Neanderthal

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## Homo neanderthalensis

Various Neanderthal paintings by Tom Björklund.

## Statistics

### Height

5 feet-5 feet 5 inches (152/156-164/168 centimeters)

### Diet

Omnivorous

The Neanderthal or Neandertal (*Homo neanderthalensis*) is an extinct species of hominin that lived across Eurasia and possibly Africa until about 40,000 years ago.

## History

In 2022, researcher Svante Pääbo won a Nobel prize for sequencing an entire Neanderthal genome. The genomes of entire individuals were then released to the Public Domain.

## Description

Neanderthals are often depicted as stockier than apomorphic humans, with large barrel-shaped ribcages, a wider pelvis and proportionally shorter arms and legs. The body mass index for adult Neanderthal males (26.9-28.2) is overweight when compared with the average human index. This indicates that Neanderthals were very robust in build. The LEPR gene in Neanderthals, tasked with storing heat and fat production, is very similar to a woolly mammoth's, possibly correlated to cold adaptations. The average height of Neanderthals (5 feet, 152-156 centimeters (females) and 5 feet 5 inches, 164-168 centimeters (males) based on 45 long bones, 14 men and 7 woman) decreased by about 10 centimeters (4 inches) near the end of Late Paleolithic. Their weight, 66.4 kilograms (146 pounds) female and 77.6 kilograms (171 pounds) male are based on 26 specimens. Neanderthal neck vertebrae are thicker than most humans, likely to stabilize the larger and differently-sized head. The Neanderthal's thorax is similar in size to a humans, but longer and straighter ribs would have made the mid-lower thorax be wider and having stronger breathing, possibly having a larger diaphragm and maybe even a larger lung capacity. Femoral robusticity in Neanderthals is different than *Homo sapiens*, which indicates a different level of mechanical stimulus to produce morphological response in the femur, either caused by migration-caused gene flow and/or physical activity in populations.

The estimated lug capacity of Kebara 2 is 9.04 liters (2.39 US gallons). The average Neanderthal chest is antero-posteriorly pronounced and the sacrum is more vertically inclined. This lowers lordosis, enhancing mediolateral flexion and supporting a wide lower thorax. This may be common in Homo, with the thinner thorax in apomorphic humans being abnormal in the genus. Neanderthal proportions are often described as "hyperarctic", having similar adaptations than other cold-based humans. Among apomorphic humans, Neanderthals are built similar to Inuit and Siberian Yupik populations, as lower limb length equals greater heat retained. However, individuals from more temperate environments, like Iberia, keep this physique. John Stewart et al. in 2019 theorized that Neanderthals were adapted to sprinting, because they preferred warmer areas rather than the colder Mammoth Steppe. DNA analysis also shows a higher concentration of fast-twitch muscle fibers. He also pointed out that the higher amount of muscles opposed the endurance-running hypothesis, as persistence hunting may only be an effective strategy in warmer climates, where hunters can chase they prey to hypothermia. Longer heel bones reduced Neanderthals persistence hunting capabilities, were shorter limbs allowed them to reduce moment arm at the limbs and causing rotational force at the wrists and making joint muscles work harder, accelerating in speed. In 1981, Erik Trinkaus noted this explanation, deeming it less likely.

Neanderthal faces are known to have a reduced chin, large and further placed nose, a dramatic brow ridge and a large forehead. Neanderthal skulls are also more elongated and globular than in apomorphic humans, with an occipital bun that protrudes to the rear of the skull, differing in the individual. This is caused by the cranial base and temporal bone arranged higher towards the front of the skull and a flat skullcap. Neanderthals also have larger eyes, likely a low-light adaptation. The enlarged nose and paranasal sinuses are thought to have evolved to warm air before it entered the lungs and to retain moisture (as the "nasal radiator hypothesis"). However, these features are usually reduced in cold-adapted animals, maybe the nose instead caused by genetic drift. The sinuses are not extremely large, comparable to apomorphic humans. However, the sinus-size is not necessary to breathe in cold air, and their function and reason to evolve such a large nose is unknown. Neanderthals are noted as not a good example of evolutionary pressure to evolve drastic features. Additionally, a computer reconstruction of predicted Neanderthal nose soft tissues show that they would likely be similar to Arctic human populations. This means, these may have evolved convergently as both separated groups breathe cold, dry air. Neanderthals also have jaw prognathism, outdatedly cited as a response to a great bite force referenced by wear on the front teeth (as the "anterior dental loading hypothesis"), however similar trends are seen in apomorphic humans with a lesser bite force. This protrusion may have evolved to fit larger teeth in the mandible, resisting abrasion and wear in the back teeth. The wear in the front teeth likely resulted from repeated use. Similar patterns arise in apomorphic Inuit populations. Neanderthal incisors are large and shovel-shaped, and have a high level of taurodontism compared to lower levels in apomorphic humans. This condition was thought to have evolved due to demand but is likely genetic drift. Bite force is now thought to be the same than in apomorphic humans, with male and female 285 newtons (64 pound-force) and 255 newtons (57 pound-force) respectively.

Neanderthals have many distinctive traits. In the skull: a sloping forehead, a suprainiac fossa, an occipital bun, midsagittal, maxillary and mandibular prognathism, a less neotenized skull, a low elongated skull with a flat lambdoid region, a broad cranial vault with "en bombe" parietal morphology, a flat basicranium, a supraorbital ridge, 1500-1740 centimeters cubed cranial capacity, reduced chin, mastoid process crest, no groove on the canines, a retromolar space posteriorly positioned to the third molar, a broad and large nose, bony projections at the nasal opening's side, a bony labyrinth in the ear and a larger mental foramen to supply blood to the face. In the postcranials: a more robust build, proportionally long clavicles, broad shoulders and pelvis, large and wide shoulder blades, a broad chest, an unclear barrel-shaped ribcage with no waist, more laterally curved radius with a radial tuberosity placed more medially, a longer radial neck and ovoid head, a well-developed interosseous crest, smaller curved limbs, a trochlear notch facing more anteriorly on the ulna with brachialis inserted lower with a larger and more sinusoidal mid-shaft, thick joints, large hands and feet, large and round fingertips, large kneecaps, thick bowed thigh bone shaft and femur, short tibiae and fibulae, a large calcaneus and a long slender superior pubic ramus. Though the foot is near identical in structure to *Homo sapiens*, unique foot posture indicates adaptation to mobility strategy and phylogenesis rather than *Homo sapiens* who are also adapted for footwear. A horizontal pterygoid oriented similar to apomorphic human infants is autapomorphic, which may suggest an inferior, low placement of the torus tubarius and Eustachian tube orifice at the lateral nasopharyngeal wall.

## Classification

Neanderthals are classified in the genus *Homo*, but it is unsure if it is *H. neanderthalensis* or *H. sapiens neanderthalensis*. If it is *H. s. neanderthalensis*, this would necessitate the need to make *H. sapiens* to be *H. s. sapiens*. Most of the controversy derives from the definition of 'species', as the traditional definition defines it as two geographically isolated specimens, but admixture of the two species is known to have happened. However, the apparent absence of the Neanderthal gene patrilineal Y-chromosome and matrilineal mitochondrial DNA (mtDNA) in humans, along with the underrepresented Neanderthal X-chromosome could mean that hybrids had infertility or reduced fertility rates. In 2014, geneticist Svante Pääbo described 'taxonomic wars' as unresolvable "since there is no definition of species perfectly describing the case". Neanderthals are thought to be closer-related to the Denisovans than to apomorphic humans. Based on nuclear DNA (nDNA) Neanderthals and Denisovans have the most recent last common ancestor than apomorphic humans. However, Neanderthals and apomorphic humans share last common ancestor based on mtDNA. This likely arose because of a hominin interbreeding event after the Denisovan/Neanderthal spit event that introduced a new line of mtDNA. This either involved an introgression from an unknown plesiomorphic human into the Denisovans, or an introgression from an unidentified African human population into Neanderthals.

A controversial study from 2022 suggests that Neanderthals should not be considered synonymous, since there is no reason to do so according to available evidence. Additionally, it calls for paleoanthropological taxonomy to reflect on the systematics of related zoological and biological fields.

### Evolution

It is commonly thought that *Homo heidelbergensis* is the last common ancestor to the Neanderthals, Denisovans and apomorphic humans, before populations got stranded in Europe, Asia and Africa, respectively. The taxonomic distribution between *H. heidelbergensis* and Neanderthals is based on a gap in the fossil record occurring in Europe 300-243 thousand years ago during marine isotope stage 8. Neanderthals date after this gap However, the 430 thousand year-old fossils at Sima de los Huesos could represent basal Neanderthals or a closely-related population and the 400 thousand year-old Aroerira 3 could be a transitional form. Ancestral and more derived forms may have coexisted. It is also possible a gene flow between Western Europe and Africa during the Middle Pleistocene existed, and obscured Neanderthal characteristics, like in Ceprano, Italy and Sićevo Gorge, Serbia. The fossil record is more plentiful from 130 thousand years ago and onwards, and specimens from then and onwards make up the vast majority. Dental elements from the Visogliano and Fontana Ranuccio sites of Italy indicate that the signature Neanderthal features had occurred 450-430 thousand years ago, Middle Pleistocene.
The two main hypotheses regarding Neanderthal after their split with their concestor:

* The two-phase model states that after a major event, possibly the Saale glaciation, caused European *H. heidelbergensis* to quickly become robust and grow in size, enlengthening the head (phase 1), which led to other developments in the head and body (phase 2). Neanderthal anatomy may not have been driven by cold environment.
* The accretion model states Neanderthals slowly evolved from an ancestor, *H. heidelbergensis*, and divided into four stages: early pre-Neanderthal (MIS 12, Elster glaciation), pre-Neanderthals (MIS MIS11-9 Holstein interglacial) early Neanderthals (MIS 7-5, Saale glaciation/Eemian) and classic Neanderthals (MIS 4-3, Würm glaciation).

Many dates of the Neanderthal/human split have been proposed. One, 250,000 thousand years ago sites "*Homo helmei*" as the last common ancestor, with the subsequent split associated with the Levallois technique when making stone tools. The 400,000 date uses *Homo heidelbergensis* as the last common ancestor, and 600,000 thousand years ago states that *Homo rhodesiensis* is the last common ancestor, but variations of this model push the date to 1 million years ago. A 2020 study of Neanderthal enamel proteomes states that *Homo antecessor* is related, but not directly ancestral. DNA results show various dates of the Neanderthal/human split: 538-315, 553-321, 565-503, 654-475, 690-550, 765-550, 741-317 and 800-520 thousand years ago, with an additional dental analysis concludes before 800,000 thousand years ago. Neanderthals and Denisovans are more closely related to one-another than to apomorphic humans, meaning the Neanderthal/Denisovan occurred after humans did the same. Assuming a constant mutation rate of 1x10−9 or 0.5x10−9 per base pair per year, they split either 236-190 or 473-381 thousand years ago. Using 1.1x10−8 per generation, with a new generation born every 29 years, it is likely around 744,000 years ago. However, using 5x10−10 nucleotide site every year, the date drops to around 644,000 years ago. Using the last two dates, this split likely happened by the time hominins had spread across Europe, and Neanderthal features had arisen by 600-500 thousand years ago. Before the split, Neanderthal/Denisovan "Neandersovans" had migrated from Africa to Europe, interbreeding with a "superarchaic" human that had already inhabited the area, these would have been descended from hominins that migrated from Africa 1.9 million years ago. A tooth from a 7-12 year-old Neanderthal tooth found near tools (the Shukbah tooth) was lost in a private collection, but was rediscovered and sent to the British Museum of Natural History and studied by researchers. This tooth was discovered in Arabia, the southernmost Neanderthal location known. No Neanderthal remains are yet known from Africa, but the researchers suggest populations may have dispersed into the region on occasion. The technology found near features techniques known from *Homo sapiens*. This suggests the two species lived alongside each other, felt the need to share technology, or both.

    Oase 1 is a hominin fossil found in Peștera cu Oase and discovered in 2003 by Ștefan Milota, Ricardo Rodrigo and Mircea Gherase when the team stumbled across many human remains near the cave's surface. Among them, Oase 1 is an individual consisting of a mostly-complete left temporal bone and many frontal, parietal and occipital fragments. Lagar Velho 1, the Lagar Velho boy or Lapedo child is a complete skeleton found in Portugal, believed to have had a Neanderthal and human parent. These remains were found at the Late Paleolithic Abrigo do Lagar Velho, led by archeologist João Zilhão. This discovery provided evidence for humans west of the Iberian Peninsula: a 4-year-old child buried with a pierced shell and red ochre that lived 24,500 years ago. The skull, teeth and postcrania show a vast collection of Neanderthal and human features. Late plesiomorphic and early apomorphic people in Iberia are the likely cause of admixture, which refutes an old theory that Neanderthals were replaced by apomorphic humans, and bringing forth new ideas of human culture. Prof. Dr. C.P.E. Zollikofer disputed this idea, stating that the specimen shows no Neanderthal traits, however, later genetic studies prove that admixture did occur. An archeology museum revolving around the Lapedo child is planned to be built by the Convent of St. Augustine. American anthropologist Brian Pierson made a reconstruction of the skeleton and the specimen's face, which can be found at the Interpretation Centre of the Lagar Velho.

Oase 1, after genetic analysis, has traits from plesiomorphic and Early European apomorphic Humans (EEMH) and Neanderthals. Since October 2015, multiple genetic studies have taken place. About 6-9% of the genome is Neanderthal, having the largest percentage of plesiomorphic introgression in anatomically accurate humans, and with disequilibrium patterns indicating a Neanderthal ancestor 4-6 generations earlier. The specimen is male, and Fu et al. in 2015 propose the individual sharing more alleles with apomorphic East Asian populations than with Europeans. However, he shared equal alleles with Mesolithic Europeans and Eastern Eurasians, which indicates a non-pre LGM-European admixture in today's Europeans. Part of this stems from Basal Eurasian ancestry that was brought to Europe by Yamnaya pastoralists and Anatolian farmers. Additionally, Oase 1 belongs to the Y-DNA haplogroup and mitochondrial DNA haplogroup. Poznik et. al. in 2016 suggest Oase's Y-DNA haplogroup to belong to K2a. He possesses similar SNPS to the Ust'-Ishim man (another hybrid), a rare lineage found in two living men (of Tegulu and Malay backgrounds, where Poznik suggested a new subclade; K2a1) and upstream from haplogroup NO. Earlier research by Fu et al. states Oase belongs to a subclade of Y-DNA haplogroup F (rather than G, H , I and J) and possibly macrohaplogroup K. Fu also states Oase belongs to a maternal lineage that relates to mitochondrial DNA haplogroup N, diverging from N before one-another[*citation needed*]. A genetics study in 2022 detects a small amount of Neanderthal DNA in living populations, which either indicates an introgression event from Africans and into Neanderthals or where Neanderthals introgressed into Africans that migrated back. Many modern mutations caused by Neanderthal introgression can be seen altering the development of the immune system and metabolism.

## Paleobiology

Neanderthal braincases for males and females respectively average 1600 centimeters3 (98 inches3) and 1300 centimeters3 (79 inches3). This within the range of apomorphic human men and woman: 1270 centimeters3 (78 inches3) for males and 1130 centimeters3 (69 inches3) respectively. Between 28 human specimens 190-25 thousand years ago, disregarding sex, is 1478 centimeters3 (90.2 inches3). Apomorphic human brain sizes are suggested to have decreased since the Late Paleolithic. The largest recorded Neanderthal brain, Amud 1, is calculated to 1736 centimeters3 (105.9 inches3), the largest of all hominids recorded. Both Neanderthal and sapiens enfants measure 400 centimeters3 (24 inches3). In Neanderthals, the occipital lobe that operates vision was much larger than in apomorphic humans and had larger eyes to deal with lower light in Europe. More brain tissue is reported to control the body and maintenance, as the cognitive areas were proportionally smaller. This includes the cerebellum that operates muscle memory, possibly language, attention, working memory, social capabilities and thought, the parietal lobes controlling visuospatial function and episodic memory, the temporal lobes associated with spoken comprehension and emotional association, the orbitofrontal cortex controlling decision making and the olfactory bulb that controls smell. In 2021, researchers created a 3D reconstruction of *H. sapiens* and Neanderthals ear canals, The results entered the information into a software, which developed a further model. They found that Neanderthals could hear up to 5 kHz, which is the standard speech frequency in *H. sapiens*. They also compared the latter two were compared with Atapuerca specimens, with the Neanderthals hearing a 4-5 kHz range, better than the recorded Atapuerca specimens. Each amount of fossils revealed an estimated occupied bandwidth, with Neanderthals having a wider bandwidth, matching *H. sapiens* than their Atapuerca ancestors. The study found that Neanderthals were tuned to produce and perceive *H. sapiens*' speech, leading way to increasingly difficult interspecies behaviour . One study found that different morphological variables in the Neanderthal mandible were cause for a distinct style of speaking than contemporaneous *Homo*. More evidence point out that the Neanderthals could practice/shown much more 'humane' behavior or cultural imprints, than previously thought, as a study found out the species even practiced primitive form of oral hygiene with toothpicks.

Lack of sunlight in Paleolithic Europe likely led to the proliferation of lighter skin in Neanderthals, although lighter skin was not common in Europe until possibly the Bronze Age. BNC2 is known to have existed in Neanderthals, which is often associated with with light skin, although the BNC2 variant that also exists in them shows individuals could have also had darker skin, as recorded in the UK Biobank. Skin tone likely varied depending on region. DNA from three Croatian Neanderthals show that individuals from the latter region had darker skin, eyes and hair; their eyes close in colour to apomorphic Europeans. In apomorphic humans, skin and hair tone is regulated by melanocyte-stimulating hormones, increasing eumelanin to phaeomelanin proportions, which is encoded by the MC1R gene. 5 known variants in apomorphic humans that cause light hair and skin exist, with the unknown R307G variant in Neanderthals producing light skin and red hair. This was identified from a specimen from Monti Lessini, Italy and possibly another from Cueva del Sidrón, Spain. As in humans, red pigment was likely not common, as it is not present in many genome-sequenced individuals. Some researchers argue the large nose of Neanderthals were an adaptation to the cold, but primate-related Arctic studies show that sinus reduction happens more often than enlargement, according to Allen's rule. Todd C. Rae summarizes Neanderthal anatomy as attempting to find an explanation for a "paradox" as to why these features evolved. Rae concludes that the larger nose likely evolved in the Middle East, which was warmer, and did not change as they entered Europe. However, individuals in Spain date to at least 700,000 years ago, prior to leaving the Middle East. Rae proposes that the increased physical activity and enlarged muscle mass led to a higher oxygen intake demand. Levantine Neanderthals show phenotypes drastically more similar to apomorphic humans than to European classic Neanderthals. This may be caused by a gene flow from apomorphic humans to the Levantine corridor, or possibly the classic stage is specialized to climate. It is postulated that Neanderthals still visited northern settlements, but most often in the warmer months. However, Neanderthal settlements are also found in regions that would have been more temperate as well.

The maximum lifespan, menopause and gestation time in Neanderthals was likely similar to apomorphic humans. Models on caloric requirements in Neanderthals are mostly higher, due to their larger body mass, higher physical activity levels (PALs, larger distances traveled while foraging) and heat production and faster basal metabolic rates (BMRs). Using BMRs and PALs, American archeologist Bryan Hockett estimated a pregnant Neanderthal would need to consume 5500 calories per day, relying on big game meat and causing numerous deficiencies and nutrient poisonings. He concludes that these are not good assumptions to make, and are not practical. Neanderthals would have been active during the dimmer light, since daylight would have been reduced. They hunted large game, usually predators who are active and stalking at night. They also have larger eyes and more complex neural processing. Colour blindness (possibly enhancing mesopic vision) typically correlates with northern-based populations. Neanderthals from Vindija Cave, Croatia have Opsin gene substitutes, which can influence seeing in colour. However, the functions of these substitutions are unknown. Neanderthal-derived alleles such as ASB1 and EXOC6 are typically associated with the evening type, narcolepsy and daytime naps. The neocortex was likely smaller than contemporaries, as implied by the TKTL1 protein, and thus the frontal lobe had lesser neurogenesis.

### Paleopathologies

Neanderthals, in life, suffered many traumas, 79-94% of specimens healing, 37-52% severely injured and 13-19% injured before maturity. An extreme example of injury is in Shanidar 1, who's right arm is amputated after nonunion occurred (a failure in healing after bone breakage in adolescence), osteomyeitis on the left clavicle, an abnormal gait, visual problems in the left eye and possibly even lost hearing (swimmer's ear?). Trinkaus estimates in 1995 that 80% of Neanderthal lives were lost from injury before the age of 40, and that Neanderthals used risky hunting tactics (as the "rodeo rider hypothesis"). However, cranial trauma rates between Neanderthals and Paleolithic humans, though Neanderthals have a higher injury rate, but there are few specimens from either species that survived until 40 years of age and similar injury patterns exist. Trinkaus in 2012 concluded that Neanderthals injured one-another via interpersonal violence. A 2016 research study looks at 124 Neanderthal samples and argues trauma is caused most often by animal attacks; 36% bear attacks, 21% big cats and 17% wolves, this totals to 92 positive cases and 74% rate. No reported cases from *Hyaena*s exist, although they likely opportunistically attacked Neanderthals. Such high rates of confrontation likely occurred after disputed over caves, food or even Neanderthals hunting them. Low populations likely caused low genetic diversity and inbreeding, which prompted inbreeding depression. It is unknown how these environments strained an individuals genetic burden, likely having a higher birth defect rate than in apomorphic humans. However, it is known that all 13 inhabitants of the Sidrón Cave exhibited 17 birth defects due to inbreeding or recessive disorders. La Chapelle-aux-Saints 1, likely due to being 60-70 years old, showed signs of Baastrup's disease and osteoarthritis. Shanidar 1 shows the most prehistoric case of diffuse idiopathic skeletal hyperostosis (DISH), and likely died between 40 and 50. DISH can restrict movement, and could indicate a high incident rate for elderly Neanderthals. Possible vertebral brucellosis nay be present in La Chapelle-aux-Saints 1.

Like other animals, Neanderthals were at risk to infectious disease, alongside parasites. apomorphic humans likely transmitted disease to them, a possible candidate the stomach bacteria *Helicobacter pylori*. Derived human papillomavirus variant 16A may descend from Neanderthals, specifically from introgression. One individual from Cueva del Sidrón, Spain shows sign of gastrointestinal *Enterocytozoon bieneusi*. Leg bones from French specimen La Ferrassie 1 adorn lesions similar to periostitis, an inflammation of the tissue surrounding bone, likely resulting from hypertrophic osteoarthropathy, which is caused by chest infection or lung cancer. Neanderthals have a significantly lower cavity rates than in apomorphic humans, despite consuming cavity-causing foods in great quantity, which would indicate a lack of oral bacteria, such as *Streptococcus mutans*. Two 250,000 year old Neanderthal children from a site in Payré, France are the earliest cases of lead poisoning in the fossil record. This was likely caused by ingesting contaminated substances or breathing in lead-laced smoke, additionally, two lead mines sit within 25 kilometers (16 miles) of the discovery site.

### Ontogeny

Neanderthals may have grown ore extensively than in apomorphic humans, the latter having the slowest body growth in any mammal (with a lack of growth between infancy and puberty, which is made up for in the adolescent growth spurt later on). This was first questioned by excavators in 1928 in the Mousterian rock-shelter while excavating a juvenile individual. In 1931, Arthur Keith writes "Apparently Neanderthal children assumed the appearances of maturity at an earlier age than apomorphic children.". The maturity can be estimated based on body maturation rate of the individual and age at death. Additionally, the age of immature individuals can be estimated based on tooth morphology, including development and emergence. These methods have been disputed, both in favour, and not for maturation differences between Neanderthals and humans. In 2007, it was found age can be found by scanning the item via an x-ray synchrotron microtomography to detect patterns. This is noninvasive to the fossils themselves. September 2017 research based on a 7.7 year old (at death) Neanderthal juvenile found in Spain and dating to 49,000 years ago indicates that juveniles may have actually grew similarly to humans. The team examined dental, cranial and postcranial material, and documented development rates compared to age. The main differences between the two species are based in the spinal column, and several features hinted towards brain continuing growth. Vertebral and brain maturation is noted to likely have been to support thee larger bodies. The difference is less fundamental than previously thought.

### Culture

Neanderthals lived in more sparse groups, probably at 10-30 individuals average. It was found adolescents were forced to join hunting parties right at the time they turn adolescent; however, bone trauma is similar to apomorphic Inuits, suggesting a similar childhood with injuries caused by winter and low food. Groups formed nuclear families of no more than 3, likely moving between caves based on the season. These caves may have been used for up to and over 100 years. Cave bears competed with Neanderthals for these spaces, with a decline in their populations starting 50,000 years ago, though they went extinct long after Neanderthals did. Open-air sites are considered to have been used for killing and butchering prey rather than a home. Sometimes the amount of individuals needed to retain a stable population were not present, so some Neanderthals show signs of inbreeding and the resulting defects caused by this. Recessive disorders are also an explanation for these. Bands of Neanderthals probably did not interact often, since they tended to stay out of the way of other Neanderthal groups and their smaller size limited interaction. Genetically, 3 groups of Neanderthals are mapped; Western Europe, Mediterranean coast and east of Caucasus, with some migrating among these. Macro-regions also exist; Acheulean-tradition Mousterian in the southwest, Micoquien in the northeast, and Mousterian with bifacial tools between the former. The latter may be the interactions and fusion of the other 2. Some have found Neanderthals to often interact with neighbouring communities but not far-out communities. Over long periods of time, cross-continental migrations occurred. In some places, intraspecific group conflicts are noted, specifically with gashes to the skull caused by a bladed weapon and a rib lesion caused by a projectile. Some of the oldest preserved art are the ochre hand stencils from the 65,000-year-old site of Maltravieso Cave.

Some people suggest no sexual division labour was present, however, some Neanderthals have tooth wearing suggesting carrying items in their mouths, with men having more wearing on the upper and women on the lower teeth, suggesting a divide in labour. Controversially, Neanderthals may have worn decorative items such as clothes or jewelry to display their hierarchy in the group. Some also suggest the small amount of preserved burials is due to higher ranking members receiving elaborate burials; it was also suggested elders got a similar treatment due to how long they lived given mortality rates. Alternatively, it was suggested more Neanderthals were buried but their graves were vandalized and/or destroyed by bears. Dead children may have received proper burial rather than any other age due to the low rarity of Neanderthal burials under the age of 4. It was once suggested providing food boosted worth and anyone who failed to keep up with their group would be left for dead, including the disabled. However, there is evidence for Neanderthals caring for their sick and disabled, with this dating as far back as *Homo heidelbergensis*. If this is true, this would explain how Neanderthals survived for so long. Some claim evidence for Neanderthal funeral practices but there are all controversial and can not be 100% proven, since the intent of the individuals can not be obtained from fossils. It has sometimes been proposed that Neanderthal cults existed, such as cave bear skulls arranged could be a special form of worship, but this can also be explained by natural phenomena. Additionally, it was once thought they killed and cannibalized others to use their skulls in ceremony, but there is no evidence to state cannibalism was ritualistic. It was also proposed a "Cult of the Sun Bird" for a supposed Neanderthal cult worshiping golden eagles, since their bones were modified more than other birds when in possession.

### Diet

Neanderthals were once thought to have been scavengers but were likely apex predators. They ate mammoths, birds, ungulates, edible plants such as nuts, moss and legumes, mushrooms, boars, elephants, rhinoceros, fish, dolphins, seals, urchins, shellfish, crabs, and sometimes bears. All of these depend on where the group lived and how they gathered food. Neanderthals were able to prepare food with a wide range of techniques, but were unable to boil soup, stew or animal stock. However, they did boil bone marrow to make fat stocks. Neanderthal teeth decreased in size from 100,000 years and later, which suggests they had depended in softened food as a result of cooking advancement. Neanderthals were probably preserving their food, since many game were brought back to their home within hunting season and it would be difficult to consume the game before spoilage. They were also capable of food storage. Food such as shellfish spoils quickly, so they would have had to preserve them quickly after gathering, such as storage in water-soaked algae. Neanderthals had high competition with other Ice Age predators that hunted similar prey. Neanderthals are occasionally known to be prone to cannibalism, processing other Neanderthals similar to how large ungulate game was processed. This is explained as ritual, pre-burial to deter foul stenches or scavengers, an act of war or for food. It is possible cannibalism was not common and only happened during food sources, which is is similar to many recorded incidents of cannibalism in apomorphic human history. A 76,000 year-old Neanderthal hunting camp was described in 2021 from the Navalmaíllo Rock Shelter. Here, tools and fire were used to process the large bovids and cervids this group ate. The Neanderthals from Sidrón Cave are known to have lived not long before the Neanderthal extinction event, showing signs of starvation and they themselves were cannibalized by other Neanderthals. It is thought that the Neanderthals of this time had difficulty gathering food, and since they require more food, and in turn, calories than Homo sapiens, groups had to compete with each other groups and eventually assimilated. Italian Neanderthals bear injuries consistent with swimmer's ear and thus diving 3-4 meters for clamshells. However, Stringer notes that Inuit, who do not often swim, have the same injuries, which are due to wet ears and wind chill. Neanderthals and *Homo sapiens* from Franchthi and Shanidar Cave, revealed using analysis of charred food, suggests that plants were frequently combined in complex cooking steps to make food more palatable. DNA traces of penicillium has been found in Neanderthal plaque from El Sidrón, suggesting the use of fungal antibiotics; individuals here were also found to have ingested chamomile and yarrow to aid in digestion, as well as mushrooms, nuts, moss and poplars.

### Technology

There are numerous cases of Neanderthals creating jewelry and art with available materials. Neanderthals and other ancient *Homo* have also bee found creating cave murals depicting ancient fauna, teaching other Neanderthals and abstract patterns including dots, lines and hand prints. Some cases of prehistoric music have also been documented, including a flute made of bone that may or may not have been created by Neanderthals. In terms of tools, Neanderthals made Mousterian tools, which had various geographical sub-types. They also used natural elements as tools. Some populations may show signs of migrating *Homo sapiens* sharing their techniques with local Neanderthals. It is unknown of Neanderthals used ranged weapons. Several cases of fires used by Neanderthals have been recorded but it is unknown if they created the fires or were simply taking advantage of their natural occurrence. Bark tar was used for hafting. Evidence suggests Neanderthals were capable of creating boats and sailing to islands. Since Neanderthals lived lives of extensive traumatic injury, many of which had high mortality rates, they likely treated wounds to some extent. They probably had knowledge of medicinal plants similar to apomorphic human. The Ndutu skull was found at Lake Ndutu, which was full of faunal and lithic remains, of which 20 were definitely tools comprised of spheroids, 3 regular flakes, 2 triangular flakes, 1 rectangular flake and hammerstones. Later, Acheulean hand axes were discovered.

## Paleoecology

Analysis of the environment in southwestern France shows that Neanderthal populations were likely overturned by *H. sapiens* through forest opening, placing ecological pressure on both parties.

## Cryptozoology

Shackley (1982) presented evidence to suggest that Neanderthals had survived to the present day in high-altitude Russia and Mongolia to explain some Mousterian-grade tools found in these places despite a lack of biological evidence. She classifies these as "unidentified hominids", although Walls (1984) considers her claims "neither new nor unique". Russian and North American a anthropologists find the idea of living Neanderthals controversial. Walls notes that Shackley did not consider research presented at the University of British Columbia's Sasquatch and Similar Phenomena conference that defined "wildmen" as myth, that her idea shares many features with the North American ape cryptids such as sasquatch, the folklorist's work is ignored and there is no skeletal remains to support this. Cryptohominids termed "Neanderthaloid wildmen" include *Dinanthropoides nivalis* (the yeti), *H. pongoides* or *H. n. pongoides* (wildmen of the Almas and Yeren), *Primihomo asiaticus* (Ksy-Gyik), *H. s. almas* (wildmen of the Almas), *Troglodytes recens* and *Paleanthropus* or *Troglodytes fossilis* (a postulated direct ancestor that survives in the present day). In western Matto Grosso, there are reported "Neanderthaloid Submen" called the and reported by Col. P.H. Fawcett in 1914 and published by his son Brian in 1953. They describe hairy hominids using bows, lacking "acceptable speech"; Sanderson (1962) presumed them plausible but did not assume that they were extant.

## Subspecies

### Reassigned Subspecies

* *H. n. steinheimensis* (=Steinheim)
* *H. n. neanderthalensis*
* *H. n. reilingensis* (=Reilingen)
* *H. n. pongoides* (="Neanderthaloid wildmen")
* *H. n. soloensis*
* *H. n. krapinensis* (=Krapina)
	+ *H. n. var. krapinensis*
* *H. n. predmostensis*/*predmosti* (=Předmostí)
* *H. n. palaestinus*
* *H. n. pekinensis*
* *H. n. javanensis*
* *H. n. rhodesiensis*
* *H. n. njarasensis*
* *H. n. talgajensis*
* *H. n. lagoensis*
* *H. n. aniensis* (=Aniene River)
* *H. n. recens*
* *H. n. fossilis*

## Synonyms

* *Palaeoanthropus*
* *Protanthropus*
* *Acanthropus*
* *Azykhanthropus*
* *Homo-proscopinus*
* *Pithecanthropus*
* *H. sapiens krapinensis* (=Krapina)
* *H. sapiens neanderthalensis*
* *H. sapiens neandertalensis* (lapsus)
* *H. sapiens shanidarensis* (=Shanidar)
* *H. sapiens steinheimensis* (=Steinheim)
* *H. sapiens neanderthalensis*
* *H. sapiens praeneanderthalensis* (=Pre-Neanderthal)
* *H. sapiens hungaricus* (=Vértesszőlős)
* *H. sapiens anteneanderthalensis*
* *H. sapiens stupidus*
* *H. sapiens aniensis* (=Aniene River)
* *H. sapiens krapinensis* (=Krapina)
* *H. erectus reilingensis* (=*H. erectus*, =Reilingen)
* *H. erectus palestinus*
* *H. erectus hungaricus*
* *H. heidelbergensis steinheimensis*
* *H. stupidus*
* *H. primigenius*
	+ *H. p. galilaeensis* (=Amud)
	+ *H. p. faber*
	+ *H. p.* "alalus"
	+ *H. p. asiaticus*
	+ *H. p. krapinensis*
	+ *H. p. var. spyensis*
	+ *H. europaeus primigenius*
* *H. palestinensis/palestinus*
	+ *H. sapiens palestinus* (=Palestinean Man)
* *H. antiquus* (=Cro-Magnon, Neanderthal)
* *H. transprimigenius mousteriensis*
* *H. mousteriensis hauseri* (=Le Moustier)
* *H. priscus*
* *H. chapellensis* (=La Chapelle-aux-Saints 1)
* *H. acheulensis moustieri*
* *H. lemousteriensis* (=Le Moustier)
* *H. naulettensis* (=Naulette Man)
* *H. heringsdorfensis* (=Ehringsdorf)
* *H. galilensis* (=Amud)
* *H. kiikobiensis* (=Kiik-Koba)
* *H. neanderthalis* (lapsus)
* *H. neandertalensis* (lapsus)
* *H. steinheimensis* (=Steinheim)
* *H. breladensis* (=La Cotte de St Brelade)
* *H. azykhensis* (=Azykh Cave)
* *H. proscopinus*
* *H. paleohungaricus* (=Vértesszőlős)
	+ *H. erectus seu sapiens paleohungaricus*
	+ *H. "paleo-hungaricus"*
* *H. reilingensis* (=Reilingen)
* *H. calpicus*/*calpensis*/*gibraltarensis* (=Gibraltar)
* *H. aniensis* (=Aniene River)
* *H. spyensis* (=Spy)
* *H. praeneanderthalensis*/*präneanderthalensis*/*preneanderthalensis* (=Pre-Neanderthal)
* *H. camerotensis* (=Camerota)
* *H. transprimigenius* (=Le Moustier, La Chapelle, La Ferrassie, La Quina)
* *H. anteneanderthalensis*
* *H. faber*
* *H. incipiens*

## Notable Specimens

* Krapina: Krapina is a Croatian site documenting many Neanderthal specimens including material culture of various mixed classes, many skeletal specimens and even jewelry such as an eagle talon modified, dyed and strung on rope. Estimates place the site at 130,000 years old but unpublished faunal data from Chris Stringer and Rainer Grun would suggest 250,000 years. Krapina 4. 5 20 and 31 have small blunt trauma; 34.7 has a depressed skull fracture; 106 and 110 have extremely degenerated C4 and C7 cervicals; 120.71 has a fragmented rib with fibrous dysplasia; 149 has a right clavicle that was fractured and well-healed; 180 has a fracture and nounion in the right ulna; and 188.8 has a fracture and bowing in the left ulna. All of these would have been fatal since they caused massive blood loss, limited mobility or would have caused disability if not for the care of fellow group members. On the contrary, evidence of cannibalism is present since the specimens were defleshed and presumably consumed. However, defleshing may be a burial ritual rather than cannibalism. 80 individuals from crania and dental material were recovered. They average cranial capacities are 1326-1359. Postcrania are also known, including parts of the spine. Teeth show evidence as being used to carry an item in their mouths. Neanderthals numbed tooth pain by using a direct approach, as evidenced by grooving and scratching of the enamel of KDP (Krapina Dental Person) 20. They were once known as *H. (sapiens) krapinensis*.
* La Cotte de St Brelade Individual A-B: From this cave are 48,000-year-old Neanderthal teeth with traits intermediate of Neanderthals and *Homo sapiens*. While they belong to 2 Neanderthals, the teeth bear a cervix with apomorphic morphology and other features such as a lacking transverse crest with Neanderthal morphology. They were discovered in La Cotte de St Brelade from Jersey, which bears more than 20,000 tools only 8000 years before Neanderthals went extinct. These are thus the most recent Neanderthal sites. The teeth are planned to be DNA tested to test this hybrid hypothesis. They were once called *H. breladensis*.
* Steinheim skull: A skull, half being deformed and caved in via taphonomic processes. This specimen was restored in a 3D modelling program in 2021 using mirroring and retrodeformation to find the original shape. With this, it is assigned to Neanderthals. It was first named *Homo (sapiens) "steinheimensis"*, which was abandoned in favour of assignment to either *Homo heidelbergensis* or Neanderthals. The specimen was discovered in a gravel pit among animal bones and no artifacts; the excavators suspected human remains may be present. A slow-growing meningioma 51 × 43 × 25 millimeters with a volume of 29 milliliters was present. It did not impair the individual but did cause headaches. However, postcrania would be needed to investigate the cause of death. This is the earliest evidence of this type of tumour, since meningiomas are exceedingly rare.
* Denisova 17 (DC4969): A specimen described in 2021 by Brown et al., aged and genetically examined. It is an undiagnostic fragment identified as Neanderthal.
* Bawa Yawan: A milk tooth belonging to a toddler individual from one of the last Neanderthal populations in the world, 42,000 years old. It is either from an individual who died and only the tooth preserved, or it fell from the mouth of an alive individual. It was discovered in the Iranian rock shelter of Bawa Yawan, a region that was also inhabited by *Homo sapiens* and that they competed for thousands of years. It's morphology is distinctly Neanderthal.
* "Vertesszolos Man/occipital", "Sámuel" or "Samu": Found in Vértesszőlős, Central Transdanubia, Hungary in 1962 by Márton Pécsi, it was named Sámuel (the name day of the biblical Samuel) and later shortened to Samu due to the Hungarian abbreviation of "Samuel". Hungarian anthropologist Andor Thoma named the specimen as "*Homo erectus seu sapiens paleohungaricus*", believing it to be 500,000 years old; analyses in the 1990s determine the specimen is 250,000-300,000 years old (Middle Pleistocene), and the fossil was lumped into *Homo erectus* and then *Homo heidelbergensis* or *H. erectus*. Two immature teeth and Abbevilian tools are found here, and the site is now open to the public with a cast presented at a local museum, with the original deposited at the Hungarian National Museum with animal footprints and tools. "Samu" is sometimes used as Hungarian slang for plastic skeletons used in classrooms.
* Scladina Cave tooth: A 40,000 year old tooth discovered in 2022 near Scalyn, Belgium. At this time, it was one of the most significant find here. The tooth preserves tartar and other histological details necessary for studying the life history of this individual. They are searching for further remains at the site in in their collections, and a life-sized replica was displayed at a museum in Ardenne, with the original stored in an air-conditioned room.
* Megalopolis molar: An upper third molar that is likely to be a *Homo neanderthalensis* or of a more derived Neanderthal population (though it was once considered to overlap with the *Homo erectus* hypodigm) akin to Steinheim from the Megalopolis Basin of Greece. It first appeared in a 2016 encyclopedia and was noted to be undescribed. It was described by Röding et al. (2021). It was briefly mentioned in Xirotiris et al. (1979). The crown is eroded and the overall specimen is notably small. Some attributes are basal and indicates a very early hominin presence at Peloponnese, Greece 400-500,000 years ago.
* Azykh: A mandibular fragment of an 'elderly" (for the time) woman from Azykh Cave, dated from 300-400 kya. It is one of the oldest fossils from the Caucasus, and her identity as a Neanderthal or *Homo heidelbergensis* is debated. The fossil is also known as *Azykhanthropus* and *Homo azykhensis*.
* Cova Foradada: A skull, thorax and one arm were discovered in 2010, found articulated in a small cave hollow. These remains remain undescribed, and may be evidence of burial. Other Neanderthals from the site include lithics and a maxilla and corresponding cranial fragments found in 2000.
* Ndutu cranium: A 400,000 year-old skull found in Tanzania. R.J. Clarke in 1976 referred it to *H. erectus*, and has been viewed as such since, with *H. sapiens* attributes recognized. Due to comparative research and similar finds, the latter seems most appropriate. Indirect cranial capacity suggests 1100 ml. The morphology of the sulcus and a protuberance existence leads Phillip Rightmire to state the skull is unlike *H. erectus*. However, Stinger (1986) suggests a thickened illiac pillar is in *H. erectus*. Clarke (1989) explains that an expanded parietal and occipital regions of the brain make it of *H. sapiens*. In 2021, it was referred to *Homo bodoensis*.
* Ceprano Man, Argil or Ceprano Calvarium: This hominin is known from a single calvaria unearthed during highway construction near Ceprano, Italy in 1994. Although the fossil was initially damaged by a bulldozer, it was studied by Italo Biddittu who was near when the skull cap was found. Mallegni et al. in 2003 proposed a new name for the fossil, *Homo cepranensis*, which has stuck since. However, some consider the fossil an *H. heidelbergensis*, but Mounier et al. (2011) state the fossil is an appropriate ancestor to *H. heidelbergensis*, and proceed unique traits in the region. Of the holotype, the sulcus supraorbitalis is partial, a frontal tuber is half-developed and shifts medially, the supraorbital region concaves medially and the external auditory meatus is positioned intermediate regarding the processus zygomaticus temporalis are all exclusive characteristic of Mid-Pleistocene individuals. The torus occipitalis transversus is straight and articular tubercle is medio-laterally concaved, which are defined as derived. The petro-tympanic crests downward position, the opisthocranion coexists with the inion, the processus retromastoideus and torus angularis parietalis are all considered primitive. Based on regional correlations and numerous absolute dates, the fossil was dated 690,000-900,000 years old. In 2021, it was suggested to possibly be *Homo bodoensis*.
* Castel di Guido: One "*H.* *erectus**"* from Castel di Guido was initially thought to represent a hyoid showing a lack of speech capabilities. However, later analysis shows that the bone is a vertebral rim. The remains from this site are around 400kya and represent early Neanderthals.

### Artifacts

* Tepe Hesar: An Iranian Chalcolithic and Paleolithic (40-70,000 years) site preserving Levallois Neanderthal tools. They are coated in a shiny black of brown substance called desert polish. They were created by Neanderthals.
* Aranbaltza II: Over 5000 Châtelperronian tools from the Iberian Peninsula, indicating that Neanderthals had been replaced and gone extinct before their total extinction date. These populations likely originated from French populations.
* Bešeňová Báňa: A Slovakian site first described in 2022. From this site are mollusc and mammal fossils near an abandoned travertine quarry. Herbivore remains bear modification, and with other stratigraphical, paleoecological, taphonomical and taxonomical evidence, they suggest that this Middle Pleistocene site was an important Neanderthal settlement.
* Plakias: Artifacts from Crete, Greece were found by a US-Greek archeological team while combing a Cretan gorge hoping to find small stone tools from *Homo sapiens* when they sailed over about 11,000 years ago. However, Thomas Strasser and team discovered a 5-inch-long handaxe that was knapped from a local quartz cobble and being archaic, a seemingly out-of-place artifact would suggest that *Homo* had reached here long before. This would suggest that the people, first attributed to *Homo heidelbergensis* or *Homo erectus*, from this time were capable sailors. Four US universities and the Directorate of Paleoanthropology and Speleology continued combing and returned with 30 handaxes and similar tools embedded in nine different locations of the southwest coast near Plakias. They may have eroded from caves into sea cliffs and falling into beach deposits. The youngest terraces bearing there are 45,000 years old using radiocarbon and the oldest is at least 130 kya, though these are not concrete. If these Neanderthals left from Turkey or Greece, they would have traveled 12-24 miles (19-39 kilometers) respectively or would have sailed over 125 miles (200 kilometers) from Africa. This trace suggests that the island held a sustainable population. These may suggest that other populations sailed as well.
* Arenero de Pedro Jaro I: Acheulean lithics from the type locality of *Dama celiae*, having bones with cut marks that suggest hunting and consumption by local Neanderthals.

## Gallery

### Fossils

Various fossils from Krapina, including Krapina 3 (top left), modified eagle jewelry (bottom middle) and other assorted Neanderthal material (remaining boxes).La Cotte de St Brelade teeth A and B.Steinheim skull.Denisova 17 is a Neanderthal, Denisova 18 is H. sp. and Denisova 19-21 are Denisovan.Vertesszolos occipital.Megalopolis molar.
### Artifacts

Tepe Hesar.Aranbaltza II.Plakias.Arenero de Pedro Jaro I (Orcasitas).
### Reconstructions

**Credit: Tom Björklund.****Credit: Tom Björklund.****Credit: Tom Björklund.****Credit: Tom Björklund.****Credit: Tom Björklund.****Credit: Tom Björklund.****Credit: Britannica.****Credit: Tom Björklund.**

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