Neanderthal


Neanderthals are an extinct group of archaic humans who inhabited Europe and Western and Central Asia during the Middle to Late Pleistocene. Neanderthal extinction occurred roughly 40,000 years ago with the immigration of modern humans, but Neanderthals in Gibraltar may have persisted for thousands of years longer.
The first recognised Neanderthal fossil, Neanderthal 1, was discovered in 1856 in the Neander Valley, Germany. At first, Neanderthal 1 was considered to be one of the lower races in accord with historical race concepts. As more fossils were discovered through the early 20th century, Neanderthals were characterised as a unique species of underdeveloped human, in particular by Marcellin Boule. By the mid-twentieth century, it was believed that human evolution progressed from an ape-like ancestor through a "Neanderthal phase" to modern humans. This gave way to the "Out of Africa" theory in the 1970s. With the sequencing of Neanderthal genetics first in 2010, it was discovered that Neanderthals interbred with modern humans.
Neanderthal anatomy is characterised by a long and low skull, a heavy and rounded brow ridge, an occipital bun at the back of the skull, strong teeth and jaws, a wide chest, and short limbs. These traits gradually became more frequent through the Middle Pleistocene of Europe, possibly due to natural selection in a cold climate, as well as genetic drift when populations collapsed during glacial periods. Neanderthals would also have been effective sprinters. Neanderthal specimens vary in height from, with average male dimensions estimated at and. While Neanderthal brain volume and ratio to body size averaged higher than any living human population — for males and for females — their brain organisation differed from modern humans in areas related to cognition and language, which could explain the comparative simplicity of Neanderthal behaviour to Cro-Magnons in the archaeological record.
Neanderthals maintained a low population and suffered inbreeding depression, which may have impeded their ability to progress technologically. They produced Mousterian stone tools and possibly wore blankets and ponchos. They maintained and might have created fire. They predominantly ate whatever was abundant close to home, usually big game as well as plants and mushrooms. Neanderthals were frequently victims of major physical traumas and animal attacks. Examples of Palaeolithic art have been inconclusively attributed to Neanderthals, namely possible ornaments made from bird claws and feathers; collections of unusual objects including crystals and fossils; and engravings. Neanderthals buried their dead, but there is no clear indication that they believed in life after death.

Taxonomy

Etymology

Neanderthals are named after the Neander Valley in which the first identified specimen was found. The valley was spelled Neanderthal and the species was spelled Neanderthaler in German until the spelling reform of 1901. The spelling Neandertal for the species is occasionally seen in English, even in scientific publications, but the scientific name, H. neanderthalensis, is always spelled with th according to the principle of priority. The vernacular name of the species in German is always Neandertaler, whereas Neandertal always refers to the valley. The valley itself was named after the late 17th century German theologian and hymn writer Joachim Neander, who often visited the area. His grandfather, a musician, had changed the family name from the original German Neumann "new man" to the Graeco-Roman form Neander, following the fashion of the time.
The th in Neanderthal can be pronounced as following the German convention or Anglicized as fricative /θ/, as per the standard English pronunciation of th.
Neanderthal 1, the type specimen, was known as the "Neanderthal cranium" or "Neanderthal skull" in anthropological literature, and the individual reconstructed on the basis of the skull was occasionally called "the Neanderthal man". The binomial name Homo neanderthalensis was first proposed by Irish geologist William King in a paper read to the 33rd British Science Association in 1863. He extended the name "Neanderthal man" from the individual specimen to the entire species and formally recognised it as distinct from modern humans. However, in 1864, he recommended that Neanderthals and modern humans be classified in different genera as he compared the Neanderthal braincase to that of a chimpanzee and argued that they were "incapable of moral and conceptions".

Discovery

A number of Neanderthal fossils had been discovered before their antiquity was fully understood. The first Neanderthal remains—Engis 2 —were discovered in 1829 by Dutch/Belgian prehistorian Philippe-Charles Schmerling in the Grottes d'Engis, Belgium. He concluded that these "poorly developed" human remains must have been buried at the same time and by the same causes as the co-existing remains of extinct animal species. In 1848, Gibraltar 1 from Forbes' Quarry was presented to the Gibraltar Scientific Society by their Secretary Lieutenant Edmund Henry Réné Flint, but was thought to be a modern human skull.
In 1856, local schoolteacher Johann Carl Fuhlrott recognised bones from Kleine Feldhofer Grotte in Neander Valley—Neanderthal 1—as distinct from modern humans, and gave them to German anthropologist Hermann Schaaffhausen to study in 1857. It comprised the cranium, thigh bones, right arm, left humerus and ulna, left ilium, part of the right shoulder blade, and pieces of the ribs.

Research history

Following Charles Darwin's 1859 On the Origin of Species, Fuhlrott and Schaaffhausen argued that Neanderthal 1 represents a primitive lower human form, aligning more closely with non-human apes as well as Negroids, Eskimos, and Aboriginal Australians. The uniqueness of Neanderthal Man met opposition namely from the pathologist Rudolf Virchow, who argued against defining new species based on only a single find. In 1872, Virchow erroneously interpreted Neanderthal characteristics as evidence of senility, disease, and malformation instead of archaicness, which stalled Neanderthal research until the end of the century.
By the early 20th century, numerous other Neanderthal discoveries were made, establishing H. neanderthalensis as a legitimate species. At first, many palaeontologists considered Neanderthals to be an intermediary phase between modern humans and more apelike ancestors, as suggested by German anatomist Gustav Albert Schwalbe. This hypothesis was opposed by French palaeontologist Marcellin Boule, who authored several publications starting in 1908 describing the French Neanderthal specimen La Chapelle-aux-Saints 1 as a slouching, ape-like creature distantly related to modern man. Boule's ideas would define discussions of Neanderthals for some time.
Boule suggested two different lineages existed in Ice Age Europe: a more evolved one descending from the British Piltdown Man to the French Grimaldi Man which would culminate with modern Europeans; and a less evolved dead-end lineage leading from the German Heidelberg Man to Neanderthal Man. As the focus of human origins shifted from Europe to East Asia by the 1930s and 40s with discoveries such as Java Man and Peking Man, the question of a "Neanderthal phase" in human evolution once again became a topic of discussion. The definition of "Neanderthal" expanded to include several anatomically variable specimens around the Old World. Some specimens were described as "progressive" Neanderthals which would evolve into some local subspecies of H. sapiens, while the "classic" Neanderthals of the Western European Würm glaciation would not.
In the 1970s, with the formulation of cladistics and the consequent refinement of the anatomical definitions of species, this "global morphological pattern" fell apart. The "Neanderthaloids" of Africa and East Asia were reclassified as distant relatives to H. neanderthalensis. At around the same time, the "Out of Asia" hypothesis was overturned by the "Out of Africa" hypothesis, which posited that all modern humans share a fully modern common ancestor. There were two main schools of thought: modern humans competitively replaced all other archaic humans, or extensively interbred with them while dispersing throughout the world. In 2010, the first mapping of the Neanderthal genome demonstrated that there was at least some interbreeding between archaic and modern humans. Subsequent genetic studies continue to raise questions on how Neanderthals should be classified relative to modern humans.

Classification

Neanderthals can be classified as a unique species as H. neanderthalensis, though some authors argue expanding the definition of H. sapiens to include other ancient humans, with combinations such as H. sapiens neanderthalensis. The latter opinion has generally been justified using Neanderthal genetics, as well as inferences on the complexity of Neanderthal behaviour based on the archaeological record. While there seems to have been some genetic contact between these two groups, there are potential indicators of hybrid incompatibility, which if true could justify species distinction. The crux of the issue lies in the vagueness of the term "species".
Among identified archaic humans, Neanderthals are most closely related to Denisovans based on nuclear DNA analyses. Denisovans are an enigmatic group of Late Pleistocene humans only recognisable by a genetic signature rather than anatomical landmarks. Likely due to more recent interbreeding episodes, the mitochondrial DNA and Y-chromosome DNA are more similar between Neanderthals and modern humans than between Neanderthals and Denisovans. Similarly, 430,000 year old fossils from the Sima de los Huesos are more closely related to Neanderthals in their nDNA, but their mtDNA aligns more closely with Denisovans.
A 2021 phylogeny of some Middle Pleistocene and Neanderthal fossils using tip dating: