Late Pleistocene Assemblages Research Articles

Beyond Large-Shaped Tools: Technological Innovations and Continuities at the Late Early Pleistocene Assemblage of El Barranc de la Boella (Tarragona, Spain)

The emergence of the mode 2 technocomplex, traditionally characterized by the development of large cutting tools (LCTs) or standardized large-shaped tools, has also implied a range of new technological behaviours. These include enhanced raw material management, more sophisticated knapping strategies, and transport patterns, among other innovations. Unit II of the localities of La Mina, El Forn, and Pit 1 in El Barranc de la Boella (Tarragona, Spain) — dating back to between 0.99–0.78 million years ago — contains the oldest mode 2 assemblages in the European subcontinent and represents an exceptional opportunity to examine the technological features associated with the appearance of LCTs in Europe and to provide clues to shed light to different hypotheses about their appearance. The presence of functionally diverse but environmentally similar localities enables a study of behavioural flexibility and technological variation, marking a critical contribution to understanding early human technological evolution. Our research focuses on the technological behaviours of hominins in unit II of Pit 1, La Mina, and El Forn at Barranc de la Boella, examining five keys technological aspects such as (1) raw material management, (2) core reduction sequences, (3) reduction intensity, (4) large flake production, and (5) the spatiotemporal integrity of reduction sequences. We compare these behaviours with those observed in mode 1 (~ 1.8–0.9 Ma) and early Middle Pleistocene (~ 0.78–0.6 Ma) assemblages across western Europe. Our findings reveal significant advancements and anticipatory behaviours at El Barranc de la Boella, such as the use of procedural templates for core reduction strategies and the production of large-shaped tools governed by principles of gestural economy, and also the existence of different reduction sequences in different raw materials according to specific purposes. The technological behaviours observed at El Barranc de la Boella seem to be more indicative of population dispersals rather than local evolutionary developments from mode 1 technologies. The non-linear evolution of core knapping strategies, along with the variability in large-shaped tool types, suggests multiple waves of hominin dispersals into Europe during this crucial period. We propose that El Barranc de la Boella may represent an early dispersal of the Acheulean from Africa around 1.4 million years ago, potentially connected to assemblages such as ‘Ubeidiya. This study underscores the complexity of lithic technology during this period and contributes significantly to our understanding of the complex emergence and adoption of new technological behaviours in European early mode 2 assemblages, extending beyond the mere appearance of LCTs.

Palynological characteristics of plant communities in the eastern Arctic during the Early to Middle Calabrian Age

Palynological analysis of Early Pleistocene sediments from Lake El'gygytgyn indicate that climate was warmer than present between c. 1.2860 and 1.6975 Ma (late Gelisian–early Calabrian ages), although variations in the paleovegetation indicate fluctuations between relatively cool and warm conditions. During the coolest intervals, the vegetation on the Anadyr Plateau was a mix of Betula-Salix shrub tundra and Larix forest-tundra. Larix forests, which probably included trees species of Betula and Alnus, characterized the regional vegetation during the warmest times. Slightly cooler interglaciations are indicated by the presence of Larix-forest tundra. Pinus pumila pollen is not consistently present in all interglaciations, indicating that variations in the extent and/or duration of snow cover occurred during these warm intervals. The absence of pollen from this evergreen shrub contrasts with Holocene and Late Pleistocene assemblages, where the taxon is a hallmark of warm conditions. During the Early Pleistocene, Betula pollen indicates the plant's presence during warm and cool intervals, in contrast to Late and Middle Pleistocene spectra where shrub Betula is absent or rare. The pollen data from MIS 55 suggest that it was the coolest of the Pleistocene “super” interglaciations and that the MIS 43 climate was as warm as that of MIS 55. The El'gygytgyn palynological record is a powerful reminder that the distribution of arctic communities can be vastly reduced or eradicated during warm Earth scenarios.

A comparative study of the Early Pleistocene carnivore guild from Dmanisi (Georgia)

The carnivore guild of the Early Pleistocene site of Dmanisi is among the most diverse of the Early Pleistocene of the entire Old World. It includes 14 carnivoran taxa: Homotherium latidens, Megantereon whitei, Panthera onca georgica, Acinonyx pardinensis, Lynx issiodorensis; Pachycrocuta brevirostris; Canis (Xenocyon) lycaonoides, Canis borjgali, Vulpes alopecoides; Ursus etruscus; Lutra sp., Martes sp., Meles sp., and Pannonictis sp. The analysis of this rich carnivore guild was carried out under different methodological approaches to compare the assemblage with other chronological coeval European, Asian, and African sites from a paleobiological perspective. To achieve the goal, we used a permutational hierarchical method called boostrapping cluster analysis based on taxonomic absence/presence matrices (at both generic and specific level) and on ecological matrices (considering dietary preferences/hunting strategies of each carnivoran) and carried out Mantels tests assessing magnitude of time, space, ecology, and taxonomy as source of difference between guilds. Our results suggest a close similarity among the Dmanisi carnivore assemblage and other guilds recorded from European late Villafranchian sites such as Pirro Nord, Venta Micena, and Apollonia 1 and, in a lesser extent, to European Epivillafranchian sites as Vallonnet, Untermassfeld, or the Vallparadís Section. Early to Middle Pleistocene Asian carnivore assemblages display several similarities with the Dmanisi guild mainly in the record and diversity of felid and the canid ecomorphotypes. Eastern African sites such as Olduvai and Omo, as well as South African sites, display a lower similarity with the studied sample, basically for the most diverse hyenid taphocoenoses. To sum up, the present study suggests a close similarity between the Dmanisi carnivore guild and other European Late Early Pleistocene assemblages without close parallels with African or Asian assemblages.

Analysis of Late Pleistocene megafauna and puparia from the Lent dredging site, province of Gelderland (the Netherlands)

Abstract More than 900 vertebrate bones, ranging from Late Pleistocene to Holocene in age, have been identified in a collection that was recovered by a single dredging operation for the construction of artificial lakes near Lent (Nijmegen, province of Gelderland, the Netherlands). The Late Pleistocene assemblage comprises mainly Weichselian glacial fauna such as mammoths, reindeer and bison. Some Eemian fauna is represented as well, e.g. straight-tusked elephant. The abundance of certain species over others suggests that preservation bias had a considerable impact on this assemblage, while its time-averaged nature resulted in overrepresentation of certain species. A case study is here conducted on a fragmentary skull of a subadult woolly mammoth bull with embedded blowfly puparia. Some of these puparia are fully developed, indicating prolonged exposure of the mammoth carcass.

Frugivore-fruit size relationships between palms and mammals reveal past and future defaunation impacts

Mammalian frugivores are critical seed dispersers, but many are under threat of extinction. Futhermore, the impact of past and future defaunation on plant assemblages has yet to be quantified at the global scale. Here, we integrate palm and mammalian frugivore trait and occurrence data and reveal a global positive relationship between fruit size and frugivore body size. Global variation in fruit size is better explained by present-day frugivore assemblages than by Late Pleistocene assemblages, suggesting ecological and evolutionary reorganization after end-Pleistocene extinctions, except in the Neotropics, where some large-fruited palm species may have outlived their main seed dispersers by thousands of years. Our simulations of frugivore extinction over the next 100 years suggest that the impact of defaunation will be highest in the Old World tropics, and an up to 4% assemblage-level decrease in fruit size would be required to maintain the global body size–fruit size relationship. Overall, our results suggest that while some palm species may be able to keep pace with future defaunation through evolutionary changes in fruit size, large-fruited species may be especially vulnerable to continued defaunation.

Extinction of North AmericanCuvieronius(Mammalia: Proboscidea: Gomphotheriidae) driven by dietary resource competition with sympatric mammoths and mastodons

AbstractThe gomphotheres were a diverse and widespread group of proboscideans occupying Eurasia, North America, and South America throughout the Neogene. Their decline was temporally and spatially heterogeneous, and the gomphotheres ultimately became extinct during the late Pleistocene; however, the genusCuvieroniusis rarely represented in late Pleistocene assemblages in North America. Two alternative hypotheses have been invoked to explain this phenomenon: (1) competitive exclusion by sympatric mammoths and mastodons or (2) ecologic displacement due to an environmental transition from closed forests to open grasslands. To test whether competition for resources contributed to the demise of North AmericanCuvieronius, we present herein a large collection of stable isotope and dental microwear data from populations occupying their Pleistocene refugium in the Atlantic Coastal Plain. Results suggest thatCuvieroniusconsumed a wide range of resources with variable textural and photosynthetic properties and was not specialized on either grasses or browse. Further, we document evidence for the consumption of similar foods between contemporaneous gomphotheres, mammoths, and mastodons. The generalist feeding strategy of the gomphotheres likely facilitated their high Miocene abundance and diversity. However, this “jack of all trades and master of none” feeding strategy may have proved challenging following the arrival of mammoths and likely contributed to the extirpation ofCuvieroniusin North America.

Трасологические исследования комплексов позднего плейстоцена и раннего голоцена Северо-Восточной Азии. История и современное состояние

Трасологические исследования комплексов позднего плейстоцена и раннего голоцена Северо-Восточной Азии. История и современное состояние

Molluskan Grazing Traces (Ichnogenus Radulichnus Voigt, 1977) on a Pleistocene Bivalve from Southern Brazil, With the Proposal of a New Ichnospecies

The ichnogenus Radulichnus Voigt, 1977 is recorded for the first time from a bivalve, Anomalocardia brasiliana (Gmelin, 1791), in a late Pleistocene molluskan assemblage from the southern Brazilian coastal plain. Grazing traces comprise short (<1 mm), parallel furrows, arranged in rows on the inner (concave) shell surface and mostly concentrated in its central area. Radulichnus accommodates scratches on hard substrates produced by the radula of grazing gastropod or polyplacophorans. Our literature survey on fossil and extant traces, as well as studies on the grazing mechanism in living mollusks, document at least two distinct morphotypes that are related to differences in the feeding modes of the producers. We propose to distinguish a second ichnospecies of Radulichnus, in addition to the type, R. inopinatus Voigt, 1977 (produced by gastropods), which is named R. transversus isp. nov., and attributed to polyplacophorans. Grazing traces on the shell of A. brasiliana match the morphotype produced by polyplacophorans mollusks, and are indicative of its complex taphonomic history in comparison with other shells in this assemblage.

Late Pleistocene lithic technology and human adaptation in Northeast China: A case study from Taoshan site

Late Pleistocene witnessed significant changes in lithic technological behaviors, mobility patterns and subsistence strategies in northeastern Eurasia, which are also correlated with paleoclimatic and paleoenvironmental changes. Integrating artifact analysis and paleoenvironmental evidence, Taoshan, a newly excavated archaeological site, provides a crucial case study for evaluating human adaptation in Northeast China. Taoshan site was excavated in 2013–2014, uncovering three prehistoric layers and abundant remains. Here we present the results of a systematic techno-typological analysis of the Late Pleistocene assemblages of the site, and place special emphasis on diachronic cultural developments and underlying dynamics.

Ungulate dietary traits and plasticity in zones of ecological transition inferred from late Pleistocene assemblages at Jou Puerta and Rexidora in the Cantabrian Region of northern Spain

The diet and habitats of Late Pleistocene ungulate taxa were analyzed in the caves of Jou Puerta and Rexidora (Marine Isotope Stage, MIS 3), in the Cantabrian Region of northern Spain, a zone of ecological transition between mainland Europe and Central-Southern Iberia. The two ungulate assemblages, which are composed mainly by cold-adapted species, were analyzed through tooth mesowear and microwear. The Jou Puerta community is dominated by the leaf browsers while at Rexidora the dominant taxa are grazers that suggest an open landscape with availability of grass, but also with significant presence of trees and/or forbs. Our results suggest an open landscape with presence of arboreal plant taxa during the coldest events of the MIS3. Concerning dietary plasticity, the diets for the cold-adapted woolly rhino and the reindeer in populations that lived in the southwestern boundary of their Eurasian distribution fit within the plasticity observed for other populations during the Pleistocene. It is proposed here that diet was not a limiting factor for the populations that inhabited the limit of their known range of distribution in the Cantabrian Region.

Lithic miniaturization in Late Pleistocene southern Africa

Lithic miniaturization refers to the systematic production and use of small tools made from small cores. Archaeologists typically associate the production of miniature toolkits with small retouched tools or ‘microliths’. Yet, ethnographic accounts and archaeological data show that miniature toolkits also comprised small unretouched flakes and bladelets. Because it proliferates during the Late Pleistocene and is associated with the Later Stone Age in Africa, archaeologists associate lithic miniaturization with ‘modern human behavior’. However, toolkit miniaturization is no more consistent a feature of Late Pleistocene assemblages than bone point production, rock art, or engraved ochre. Lithic miniaturization was one of many strategic options our Pleistocene ancestors employed. The paper reports a miniaturized lithic assemblage marked by the systematic production of small unretouched flakes and bladelets from small cores (<5g) during the Late Pleistocene (c. 14–12ka calBP) at Ntloana Tšoana rock shelter in Lesotho. This assemblage was made on a range of locally available rock types using freehand and bipolar reduction. Analogous lithic assemblages are reported from elsewhere in southern Africa suggesting that unretouched miniaturized toolkits were widespread. The process of systematic unretouched small tool production could be ancient, but the extent of these processes remain unknown. Using appropriate terminology combined with contextual approaches to defining and quantifying lithic miniaturization while demonstrating the intent to miniaturize toolkits provide initial solutions to this problem.

Bone lesions in a Late Pleistocene assemblage of the insular deer Candiacervus sp.II from Liko cave (Crete, Greece)

Candiacervus sp.II is one of the deer species that inhabited the island of Crete during the Late Pleistocene. The species evolved on the island under a prolonged period of isolation and, as a consequence, developed a high degree of endemism. Fossils of this species have been discovered at many Cretan sites, including Liko cave (an attritional accumulation of several thousand fossils). In this paper, we present the results of a systematic analysis of the prevalence and anatomical distribution of bone lesions of Candiacervus sp.II, from that cave. We identified one metapodial with a healed fracture and nine (various) specimens with moderate to severe degenerative lesions of osteoarthritis. The lesions were evaluated macroscopically and radiographically, and they were classified as traumatic or degenerative. Degenerative lesions that affected adult individuals had prevalence rates below 5% and were attributed to environmental or nutritional causes. Representative bones were sampled for histological evaluation, to provide essential baseline data on possible underlying disorders. The aims of this study are to provide evidence for bone disease contributing to species morbidity, and to shed new light on causes and potential palaeoecological significance.

Is the enamel differentiation quotient (SDQ) of the narrow-headed vole (Microtus gregalis) useful for Pleistocene biostratigraphy?

In order to explore temporal trends of morphological variation in the narrow-headed vole (Microtus gregalis) traditional morphometric methods were used to quantify enamel differentiation quotient (SDQ), tooth size (L) and tooth proportion (A/L) in over 800 first lower molars from two sites in Central Europe. Our results suggest that SDQ values in late Middle Pleistocene assemblages are significantly lower than those in Late Pleistocene populations. A greater chronostratigraphic variation in SDQ occurs across the Late Pleistocene compared to the late Middle Pleistocene. This probably indicates repeated extinctions and re-colonisations, most likely from Eastern Europe or Asia, during MIS 3 and MIS 2. However, it is unclear whether this pattern represents a local phenomenon or whether the SDQ changes took place across a large geographical scale. Further studies on fossil and recent populations of the species are necessary to identify the determinants of SDQ variation in Microtus gregalis and to confirm the usefulness of the index for the Pleistocene stratigraphy.

Post mortem modifications (pseudopaleopathologies) in middle-late Pleistocene mammal fossils from southern Brazil

Here are described post mortem modifications (pseudopaleopathologies) in 762 specimens of extinct mammals from three middle to late Pleistocene assemblages (Campanha, Chui Creek and continental shelf) of the state of Rio Grande do Sul, southern Brazil. These assemblages include mostly large-sized herbivores preserved in fluvial facies, although the assemblage from the shelf consist of remains that were reworked several times by sea-level oscillations. Although some patterns of modification are similar between the assemblages (e.g. post-fossilization plain fractures), others such as the grade of abrasion or weathering varies according to the depositional setting. The scarcity of modifications that could be attributed to predators or scavengers indicate that carnivory was not a major process driving the formation of the assemblages. Most of the fossils from the Campanha are complete, but a relatively high proportion of them exhibits advanced stage of weathering, whereas those from Chui Creek and the continental shelf are mostly unweathered. Most of the remains from the Campanha and Chui Creek are unabraded, whereas all fossils from the shelf were subject to abrasion. Few fossils exhibit signs of modification by organisms such as plants and insects. Fossils from the continental shelf have crusts of beach rock, and some specimens from the Campanha and Chui Creek have crusts of calcium carbonate or iron oxide. Besides the abrasion, the main difference between fossils from the continental and marine assemblages is the dark color and higher mechanical resistance of the latter. This difference is related to post depositional geochemical processes in the marine environment. The chemical composition of fossils from Chui Creek and continental shelf was qualitatively analyzed using energy dispersive X-ray spectrometry (EDX) and compared with recent bones. The results show the presence in the fossils of several elements not found in recent bones, incorporated through diagenetic processes controlled by specific conditions of each depositional environment.

What is the taphonomic agent responsible for the avian accumulation? An approach from the Middle and early Late Pleistocene assemblages from Payre and Abri des Pêcheurs (Ardèche, France)

The acquisition of quick-flying small prey has been a widely discussed topic over the last decades, especially since the emergence of archaeological bird collections more ancient than expected, with evidence of anthropogenic processing. Residue analyses carried out on stone tools at the Payre site in France (MIS 8–5) suggest human activity on birds; however, no direct analyses have been carried out on avian specimens from the site. Similarly, no detailed data from numerous bird bones have been obtained from a nearby site, Abri des Pêcheurs (MIS 5–3), which could also provide important information about subsistence strategies in the region. Bird bones from both sites were analyzed here within the framework of the taphonomic methodology and identified a wide variety of avian specimens, suggesting nocturnal raptors and small mammal carnivores as main accumulator agents. Despite the anthropogenic activity at Payre proposed from lithic residue and use-wear analyses, bird bones do not reveal any hominid intervention. Our attempt here is to explain this apparent contradiction and discuss possible hypotheses regarding the presence of avian specimens at both archaeological sites.

Differential insect and mammalian response to Late Quaternary climate change in the Rocky Mountain region of North America

Of the 200 beetle species identified from Rocky Mountain Late Pleistocene insect faunal assemblages, 23% are no longer resident in this region. None of the 200 species is extinct. In contrast to this, only 8% of 73 identified mammal species from Rocky Mountain Late Pleistocene assemblages are no longer resident in the Rockies, and 12 species are now extinct. Since both groups of organisms are highly mobile, it would appear that their responses to the large-scale fluctuations of climate associated with the last 125,000 years have been considerably different. Most strikingly contrasting with the insects, there are no mammals in the Rocky Mountain Late Pleistocene fossil record that are found exclusively today in the Pacific Northwest (PNW) region. The PNW does have a distinctive modern mammalian fauna, but only one of these, Keen's Myotis, has a fossil record outside the PNW region, in the eastern and central United States. No modern PNW vertebrate species have been found in any Rocky Mountain fossil assemblages. Based on these data, it appears that there has been little or no mammalian faunal exchange between the PNW region and the Rocky Mountains during the Late Pleistocene or Holocene. This is in stark contrast to the fossil beetle record, where PNW species are a substantial component in many faunas, right through to the Late Holocene.

Continental-style avian extinctions on an oceanic island

Abstract The Indonesian islands have long been recognized for their rich and unique avifaunas, but little is known regarding how past processes and events have shaped current avian distributions. Here we compare the modern non-passerine avifauna of Flores with the Late Pleistocene non-passerine fossil assemblage from the cave site of Liang Bua to assess whether the Late Pleistocene assemblage differs from the modern avifauna. Randomized permutation tests failed to detect a statistically significant difference in body size distributions, but a significant difference in dietary guild was found, as the modern fauna lacks scavengers. The emerging pattern of avian extinctions on Flores is characterized by a low proportion of extinct species, a loss of large-bodied species, and apparently minor effects on avian community structure. This is in contrast to other oceanic islands, which experienced dramatic changes in avifauna after the arrival of modern humans. Flores' close proximity to other islands and landmasses likely allowed for population connectivity that buffered populations from extinction. Widespread species may also have been able to recolonize if local extirpations took place. The extinction of the large-bodied avian scavengers Leptoptilos robustus and Trigonoceps sp. on Flores is consistent with the pattern of human-caused extinctions on other oceanic islands. However, the loss of these two large scavenging species may be linked to the extinction of the pygmy proboscidean ( Stegodon florensis insularis ). Such a dependence of avian species on mammalian megafauna, leading to extinction by trophic cascade, is characteristic of continental Late Pleistocene extinctions.

Constraint envelope analyses of macroecological patterns reveal climatic effects on Pleistocene mammal extinctions

Quantitative analysis of macroecological patterns for late Pleistocene assemblages can be useful for disentangling the causes of late Quaternary extinctions (LQE). However, previous analyses have usually assumed linear relationships between macroecological traits, such as body size and range size/range shift, that may have led to erroneous interpretations. Here, we analyzed mammalian datasets to show how macroecological patterns support climate change as an important driver of the LQE, which is contrary to previous analyses that did not account for more complex relationships among traits. We employed quantile regression methods that allow a detailed and fine-tuned quantitative analysis of complex macroecological patterns revealed as polygonal relationships (i.e., constraint envelopes). We showed that these triangular-shaped envelopes that describe the macroecological relationship between body size and geographical range shift reflect nonrandom extinction processes under which the large-bodied species are more prone to extinction during events of severe habitat loss, such as glacial/interglacial transitions. Hence, we provide both a theoretical background and methodological framework to better understand how climate change induces body size-biased species sorting and shapes complex macroecological patterns.

The avian fossil record in Insular Southeast Asia and its implications for avian biogeography and palaeoecology

Excavations and studies of existing collections during the last decades have significantly increased the abundance as well as the diversity of the avian fossil record for Insular Southeast Asia. The avian fossil record covers the Eocene through the Holocene, with the majority of bird fossils Pleistocene in age. Fossil bird skeletal remains represent at least 63 species in 54 genera and 27 families, and two ichnospecies are represented by fossil footprints. Birds of prey, owls and swiftlets are common elements. Extinctions seem to have been few, suggesting continuity of avian lineages since at least the Late Pleistocene, although some shifts in species ranges have occurred in response to climatic change. Similarities between the Late Pleistocene avifaunas of Flores and Java suggest a dispersal route across southern Sundaland. Late Pleistocene assemblages of Niah Cave (Borneo) and Liang Bua (Flores) support the rainforest refugium hypothesis in Southeast Asia as they indicate the persistence of forest cover, at least locally, throughout the Late Pleistocene and Holocene.

Late Pleistocene-Holocene non-passerine avifauna of Liang Bua (Flores, Indonesia)

ABSTRACT Liang Bua, a limestone cave on the Indonesian island of Flores, has a depositional sequence that spans the last 95,000 years and includes well-preserved faunal remains. Birds are well represented throughout the stratigraphic sequence at Liang Bua. Here, we present the results of the first comprehensive study of avian remains retrieved from Sector XI, a 2 m by 2 m archaeological excavation along the east wall of the cave. A total of 579 specimens were identified as avian, with 244 belonging to at least 26 non-passerine taxa in 13 families. The late Pleistocene assemblage (23 taxa) includes the first recorded occurrence of vultures in Wallacea, as well as kingfishers, snipes, plovers, parrots, pigeons, and swiftlets. Together, these taxa suggest that during this time the surrounding environment was floristically diverse and included several habitat types. Two of these taxa, the giant marabou Leptoptilos robustus and the vulture Trigonoceps sp., are extinct. Eight taxa were identified in the Holocene assemblage, and five of these were also present in the late Pleistocene. Imperial pigeons Ducula sp. and the Island Collared Dove Streptopelia cf. bitorquata appear only in the Holocene assemblage. The differences in faunal composition between the late Pleistocene and Holocene assemblages may reflect a change in avifaunal composition due to climatic and environmental changes near the Pleistocene–Holocene transition, possibly amplified by impacts associated with the arrival of modern humans; however, the small Holocene sample prevents a firm conclusion about faunal turnover from being made.