examined the remains of 6 Cardials buried in Spain c. 5470–5220 BC. The 6 samples of
mtDNA extracted belonged to the maternal haplogroups
K1a2a,
X2c,
H4a1a (2 samples),
H3 and
K1a4a1. The authors of the study suggested that the Cardials and peoples of the
Linear Pottery Culture were descended from a common farming population in the
Balkans, which had subsequently migrated further westwards into Europe along the
Mediterranean coast and
Danube river respectively. Among modern populations, the Cardials were found to be most closely related to
Sardinians and
Basque people. The Iberian Cardials carried a noticeable amount of hunter-gatherer ancestry. This hunter-gatherer ancestry was more similar to that of
Eastern Hunter-Gatherers (EHGs) than Iberian hunter-gatherers, and appeared to have been acquired before the Cardial expansion into Iberia. found traces of maternal genetic affinity between people of the Linear Pottery Culture and Cardium pottery with earlier peoples of the Near Eastern
Pre-Pottery Neolithic B, including the rare mtDNA (maternal) basal haplogroup
N*, and suggested that Neolithic period was initiated by seafaring colonists from the Near East. examined three Cardials buried at the Zemunica Cave near
Bisko in modern-day
Croatia c. 5800 BC. The two samples of
Y-DNA extracted belonged to the paternal haplogroups
C1a2 and
E1b1b1a1b1, while the three samples of
mtDNA extracted belonged to the maternal haplogroups
H1,
K1b1a and
N1a1. The team further examined two Cardials buried at Kargadur in modern-day Croatia c. 5600 BC. The one male carried the paternal haplogroup
G2a2a1, and the maternal haplogroup
H7c, the female carried
H5a. All three belonged to the
Early European Farmer (EEF) cluster, thus being closely related to earlier Neolithic populations of north-west Anatolia, of the Balkan Neolithic, contemporary peoples of the Central European Linear Pottery culture, and later peoples of the Cardial Ware culture in
Iberia. This would suggest that the Cardial Ware people and the Linear Pottery people were derived from a single migration from Anatolia into the Balkans, which then split into two and expanded northward and westward further into Europe. Five individuals buried in two sites linked to Impressa ware were tested geneticaly (Grotta Continenza in Trasacco, and Ripabianca di Monterado in Ancona), the males had Y-chromosomes G-L91 (G2a2a1a2), R-M343 (R1b), J-L26 (J2a1) and J-M304 (J*). These Neolithic individuals could be modeled as a mixture of ~5% Western hunter-gatherer and ~95% Anatolian farmers (who carried an additional Caucasian HG ancestry). Five herders with Cardium pottery were buried in a cave of the Aragonese
Pre-Pyrenees (Cueva de Chaves, Bastarás, Huesca province), the genetic analysis found that the Y-haplogroup of two males was I2a1b, being the other male assigned to
R1b-M343. Admixture models found that their ancestry was 4/5 Anatolian-like and 1/5
Villabruna-like. The remains of three transhumant herders found in Cova dels Trocs (Sant Feliu de Veri, Bisaurri, in the Spanish Pyrenees) were analyzed, the Y-chomosomes were: R1b1, F*. and I2a1a. Three individuals buried in the Pendimoun rock-shelter (Castellar,
Alpes-Maritimes) were tested geneticaly, the male individual carried Y-haplogroup I-M423 (I2a1a2b). Two individuals from the Cardial cave Gruta do Caldeirão (municipality of Tomar, in central Portugal) were assigned to Y-chromosome haplogroup I2a1a. Four individuals from the Kaf Taht el-Ghar site (a cave near Tétouan, in the
Rif) were analyzed, the only Y-haplogroup found in the two males was G2a (subclade G2a2b2a1a1c1a); the autosomal components of the buried were Anatolian Neolithic ancestry (72%), Western Hunter-Gatherer ancestry (10%) and local Maghrebi ancestry (18%). The late-Neolithic
Kehf el Baroud inhabitants in present-day
Morocco (c. 3700 BC) were modelled as being of about 50% local North African ancestry and 50% Early European Farmer (EEF) ancestry. It was suggested that EEF ancestry had entered North Africa through Cardial Ware colonists from Iberia sometime between 5000 and 3700 BC. They were found to be closely related to the
Guanches of the
Canary Islands. The authors of the study suggested that the
Berbers of Morocco carried a substantial amount of EEF ancestry before the establishment of
Roman colonies in Berber Africa. According to Simões (2023) human remains from the earliest Neolithic contexts in northwestern Africa had European Neolithic ancestry (c. 5400 BC), indicating that the first stages of the Neolithisation process in northwestern Africa were started by the migration of Neolithic farmers from Iberia. The earliest pottery in the
Tingitan peninsula (the African portion of the
Gibraltar Strait) was also of Cardial type, with clear affinities to archaic Cardial pottery from
Catalonia–
Valencia. ==See also==