adapted to the open environments of the late Miocene. However, both were extinct by 7 million years ago. In 2017, a reanalysis of Graecopithecus fossils from Greece and Bulgaria, previously associated with Ouranopithecus, concluded that the species was in fact a hominin dating to just after the last common ancestor of humans and chimpanzees (about 7.2 million years ago). The authors suggested that the origins of the human lineage were therefore in the Mediterranean, not Africa. Others are sceptical of their claims. Although subtropical conditions returned to Europe in the Pliocene (5.33–2.58 million years ago), there are no known fossil hominids from this period. Archaic humans Homo erectus populations lived in southeastern Europe by 1.8 million years ago. The most archaic human fossils from the Middle Pleistocene (780,000–125,000 years ago) have been found in Europe. Remains of Homo heidelbergensis have been found as far north as the Atapuerca Mountains in Gran Dolina, Spain, and the oldest specimens can be dated from 850,000 to 200,000 years ago. Neanderthals evolved from a branch of Homo heidelbergensis that migrated to Europe during the Middle Pleistocene. Neanderthal populations date back at least as far as 400,000 years ago in the Atapuerca Mountains, Spain. While lacking the robustness attributed to west European Neanderthal morphology, other populations did inhabit parts of eastern Europe and western Asia. Between 45,000–35,000 years ago, modern humans (Homo sapiens) replaced all Neanderthal populations in Europe anatomically and genetically. This is evident in the transfer and combination of technology and culture. Anatomically modern humans The recent expansion of anatomically modern humans reached Europe around 40,000 years ago, from Central Asia and the Middle East, as a result of cultural adaption to big game hunting of sub-glacial steppe fauna. Neanderthals were present both in the Middle East and in Europe, and the arriving populations of anatomically modern humans (also known as "Cro-Magnon" or European early modern humans) have interbred with Neanderthal populations to a limited degree. Modern human remains dating to 43–45,000 years ago have been discovered in Italy and Britain, with the remains found of those that reached the European Russian Arctic 40,000 years ago. The composition of European populations was later altered by further migrations, notably the Neolithic expansion from the Middle East, and still later the Chalcolithic population movements associated with Indo-European expansion. The modern indigenous population of Europe is composed of three major foundational populations, dubbed "Western Hunter-Gatherers" (WHG),