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Post by Admin on Jan 18, 2020 20:34:12 GMT
Synthesis of Population Genetic Analyses The aDNA data from a range of Mesolithic hunter–gatherer samples from regions neighboring the LBK area have been shown to be surprisingly homogenous across space and time, with an mtDNA composition almost exclusively of hg U (∼80%), particularly hg U4 and U5, which is clearly different from the LBK dataset as well as the modern European diversity (Table 2) [20]. The observation that hgs U4 and U5 are virtually absent in the LBK population (1/42 samples) is striking (Table 2). Given this clear difference in the mtDNA hg composition, it is not surprising that the pairwise FST between hunter–gatherers and the LBK population is the highest observed (0.09298) when we compared ancient populations with representative population pools from Central Europe and the Near East (Table 3; see also [20]). If the Mesolithic data are a genuine proxy for populations in Central Europe at the onset of the LBK, it implies that the Mesolithic and LBK groups had clearly different origins, with the former potentially representing the pre-Neolithic indigenous groups who survived the Last Glacial Maximum in southern European refugia. In contrast, our population genetic analyses confirm that the LBK shares an affinity with modern-day Near East and Anatolia populations. Furthermore, the large number of basal lineages within the LBK, a reasonably high hg and haplotype diversity generated through one- or two-step derivative lineages, and the negative Tajima's D values (Tables 1 and 2) indicate a recent expansion. These combined data are compatible with a model of Central Europe in the early Neolithic of indigenous populations plus significant inputs from expanding populations in the Near East [4],[12],[34]. Overall, the mtDNA hg composition of the LBK would suggest that the input of Neolithic farming cultures (LBK) to modern European genetic variation was much higher than that of Mesolithic populations, although it is important to note that the unique characteristics of the LBK sample imply that further significant genetic changes took place in Europe after the early Neolithic.
aDNA data offers a powerful new means to test evolutionary models and assumptions. The European lineage with the oldest coalescent age, U5, has indeed been found to prevail in the indigenous hunter–gatherers [12],[35]. However, mtDNA hgs J2a1a and T1, which because of their younger coalescence ages have been suggested to be Neolithic immigrant lineages [8],[12], are so far absent from the samples of early farmers in Central Europe. Similarly, older coalescence ages were used to support hgs K, T2, H, and V as “postglacial/Mesolithic lineages,” and yet these have been revealed to be common only in Neolithic samples. The recent use of whole mitochondrial genomes and the refinement of mutation rate estimates have resulted in a general reduction in coalescence ages [8], which would lead to an improved fit with the aDNA data. However we advise caution in directly relating coalescence ages of specific hgs to evolutionary or prehistoric demographic events [36]. Significant temporal offsets can be caused by either observational bias (the delay between the actual split of a lineage and the eventual fixation and dissemination of this lineage) or calculation bias (incorrect coalescent age estimation). aDNA has considerable value not only for directly analyzing the presence or absence of lineages at points in the past but also for refining mutation rate estimates by providing internal calibration points [37].
Archaeological and anthropological research has produced a variety of models for the dispersal of the Neolithic agricultural system (“process of Neolithization”) into and throughout Europe (e.g., [1],[2],[38]). Our findings are consistent with models that argue that the cultural connection of the LBK to its proposed origin in modern-day Hungary, and reaching beyond the Carpathian Basin [23],[32],[38],[39], should also be reflected in a genetic relationship (e.g., shared haplotype analyses; Table S4). Therefore at a large scale, a demic diffusion model of genetic input from the Near East into Central Europe is the best match for our observations. It is notable that recent anthropological research has come to similar conclusions [40],[41]. On a regional scale, “leap-frog” or “individual pioneer” colonization models, where early farmers initially target the economically favorable Loess plains in Central Europe [33],[42], would explain both the relative speed of the LBK expansion and the clear genetic Near Eastern connections still seen in these pioneer settlements, although the resolving power of the genetic data is currently unable to test the subtleties of these models.
In conclusion, the new LBK dataset provides the most detailed and direct genetic portrait of the Neolithic transition in Central Europe; analysis of this dataset reveals a clear demonstration of Near Eastern and Anatolian affinities and argues for a much higher genetic input from these regions, while also identifying characteristic differences from all extant (meta-)populations studied. Ancient genetic data from adjacent geographic regions and time periods, and especially from the Near East and Anatolia, will be needed to more accurately describe the changing genetic landscape during and after the Neolithic, and the new multiplexed SBE assays offer a powerful means to access this information.
Haak W, Balanovsky O, Sanchez JJ, Koshel S, Zaporozhchenko V, Adler CJ, et al. (2010) Ancient DNA from European Early Neolithic Farmers Reveals Their Near Eastern Affinities. PLoS Biol 8(11): e1000536.
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Post by Admin on Jan 21, 2020 19:10:21 GMT
The driving force behind the transition from a foraging to a farming lifestyle in prehistoric Europe (Neolithization) has been debated for more than a century [1, 2, 3]. Of particular interest is whether population replacement or cultural exchange was responsible [3, 4, 5]. Scandinavia holds a unique place in this debate, for it maintained one of the last major hunter-gatherer complexes in Neolithic Europe, the Pitted Ware culture [6]. Intriguingly, these late hunter-gatherers existed in parallel to early farmers for more than a millennium before they vanished some 4,000 years ago [7, 8]. The prolonged coexistence of the two cultures in Scandinavia has been cited as an argument against population replacement between the Mesolithic and the present [7, 8]. Through analysis of DNA extracted from ancient Scandinavian human remains, we show that people of the Pitted Ware culture were not the direct ancestors of modern Scandinavians (including the Saami people of northern Scandinavia) but are more closely related to contemporary populations of the eastern Baltic region. Our findings support hypotheses arising from archaeological analyses that propose a Neolithic or post-Neolithic population replacement in Scandinavia [7]. Furthermore, our data are consistent with the view that the eastern Baltic represents a genetic refugia for some of the European hunter-gatherer populations. Results and Discussion By 6,700 years before present (BP) the Neolithization process had influenced most of northern Europe [9]. However, Scandinavia (including Denmark) was still occupied by highly mobile hunter-gatherer groups. Although the hunter-gatherers of Denmark and southern Sweden adopted pottery early on, the Neolithization first took real shape with the appearance of the Funnel Beaker Cultural complex (FBC, also known as the Trichterbecher Kultur [TRB]) some 6,000 years BP (the oldest evidence possible dating back some 6,200 years BP [9]). At this time domestic cattle and sheep, cereal cultivation, and the characteristic TRB pottery were introduced into most of Denmark and southern parts of Sweden [6]. Nevertheless, the Neolithization process was slow in Scandinavia, and large areas remained populated by hunter-gatherer groups until the end of the 5th millennium BP. One of these last hunter-gatherer complexes was the Pitted Ware culture (PWC), which can be identified by its single-inhumation graves distributed over the coastal areas of Sweden and the Baltic Sea islands that lie closest to the Swedish coast. Intriguingly, the PWC first appears in the archaeological record of Scandinavia after the arrival of the TRB (some 5,300 years BP) and existed in parallel with farmers for more than a millennium before vanishing about 4,000 years BP (Figure 1). This prolonged coexistence of hunter-gatherers and farmers in Scandinavia has been a focal point of debate within archaeology since 1909, when PWC human remains were used to argue for an early eastern influence on Neolithic Scandinavia, thus favoring relations to modern Saami people [10]. However, it has mainly been used as a key argument against both a rapid Neolithic transition and a large-scale population replacement between the Mesolithic and the present [7, 8]. Figure 1 Scandinavia with the PWC and the Architectural Structures of the TRB Displayed Three main hypotheses have been proposed to explain the origin of the PWC: (1) it has an origin in the late Mesolithic hunter-gatherer complexes of northern Europe [11] that, given that Neolithic or post-Neolithic population replacement took place, would make them genetically distinct from modern Scandinavians; (2) the PWC arose from a reversion to the hunter-gathering subsistence strategy among TRB peoples [12], and, given that no population replacement took place in Scandinavia during the Neolithization process, PWC peoples are the direct ancestors of modern Scandinavians; and (3) the PWC originated in populations ancestral to modern Saami people of present-day northern Scandinavia [10, 13]. To investigate PWC ancestry components in modern Scandinavians and peoples of the Baltic region, we recovered ancient mitochondrial (mtDNA) sequences (316 bp of the D-loop) from the skeletons of 22 individuals deriving from the two different cultures (see Table S1 available online). Three of these were TRB (all from one passage tomb, Gökhem, dated to 5,500–4,500 years BP, Figure 1), and 19 belong to the PWC (recovered from three different sites on the Baltic island of Gotland dated to 4,800–4,000 years BP, Figure 1). Quantitative real-time PCR was used to assess the total human mtDNA content in all samples (Tables S2 and S3, Figure S1) and to screen for appropriate molecular behavior (degradation ratio [14], Table S4). Amplicons were sequenced with the Roche Genome Sequencer FLX platform to retrieve synthetic clones [15] (Table S5). Sequences were regarded as authentic if they (1) originated from DNA extracts containing more than 1000 molecules of the quantified 80 bp fragment, (2) were supported by two independent extractions, (3) were based on a minimum of 20 synthetic FLX clone sequences, and (4) expressed a degradation ratio higher than 1 (Supplemental Data). Reduced median networks [16] were used to graphically illustrate substitution differences among sequences and to enable sequence assignation to previously defined haplogroups [4, 17]. Haplogroups U4/H1b, U5, and U5a were found to have high incidence among the PWC but are all rare among contemporary Scandinavians and Saami (Figures 2A–2C). It is noteworthy that a high frequency of U lineages, especially U5, has been inferred for pre-Neolithic Europeans with the use of modern mtDNA data [18]. Interestingly, compared to the rest of Europe, the U haplogroups have relatively high frequencies among populations in the eastern Baltic region such as the Latvians and the Lithuanians (Figure 2C).
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Post by Admin on Jan 22, 2020 17:44:39 GMT
Figure 2 Haplogroup Distributions in Investigated Populations Analysis of molecular variance [19] (AMOVA) revealed that the PWC sequences are significantly differentiated from samples of contemporary Swedes [20] (n = 289, FST = 0.05174, p < 0.001), Saami [20] (n = 38, FST = 0.25037, p < 10−6), Norwegians [21] (n = 323, FST = 0.06148, p < 0.001), Finns [22] (n = 79, FST = 0.05327, p < 0.005), Estonians [22] (n = 117, FST = 0.04745, p < 0.003), Lithuanians [22] (n = 163, FST = 0.04022, p < 0.004), and Latvians [22] (n = 114, FST = 0.03622, p < 0.011). To examine whether population differences could be accounted for by drift alone under the null hypothesis of population continuity, we performed coalescent simulations assuming a wide range of combinations of ancestral population size at the Upper Paleolithic colonization of Europe, 45,000 years ago, and the time of arrival of farming in Scandinavia, 6,200 years ago. As a conservative measure, we assumed a relatively high (compared to other published estimates) mutation rate of 7.5 × 10−6 per site per generation [23] to ease the burden of explaining high FST values. We sampled sequences from each simulation according to the numbers and dates of the data considered here and calculated the proportion of simulated FST values that were greater than those observed (Supplemental Data). The null hypothesis of population continuity between the PWC and modern Swedes can be rejected under a range of assumed ancestral population size combinations (including almost all that assume a Neolithic effective population size > 15,000), as can population continuity between the PWC and Norwegians under most assumed ancestral population size combinations (including almost all that assume a Neolithic effective population size > 6,000) (Figure 3). Population continuity between the PWC and modern Saami can be rejected under all assumed ancestral population size combinations. However, population continuity between the PWC and contemporary Baltic populations cannot be rejected under most assumed ancestral population size combinations (Supplemental Data). Figure 3Probabilities of Obtaining the Observed Genetic Differences, as Measured by FST, between Ancient PWC and Modern Population Samples under a Model of Population Continuity These results indicate that the PWC hunter-gatherers are unlikely to be the main ancestors of either modern Scandinavians or Saami, despite their presence in Scandinavia at the early stages of Neolithization. On the contrary, the observed FST values indicate greater similarity between the PWC and modern eastern Baltic populations, and coalescent simulations confirm that those non-Scandinavian populations could plausibly be the direct descendents of PWC hunter-gatherers. Having only obtained three TRB sequences, we cannot exclude continuity with any of the modern populations. Although complex demographic scenarios such as local population structuring, or sampling problems including close relatedness among individuals from the same site, might also explain the patterns of differentiation that we observe, we found no significant differentiation among the three PWC sites that we sampled (AMOVA pairwise FST = −0.0189; p = 0.54733; exact test of population differentiation global p value = 0.43421) and also note that the ubiquity of U4 and U5 types at those sites suggests that we are looking at patterns of genetic variation that extend beyond the local scale. It is noteworthy, however, that our interpretation is highly dependent on the assumed effective population size (Ne) at the onset of the Neolithic in Scandinavia 6,200 years BP. If Ne at this time were low (< 6,000 if modern Norwegians and Swedes share a common ancestry, < 15,000 for the unlikely event that the two populations have different ancestry in the region), then drift would be sufficient to explain the FSTs for both modern Swedes and northeastern Baltic peoples. Furthermore, it may be possible that a relatively low level of admixture in Scandinavia between the PWC and the TRB could be sufficient to explain the differences observed between the PWC and modern Scandinavians. Currently, however, this remains untestable, because we lack an appropriate proxy for early farmers. Given our results, it remains possible that the PWC represent remnants of a larger northern European Mesolithic hunter-gather complex. However, it appears unlikely that population continuity exists between the PWC and contemporary Scandinavians or Saami. Thus, our findings are in agreement with archaeological theories suggesting Neolithic or post-Neolithic population introgression or replacement in Scandinavia. To what extent this holds true for other parts of Europe requires further direct testing, although morphological [24, 25], ancient [26], and modern [4, 5] genetic data suggest that this is probably the case. Thus, theories favoring a Neolithization process that involved population continuity and was mediated by culture exchange only appear increasingly unlikely. Interestingly, however, the data analyses are consistent with a view that the eastern Baltic area remained a genetic refugia for some of the European hunter-gatherer populations. This is in agreement with findings of Mesolithic to Neolithic continuity among Latvian cemeteries [27]. Although the hunter-gatherer lifestyle was culturally replaced here, as in Scandinavia, the populations of the eastern Baltic area may have kept a certain level of population continuity. Experimental Procedures Skeletal remains from 74 individuals of eight middle-Neolithic sites were initially selected. Of these, 41 yielded sequence data, but only 22 (19 PWC and 3 TRB) met all requirements demanded for authenticity (Supplemental Data). A set of nonhuman samples, mainly harp seals (Phoca groenlandica, n = 31), of the same age and from the same sites as the human remains were used as controls and screened for human DNA (contamination) as well as for putative animal DNA (preservation). The material was extracted in duplicates via a silica spin-column method including chemical decontamination [14] (Supplemental Data). Quantitative real-time PCR of an 80 bp and a 136 bp coding-region fragment was used to assess the total human mtDNA content in all samples and to screen for appropriate molecular behavior (degradation [14]) (Supplemental Data, Table S1). We amplified the D-loop in seven overlapping fragments of varying size and sequenced them on the Roche Genome Sequencer FLX System to retrieve synthetic clones. Tagged primers were used to provide for individual identification after sequencing [15] (Supplemental Data). For analyses, we used 316 bp of the D-loop, spanning 16,051–16,383 (16 positions were removed, Supplemental Data). Analysis of molecular variance and Fst calculations was carried out with the Arlequin 3.1 software [19]. Networks were constructed with the NETWORK software and the reduced median algorithm [16], with the threshold set for 1. Coalescent simulations were performed assuming a wide range of combinations of ancestral population size at the Upper Palaeolithic colonization of Europe, 45,000 years ago (Ne = 10–4,959, and time of arrival of farming in Scandinavia 6,200 years ago, Ne = 500–20,350). Sequences were sampled from each simulation according to the numbers and dates of the data considered, and the proportion of simulated FST values that were greater than those observed were calculated (details on the evolutionary models are provided in Supplemental Data). Published:September 24, 2009 DOI:https://doi.org/10.1016/j.cub.2009.09.017
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Post by Admin on Jan 23, 2020 18:10:59 GMT
During the 4th and 3rd millennia BC, Italy was home to complex networks of metalwork exchange, according to a study published January 22, 2020 in the open-access journal PLOS ONE by Andrea Dolfini of Newcastle University (UK), and Gilberto Artioli and Ivana Angelini of the University of Padova (Italy). Research in recent decades has revealed that copper mining and metalwork in Italy began earlier and included more complex technologies than previously thought. However, relatively little is known about metalwork exchange across the country, especially south of the Alps. In this study, Dolfini and colleagues sought to understand how commonly and how widely copper was imported and exchanged throughout Late Neolithic (Copper Age) Italy. The researchers conducted an analysis of 20 copper items, including axe-heads, halberds, and daggers, from central Italy dating to the Copper Age, between 3600 and 2200 BC. Comparing archaeological data and chemical signatures of these items to nearby sources of copper ore, as well as to other prehistoric sites, they were able to determine that most of the examined objects were cast from copper mined in Tuscany, with the rest sourced from the western Alps and possibly the French Midi. These results not only confirm the importance of the Tuscan region as a source of copper for Copper Age communities in Italy, reaching as far as the Tyrolean area home of the Alpine Iceman, but also reveal the unexpected finding that non-Tuscan copper was a significant import to the region at this time. These data contribute to a growing picture of multiple independent networks of Copper Age metal exchange in the Alps and neighboring regions. The authors note that future research might uncover other early sources of copper, as well as more details of the interactions between these early trade networks. The authors add: "The first systematic application of lead isotope analysis (a geological sourcing technique) to Copper Age metal objects from central Italy, 3600-2200 BC, has shed new light on the provenance of the copper used to cast them. The research has revealed that, while some of the copper was sourced from the rich ore deposits of Tuscany, as was expected, some is from further afield. This unforeseen discovery demonstrates that far-reaching metal exchange networks were in operation in prehistoric Europe over a thousand years before the Bronze Age." Introduction The last two decades have witnessed a major surge of interest in the origins of copper mining, smelting, and working in the Italian peninsula. The new research season was inaugurated by fieldwork and radiocarbon dating at Libiola and Monte Loreto, two copper mines from eastern Liguria (Fig 1). Investigations brought to light prehistoric workings and galleries for the extraction of chalcopyrite (presumably supplemented by near-surface deposits of copper oxides/carbonates) dating from the mid-4th millennium BC. This pushed back significantly the beginnings of copper mining south of the Alps [1–2]. Such a surprising discovery led to reconsidering the chronology of early Italian metals, which at the time were overwhelmingly dated to the 3rd millennium BC [3–4]. A review of the evidence showed instead that Italian copper production likely commenced in the late 5th or early 4th millennium BC (i.e. the late/final Neolithic). It also showed that Neolithic metalworkers did not just manufacture small awls and points, as was until then believed, but also large, technologically complex axe-heads [5]. The discovery of final Neolithic smelting evidence from Orti Bottagone, Tuscany, further corroborated the new picture, showing that copper was not only cast and worked, but also smelted (and presumably mined) in the area from the early 4th millennium BC [6–7:p.147]. Fig 1. Map of the sites mentioned in the article. Inset: find spots of the metal objects analysed. 1: Libiola; 2: Monte Loreto; 3: Orti Bottagone; 4: San Carlo-Cava Solvay; 5: Grotta San Giuseppe; 6: Grotta del Fontino; 7: Grotta della Spinosa; 8: Fontenoce di Recanati; 9: Ponte San Pietro; 10: Selvicciola; 11: Rinaldone; 12: Lucrezia Romana; 13: Romanina; 14: The Iceman; 15: Zug-Riedmatt; 16: Saint Blaise/Bains des Dames; 17: Hornstaad; 18: Col del Buson; 19: Arene Candide; 20: Grotta della Pollera; 21: Lipari; 22: Saint Véran; 23: Fontaine-le-Puits. Hollow circles: contemporary cities. Inset: squares: axe-heads; dots: daggers; triangles: halberds. Objects ID 7–8, 38–39 (plus hafting rivet ID 39A): Pianizzoli; 55: Garfagnana; 71: Vetralla; 85, 88, 355: Province of Florence; 100: Province of Grosseto; 126–127: Il Teso; 170: Querceto; 357–358: Near Terni; 360: Province of Siena; 385: Selvena; 412: Abruzzo; 422: Corneto (modern-day Tarquinia); 425: Sarteano, Palazzone. Note that the locations of objects ID 55, 85, 88, 100, 355, 357, 358, 360 and 412 are approximate (image: Andrea Dolfini; base map: U.S. Geological Survey). Further research indicated that metal production and use surged dramatically in the early Copper Age, 3600–3350 BC. In the ore districts of Tuscany, in particular, the mid-4th millennium BC marked the inception of a mature technological tradition that, somewhat inappropriately, has come to be known as ‘Rinaldone metallurgy’ after a central Italian burial custom [8]. So-called ‘Rinaldone metallurgy’ featured: (a) the mastering of copper and arsenical-copper alloys (including a ternary Cu-As-Sb alloy that is typically found in central Italy) [9–10]; (b) early experimentation with silver and antimony extraction and casting; (c) the manufacture of distinctive objects including personal ornaments, axe-heads, daggers, and halberds; (d) technological improvements in metal casting and working; and (e) important changes in the cultural signification of metals, which entered the funerary domain to mark new ideas of gender, age, and perhaps status [11–13]. Such uses of metal objects are most apparent in iconic ‘Rinaldone’ warrior burials [8,14–15]. More recently, slag analysis from the now-destroyed settlement site of San Carlo-Cava Solvay has enabled further insights into early extractive metallurgy. Nestled at the edge of the Tuscan Colline Metallifere (or ‘Ore Mountains’), the site was excavated in the 1990s under rescue conditions [7]. The fieldwork brought to light several fired-clay platforms partly surrounded by curvilineal limestone walls, similar to early smelting installations from the eastern Alps and southern France [16–18]. Several copper-based compounds including sulphides and sulphosalts were smelted at the site using a surprisingly efficient reduction technology. This seemingly involved the crucible smelting of mixed charges of copper sulphides and oxides/carbonates, aided by blowpipes and/or bellows [19]. The process generated mature silica slags lacking the unreacted ore and entrapped copper prills typically found in Chalcolithic metallurgical residues [20]. This is all the more remarkable considering that the site dates to the late 4th millennium BC [7:p.142]; this makes it one of the earliest metallurgical sites in Western Europe. Despite these advances, a crucial field of research has remained woefully underdeveloped in Italian archaeometallurgy, namely the study of early metalwork exchange. The topic was recently addressed for the Alpine region using, alternatively, lead isotope analysis (LIA) [21–22] and the so-called ‘Oxford system’ charting changes in trace element composition and alloy type over time and geography [23–24]. South of this area, however, the provenance and circulation dynamics of early metals have never been researched on a meaningful scale. Consequently, fundamental questions concerning the role of metals in the major social transformations affecting the Italian peninsula in the 4th and 3rd millennia BC are still unanswered [25]. Some are surprisingly basic, viz. was all central Italian metalwork fashioned from local copper? How far did metal objects travel from the ore districts of Tuscany and Liguria? And was alien copper imported into the region? Answering these questions would enable us to interrogate the record with increasingly sophisticated queries regarding exchange modes and mechanisms. Was metal primarily circulated over short distances, perhaps by direct exchange? Or did it travel further afield through multiple handovers? If so, what was the reach of the network? Did early metals ‘make the world go round’, as Pare [26] and many others suggest they did in the Bronze Age, or did they perform more restricted social functions prior to this time? This article provides a science-based, and theoretically informed, pathway towards addressing these questions. Its aim is to assess metalwork circulation and exchange in 4th and 3rd millennia BC central Italy through the critical application of LIA [27–29]. Aside from sporadic attempts based on very few samples [30–31], this analytical method has not yet been applied to this region. Importantly, the isotopic data will be cross-referenced with, and augmented by, archaeological and copper chemistry datasets, which are indispensable for testing the provenance hypotheses generated by LIA [22,28–29,32]. As will be shown below, this multi-pronged approach has yielded original insights into the social dynamics of copper procurement and displacement in one of the most important mining and metalworking districts of early Europe.
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Post by Admin on Jan 24, 2020 20:00:53 GMT
Cultural context In the late and final Neolithic (4500–3600 BC), Italy partook in the wide-ranging communication networks and distinctive social phenomena characterising much of the central Mediterranean region (Table 1). These encompassed the long-distance exchange of obsidian from Lipari and other island sources; the emergence of a more mobile economic regime and lifestyle, which saw the abandonment of early/middle Neolithic nucleated villages and the transformation of the inhabited landscape towards smaller and shorter-lived settlements; the birth of formalised burial practices manifesting themselves in new funerary structures (e.g. rock-cut tombs) and mortuary customs (e.g. increasingly elaborate bone manipulation rituals); and a suite of technological innovations including the extractive metallurgy of copper [25,33–34]. Furthermore, new techniques of flint pressure-flaking were introduced at this time for manufacturing tanged-and-barbed arrowheads and daggers, and new stone materials such as jasper and steatite (also known as soapstone) replaced long-revered ‘greenstone’ and obsidian. Finally, the horse, plough, and wheel were introduced into the peninsula in the late 4th millennium BC, increasing agricultural output and perhaps triggering population growth [25,35]. Pottery styles provide one of the best indicators for changing communication networks from the late Neolithic to the Copper Age. In the former period, central Italy featured a marked east-west split in its cultural manifestations. East of the Apennines, late Neolithic potters fashioned bowls and plates in the Late Ripoli style, which is characterised by either plain or burnished surfaces. West of the range, however, craftspeople shaped and decorated their wares based on original combinations of the eastern Late Ripoli, north-western Chassey-Lagozza, and southern Diana ceramic styles [36–39] (Fig 2). This highlights the role of west-central Italy as a crossroads connecting north-western Italy, the mid-Adriatic coast, and the southern peninsula. Significantly, of all major ceramic styles documented in late Neolithic Italy, north-eastern Square-Mouthed Pottery, phase III (SMP III), is the sole not to be found in the region. This provides important evidence regarding the lack of communication networks linking north-east and west-central Italy. As we shall see below, this bears important implications for the unfolding of metalwork exchange in the Copper Age. Fig 2. Late Neolithic ceramic styles from west-central Italy. A: Late Ripoli pottery from Casale di Valleranello, Rome. B: Pottery from Quadrato di Torre Spaccata, Rome; this is mostly in the northern Chassey-Lagozza style, but the bottom left vessel has a rocchetto (spool-shaped) handle typical of southern Diana wares (image: Giovanni Carboni). The mid-4th millennium BC marked a ‘perfect split’ between domestic and funerary pottery production on both sides of the peninsula. On the one hand, the late Neolithic predilection for glossy surfaces was carried into the realm of mortuary practices, which acquired unprecedented social centrality in the Copper Age [34,40–41]. On the other hand, the taste for coarse impasto wares with textured surfaces, which had gained momentum in the final Neolithic, continued to thrive at open-air villages and other habitation sites. Extending previously localised trends to the entire central peninsula, Copper Age domestic ceramics were shaped and ornamented according to a bewildering array of stylistic traits, which household and village potters blended into idiosyncratic material assemblages [42]. As a counterpoint to the pulverisation of domestic pottery styles, superregional funerary styles emerged at this time throughout Italy. The process is mirrored by the burial customs themselves, which, from the mid-4th millennium BC, increasingly shared common traits over wide areas. Two such customs emerged in central Italy: interment in caves, or ‘Vecchiano’ tradition, and burial in small hypogeal chamber graves (occasionally supplemented by trench and cist tombs), or ‘Rinaldone’ tradition. The former is mainly found along the Tyrrhenian littoral from eastern Liguria to southern Tuscany, while the latter clusters at three principal foci in southern Tuscany/northern Latium, the lower Tiber valley, and the Adriatic lowlands [43–47]. In burial caves, mortuary rites centred on the inhumation of articulated bodies followed by bone manipulation and fragmentation, once the flesh had naturally decayed. The archaeological outcome of such processes are thick deposits in which large amounts of bone fragments are mingled with potsherds, dismembered ornaments, and (mostly unbroken) flint and metal weapons. Articulated bodies are all but absent. Copper-alloy daggers and halberds were deposited with relatively high frequency at these sites, along with (necklace?) beads made from metallic silver and antimony, and also steatite and animal bone [48–49]. Metal axes, on the other hand, are relatively rare, as they were preferentially deposited in the open landscape [50]. Burial caves such as Grotta del Fontino, Grotta della Spinosa, and Grotta San Giuseppe provide prime examples of this funerary behaviour [51–54]. Fig 3. Articulated burial and dismembered human remains from Ponte San Pietro, tomb 22. The chamber tomb is typical of the Rinaldone burial custom, central Italy, c.3600-2200 BC. Reprinted from Miari 1995 under a CC BY licence, with permission from Monica Miari, original copyright 1995. The Rinaldone burial tradition is typified by small hypogeal cemeteries comprising clusters of rock-cut chambers accessed through short shafts or entrance corridors. The tombs normally hosted small numbers of bodies, both articulated and reorganised. Stereotyped grave sets were used to mark out gender, age, and other aspects of personal identity [34,40,55]. Typically, adult males were given copper-alloy daggers and halberds (along with flint and hardstone weapons), while women and children were buried with silver and antimony beads, as well as non-metal ornaments [56–57]. Copper awls were placed with all gender and age categories, and so were flask-shaped funerary vessels [14]. Numerous Rinaldone-style burial sites are known in central Italy. Among the most representative of the funerary behaviour described above are Fontenoce di Recanati in Adriatic Italy and Ponte San Pietro (Fig 3), Selvicciola, Rinaldone, Lucrezia Romana, and Romanina in Tyrrhenian Italy [8,43,58–65].
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