There is no reason, therefore, to question that the association pertains in those other parts of Iberia where stratigraphy and dating support persistence of the Middle Paleolithic into the same time range: Gibraltar and Portugal.


The earliest Upper Paleolithic south of the Ebro

The persistence of a Neandertal-associated Middle Paleolithic from Iberia’s Mediterranean Southeast to its Atlantic seaboard implies that archeological manifestations of the modern human-associated Aurignacian I not be found across the same territory. Such is indeed the case. Neither stratigraphic units containing diagnostic assemblages nor isolated index fossils of the Early Aurignacian have been identified in the long cave sequences spanning the MP-UP transition known in those parts of the peninsula: Cova Beneito (Valencia), Cueva Bajondillo (Andalusia), Gorham’s Cave (Gibraltar), and Gruta do Caldeirão (Portugal) (Zilhão, 2006a). At these sites, and at others that are either open-air, single-occupation localities, or lack a basal Middle Paleolithic, the earliest Upper Paleolithic is the Aurignacian II (Evolved Aurignacian) or III–IV (a.k.a. Late Aurignacian).

The Mula basin sites suffice to demonstrate that, by 36.5–37.1 ka, the Aurignacian II was already present in Iberian regions to the South of the Ebro basin. This interval is the same during which, based on Bayesian modeling of available dates, Banks et al. (2013b) found that the transition from the Early to the Evolved Aurignacian had occurred to the North. This technological transition would therefore seem to have been concomitant with a process of settlement expansion: in Northern Europe, toward the British Isles and equivalent latitudes of Germany and Poland that, during the previous phase, had become devoid of human occupation; in Iberia, toward the lands beyond the Ebro basin, eventually leading to replacement of their late-persisting Mousterian and the assimilation of its Neandertal makers. The “Ebro Frontier” model provides a biogeographical and paleoecological framework for the interpretation of these developments in terms of population history.


Conclusions The technological and use-wear evidence rejects interpreting layer I-k of Cueva Antón and occupation horizons OH20 and OH19 of La Boja as distinct structural poses of a single, multifaceted system. Put another way, the small lithic assemblage in layer I-k of Cueva Antón cannot be interpreted as a functionally specialized, or activity-specific facies of the region’s Evolved Aurignacian. Instead, layer I-k of Cueva Antón and occupation horizons OH20 and OH19 of La Boja stand for concrete manifestations of mutually exclusive, long-lasting technologies whose succession, rather than a gradual transition, truly consisted of an abrupt replacement. As the efficiency of stone tool production in terms of cutting edge per unit of mass is identical in both technologies (Muller and Clarkson, 2016), the parsimonious reading of this replacement process is that it represents a major break, with demic underpinnings, in regional cultural trajectories.

The evidence from stone tool technology and the stratigraphic layout of sites is that the pattern derived from the high-precision Mula basin data can be extrapolated to all Iberian regions to the South of the Ebro basin. In these regions, artefact assemblages attributable to the earliest phases of Western Europe’s Upper Paleolithic are missing from stratified sites that contain deposits spanning the MP-UP transition, and have never been found as single-component, open-air contexts. In addition, no isolated occurrences of their index fossils (e.g., Châtelperronian points/knives, or Aurignacian split-based bone points) have ever been reported among surface, mixed, or post-depositionally disturbed deposits. From the basics of Prehistoric Archeology, i.e., from the culture-stratigraphic reasoning providing the framework for all its chronologies, the only inference that one can derive from this pattern is that, southward of the Ebro basin, a late-persisting Mousterian occupies the time slot in which the Aurignacian I is found elsewhere. The radiocarbon evidence is entirely consistent with this notion, which available luminescence and U-series independently support, and which no other kinds of radiometric dating results have so far countered.

A corollary of these findings is that Neandertals persisted until ca.37 ka across Southern and Western Iberia — which carries implications for the authorship of all other aspects of these regions’ archeological record. For instance, given their dating and archeological associations, there can be no question that the painted/perforated shells from Cueva Antón and Cueva de los Aviones, as well as the abstract engraving and ornamental use of raptor feathers documented at Gorham’s Cave, stand for manifestations of Neandertal symbolism (Zilhão et al., 2010a; Finlayson et al., 2012; Rodríguez-Vidal et al., 2014). Knowing that minimum ages of 40.8 ka for a red disk and 37.3 ka for a hand stencil have been obtained at El Castillo cave (Cantabria) (Pike et al., 2012), and that such motifs exist in Extremaduran and Andalusian sites, it is easy to see how the “Ebro Frontier” pattern may also bear implications for the authorship of cave paintings.

Recent advances in the field of Genetics increasingly make it clear that, in the Late Pleistocene of Eurasia, the continental extension of rather homogeneous archeological cultures is superimposed on complex ancestry patchworks (Mallick et al., 2016; Pagani et al., 2016). This can be explained by a pattern of long-distance diffusion and cultural resilience, which maintained networks over the long-term, combined with extended periods of geographical isolation, which conserved regional genetic variants. The “Ebro Frontier” effect makes this mechanism apparent even in the refugia of Southern Europe and especially so at the time of the MP-UP transition. This visibility is due to when the frontier formed and for how long it lasted, both allowing the effect to be picked-up with the current resolution of dating techniques. Likely, however, similar, broadly coeval but chronometrically less visible Late Pleistocene frontiers must have existed in other parts of Asia and Europe, as well as during the earlier phases of the process of modern human dispersal into these continents.

The results we report here highlight the need for proper integration of the biological and the archeological evidence when reconstructing Late Pleistocene population histories. All lines of evidence are now converging to support replacement-through-admixture, or Assimilation, as the best explanation for the disappearance of the Neandertal and other archaic phenotypes. The Iberian evidence suggests this was a time-transgressive evolutionary outcome stemming from dynamic, complex and geographically uneven processes — a punctuated history in which the long-term maintenance of pan-continental networks of gene flow and cultural exchange did not exclude the occurrence of extended periods of significant geographical isolation.

(Source: “Precise dating of the Middle-to-Upper Paleolithic transition in Murcia (Spain) supports late Neandertal persistence in Iberia”, by João Zilhão et al., November 2017)


Research-Selection for NovoScriptorium: Maximus E. Niles


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