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6 v.

PERSPECTIVES ON THE BIOGEOGRAPHIC AND CULTURAL ADAPTATIONS OF EARLY HUMAN S DURING THE FIRST INTERCONTINENTAL DISPERSALS

Ignacio de la Torre / Alfonso Benito-Calvo / Faysal Bibi / Jackson Njau / Shuwen Pei / Florent Rivals Haowen Tong / Kevin Uno / Sara Varela / Xiujie Wu

Abstract que retrotraen ambos eventos varios cientos de Our understanding of the emergence and miles de años antes de lo que se pensaba. A me- dispersal of the earliest tool-making hominins nudo, los factores climáticos y medioambientales has been revolutionised in the last decade, with se han considerado como los actores principales sites in eastern Africa and China pushing records que guiaron la mayoría de los eventos evolutivos of both events several hundred thousand years en nuestra especie. Sin embargo, los modelos que earlier than previously thought. In recent years, vinculan dinámicas terrestres climáticas con espe- climate and environmental factors have been ciación biológica, innovación cultural y eventos considered by many as primary drivers of these migratorios requieren una validación más sólida, evolutionary events in human history. However, y necesitan de nuevas aproximaciones a la bio- models linking Earth's dynamics with biological geografía y conductas adaptativas de los primeros speciation, cultural innovation and migration humanos. En el presente artículo proponemos la events with climate require further testing, and necesidad de adoptar una aproximación geográ- recent discoveries suggest that the picture of the fica amplia al estudio de las dinámicas de ocu- earliest human colonization across the Old World pación de los primeros humanos, basada en la is far more complex, demanding new approach- comparación de Africa oriental y China, regiones es to the biogeography and adaptive behaviours que contienen dos de las secuencias más largas of early humans. In this paper, we argue for a de primeros yacimientos arqueológicos en todo broader geographic approach to the study of ear- el mundo. Repasamos así los temas de investiga- ly human occupation dynamics comparing long ción principales relacionados con el estudio de archaeological sequences from eastern Africa las primeras migraciones humanas, y propone- and China. We thus review major research ques- mos una hoja de ruta para la comprensión de las tions involved in the investigation of the earliest trayectorias evolutivas alternativas adoptadas por human migrations and propose a route map to homininos que, aun compartiendo un fondo bio- better understand the alternative evolutionary lógico y cultural común, se enfrentaron a retos y trajectories adopted by hominins that shared an oportunidades climáticas y biográficas diferentes. overarching biological and cultural background, but that faced different climatic and biogeo- Palabras clave: primeros talladores, migrasiones graphic challenges and opportunities. humanas, Pleistoceno inferior en China y Africa.

Keywords: early makers, hominin mi- grations, early Pleistocene in China and Africa.

Resumen La comprensión de la aparición y dispersión de los primeros homininos que usaron herramientas ha experimentado una rev?lución en la última década, con yacimientos en Africa oriental y China

73 5.1. INTRODUCTION chaeological record; as the gaps between known sites become wider in time and space, it becomes Timing of the emergence of the earliest tech- crucial to go beyond first-appearance discussions no-complexes (De la Torre 2011) and the first and focus on the broader biogeographic and cul- hominin dispersals from Mrica into Europe (Roe- tural patterns involved. In addition, research on broeks 2006) and Asia (Dennell and Roebroeks the premodern human occupation of the Old 2005) are regularly pushed back. This pace has World can no longer be compartmentalised re- accelerated in the last decade, when our views gionally, and any comprehensive study should be have changed radically with the earliest artefacts based on a comparative analysis of the alternative reported at 3.3 million years ago (Ma) in Mrica evolutionary and cultural pathways observed in (Harmand et al. 2015), 1.85 Ma in Georgia (Fer- each area. The need for drawing similarities and ring et al. 2011), 1.2 Ma in Southwestern Europe differences between the earliest archaeological (Carbonell et al. 2008), and potentially at 2.1 sequences - those of Mrica and Asia - has be en Ma in China (Zhu et al. 2018). While the impor- recognised in recent years, but efforts have fo- tance of such discoveries cannot be overstated, cused on either general literature reviews of the it is also true that the field is dominated by a evidence in each region (Schick 1994; Dennell Joining-up-the-dots' perspective (Dennell 2017) and Roebroeks 2006; Norton and Braun 2010), focused on first appearance data, which is by de- or comparisons of particular attributes of sorne fault provisional and is proving to be unreliable stone tools (Norton and Bae 2008; Petraglia and as an explanatory device for human biological Shipton 2008; Lycett and Bae 2010). and cultural adaptations. In the discipline of as a whole, sub- Pliny's motto "ex Mrica semper aliquid novi" stantial fieldwork efforts in the last few decades drives a paradigm in which this continent saw the have produced extensive datasets, and the time emergence of consecutive species of Early Pleisto- is ripe for large-scale synthetic research (Kintigh cene (Antón and Swisher 2004) and Middle Pleis- et al. 2014). The archaeology of premodern hu- tocene (Lahr and Foley 2001) premodern humans mans is no exception to this trend, particularly in and Hamo sapiens (Stringer and Galway-Witham the two regions concerning this paper; triggered 2018), who then colonised the Old World. Despite by the discoveries at Olduvai Gorge, fieldwork in the problems of this paradigm (Dennell and Roe- eastern Mrica since the 1960s has produced an broeks 2005), Mrica is generally seen as the only unparalleled dataset of Early sites. Sim- region in the world with continuous human oc- ilarly, the exponential growth of scientific research cupation throughout the entire Early and Middle in China sin ce the 1990s has multiplied the num- Pleistocene (Dennell 2003). Massive chronologi- ber of archaeological sites and revolutionised our cal gaps between the earliest human evidence in understanding of human evolution in Asia. eastern latitudes of Europe (Ferring et al. 2011) The synthetic perspective advocated by Kintigh and in the western Mediterranean (Carbonell et et al. (2014) for archaeology - also identified as al. 2008; Toro-Moyano et al. 2013) may suggest one of the grand challenges in the study of cul- failed initial migrations and more than one "Out tural evolution as a whole (Brewer et al. 2017) - of Mrica" event, while the fragmentary evidence in has indeed be en substantiated in Antonio Gil- northern Europe could be linked to intermittent man's career, in which a comparative approach human occupation only during interglacial peri- has proved to be a highly productive research ods (Roebroeks 2005). An even more complicated strategy (e.g., Gilman 1991). Such perspective is picture emerges from Asia, where new discoveries now possible and greatly needed to properly un- (Zhu et al. 2018) suggesting human presence at derstand the ecological and cultural patterns of 2.1 Ma would necessarily imply that Hamo ergaster/ early humans. Nonetheless, such research should erectus was not the first colonizer of Eurasia, as be based on the systematic analysis of quantita- often thought (e.g., Antón 2003), and where the tive datasets (thus superseding the current stage Early and Middle Pleistocene archaeological re- of superficialliterature review comparisons) and cord is so sparse that a 'source-and-sink model' include computational modelling - also identi- (Martinón-Torres et al. 2018) has be en invoked fied as a general priority for the advance of the to interpret a discontinuous pattern of occupation discipline (Kintigh et al. 2014). Given the lack for the continent during this time. of systematic comparisons between the eastern A compendium of the 21st century grand Mrican and Chinese sites, metadata research challenges for archaeology (Kintigh et al. 2014) must also be accompanied by a direct analysis recognises, however, that key questions of the dis- of archaeological assemblages. cipline are no longer concerned with the earliest, In summary, the confirmation of eastern M- the longest, or the otherwise unique. If anything, rica and China as the regions with the longest the ever-increasing age depth for the emergence record of early archaeological sequences and the and dispersal of the first human cultures only substantial datasets produced in recent years, exacerbates the complexities of the early ar- present a unique opportunity to examine the be-

74 havioural ecology of premodern humans during see species turnovers in the context of climate the earliest colonization of the Old World, but change (Vrba 1996). These climatic hypothe- also a challenge due to the disparity of archives, ses have used the eastern Africa record to link research traditions and the lack of systematic aridification even ts (De Menocal 2011), lake intra and inter-regional comparisons. Such ex- pulses (Trauth et al. 2005) or increasing envi- ceptional challenges and opportunities should ronmental variability (Potts 1996) with species constitute the basis of new research programmes turnover and migration. to compare early human cultural and ecological In the northern latitudes of Eurasia hominin adaptations in what sorne would consider the speciation events might not been as recurrent source of most hominin species (eastern Mrica) as in Mrica, but climatic variations must have and in the recipient area for the first human mi- played a fundamental role in shaping human gration/s (namely eastern Asia). Stemming from biogeographic ranges, due to the constraints im- this all-embracing objective to understand the posed by the advance of glaciers during stadial alternative evolutionary trajectories adopted by stages. This process is relatively understood hominins that shared an overarching biological in Europe for modern human occupation dur- and cultural background (i.e., theywere premod- ing the Last Glacial Maximum (Tallavaara et al. ern humans using Early Stone Age technolog- 2015; Burke et al. 2017), the local extinctions ical solutions), research questions that deserve of Neanderthals in northern latitudes (Hublin special attention include: how can we identify and Roebroeks 2009) and across the continent migration waves, and how many were there in the (Tzedakis et al. 2007), and the punctuated na- Early and Middle Pleistocene, between the two ture of Early Pleistocene hominin colonization regions? How continuous is human occupation (Hosfield and Cole 2018). A similar latitudinal in the supposed evolutionary centre in eastern expansion and contraction of hominin popula- Mrica as opposed to the discontinuous record tions dictated by climatic fluctuations is consid- of China? To what extent do biotic versus abiotic ered for Asia (Dennell 2003, 2017a), although evolutionary factors dictate divergent trajectories here the model requires testing due to the near- in the colonization of Mrica and Asia? Can risks ly complete absence of environmental proxies and challenges faced by early humans in each re- from archaeological contexts. gion explain variations in hominin adaptations? While the influence of abiotic mechanisms Such questions are crucial to our understand- in biogeographic distributions both in eastern ing of Early Stone Age behavioural ecology, and Mrica and in China is evident, they do not satis- structure the following themes. factorily explain patterns observed in the archae- ological and paleontological record. In eastern Mrica, Bibi and Kiessling (2015) emphasized 5.2. EARLY HUMAN EXPANSION DYNAMICS the importance of continuous (and likely biotic) ACROSS THE OLD WORLD - THE PREMISES factors in modulating faunal change across the Pliocene and Pleistocene, whereas comparison of a tightly-constrained part of the Early Pleisto- cene sequence at Olduvai detected no changes 5.2.1. THE ROLE OF THE RED QUEEN AND THE in the faunal community despite intervening cli- COURT JESTER IN EARLY HUMAN DISPERSALS AND matic fluctuations (Bibi et al. 2018). This sug- FAUNAL TURNOVERS IN EAST AFRICA AND CHINA gests that mammal communities might be more robust than expected and that their foodwebs According to Benton (2009), paleoecologists remained relatively stable (Nenzén, Montoya and typically contrast evolutionary events through Varela 2014). a 'Court Jester' perspective (one where extrin- In the case of China, the biogeographic divi- sic, abiotic factors such as tectonic episodes sion of modern faunal communities (Xie, MacK- or climate change are responsible for extinc- innon and Li 2004) is broadly reproduced in the tion and speciation), whereas biologists - who Pleistocene (Tong 2006) and there is paleonto- normally are less concerned with time depth logical evidence for shifts following climatic oscil- - often privilege the classic Darwinian theo- lations (Jablonski et al. 2000; Norton et al. 2010), ry (nowadays summarised in the 'Red Queen' so it is assumed that hominin ranges also fluc- hypothesis) that invokes intrinsic factors of tuated accordingly (Dennell 2013). Despite the species competition (see also Van Valen 1973; appeal of this model, the archaeological evidence Barnosky 2001). While both perspectives co- do es not entirely follow predictions. For example, exist in human evolutionary studies, there is Zhoukoudien is the only site where human pres- an overwhelming preference for abiotic expla- ence is recurrently recorded throughout the Mid- nations, from models that highlight the role dIe Pleistocene (Shen, Gao and Granger 2009), of geographic features in shaping speciation and yet is located at quite a high altitude. In (Bailey, Reynolds and King 2011) to those that addition, the lack of environmental proxies to

75 test if the occupation of northern latitudes was have yet been indisputably reported in the Early indeed limited to interstadials is accompanied by Pleistocene of China, so it is also assumed that a strikingly low density of human presence in the the founding Homo populations in East Asia re- central and southern parts of China through the mained isolated from cultural and genetic flows entire Pleistocene. While it has be en proposed operating in the rest of the Old World. There is, that premodern humans were ill-adapted to the however, no archaeological evidence that con- subtropical forests of southern China (e.g., Cio- clusively supports (or falsifies, for that matter) chon and Bettis 2009), low density of hominin this hypothesis. Pei et al. (2017) have highlight- occupation in the central regions - which would ed how little we know about the technological be expected to act as refugia during colder peri- strategies ruling artefact production in the ear- ods (Denne1l2017b) - cannot be attributed only liest Chinese sites, which makes it even more to sampling and/ or taphonomic bias, given that pertinent to consider these in relation to eastern paleontological assemblages in this area are rea- African 'handaxe-free assemblages' (cf. De la sonably abundant. Therefore, we argue that cli- Torre 2009) of a similar age. Equally important matic factors alone do not explain hominin bio- is the gap in our knowledge of mammal taxa geographic dynamics. We hypothesize that biotic other than hominins; while comparative anat- interactions (e.g., competition), which are known omy of African and Asian early human fossils to be essential in shaping biodiversity and bioge- has received substantial attention (Antón 2003; ographic patterns (Araujo and Rozenfeld 2014), Rightmire 2008), attempts to insert earliest hu- may have played a fundamental role in hominin man dispersal waves into the context of wider occupation dynamics, and therefore need to be mammal migrations are limited (O'Regan et al. properly addressed. This is particularly relevant 2011; Tong et al. 2011) and have not involved when considering that early Homo had recently direct comparisons of the faunal assemblages. encroached on the carnivore guild (Pobiner and The archaeological evidence of the Middle Blumenschine 2003), and therefore ecological Pleistocene is also contentious. Evidence for a competition with resident carnivore species may new dispersal of hominins from Africa have be en significant. is reported at Ma (Goren-Inbar et al. 2000), but there is no agreement whether it reached eastern Asia. In recent years, handaxe-bearing 5.2.2. CULTURAL AND BIOLOGICAL archaeological sequences have been discovered CONNECTIVITY BETWEEN TWO CONTINENTS? in central China (see a review in Li, Kuman and Li 2018), thus forcing a reconsideration of the The first appearance datum of hominins in meaning of the Movius Line. For sorne, eastern China has steadily been pushed back from 1 Asian handaxes are the result of independent Ma (Schick and Toth 2000) to l.36 Ma (Zhu et innovations and technological convergence with al. 2001), l.66 Ma (Zhu et al. 2004) and now to the West (Corvinus 2004; Lycett and Bae 2010). potentially 2.1 Ma (Zhu et al. 2018). The discus- Alternatively, other authors emphasize similari- sion of human fossils has played an essential role ties with the African and western Asian Acheu- in framing both the ecomorphological potential lean (Petraglia and Shipton, 2008; Li, Kuman of early Homo to colonize new regions (Antón, and Li 2018), implying the existence of Middle Leonard and Robertson 2002) and in ascertaining Pleistocene cultural interactions across the en- the evolutionary links between the African and tire Old World. Most of the discussion, however, Asian premodern human demes (Antón 2003; has revolved around the comparison of handaxe Kaifu 2017), although they have often reached metrics from published sources, with no consider- opposite conclusions (e.g., Martinón-Torres et al. ation of handaxe technical features or the rest of 2007; Rightmire 2008). the elements of the reduction sequences. These If paleoanthropological discussions are in- latter lines of inquiry, based on a first-hand com- conclusive regarding the timing and extent parison of materials from Africa and China, may of gene flow between the two continents, the be more effective in deciphering the overarching interpretation of archaeological patterns is no knapping schemes (De la Torre and Mora 2009), less challenging. This essentially revolves around could facilitate a more solid assessment of ho- the controversial 'Movius Line', which arguably mologies, technological convergen ces or shared separates assemblages without handaxes to the technological principIes, and could provide a east of India from those with handaxes in the methodology to evaluate the extent of continen- rest of the Old-World Lower Palaeolithic (Schick tal interactions. Additionally, and although the 1994). Given the ever-increasing age of the ear- recently discovered handaxe sequences have rare- liest sites in China, it is now accepted that early ly yielded fossils, many other Middle Pleistocene Homo arrived in East Asia equipped with an Old- sites in China contain abundant bone assemblag- owan , and before the emergence of es (Tong 2006), which so far have played no role the Acheulean in eastern Africa. N o handaxes in a revaluation of the Movius Line.

76 5. 2 .3. CENTRE AND PERIPHERY IN THE EARLY historie discipline is setting an agenda to evaluate AND MIDDLE PLEISTOCENE? CONTINUITIES the impact of discontinuities in the evolution of AND DISCONTINUITIES OF THE EARLY past societies (Barberena et al. 2017), an in-depth ARCHAEOLOGICAL RECORD analysis of the tempo and modo of Mrican and Asian Early and Middle Pleistocene sequen ces is Eastern Mrica may have acted as a source area essential to test their roles as centre and periph- in which hominins - and many other mammal ery during the age of premodern humans. lineages - emerged and evolved. During the Early Pleistocene, eastern Mrica is often considered as the main source are a for technological innova- 5.2-4. RISKS AND EARLY HUMAN RESILIENCE tions and dispersals into Eurasia. The translation IN CHALLENGING ENVIRONMENTS ofWallerstein's (1974) concepts into Prehistoric research (Rowlands et al. 1987) has also be en Early Hamo (sensu Antón, Potts and Aiello applied to the early archaeological record by 2014) and Middle Pleistocene premodern hu- Dennell (2003), who portrays eastern Mrica as mans - who may include more than one deme the only part of the Old World with continuous (Antón 2003; Rightmire 2008; Kaifu 2017) - show human occupation throughout the entire Pleis- a trend towards increased body and brain size, tocene. In contrast, most of Eurasia remained developmental plasticity and dietary expansiono unoccupied - or only sporadically inhabited - Yet as an Mrican lineage, early Hamo would have by premodern humans, with ephemeral coloni- encountered thermoregulatory challenges in the zation conditioned by latitudinal variations in of northern latitudes of Eurasia (Ruff 1993), chal- ecosystems between glacial and interglacial peri- lenges that would have needed to be met with ods (ibid.; Roebroeks 2006). Recent genetic stud- cultural insulation solutions (Hosfield 2016). The ies estimate that effective human population size largest concentration of Asian Early Pleistocene in the entire Old World before 1.2 Ma was under sites is found in the Nihewan basin, where today 26,000 people (Huff et al. 2010), which suggests (i.e., art interstadial period) winter temperatures hominins would have be en an exceedingly rare drop below -10°C. Given the rare evidence offire proportion of the mammal communities in Eur- use before the end of the Middle Pleistocene asia (Dennell 201 7b), and provides further sup- (Roebroeks and Villa 2011), it is plausible to de- port to the notion of essentially deserted land- pict the settlement of the northern latitudes of scapes throughout the Early Pleistocene. China as episodic (only during interstadials) and Populations may have not be en much larg- seasonal - during the warm months of the year er in the Middle Pleistocene, particularly in the (Dennell 2013). This model, however, remains case of Asia, where low demographic density has entirely hypothetical, which is unfortunate given been used to explain the Movius Line (Lycett that we now know Early Pleistocene hominins and Bae 2010; Lycett and Norton 2010). Since did occupy northern European latitudes during the loss of cultural traits and a lack of innovation cold periods (Parfitt et al. 2010), and arguments transmission occur when small demographic lev- exist in favour ofyear-round (rather than season- els (Shennan 2000) and local extinctions (Pre- al) occupation (Hosfield 2016). Nonetheless, the mo and Kuhn 2010) predominate, low hominin archaeological evidence in Europe is too sparse, density and weakness of social networks could and a proper assessment of early human resil- have precluded cultural transmission of innova- ience to extreme environments should include tions in Asia, whereas higher interconnectedness the richer record of northeast China. in Mriea enabled the spread of the Acheulean Early Homo's encroachment on the carnivore (Lycett and Norton 2010). But alternative views guild (Pobiner and Blumenschine 2003) repre- are also possible, in which Mrica too could have sented an adaptive opportunity with paramount be en the scenario of independent emergence evolutionary implications (Aiello and Wheeler and disappearance of early where 1995), but it may have also presented significant cultural transmission was minimal (Tennie et al. risks. If australopithecines regularly faced preda- 2017), and therefore where a continuity of the tion by Mriean carnivores (Brain 1981) before archaeological record is not a prerequisite. hominins began to compete for meat resources, Once again, these models remain mostly the- once early Hamo became part of the carnivore oretical, but the dramatic time depth gained by guild at Ma, risks were unlikely to have the earliest archaeological record both in Mrica decreased (Van Valkenburgh 2001). Njau and and China in the last few years, and the signifi- Blumenschine (2012) showed that early Hamo cant number of archaeologieal sites now reported faced high predation risk from predators and in both regions - many of them in stratigraphic competition with carnivores (Pante et al. 2018). succession in various sedimentary basins - offers Still, carnivore paleoguilds of Mrica show a clear new opportunities for testing such hypotheses. acceleration of extinctions after 1.5 Ma, which In the current context where archaeology as a has be en correlated with the progressively higher

77 position of Homo in trophic chains (Lewis 1997; ing of these models, however, should be tested Werdelin and Lewis 2013). with first-hand analyses; taxonomic systematics of While a similar pattern of hominin-induced selected geographic regions across wide temporal extinction of carnivores may have occurred in Eu- spans can be used, for example to portray chang- (Croitor and Bruga120l0; Meloro and Clauss es in community structure before and after stone- 2012), the evidence in Asia is ambiguous, and tool making emerged and hominins encroached there is a pressing need for dedicated taphonomic on the carnivore guild. studies of Early Pleistocene sites (Pei et al. 2017). Early archaeological sites in China are secular- Additionally, a significant number of Chinese Mid- ly branded as -like or Mode I (see recent dIe Pleistocene have yielded hominin fossils reviews in Pei et al. 2017, 2019), thus emphasiz- alongside remains of other animals, but no (or ing their technological affinities with the African very few) artefacts (Bakken 1997; Liu, Zhang and Oldowan, but no direct, systematic comparisons Wu 2005; Wu et al. 2019). Even the richest Middle of assemblages have ever be en conducted. There- Pleistocene archaeological assemblage in China, fore, it is unclear what their shared features are, Zhoukoudian, could be a palimpsest ofhyaena-den apart from being core-and-flake technologies, accumulations (Binford and Ho 1985), where and thus the alleged close cultural phylogeny >60% of human fossils bear tooth marks (Boaz et is an assumption rather than a proof that the al. 2004). AH of this leads us to hypothesize that earliest colonization of China was made by Old- many deposits in Middle Pleistocene China owan hominins. First-hand comparison of stone were formed by carnivores, rather than hominins. tool collections from African Oldowan sites and This may be pointing at a very different pattern of those with a similar age in China (particularly carnivore-hominin interactions from that observed the rich Nihewan archaeological sequence) is in Africa and Europe, but needs validation via first- therefore essential to establish a systematic set hand analysis of the evidence. of defining features for both continents. In a sim- ilar vein, the Movius Line controversy has only recentIy prompted a quantitative approach, but 5.3. DISCUSSION - A ROADMAP the debate is still based on published material TO THE INVESTIGATION and grounded on metrical attributes that are not OF BIOGEOGRAPHIC AND CULTURAL necessarily informative of similarities and differ- PATTERNS DURING THE EARLIEST HUMAN ences between East and West handaxes. Again, DISPERSALS first-hand comparisons between African and Chi- nese Middle Pleistocene sites are vital, and new Competing hypotheses on the character and studies should go beyond formal comparisons of influence of clima tic fluctuations in East Afri- metrical features and focus on the technological can and Chinese mammal communities require principIes governing artefact production. None- validation through acquisition of new empirical theless, comparisons should not be limited exclu- evidence from environmental proxies but also sively to stone artefacts; the lack of comparative through computational modeling, to produce analysis between African and Chinese faunas has Early and Middle Pleistocene climate models for severely limited our understanding of mamma- both regions. Nonetheless, considering that the lian biogeography throughout the Pleistocene. impact of Pleistocene climatic changes on long- Much primary work - in the form of direct com- term mammalian community structure appears parative taxonomic analysis - is needed to test the to have been minimal in sorne regions (e.g. Bibi alleged biogeographic isolation of China during and Kiessling 2015), other factors may induce the Early and Middle Pleistocene, and to under- community collapse. For instance, the impact of stand whether human dispersals took place with- early humans in local extinctions is suspected in a broader context of faunal dispersals. (Petraglia 2017), particularly during the coloniza- Concerning the explanatory potential of the tion of new territories (Dennell 2017), but is yet centre and periphery concepts to depict earli- to be properly investigated. Given the existence est human dispersals, the main assumption to of long paleontological records before and after be tested is that discontinuities in the archaeo- the emergence of stone tool technology in Africa logical record of local sequen ces are linked to and the arrival of humans in China, modeling the increased aridity (eastern Africa) and cold con- impact of hominin encroachment on mammal ditions (China), and that the opposite will be species biogeography and turnover is fe asible and true during benign periods. However, despite should be a priority of future research, particular- the considerable intensity of research in east- ly by analysing changes at a local scale using fau- ern Africa, and the growing record in China, nal assemblages from African and Chinese fossil no comprehensive database yet exists of all the sites, and by running experiments on the recon- Early-Middle Pleistocene sequences and the ar- structed foodwebs to model hominin impact on chaeological proxies therein, so that they can be the fossil mammal communities. Ground-truth- contrasted with the paleoenvironmental record.

78 An assessment of the succession of archaeological during the first hominin dispersals across the Old sites within each sequence and an evaluation of Word. Significant efforts have been made earlier density peaks, troughs and gaps is also due, and in the discipline (Movius 1948; Clark 1969) and is now fe asible thanks to the relatively detailed in recent years (e.g., Petraglia and Korisettar 1998; availability of published reports. Denne1l2003) to decipher global patterns ofEarly Finally, the study of abiotic and biotic risks Stone Age occupation, but clearly more work can faced by early humans requires the formulation be done, particularly now, when the hard evidence of new conceptual premises but also the appli- - especially in China - is increasing exponentially. cation of new analytical techniques. As for the We have argued in this paper that such work latter, speculative characterization of extreme requires new and unconventional approaches to environments conditioning human occupation decipher archaeological patterns, which should should be tested through direct paleoecolog- go beyond paleoanthropological considerations ical analyses of relevant sites. Regarding biotic and evaluate the role of natural phenomena in risks and inter-species competition, after three shaping our understanding of Pleistocene data. decades the hunting-scavenging debate of early New perspectives should also be adopted in the Hamo in Mrica shows signs of having reached a selection of units and scales of analysis, in or- conceptual and methodological impasse (Sahle, der to embrace a myriad of temporal and geo- El Zaatari and White 2017). We thus advocate graphical resolutions comprising both metadata for a shift of focus where comparative analysis and primary sources, and analyses ranging from of bone surface modifications is coupled with a macro to micro scales. This kind of approach systematic assessment of community structures should be based on international collaborations through time and space (e.g., Patterson et al. where new data sets are produced, compared 2017). Zooarchaeological analysis of assemblages and synthesised, and be grounded in multidis- from different periods using the same protocols ciplinary perspectives that combine standard ar- would guarantee the application of standard- chaeological approaches with expertise ranging ized analytical methods (e.g., Njau and Gilbert from geochemistry to computational modelling. 2016), and therefore that results are comparable The application of new analytical techniques to (i.e., that hominin/ carnivore involvement in the both legacy data and primary sources and their formation of each faunal assemblage can be as- consolidation into models linking archaeological sessed relatively to one another). More important- and paleoclimatic data, alongside the considera- ly, by adding a community structure dimension tion of archaeological patterns in their biotic and to studies, changes in the diversity, abundance, abiotic context, wiIl enable validation of previous size and ecological preferences of potential prey theories and formulation of novel hypotheses on and predator competitors can provide insights the tempo and modo of earliest human occupation about human-carnivore interactions (e.g., Blu- in the Old Word, and particularly of its longest menschine et al. 2012). sequences in East Mrica and China.

5.4. CONCLUSIONS ACKNOWLEDGEMENTS

Regionalization of the archaeological record We thank Pedro Díaz-del-Río, Inés Sastre and from the Middle Palaeolithic / Middle Stone Katina Lillios for inviting us to participate in Age onwards requires a careful consideration of this volume, and we are honoured to be part of biological and cultural local trajectories to assess this festschrift to Antonio Gilman, who has con- variability in a global context. In contrast, the tributed so importantly to comparative research Lower Palaeolithic / Early Stone Age (and par- in prehistory. This project has received funding ticularly its earlier stages) shares sorne essential from the European Research Council (ERC) un- characteristics - e.g., it is a hand-held stone tool der the European Union's Horizon 2020 research technology produced by premodern humans - and innovation programme (BICAEHFID, grant that may promote searching for global patterns agreement No. 832980).

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