Gene-flow from steppe individuals into Cucuteni-Trypillia associated populations indicates long-standing contacts and gradual admixture

The Cucuteni-Trypillia complex (CTC) flourished in eastern Europe for over two millennia (5100–2800 BCE) from the end of the Neolithic to the Early Bronze Age. Its vast distribution area encompassed modern-day eastern Romania, Moldova and western/central Ukraine. Due to a lack of existing burials throughout most of this time, only little is known about the people associated with this complex and their genetic composition. Here, we present genome-wide data generated from the skeletal remains of four females that were excavated from two Late CTC sites in Moldova (3500–3100 BCE). All individuals carried a large Neolithic-derived ancestry component and were genetically more closely related to Linear Pottery than to Anatolian farmers. Three of the specimens also showed considerable amounts of steppe-related ancestry, suggesting influx into the CTC gene-pool from people affiliated with, for instance, the Ukraine Mesolithic. The latter scenario is supported by archaeological evidence. Taken together, our results confirm that the steppe component arrived in eastern Europe farming communities maybe as early as 3500 BCE. In addition, they are in agreement with the hypothesis of ongoing contacts and gradual admixture between incoming steppe and local western populations.

The preservation of the skeleton is medium to good. The skeleton is almost complete, found partially articulated.
Anthropological type: Mediterranean and Dinaric features, in equal proportions.
Pathologies at the dental level: antemortem loss of 11 teeth from the lower arcade (partial mandibular edentation); advanced dental wear (attrition type).
Pathologies, non-metric traits and occupational stress markers at the postcranial level: generalized degenerative osteoarthritis manifested through osteophytes, corrosion, eburnation, and deformed joint surfaces; spondylolysis of the fifth lumbar vertebra; squatting facets of the tibiae (more pronounced on the right side); additional femoral trochanter.
KH160122: Pocrovka 5, pit 98 / 1998, skeleton 2/B, female, 20-25 years old (young adult) The preservation of the skeleton is medium to good. The skeleton is almost complete, found partially articulated.
Anthropological type: Mediterranean and Dinaric features.
Pathologies at the dental level: quite thin supragingival plaque on the upper and lower front teeth (on the vestibular and lingual sides); occupational / cultural dental wear (abrasion type) on the upper front teeth.
Pathologies and non-metric traits at the cranial level: inactive porotic hyperostosis (porotic type cribra cranii) on the occipital and parietals, more concentrated in the proximity of the sagittal and lambdoid cranial sutures; unilateral foramen parietale (left side); frontal notch (bilateral); double zygomatic foramina (left side).
Pathologies, non-metric traits and occupational stress markers at the postcranial level: hypertrophic periosteal reaction on the diaphyses of the tibiae and femora; preauricular sulcus (parturition scars); pronounced muscle attachments on the humeri and femora; additional femoral trochanter.
Anthropological type: Mediterranean and Dinaric features.
Pathologies and non-metric traits at the dental level: supragingival plaque on the front teeth (vestibular side); occlusal and cemento-enamel junction (CEJ) cavities on the mandibular teeth; antemortem loss of two teeth not long before death; two apical abscesses; congenital absence of the third molar; moderate dental wear (attrition type).
KH160121: Gordinești / 1981, grave 1, probably female, 9 years old ± 24 months (child or infans II) The preservation of the skeleton is medium to good. The skeleton is incomplete. The skull is represented through occipital, upper maxilla (partial), mandible (with deciduous and permanent teeth in the sockets and without symphysis), nine fragments from parietals, two from the temporals, one from the frontal (right side), one from the sphenoid and two isolated upper teeth (first and second right upper incisors). The postcranial skeleton includes the illia, a rib fragment and six cervical vertebrae (C1-C6).
The age at death was estimated by analysing the temporary/permanent dentition. Sex was determined by taking into consideration features of the entire skeleton, especially the illia and the skull fragments.
Pathologies at the cranial level: active porotic hyperostosis (cribrotic type cribra crania and cribra orbitalia) on the occipital, parietals and superior wall of the right orbit (left half of the frontal bone is missing).

Taphonomic data
The preservation of the remains is generally satisfactory. The skeletons of the adult females from Pocrovka V are almost complete, and the sub-adult skeleton from Gordineşti is partially represented (25-75%). The taphonomic analysis showed a very slight subaerial weathering on all remains (in form of thin calcareous depositions). No skeletal element presents traces of aerial weathering, such as cracking or peeling of the bone surface. All breakages and fissures that have led to the fragmentation of the skeletal elements occurred postmortem. No perimortem breaks were identified. All disarticulations were produced postmortem. There were no traces of slicing or scraping, chopping tool marks, impact / percussion marks, polishing or abrasion, or cut marks from defleshing activities.
It is important to mention that there are no elements that indicate decapitation. Carnivore-ravaged marks or rodent tooth scratches, gnawing, and tooth punctures are missing. Root-etching on the