The genomic study of specimens dating to the Early and Middle Pleistocene (EP and MP), a period spanning from 2.6 million years ago (Ma) to 126 thousand years ago (ka), has the potential to elucidate the evolutionary processes that shaped present-day biodiversity. Obtaining genomic data from this period is challenging, but mitochondrial DNA, given its higher abundance compared to nuclear DNA, could play an important role to understand evolutionary processes at this time scale. In this study, we report 34 new mitogenomes, including two EP and nine MP mammoth (Mammuthus spp.) specimens from Siberia and North America and analyze them jointly with >200 publicly available mitogenomes to reconstruct a transect of mammoth mitogenome diversity throughout the last million years. We find that our EP mitogenomes fall outside the diversity of all Late Pleistocene (LP) mammoths, while those derived from MP mammoths are basal to LP mammoth Clades 2 and 3, supporting an ancient Siberian origin of these lineages. In contrast, the geographical origin of Clade 1 remains unresolved. With these new deep-time mitogenomes, we observe diversification events across all clades that appear consistent with previously hypothesized MP and LP demographic changes. Furthermore, we improve upon an existing methodology for molecular clock dating of specimens >50 ka, demonstrating that specimens need to be individually dated to avoid biases in their age estimates. Both the molecular and analytical improvements presented here highlight the importance of deep-time genomic data to discover long-lost genetic diversity, enabling better assessments of evolutionary histories.

Historically, the extensive involvement of citizen scientists in palaeontology and archaeology has resulted in many discoveries and insights. More recently, machine learning has emerged as a broadly applicable tool for analysing large datasets of fossils and artefacts. In the digital age, citizen science (CS) and machine learning (ML) prove to be mutually beneficial, and a combined CS-ML approach is increasingly successful in areas such as biodiversity research. Ever-dropping computational costs and the smartphone revolution have put ML tools in the hands of citizen scientists with the potential to generate high-quality data, create new insights from large datasets and elevate public engagement. However, without an integrated approach, new CS-ML projects may not realise the full scientific and public engagement potential. Furthermore, object-based data gathering of fossils and artefacts comes with different requirements for successful CS-ML approaches than observation-based data gathering in biodiversity monitoring. In this review we investigate best practices and common pitfalls in this new interdisciplinary field in order to formulate a workflow to guide future palaeontological and archaeological projects. Our CS-ML workflow is subdivided in four project phases: (I) preparation, (II) execution, (III) implementation and (IV) reiteration. To reach the objectives and manage the challenges for different subject domains (CS tasks, ML development, research, stakeholder engagement and app/infrastructure development), tasks are formulated and allocated to different roles in the project. We also provide an outline for an integrated online CS platform which will help reach a project’s full scientific and public engagement potential. Finally, to illustrate the implementation of our CS-ML approach in practice and showcase differences with more commonly available biodiversity CS-ML approaches, we discuss the LegaSea project in which fossils and artefacts from sand nourishments in the western Netherlands are studied.

Pollen analysis of five teeth (two premolars, three molars) from a single maxilla of a giant deer (Megaloceros giganteus) found in Pleistocene deposits in The Netherlands reflects the diet and the landscape of the specimen that lived in eastern Doggerland. Apiaceae (among which Anthriscus sylvestris, Heracleum and Hydrocotyle), Asteraceae (among which Anthemis-type, Senecio-type and Cichorioideae), Filipendula, Poaceae and Symphytum were among the ingested plants. The landscape had dense, species-rich tall-herb vegetation and an open tree layer of Alnus and Betula, whereas thermophilous tree taxa were absent. Climate was probably cool-temperate, semi-dry, sub-continental. A radiocarbon date of the maxilla is beyond the 14C detection limit. We conclude that our giant deer most likely lived during the early Eemian or during an early Weichselian interstadial.

ABSTRACT Three hundred forty‐one radiocarbon dates from the Groningen radiocarbon database are compiled in this study. They show for the first time that organic sediment samples from the eastern Netherlands and mammal bones from Doggerland reflect shifts in the presence and the density of vegetation (food for herbivores) and mammal biomass during the last ice age (Weichselian Stage, ~119–14.7 ka cal bp ). Comparison with oxygen isotope curves of Greenland ice cores and geomorphological data shows that cold climate, in particular during the younger part of the Weichselian Middle Pleniglacial and during the Late Pleniglacial, and related scarcity or even absence of vegetation, were limiting factors for the carrying capacity of the landscape and thus for the population density of large herbivores during the period covered by 14 C dating (last ca. 55 000 years). A ‘fossil gap’ during the Late Pleniglacial lasted ca. 13 000 years from ca. 28 to 15 ka cal bp . Previous research from the nearby Eifel region in Germany shows that environmental conditions were less extreme (‘refugium conditions’) than in the Netherlands, taking into account the continuous presence of spores of coprophilous fungi in the Eifel, indicating uninterrupted food supply for herbivores.

Hormones in biological media reveal endocrine activity related to development, reproduction, disease and stress on different timescales1. Serum provides immediate circulating concentrations2, whereas various tissues record steroid hormones accumulated over time3,4. Hormones have been studied in keratin, bones and teeth in modern5-8 and ancient contexts9-12; however, the biological significance of such records is subject to ongoing debate10,13-16, and the utility of tooth-associated hormones has not previously been demonstrated. Here we use liquid chromatography with tandem mass spectrometry paired with fine-scale serial sampling to measure steroid hormone concentrations in modern and fossil tusk dentin. An adult male African elephant (Loxodonta africana) tusk shows periodic increases in testosterone that reveal episodes of musth17-19, an annually recurring period of behavioural and physiological changes that enhance mating success20-23. Parallel assessments of a male woolly mammoth (Mammuthus primigenius) tusk show that mammoths also experienced musth. These results set the stage for wide-ranging studies using steroids preserved in dentin to investigate development, reproduction and stress in modern and extinct mammals. Because dentin grows by apposition, resists degradation, and often contains growth lines, teeth have advantages over other tissues that are used as records of endocrine data. Given the low mass of dentin powder required for analytical precision, we anticipate dentin-hormone studies to extend to smaller animals. Thus, in addition to broad applications in zoology and palaeontology, tooth hormone records could support medical, forensic, veterinary and archaeological studies.

We studied the pollen content of a well-preserved coprolite of a Late-Glacial giant ground sloth (Mylodon darwinii) from the Mylodon Cave, province Ultima Esperanza, southern Chile. The specimen was obtained in 1909 and has been stored in a museum in the Netherlands since. It was radiocarbon dated to 13,140 +/- 55 BP (15,927-15,522 cal BP), which fits with other radiocarbon dates showing the early Late-Glacial presence of M. darwinii in the province Ultima Esperanza. Contemporaneous oxygen isotope data from Antarctic EPICA Dome C indicates that our Mylodon specimen lived during a warming phase of the Late-Glacial, ca. 1000 years before the start of the Antarctic Cold Reversal. We compared our pollen data with pollen records showing contemporaneous regional vegetation and discuss the uncertainties in the interpretation of pollen spectra fromfaeces. To expand on the pollen data, we tested ancient DNA preservation in the sample; we sequenced similar to 9.4 million DNA reads and found that the concentration of ancient plant DNA is below detectable levels. Pollen analysis confirms earlier findings that the Mylodon was a grazer, but the discovery of large amounts of Fragaria and Azorella pollen in the faeces may indicate that Mylodon was also able to select and consume specific plants, and therefore could also be regarded as a selective feeder. (C) 2021 The Author(s). Published by Elsevier B.V.

Dit artikel presenteert een onderkaakfragment van een wolvenpup (Canis lupus Linnaeus, 1758) dat gevonden werd op de Zandmotor tussen Monster en Kijkduin en met de 14C-methode gedateerd is op 47,6 - 42,1 duizend jaar geleden. Het is het derde gedateerde fossiel van een wolf uit het Nederlandse Noordzeegebied, en daarom een belangrijke bevestiging dat de wolf inderdaad deel uitmaakte van de laat-pleistocene mammoetfauna in het Noordzeegebied. Het artikel plaatst de vondst in de context van recente ontdekkingen van wolvenpups in de permafrost van Canada en Siberie en vooral van eerdere vondsten van wolven in Nederland en uit de Noordzee, waarvan de ouderdom doorgaans ingeschat werd op basis van kleur en fossilisatiegraad.

Molaren van acht soorten grote herbivoren (reuzenhert Megaloceros giganteus, edelhert Cervus elaphus, rendier Rangifer tarandus, eland Alces alces, steppewisent Bison priscus, muskusos Ovibos moschatus, wolharige neushoorn Coelodonta antiquitatis en bosneushoorn Stephanorhinus kirchbergensis) werden ex situ verzameld uit opgebaggerde pleistocene en Holocene zandafzettingen uit de Noordzee en uit Nederlandse vindplaatsen in het binnenland. In de plooien van vele molaren bevonden zich gekauwde plantenresten en daarbij ook microfossielen. We determineerden pollen (stuifmeel) en sporen en interpreteren op basis hiervan voedselkeuzes en de mogelijke effecten op de pollenspectra als een deel van het bloeiseizoen vertegenwoordigd is. We bediscussieren ook de effecten van de veranderende vegetatiesamenstelling in relatie tot het klimaat en de ouderdom van de molaren. Verschillende factoren hebben bijgedragen aan de samenstelling van het pollen. Toch geven de pollenspectra een beeld van de vegetatie en kunnen we de voedselkeuze en de ouderdom afleiden en krijgen we een beeld van de landschapstypen, van subarctische open gebieden tot interglaciaal bos. Ecologische en statistische analyses van de resultaten laten verschillen zien tussen de voedselkeuze van, en de beschikbaarheid van plantensoorten voor de verschillende zoogdiersoorten. Deze bijdrage is gebaseerd op een artikel dat eerder in Quaternary Science Reviews verscheen. -- Molars of eight large herbivore species (Megaloceros giganteus, Cervus elaphus, Rangifer tarandus, Alces alces, Bison priscus, Ovibos moschatus, Coelodonta antiquitatis and Stephanorhinus kirchbergensis) were collected ex situ from Pleistocene and Holocene sands dredged in the North Sea, and from Dutch inland sites. Folds in many molars contained compacted masticated plant remains, and also microfossils. We identified pollen, spores, and non-pollen palynomorphs and on this basis interpret food preferences, represented flowering seasons, or parts of flowering seasons. We also discuss effects of changing vegetation composition in relation to climate and age of the molars, based on the pollen spectra. Various confounding factors have contributed to the recorded pollen composition, but nevertheless the pollen spectra show valuable aspects of vegetation composition, food choice, age, and landscapes, from subarctic open areas to interglacial forest. Ecological and statistical analysis of the results shows dietary differences between the mammal species analyzed. This contribution is based on a paper that appeared in Quaternary Science Reviews.

Molars of eight large herbivore species (Megaloceros giganteus, Cervus elaphus, Rangifer tarandus, Alces alces, Bison priscus, Ovibos moschatus, Coelodonta antiquitatis and Stephanorhinus kirchbergensis) were collected ex situ from Pleistocene and Holocene sands dredged in the North Sea, and from Dutch inland sites. Folds in many molars contained compacted masticated plant remains, and also microfossils. We identified pollen, spores, and non-pollen palynomorphs and discuss and interpret food preferences, represented flowering seasons, or parts of flowering seasons, and we discuss effects of changing vegetation composition in relation to climate and age of the molars, based on the pollen spectra. Various confounding factors have contributed to the recorded pollen composition, but nevertheless the pollen spectra show valuable aspects of vegetation composition, food choice, age, and landscapes, from subarctic open areas to interglacial forest. Ecological and statistical analysis of the results shows dietary differences between the mammal species analyzed.

Food sources and techniques of food procurement in the Early Palaeolithic have long been of interest to researchers. However, this is a complex and debatable problem; its solution is complicated by a lack of relevant archaeological data. This article describes and interprets archaeological evidence from a time transgressive series of sites belonging to the Taman Early Acheulean industry (Tamanian) in the southern Azov Sea region (~2.0–1.0 Ma). Analysis of the finds leads to the conclusion that makers of the Tamanian industry were hunters of large mammals, mainly southern elephants (Archidiskodon meridionalis tamanensis) and Caucasian elasmotherium (Elasmotherium caucasicum). Active hunting of these mammals as a way to procure meat emerged thanks to the particular environmental conditions of the southern Azov region in the Lower Pleistocene and the cognitive capabilities of prehistoric people. A specific strategy was devised and applied, i.e. hunting inside calderas of mud volcanoes that were present in numbers across the region. People took into consideration the behaviour of elephants and elasmotherium which came to calderas in order to drink or ‘take a bath’ in volcanic muds. It allowed prehistoric people to effortlessly hunt animals, which were immobilized in spongy mud, and to procure substantial meat supplies. Alongside hunting of large mammals, the makers of the Tamanian industry engaged extensively in a special form of gathering, i.e. beach gathering of high-protein food like shellfish, carcasses of sea animals, and fish. This reliable source of nutrition likely was one of the main reasons why the Tamanian sites were located on beaches adjacent to coastal lagoons and estuaries.

Tusks of very young mammoths and other proboscideans are extremely rare in the fossil record. This article presents all deciduous tusks and newly-developing permanent tusks of woolly mammoths which are known from Dutch localities and the North Sea. Four deciduous tusks are from The Netherlands and one is from Germany. Five specimens of newly-developing permanent tusks are also discussed. In this article, we present several characteristics and details of those tusks such as presence/absence of roots, and cementum and enamel layers that should facilitate an identification when new findings are made. We also describe a dental element, a digitelle, that is easily mistaken for a tusk of a juvenile woolly mammoth.

(1) Background: Over the last decades, important fossil records of Late Pliocene mammals and reptiles have been unearthed in Milia, Grevena (N Greece). This assemblage shows a remarkable composition and diversity, including the partial skeletons of mastodons that reached record-breaking sizes, abundant rhinos-the first occurrence of this species in Greece-and some new species; (2) Methods: We perform a thorough quantitative analysis of the recovered assemblage, presenting the information of the various collection spots in Milia, and calculate various biodiversity indices for each spot; (3) Results: Our research has revealed the vast majority of expected taxa in the assemblage. We argue that the various sub-localities in Milia could be grouped into a larger, composite assemblage representing a short period. We analyze the diversity changes through the various localities in Milia and highlight potential barriers that could affect the distribution of taxa; (4) Conclusions: We re-affirm the Early Villafranchian affinities of the fauna. In particular, Milia should date at MN16a, with a surprising presence of some more archaic, Ruscinian taxa. The fossils of Milia depict a Late Pliocene paradise in the Southern Balkans; a paradise, unfortunately, lost with the onset of the dramatic climate changes of the Quaternary.

Artikel over vondsten van de Zandmotor die geplaatst kunnen worden in het Laat Pleistoceen en het Vroeg Holoceen. Gedetermineerd door Henk Mulder en Dick Mol met medewerking van Bram Langeveld

Middens (nests and caches) of Late Pleistocene arctic ground squirrels (Urocitellus parryii) that are preserved in the permafrost of Beringia archive valuable paleoecological data. Arctic ground squirrels selectively include the plant material placed in middens. To account for this selectivity bias, we used a multi-proxy approach that includes ancient DNA (aDNA) and macro- and microfossil analyses. Here, we provide insight into Pleistocene vegetation conditions using macrofossils, pollen, phytoliths and non-pollen palynomorphs, and aDNA collected from one such midden from the Yukon Territory (Canada), which was formed between 30,740 and 30,380 cal yr BP. aDNA confirmed the midden was constructed by U. parryii. We recovered 39 vascular plant and bryophyte genera and 68 fungal genera from the midden samples. Grass and other herbaceous families dominated vegetation assemblages according to all proxies. aDNA data yielded several records of vascular plants that are outside their current biogeographic range, while some of the recovered fungi yielded additional evidence for local occurrence of Picea trees during glacial conditions. We propose that future work on fossil middens should combine the study of macro- and microfossils with aDNA analysis to get the most out of these environmental archives.

ABSTRACTThe molar of a giant deer, Megaloceros giganteus (Blumenbach, 1799), was discovered in sandy deposits of the North Sea, about 10 km west of the present shoreline of the Netherlands. Compacted masticated plant remains were preserved in the molar's deep folds. A palaeoecological analysis of these plant remains shows the complete dominance of pollen from Artemisia (sage) and other Asteraceae Tubuliflorae (Compositae), indicating that the animal foraged in a steppe environment, and may have preferred to eat Artemisia, which contain a high level of nutrients such as calcium and phosphorous components, which are important for antler building. Radiocarbon dating indicates that the plant remains are of mid‐Weichselian age [38 570 (+300, −290) 14C a BP, Greenland Interstadial‐11]. This is the first study of the food choice of M. giganteus based on palaeoecological evidence. We hypothesize about links between the extirpation of M. giganteus at the onset of the Holocene in North‐West Europe and the transition from a landscape with highly dynamic geomorphological processes and consequently prevalence of immature and nutrient‐rich soils, to a less dynamic landscape with stable, leached and acidifying soils, and the decline of Artemisia and other calciphilous plants.

The first record of arctic fox Alopex lagopus (Linnaeus, 1758) from the Eurogeul area (North Sea) is reported based on a distal humerus fragment collected by private collectors from dredged sediments on ‘De Zandmotor’. It was radiocarbon dated to 29,900 + 550/- 490 BP (GrA-69520), which is younger than most Late Pleistocene fossils from this area. Besides adding a new species to the faunal list of the Eurogeul area, this specimen also demonstrates the value of beaches nourished with dredged sediments and highlights the importance of close cooperation between all parties interested in palaeontology: from collector to researcher. The discovery prompted re-examination of the other arctic fox specimen from the North Sea, a femur of which the identification is here confirmed and measurements are provided.