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For much of the early 20th century, Neanderthal behaviour was depicted as primitive, unintelligent and brutish. Although knowledge and perception of them has markedly changed since then in the scientific community, the image of the unevolved caveman archetype remains prevalent in popular culture.

Overall, Neanderthals maintained a low population and population density, and also mainly interacted with only nearby neighbours. Many groups suffered from inbreeding depression. Communities may have seasonally migrated between caves, but most of the raw materials Neanderthals used were collected within only 5 km (3.1 mi) of a site. Indicated by frequent evidence of stunted growth and traumatic injuries, Neanderthals lived harsh lives, which may be implicated in the 150,000 year stagnation in Neanderthal stone tool innovation.

Still, Neanderthal technology achieved a degree of sophistication. It includes the Mousterian stone tool industry as well as the abilities to maintain and maybe create fire, build cave hearths, craft at least simple clothes similar to blankets and ponchos, make use of medicinal plants, treat severe injuries, store food, and use various cooking techniques such as roasting, boiling, and smoking.

Neanderthals consumed a wide array of food, mainly what was abundant in their immediate vicinity. This was normally hoofed mammals such as red deer, but also megafauna, plants, small mammals, birds, and aquatic and marine resources. Although they were probably apex predators, they still competed with cave lions, cave hyenas and other large predators. A number of examples of symbolic thought and Palaeolithic art have been inconclusively attributed to Neanderthals, namely possible ornaments made from bird claws and feathers, shells, collections of unusual objects including crystals and fossils, and engravings. Some claims of religious beliefs have been made. The extent to which Neanderthals could produce speech and use language are debated.

Popular conceptions

Charles R. Knight's 1911 "Snowbound Neanderthals"

By the early 20th century, several Neanderthal fossils were identified across Europe, establishing H. neanderthalensis as a legitimate species. The most influential specimen was La Chapelle-aux-Saints 1 ("The Old Man") from La Chapelle-aux-Saints, France. French palaeontologist Marcellin Boule authored several publications, among the first to establish palaeontology as a science, detailing the specimen, but reconstructed him as slouching, ape-like, and only remotely related to modern humans.

Boule fuelled the popular image of Neanderthals as barbarous, slouching, club-wielding primitives; this image was reproduced for several decades and popularised in prehistoric fiction works, such as the 1911 The Quest for Fire by J.-H. Rosny aîné and the 1927 The Grisly Folk by H. G. Wells in which they are depicted as monsters. In 1911, Scottish anthropologist Arthur Keith reconstructed La Chapelle-aux-Saints 1 as an immediate precursor to modern humans, sitting next to a fire, producing tools, wearing a necklace, and having a more humanlike posture, but this failed to garner much scientific rapport, and Keith later abandoned his thesis in 1915.

By the middle of the century, the scientific community began to rework its understanding of Neanderthals based on new fossil discoveries and reevaluations of earlier material. Ideas such as Neanderthal intelligence and culture became mainstream, and a more humanlike image of them emerged. In 1939, American anthropologist Carleton Coon reconstructed a Neanderthal in a modern business suit and hat to emphasise that they would be, more or less, indistinguishable from modern humans had they survived into the present. William Golding's 1955 novel The Inheritors depicts Neanderthals as much more emotional and civilised. In modern-day, Neanderthal reconstructions are often very humanlike, but the question of Neanderthal behavioural modernity is still debated.

Still, in modern popular culture, the "caveman" archetype often mocks Neanderthals as primitive, hunchbacked, knuckle-dragging, club-wielding, grunting, nonsocial characters driven solely by animal instinct. "Neanderthal" can also be used as an insult. In literature, they are sometimes depicted as brutish or monstrous, such as Elizabeth Marshall Thomas' The Animal Wife, but sometimes with a civilised but unfamiliar culture, as in Björn Kurtén's Dance of the Tiger, and Jean M. Auel's Clan of the Cave Bear and her Earth's Children series.

Social structure

Group dynamics

Skeleton of a Neanderthal child discovered in Roc de Marsal near Les Eyzies, France, on display at the Hall of Human Origins, Washington, D.C.

Reliable evidence of Neanderthal group composition comes from:

• Cueva del Sidrón, Spain showing 7 adults, 3 adolescents, 2 juveniles and an infant;
• the footprints of Le Rozel, France, showing (based on footprint size) a group of 10 to 13 members where all but one were juveniles or adolescents,
• and Chagyrskaya Cave in the Siberian Altai Mountains showing 6 adults and 5 children (including a father, daughter, and possibly cousins). This family may have died of starvation.

A Neanderthal child's teeth analysed in 2018 showed it was born in the spring and weaned after 2.5 years, similar to modern hunter gatherers. Indicated from various ailments resulting from high stress at a low age, such as stunted growth, British archaeologist Paul Pettitt hypothesised that children of both sexes were put to work directly after weaning; and American palaeonthropologist Erik Trinkaus suggested that, upon reaching adolescence, an individual may have been expected to join in hunting large and dangerous game. Neanderthal bone trauma is comparable to modern Inuit, which could suggest a similar childhood between Neanderthals and Cro-Magnons. Stunting may have also resulted from harsh winters and bouts of insufficient food resources.

Bands likely moved between certain caves depending on the season, indicated by remains of seasonal materials such as certain foods, and returned to the same locations generation after generation. Some sites may have been used for over 100 years. Sites showing evidence of no more than three individuals may have represented nuclear families or temporary camping sites for special task groups (such as a hunting party). Cave bears may have competed with Neanderthals for cave space, and there is a decline in cave bear populations starting 50,000 years ago onwards (although their extinction occurred well after Neanderthals had died out). Neanderthals also had a preference for caves whose openings faced towards the south.

Neanderthals are generally considered to have been cave dwellers. Open-air settlements near contemporaneously inhabited cave systems in the Levant could indicate mobility between cave and open-air bases in this area. Evidence for long-term open-air settlements is known from 'Ein Qashish, Israel.

Inter-group relations

Trade

Populations may not have been as capable of inter-group interaction and trade as Cro-Magnons, with genetic data suggesting low population density, and archaeological data indicating the sourcing of artefacts from usually no farther than 5 km (3.1 mi) from the main settlement.

Nonetheless, because a few Neanderthal artefacts in a settlement could have originated 20, 30, 100 and 300 km (12.5, 18.5, 60 and 185 mi) away, there does seem to have at least limited long-distance inter-group relations. British anthropologist Eiluned Pearce and Cypriot archaeologist Theodora Moutsiou speculated that Neanderthals were possibly capable of forming geographically expansive ethnolinguistic tribes encompassing upwards of 800 people, based on the transport of obsidian up to 300 km (190 mi) from the source. Still, they conceded that Neanderthal long-distance networks would have been significantly hindered by smaller population compared to Cro-Magnons.

Breeding

Neanderthal mother with child depicted in the Anthropos Pavilion of the Moravian Museum

Low population caused a low genetic diversity and probably inbreeding, which reduced the population's ability to filter out harmful mutations (inbreeding depression). It is unclear how this affected a single Neanderthal's genetic burden and, thus, if this caused a higher rate of birth defects than in Cro-Magnons. If it did, it could have contributed to the extinction of the species (mutational meltdown).

The DNA of a Neanderthal from Denisova Cave, Russia, shows that she had an inbreeding coefficient of 1⁄8. The discovery of such an individual may indicate that inbreeding was common here. The 13 inhabitants of Sidrón Cave collectively exhibited 17 different birth defects likely due to inbreeding or recessive disorders.

When inter-group resettling did happen, at least some groups may have practiced patrilocal residency (a woman moved out of her group to live with her partner), as suggested by mtDNA of the Neanderthals of Cueva del Sidrón, Spain. Here, the three adult men belonged to the same maternal lineage, while the three adult women belonged to different ones.

Population dynamics

Genetically, Neanderthals can be grouped into three distinct regions (above). Dots indicate sampled specimens.

Genetic analysis indicates there were at least three distinct geographical groups:

• Western Europe,
• the Mediterranean coast,
• and east of the Caucasus, with some migration among these regions.

Post-Eemian Western European Mousterian stone tools can also be broadly grouped into three distinct macro-regions:

• Acheulean-tradition Mousterian in the southwest,
• Micoquian in the northeast,
• and Mousterian with bifacial tools (MBT) in between the former two. MBT may actually represent the interactions and fusion of the two different cultures.

Southern Neanderthals exhibit regional anatomical differences from northern counterparts, namely:

• a less protrusive jaw,
• a shorter gap behind the molars,
• and a vertically higher jawbone.

These all suggest Neanderthal communities regularly interacted with closely neighbouring communities within a region, but not as often beyond.

Over long periods of time, there is evidence of large-scale cross-continental migration. Early specimens from Mezmaiskaya Cave in the Caucasus and Denisova Cave in the Siberian Altai Mountains differ genetically from Western European Neanderthals, whereas later specimens from these caves both have genetic profiles more similar to Western European Neanderthals than to the earlier specimens from the same locations. These suggest large-scale population replacement over time. Similarly, artefacts and DNA from Chagyrskaya and Okladnikov Caves, also in the Altai Mountains, resemble those of eastern European Neanderthal sites about 3,000–4,000 km (1,900–2,500 mi) away more than they do artefacts and DNA of the older Neanderthals from Denisova Cave, suggesting two distinct migration events into Siberia.

Neanderthals seem to have suffered a major population decline during MIS 4 (71–57,000 years ago), and the distribution of the Micoquian tradition could indicate that Central Europe and the Caucasus were repopulated by communities from a refuge zone either in eastern France or Hungary (the fringes of the Micoquian tradition) who dispersed along the rivers Prut and Dniester.

Conflict

There is some evidence of conflict: a skeleton from La Roche à Pierrot, France, showing a healed fracture on top of the skull apparently caused by a deep blade wound, and another from Shanidar Cave, Iraq, with a rib lesion characteristic of projectile weapon injuries.

It was once believed that Neanderthals practised ritual murder, headhunting, and cannibalism, but this is no longer widely supported.

Social hierarchy

Reconstruction of an elderly Neanderthal man and child in the Natural History Museum, Vienna

It is sometimes suggested that, since Neanderthals were hunters of challenging big game and lived in small groups, there was no sexual division of labour as seen in modern hunter-gatherer societies. That is, men, women, and children all had to be involved in hunting, instead of men hunting and women and children foraging. On the other hand, with modern hunter-gatherers, typically the higher the meat dependency, the more stringent the division of labour. Further, tooth-wearing patterns in Neanderthal men and women suggest they commonly used their teeth for carrying items, but men exhibit more wearing on the upper teeth, and women the lower, suggesting some cultural differences in tasks.

It is controversially proposed that some Neanderthals wore decorative clothing or jewellery—such as a leopard skin or raptor feathers—to display elevated status in the group. Trinkaus suggested that elderly Neanderthals were given special burial rites for lasting so long given the high mortality rates. Alternatively, many more Neanderthals may have received burials, but the graves were infiltrated and destroyed by, for example, bears. Given that 20 graves of Neanderthals aged under 4 have been found—over a third of all known graves—deceased children may have received greater care during burial than other age demographics.

Looking at Neanderthal skeletons from several natural rock shelters, Trinkaus found that, although Neanderthals were recorded as bearing several trauma-related injuries, none of them had significant trauma to the legs that would debilitate movement. He suggested that self worth in Neanderthal culture derived from contributing food to the group; a debilitating injury would remove this self-worth and result in near-immediate death, and individuals who could not keep up with the group while moving from cave to cave were left behind. There are nonetheless examples of individuals with other debilitating injuries being nursed for several years. There are examples of infirming injured group members in even earlier human species. Especially given the high trauma rates, it is possible that such an altruistic strategy ensured the survival of the species for so long.

Cuisine

See also: Pleistocene human diet

Hunting and gathering

Reconstruction of a Neanderthal man butchering a goat at the Neanderthal Museum

Neanderthals were once thought of as scavengers, but are now considered to have been apex predators. Inhabiting usually a forested landscape, Neanderthals were likely ambush hunters, getting close to and attacking their target—a prime adult—in a short burst of speed, thrusting in a spear at close quarters. Younger or wounded animals may have been hunted using traps, projectiles, or pursuit.

Some sites show evidence that Neanderthals slaughtered whole herds of animals in large, indiscriminate hunts and then carefully selected which carcasses to process. In 1980, it was hypothesised that two piles of mammoth skulls at La Cotte de St Brelade, Jersey, at the base of a gulley were evidence of mammoth drive hunting (causing them to stampede off a ledge), but this is contested.

They appear to have eaten predominantly what was abundant within their immediate surroundings, with steppe-dwelling communities (generally outside of the Mediterranean) subsisting almost entirely on meat from large game, forest-dwelling communities consuming a wide array of plants and smaller animals, and waterside communities gathering aquatic resources. Nonetheless, even in more southerly, temperate areas such as the southeastern Iberian Peninsula, large game still featured prominently in Neanderthal diets. Cro-Magnons, in contrast, seem to have used more complex food extraction strategies and generally had a more diverse diet.

Nonetheless, Neanderthals still would have had to have eaten a varied enough diet to prevent nutrient deficiencies and protein poisoning, especially in the winter when they presumably ate mostly lean meat. Any food with high contents of other essential nutrients not provided by lean meat would have been vital components of their diet, such as fat-rich brains, carbohydrate-rich and abundant underground storage organs (including roots and tubers), or, like modern Inuit, the stomach contents of herbivorous prey items.

Food items

Prey items

Red deer, the most commonly hunted Neanderthal game

For meat, Neanderthals appear to have fed predominantly on hoofed mammals. They primarily consumed red deer and reindeer, as these two were the most abundant game; but they also ate other Pleistocene megafauna such as chamois, ibex, wild boar, steppe wisent, aurochs, Irish elk, woolly mammoth, straight-tusked elephant, woolly rhinoceros, Merck's rhinoceros the narrow-nosed rhinoceros, wild horse, and so on. There is evidence of directed cave and brown bear hunting both in and out of hibernation, as well as butchering. Analysis of Neanderthal bone collagen from Vindija Cave, Croatia, shows nearly all of their protein needs derived from animal meat.

Neanderthals also hunted small game. Some caves show evidence of small regular rabbit and tortoise consumption. At Gibraltar sites, there are remains of 143 different bird species, many ground-dwelling such as the common quail, corn crake, woodlark, and crested lark. Scavenging birds such as corvids and eagles were commonly exploited.

Neanderthals exploited marine resources on the Iberian, Italian and Peloponnesian Peninsulas, where they waded or dived for shellfish, as early as 150,000 years ago at Cueva Bajondillo, Spain, similar to the fishing record of modern humans. At Vanguard Cave, Gibraltar, the inhabitants consumed Mediterranean monk seal, short-beaked common dolphin, common bottlenose dolphin, Atlantic bluefin tuna, sea bream and purple sea urchin; and at Gruta da Figueira Brava, Portugal, there is evidence of large-scale harvest of shellfish, crabs and fish. Evidence of freshwater fishing was found in Grotte di Castelcivita, Italy, for trout, chub and eel; Abri du Maras, France, for chub and European perch; Payré, France; and Kudaro Cave, Russia, for Black Sea salmon.

Plant items

The archaeological record shows that Neanderthals consumed a variety of plants and mushrooms across their range. Bone traumas in the leg joints could possibly suggest habitual squatting, which, if the case, was likely done while gathering foodplants.

Neanderthals from Cueva del Sidrón, Spain, based on dental tartar, consumed mushrooms, pine nuts, and moss, indicating they were forest foragers. Dental tartar from Grotte de Spy, Belgium, indicates the inhabitants had a meat-heavy diet including woolly rhinoceros and mouflon sheep, while also regularly consuming mushrooms. Neanderthal faecal matter from El Salt, Spain, dated to 50,000 years ago—the oldest human faecal matter remains recorded—show a diet mainly of meat but with a significant component of plants.

Evidence of cooked plant foods—mainly legumes and, to a far lesser extent, acorns—was discovered at Kebara Cave, Israel, with its inhabitants possibly gathering plants in spring and fall and hunting in all seasons except fall (the cave was probably abandoned in late summer to early fall). Remnants from Amud Cave, Israel, indicates a diet of figs, palm tree fruits and various cereals and edible grasses.

At Shanidar Cave, Iraq, Neanderthals collected plants with various harvest seasons, indicating they scheduled returns to the area to harvest certain plants, and that they had complex food-gathering behaviours for both meat and plants.

Food preparation

Neanderthals probably could employ a wide range of cooking techniques, such as roasting, and they may have been able to heat up or boil soup, stew, or animal stock. The abundance of animal bone fragments at settlements may indicate the making of fat stocks from boiling bone marrow, possibly taken from animals that had already died of starvation. These methods would have substantially increased fat consumption, which was a major nutritional requirement of communities with low carbohydrate and high protein intake. Neanderthal tooth size had a decreasing trend after 100,000 years ago, which could indicate an increased dependence on cooking or the advent of boiling, a technique that would have softened food.

Yarrow growing in Spain

At Cueva del Sidrón, Spain, Neanderthals likely cooked and possibly smoked food, as well as used certain plants—such as yarrow and camomile—as flavouring, although these plants may have instead been used for their medicinal properties. At Gorham's Cave, Gibraltar, Neanderthals may have been roasting pinecones to access pine nuts.

At Grotte du Lazaret, France, a total of twenty-three red deer, six ibexes, three aurochs, and one roe deer appear to have been hunted in a single autumn hunting season, when strong male and female deer herds would group together for rut. The entire carcasses seem to have been transported to the cave and then butchered. Because this is such a large amount of food to consume before spoilage, it is possible these Neanderthals were curing and preserving it before winter set in. At 160,000 years old, it is the oldest potential evidence of food storage. The great quantities of meat and fat which could have been gathered in general from typical prey items (namely mammoths) could also indicate food storage capability. With shellfish, Neanderthals needed to eat, cook, or in some manner preserve them soon after collection, as shellfish spoils very quickly. At Cueva de los Aviones, Spain, the remains of edible, algae eating shellfish associated with the alga Jania rubens could indicate that, like some modern hunter gatherer societies, harvested shellfish were held in water-soaked algae to keep them alive and fresh until consumption.

Competition

Cave hyena skeleton

Competition from large Ice Age predators was rather high. Cave lions likely targeted horses, large deer, and wild cattle; and European leopards primarily reindeer and roe deer; which overlapped with Neanderthal diet. To defend a kill against such ferocious predators, Neanderthals may have engaged in a group display of yelling, arm waving, or stone throwing; or quickly gathered meat and abandoned the kill. At Grotte de Spy, Belgium, the remains of wolves, cave lions, and cave bears indicate Neanderthals hunted their competitors to some extent.

Neanderthals and cave hyenas may have exemplified niche differentiation, and actively avoided competing with each other. Although they both mainly targeted the same groups of creatures—deer, horses, and cattle—Neanderthals mainly hunted the former and cave hyenas the latter two. Animal remains from Neanderthal caves similarly indicate they preferred to hunt prime individuals, whereas cave hyenas hunted weaker or younger prey. Nonetheless, there is evidence that cave hyenas stole food and leftovers from Neanderthal campsites and scavenged on dead Neanderthal bodies. At Payre, France, the Neanderthals may have partitioned resources with wolves.

Cannibalism

Neandertal remains from the Troisième caverne of Goyet Caves (Belgium). The remains have scrape marks, indicating that they were butchered, with cannibalism being the "most parsimonious explanation".

There are several instances of Neanderthals practising cannibalism across their range. The first example came from Krapina, Croatia, in 1899, and other examples were found at the Spanish Cueva del Sidrón and Zafarraya; and the French Grotte de Moula-Guercy, Les Pradelles, and La Quina. For the five cannibalised Neanderthals at the Grottes de Goyet, Belgium, there is evidence that the upper limbs were disarticulated, the lower limbs defleshed and also smashed (likely to extract bone marrow), the chest cavity disembowelled, and the jaw dismembered. There is also evidence that the butchers used some bones to retouch their tools. The processing of Neanderthal meat at Grottes de Goyet is similar to how they processed horse and reindeer. About 35% of the Neanderthals at Marillac-le-Franc, France, show clear signs of butchery, and the presence of digested teeth indicates that the bodies were abandoned and eaten by scavengers, likely hyaenas.

These cannibalistic tendencies have been explained as either ritual defleshing, pre-burial defleshing (to prevent scavengers or foul smell), acts of war, or simply food preparation. Because of a small number of cases, and the higher number of cut marks seen on cannibalised individuals than animals (indicating inexperience), cannibalism was probably not a very common practice. It may have only been done in times of extreme food shortages, as in some cases in recorded human history.

The arts

See also: Prehistoric art and Art of the Middle Paleolithic

Personal adornment

Neanderthals used ochre, a clay earth pigment. Ochre is well documented from 45,000 to 60,000 years ago in Neanderthal sites, with the earliest example dating to 200,000 to 250,000 years ago from Maastricht-Belvédère, the Netherlands (a similar timespan to the ochre record of H. sapiens). It has been hypothesised to have functioned as body paint, and analyses of pigments from Pech de l'Azé, France, indicates they were applied to soft materials (such as a hide or human skin). Red ochre can also have several utilities outside of decoration, such as medicine, tanning hides, food preservative, and insect repellent. Containers apparently used for mixing ochre pigments were found in Peștera Cioarei, Romania, which could indicate modification of ochre for solely aesthetic purposes.

Decorated king scallop shell from Cueva Antón, Spain. Interior (left) with natural red colouration, and exterior (right) with traces of unnatural orange pigmentation

Neanderthals collected uniquely shaped objects and are suggested to have modified them into pendants. Such objects include:

• a fossil Aspa marginata sea snail shell possibly painted red from Grotta di Fumane, Italy, transported over 100 km (62 mi) to the site about 47,500 years ago;
• three shells from Cueva de los Aviones, Spain, dated to about 120–115,000 years ago, perforated through the umbo belonging to:
  • a rough cockle associated with red and yellow pigments,
  • a Glycymeris insubrica also associated with red and yellow pigments,
  • and a Spondylus gaederopus associated with red-to-black mix of hematite and pyrite;
• and a king scallop shell with traces of an orange mix of goethite and hematite from Cueva Antón, Spain.

The discoverers of the latter two claim that pigment was applied to the exterior to make it match the naturally vibrant inside colouration. Excavated from 1949 to 1963 from the French Grotte du Renne, Châtelperronian beads made from animal teeth, shells and ivory were found associated with Neanderthal bones. The dating is uncertain, and Châtelperronian artefacts may actually have been crafted by modern humans and simply redeposited with Neanderthal remains.

Speculative reconstruction of white-tailed eagle talon jewellery from Krapina, Croatia (arrows indicate cut marks)

Gibraltarian palaeoanthropologists Clive and Geraldine Finlayson suggested that Neanderthals used various bird parts as artistic media, specifically black feathers. In 2012, the Finlaysons and colleagues examined 1,699 sites across Eurasia, and argued that raptors and corvids, species not typically consumed by any human species, were overrepresented and show processing of only the wing bones instead of the fleshier torso, and thus are evidence of feather plucking of specifically the large flight feathers for use as personal adornment. They specifically noted the cinereous vulture, red-billed chough, kestrel, lesser kestrel, alpine chough, rook, jackdaw and the white tailed eagle in Middle Palaeolithic sites. Other birds claimed to present evidence of modifications by Neanderthals are the golden eagle, rock pigeon, common raven and the bearded vulture. The earliest claim of bird bone jewellery is a number of 130,000-year-old white tailed eagle talons found in a cache near Krapina, Croatia, speculated, in 2015, to have been a necklace. A similar 39,000-year-old Spanish imperial eagle talon necklace was reported in 2019 at Cova Foradà in Spain, though from the contentious Châtelperronian layer. In 2017, 17 incision-decorated raven bones from the Zaskalnaya VI rock shelter, Ukraine, dated to 43–38,000 years ago were reported. Because the notches are more-or-less equidistant to each other, they are the first modified bird bones that cannot be explained by simple butchery, and for which the argument of design intent is based on direct evidence.

Discovered in 1975, the so-called Mask of la Roche-Cotard, a mostly flat piece of flint with a bone pushed through a hole on the midsection—dated to 32, 40, or 75,000 years ago—has been purported to resemble the upper half of a face, with the bone representing eyes. It is contested whether it represents a face, or if it even counts as art. In 1988, American archaeologist Alexander Marshack speculated that a Neanderthal at Grotte de L'Hortus, France, wore a leopard pelt as personal adornment to indicate elevated status in the group based on a recovered leopard skull, phalanges and tail vertebrae.

Abstraction

Giant deer bone of Einhornhöhle, Germany, c. 49,000 BC

The scratched floor of Gorham's Cave, Gibraltar, c. 39,000 BC

As of 2014, 63 purported engravings have been reported from 27 different European and Middle Eastern Lower-to-Middle Palaeolithic sites, of which 20 are on flint cortexes from 11 sites, 7 are on slabs from 7 sites, and 36 are on pebbles from 13 sites. It is debated whether or not these were made with symbolic intent. In 2012, deep scratches on the floor of Gorham's Cave, Gibraltar, were discovered, dated to older than 39,000 years ago, which the discoverers have interpreted as Neanderthal abstract art. The scratches could have also been produced by a bear. In 2021, an Irish elk phalanx with five engraved offset chevrons stacked above each other was discovered at the entrance to the Einhornhöhle cave in Germany, dating to about 51,000 years ago. A flint flake at the Mousterian site of Kiik-Koba in Crimea, Ukraine, is decorated with an engraving which would have required skilled workmanship.

In 2018, some red-painted dots, disks, lines and hand stencils on the cave walls of the Spanish La Pasiega, Maltravieso, and Doña Trinidad were dated to be older than 66,000 years ago, prior to the arrival of modern humans in Western Europe. If the dating is correct, this would indicate Neanderthal authorship, and similar iconography recorded in other Western European sites—such as Les Merveilles, France, and Cueva del Castillo, Spain—could potentially also have Neanderthal origins. The dating of these Spanish caves, and thus attribution to Neanderthals, is contested.

Neanderthals are known to have collected a variety of unusual objects—such as crystals or fossils—without any real functional purpose or any indication of damage caused by use. It is unclear if these objects were simply picked up for their aesthetic qualities, or if some symbolic significance was applied to them. These items are mainly quartz crystals, but also other minerals such as cerussite, iron pyrite, calcite and galena. A few findings feature modifications, such as:

• a mammoth tooth with an incision and a fossil nummulite shell with a cross etched in from Tata, Hungary;
• a large slab with 18 cupstones hollowed out from a grave in La Ferrassie, France;
• and a geode from Peștera Cioarei, Romania, coated with red ochre.

A number of fossil shells are also known from French Neanderthals sites, such as:

• a rhynchonellid and a Taraebratulina from Combe Grenal;
• a belemnite beak from Grottes des Canalettes; a polyp from Grotte de l'Hyène;
• a sea urchin from La Gonterie-Boulouneix;
• and a rhynchonella, feather star, and belemnite beak from the contentious Châtelperronian layer of Grotte du Renne.

Music

See also: Prehistoric music

The Divje Babe Flute in the National Museum of Slovenia

Purported Neanderthal bone flute fragments made of bear long bones were reported from Potočka zijalka, Slovenia, in the 1920s, and Istállós-kői-barlang, Hungary, and Mokriška jama, Slovenia, in 1985; but these are now attributed to Cro-Magnons.

The 43,000-year-old Divje Babe flute from Slovenia, found in 1995, has been attributed by some researchers to Neanderthals, though its status as a flute is heavily disputed. Many researchers consider it to be most likely the product of a carnivorous animal chewing the bone, but its discoverer Ivan Turk and other researchers have continued to argue that it was manufactured by Neanderthals as a musical instrument.

Handheld tools

Stone

Main article: Mousterian

Mousterian point

The archaeological record demonstrates a long 150,000-year stagnation in Neanderthal lithic innovation, which may have followed from the low population density and high injury rate, preventing especially complex technologies from being taught to the next generation. Still, Neanderthal lithic technology did achieve some complexities.

Neanderthals are associated with the Mousterian industry. The Mousterian is also associated with North African H. sapiens as early as 315,000 years ago and was found in Northern China about 47–37,000 years ago in caves such as Jinsitai or Tongtiandong. In Europe, it was invented around 300,000 years ago with the Levallois technique, which developed directly from the Acheulean industry (invented by H. erectus about 1.8 million years ago). Levallois made it easier to control flake shape and size. In experiments with modern humans, the Levallois technique can be taught with purely observational learning without spoken instruction.

There are distinct regional variants of the Mousterian industry, such as:

• the Quina and La Ferrassie subtypes of the Charentian industry in southwestern France
• the Acheulean-tradition Mousterian subtypes A and B along the Atlantic and northwestern European coasts
• the Denticulate Mousterian industry in Western Europe
• the racloir industry around the Zagros Mountains
• the flake cleaver industry of Cantabria, Spain, and both sides of the Pyrenees
• the Micoquian industry of Central and Eastern Europe and the related Sibiryachikha variant in the Siberian Altai Mountains

In the mid-20th century, French archaeologist François Bordes debated against American archaeologist Lewis Binford to explain this diversity (the "Bordes–Binford debate"), with Bordes arguing that these represent unique ethnic traditions and Binford that they were a response to varying environments (essentially, form vs. function). The latter sentiment would indicate a lower degree of inventiveness compared to Cro-Magnons, adapting the same tools to different environments rather than creating new technologies. A continuous sequence of occupation is well documented in Grotte du Renne, France, where the lithic tradition can be divided into the Levallois–Charentian, Discoid–Denticulate (43,300  ±929 – 40,900 ±719 years ago), Levallois Mousterian (40,200 ±1,500 – 38,400 ±1,300 years ago), and Châtelperronian (40,930 ±393 – 33,670 ±450 years ago).

There is some debate if Neanderthals had long-ranged weapons. A wound on the neck of an African wild ass from Umm el Tlel, Syria, was likely inflicted by a heavy Levallois-point javelin, and bone trauma consistent with habitual throwing has been reported in Neanderthals. Some spear tips from Abri du Maras, France, may have been too fragile to have been used as thrusting spears, possibly suggesting their use as darts.

Bone

The Châtelperronian in central France and northern Spain is a distinct industry from the Mousterian, and is controversially hypothesised to represent a culture of Neanderthals borrowing (or by process of acculturation) tool-making techniques from immigrating modern humans, crafting bone tools and ornaments. In this frame, the makers would have been a transitional culture between the Neanderthal Mousterian and the modern human Aurignacian. The opposing viewpoint is that the Châtelperronian was manufactured by Cro-Magnons instead. Abrupt transitions similar to the Mousterian/Châtelperronian could also simply represent natural innovation, like the La Quina–Neronian transition 50,000 years ago featuring technologies generally associated with modern humans such as bladelets and microliths. Other ambiguous transitional cultures include the Italian Uluzzian industry, and the Balkan Szeletian industry.

Before modern human immigration, the only evidence of Neanderthal bone tools are animal rib lissoirs—which are rubbed against hide to make it more supple or waterproof—although this could also be evidence for modern humans immigrating earlier than expected. In 2013, two 51,400- to 41,100-year-old deer rib lissoirs were reported from Pech-de-l'Azé and the nearby Abri Peyrony in France. In 2020, five more lissoirs made of aurochs or bison ribs were reported from Abri Peyrony, with one dating to about 51,400 years ago and the other four to 47,700–41,100 years ago. This indicates the technology was in use in this region for a long time. Since reindeer remains were the most abundant, the use of less abundant bovine ribs may indicate a specific preference for bovine ribs. Potential lissoirs have also been reported from Grosse Grotte, Germany (made of mammoth), and Grottes des Canalettes, France (red deer).

Shell

Smooth clam shell scrapers from Grotta dei Moscerini, Italy

The Neanderthals in 10 coastal sites in Italy (namely Grotta del Cavallo and Grotta dei Moscerini) and Kalamakia Cave, Greece, are known to have crafted scrapers using smooth clam shells, and possibly hafted them to a wooden handle. They probably chose this clam species because it has the most durable shell. At Grotta dei Moscerini, about 24% of the shells were gathered alive from the seafloor, meaning these Neanderthals had to wade or dive into shallow waters to collect them.

At Grotta di Santa Lucia, Italy, in the Campanian volcanic arc, Neanderthals collected the porous volcanic pumice, which, for Cro-Magnons, was probably used for polishing points and needles. The pumices are associated with shell tools.

Plant

At Abri du Maras, France, twisted fibres and a 3-ply inner-bark-fibre cord fragment associated with Neanderthals show that they produced string and cordage. It is unclear how widespread this technology was because the materials used to make them (such as animal hair, hide, sinew, or plant fibres) are biodegradable and preserve very poorly. Dating to 52–41,000 years ago, the cord fragment is the oldest direct evidence of fibre technology, although 115,000-year-old perforated shell beads from Cueva Antón possibly strung together to make a necklace are the oldest indirect evidence. In 2020, British archaeologist Rebecca Wragg Sykes expressed cautious support for the genuineness of the find, but pointed out that the string would have been so weak that it would have had limited functions, such as stringing small objects.

Digging stick from Aranbaltza III, Spain

At the Italian Poggetti Vecchi site, there is evidence that they used fire to process boxwood branches to make digging sticks, a common implement in hunter-gatherer societies.

Neanderthal produced the adhesive birch bark tar, using the bark of birch trees, for hafting. It was long believed that birch bark tar was an intensely complicated procedure evincing complex cognitive skills and strong cultural transmission, until a 2019 study showed it can be made simply by burning birch bark beside smooth vertical surfaces, such as a flat, inclined rock. Though some communities may not have needed cultural processes to maintain the technology, the Neanderthals at Königsaue, Germany, employed a technically more demanding underground production method, which may demonstrate some cumulative cultural practices.

The archaeological record shows that Neanderthals commonly used animal hide and birch bark, and may have used them to make cooking containers. This is based largely on circumstantial evidence, as neither fossilises well. It is possible that the Neanderthals at Kebara Cave in Israel, used the shells of the spur-thighed tortoise as containers.

Fire and construction

Many Neanderthal sites have evidence of fire, some for extended periods of time, though it is unclear whether they were capable of starting fire or simply scavenged from naturally occurring wildfires. Indirect evidence of fire-starting ability includes pyrite residue on a couple dozen bifaces from roughly 50,000 years ago in northwestern France. This could indicate they were used as percussion fire starters. Some Neanderthals around this time were collecting manganese dioxide, which can lower the combustion temperature of wood. They may have been using fire for cooking, keeping warm, and deterring predators. They were also capable of zoning areas for specific activities, such as for knapping, butchering, hearths, and wood storage. Many Neanderthal sites lack evidence for such activity perhaps due to natural degradation of the area over tens of thousands of years, such as by bear infiltration after abandonment of the settlement.

In a number of caves, evidence of hearths has been detected. Neanderthals likely considered air circulation when making hearths as a lack of proper ventilation for a single hearth can render a cave uninhabitable in several minutes. Abric Romaní rock shelter, Spain, indicates eight evenly spaced hearths lined up against the rock wall, likely used to stay warm while sleeping, with one person sleeping on either side of the fire. At Cueva de Bolomor, Spain, with hearths lined up against the wall, the smoke flowed upwards to the ceiling, and led to outside the cave. In Grotte du Lazaret, France, smoke was probably naturally ventilated during the winter as the interior cave temperature was greater than the outside temperature; likewise, the cave was likely only inhabited in the winter.

The ring structures in Grotte de Bruniquel, France

In 1990, two 176,000-year-old ring structures, several metres wide, made of broken stalagmite pieces, were discovered in a large chamber more than 300 m (980 ft) from the entrance within Grotte de Bruniquel, France. One ring was 6.7 m × 4.5 m (22 ft × 15 ft) with stalagmite pieces averaging 34.4 cm (13.5 in) in length, and the other 2.2 m × 2.1 m (7.2 ft × 6.9 ft) with pieces averaging 29.5 cm (11.6 in). There were also four other piles of stalagmite pieces for a total of 112 m (367 ft) or 2.2 t (2.4 short tons) worth of stalagmite pieces. Evidence of the use of fire and burnt bones also suggest human activity. A team of Neanderthals was likely necessary to construct the structure, but the chamber's actual purpose is uncertain. Building complex structures so deep in a cave is unprecedented in the archaeological record, and indicates sophisticated lighting and construction technology, and great familiarity with subterranean environments.

Clothes

Two racloir side scrapers from Le Moustier, France

As opposed to the bone sewing-needles and stitching awls found in Cro-Magnon sites, the only known Neanderthal tools that could have been used to fashion clothes are hide scrapers. These could have been used to make items similar to blankets or ponchos, and there is no direct evidence they could produce fitted clothes. Indirect evidence of tailoring by Neanderthals includes the ability to manufacture string, which could indicate weaving ability, and a naturally-pointed horse metatarsal bone from Cueva de los Aviones, Spain, which was speculated to have been used as an awl to perforate dyed hides, based on the presence of orange pigments. Whatever the case, Neanderthals would have needed to cover up most of their body, and Cro-Magnons would have covered 80–90%.

Since modern human/Neanderthal admixture is known to have occurred in the Middle East, and no modern body louse species descends from their Neanderthal counterparts (body lice only inhabit clothed individuals), it is possible Neanderthals (and/or modern humans) in hotter climates did not wear clothes, or Neanderthal lice were highly specialised.

Seafaring

There are claims of Neanderthal seafaring activity. Remains of Middle Palaeolithic stone tools on Greek islands have been alleged to indicate early seafaring by Neanderthals in the Ionian Sea possibly starting as far back as 150,000 to 200,000 years ago. The oldest stone artefacts from Crete date to 107,000 to 130,000 years ago, Cephalonia 125,000 years ago, and Zakynthos 35,000 to 110,000 years ago. The makers of these artefacts may have employed simple reed boats and made one-day crossings back and forth. Other Mediterranean islands with such early remains include Sardinia, Melos, Alonnisos, and Naxos (although Naxos may have been connected to land). If they could cross major waterbodies, it is possible they crossed the Strait of Gibraltar. If this interpretation is correct, Neanderthals' ability to engineer boats and navigate through open waters would speak to their advanced cognitive and technical skills.

British archaeologist Cyprian Broodbank finds most of the evidence unconvincing. He considers Cephalonia the most convincing, and Neanderthals may have made brief visits to small islands. There are no British Neanderthal fossils dating to when the region was probably an island, only when it was likely connected to the continent.

Medicine

Neanderthals appear to have lived lives of frequent traumatic injury and recovery. Well-healed major fractures on many bones indicate the setting of splints. Individuals with severe head and rib traumas (which would have caused massive blood loss) indicate they had some manner of dressing major wounds, such as bandages made from animal skin. By and large, they appear to have avoided severe infections, indicating good long-term treatment of such wounds.

Their knowledge of medicinal plants was comparable to that of Cro-Magnons. An individual at Cueva del Sidrón, Spain, seems to have been medicating a dental abscess using poplar—which contains salicylic acid, the active ingredient in aspirin—and there were also traces of the antibiotic-producing Penicillium chrysogenum. They may also have used yarrow and camomile, and their bitter taste—which should act as a deterrent as it could indicate poison—means it was likely a deliberate act. At the Kebara Cave in Israel, plant remains which have historically been used for their medicinal properties were found, including the common grape vine, the pistachios of the Persian turpentine tree, ervil seeds and oak acorns.

Language

It is not known whether the Neanderthals had the capacity for advanced language, but some researchers have argued that a complex language—possibly using syntax—was probably necessary to survive in their harsh environment, with Neanderthals needing to communicate about topics such as locations, hunting and gathering, and tool-making techniques.

In 1971, cognitive scientist Philip Lieberman attempted to reconstruct the Neanderthal vocal tract and concluded that it was similar to that of a modern human newborn. His reconstruction of the Neanderthal vocal tract left Neanderthals incapable of producing a range of speech sounds, due to the large size of the mouth and the small size of the pharyngeal cavity, thus no need for a descended larynx to fit the entire tongue inside the mouth. He concluded that they were anatomically unable to produce the sounds /a/, /i/, /u/, /ɔ/, /g/, and /k/ and thus lacked the capacity for articulate speech, though were still able to speak at a level higher than non-human primates. Lieberman's findings have subsequently been debated, especially in light of newer fossil discoveries and re-analyses. Some subsequent studies can reconstruct the Neanderthal vocal apparatus as comparable to that of modern humans, with a similar vocal repertoire. The lack of a descended larynx does not necessarily equate to a reduced vowel capacity.

Reconstruction of the Kebara 2 skeleton at the Natural History Museum, London

The 1983 discovery of a Neanderthal hyoid bone—used in speech production in humans—in Kebara 2 is almost identical to that of humans, which could suggest Neanderthals were capable of speech. Still, is not possible to accurately reconstruct the entire vocal tract with just the hyoid.

The FOXP2 gene in modern humans is associated with speech and language development. FOXP2 was present in Neanderthals, but not the gene's modern human variant. Neurologically, Neanderthals had an expanded Broca's area—operating the formulation of sentences, and speech comprehension, but out of a group of 48 genes believed to affect the neural substrate of language, 11 had different methylation patterns between Neanderthals and modern humans. This could indicate a stronger ability in modern humans than in Neanderthals to express language.

The Sima de los Huesos individuals, which may represent the ancestors of Neanderthals, had humanlike hyoid and ear bones, which could suggest the early evolution of the modern human vocal apparatus. In 2015, Lieberman hypothesised that Neanderthals were capable of syntactical language, although nonetheless incapable of mastering any human dialect.

Religion

See also: Paleolithic religion and History of religion

Funerals

Reconstruction of the grave of La Chapelle-aux-Saints 1 at the Musée de La Chapelle-aux-Saints

Although Neanderthals did bury their dead, at least occasionally—which may explain the abundance of fossil remains—the behaviour is not indicative of a religious belief of life after death because it could also have had non-symbolic motivations, such as great emotion or the prevention of scavenging.

Estimates made regarding the number of known Neanderthal burials range from thirty-six to sixty. The oldest confirmed burials do not seem to occur before approximately 70,000 years ago. The small number of recorded Neanderthal burials implies that the activity was not particularly common. The setting of inhumation in Neanderthal culture largely consisted of simple, shallow graves and pits. Sites such as La Ferrassie in France or Shanidar in Iraq may imply the existence of mortuary centers or cemeteries in Neanderthal culture due to the number of individuals found buried at them.

The debate on Neanderthal funerals has been active since the 1908 discovery of La Chapelle-aux-Saints 1 in a small, artificial hole in a cave in southwestern France, very controversially postulated to have been buried in a symbolic fashion. Another grave at Shanidar Cave, Iraq, was associated with the pollen of several flowers that may have been in bloom at the time of deposition—yarrow, centaury, ragwort, grape hyacinth, joint pine and hollyhock. The medicinal properties of the plants led American archaeologist Ralph Solecki to claim that the man buried was some leader, healer, or shaman, and that "The association of flowers with Neanderthals adds a whole new dimension to our knowledge of his humanness, indicating that he had 'soul' ". It is also possible the pollen was deposited by a small rodent after the man's death.

The graves of children and infants, especially, are associated with grave goods such as artefacts and bones. The grave of a newborn from La Ferrassie, France, was found with three flint scrapers, and an infant from Dederiyeh Cave, Syria, was found with a triangular flint placed on its chest. A 10-month-old from Amud Cave, Israel, was associated with a red deer mandible, likely purposefully placed there given other animal remains are now reduced to fragments. Teshik-Tash 1 from Uzbekistan was associated with a circle of ibex horns, and a limestone slab argued to have supported the head. Nonetheless, these contentiously constitute evidence of symbolic meaning as the grave goods' significance and worth are unclear.

Cults

It was once argued that the bones of the cave bear, particularly the skull, in some European caves were arranged in a specific order, indicating an ancient bear cult that killed bears and then ceremoniously arranged the bones. This would be consistent with bear-related rituals of recent Arctic hunter-gatherers. The alleged peculiarity of the arrangement could also be sufficiently explained by natural causes. There may also be confirmation bias here with the idea that totemism was the earliest religion.

It was also once thought that Neanderthals ritually hunted, killed, and cannibalised other Neanderthals, with the skull becoming the focus of some ceremony. In 1962, Italian palaeontologist Alberto Blanc believed a skull from Grotta Guattari, Italy, had evidence of a swift blow to the head—indicative of ritual murder—and a precise and deliberate incising at the base to access the brain. He compared it to the victims of headhunters in Malaysia and Borneo, putting it forward as evidence of a skull cult. It can also be sufficiently explained by cave hyaena scavengery. Although Neanderthals are known to have practised cannibalism, there is unsubstantial evidence to suggest ritual defleshing.

In 2019, Gibraltarian palaeoanthropologists Stewart, Geraldine and Clive Finlayson and Spanish archaeologist Francisco Guzmán speculated that the golden eagle had iconic value to Neanderthals like in some recent societies. They reported that golden eagle bones had a conspicuously high rate of evidence of modification compared to the bones of other birds, forwarding a "Cult of the Sun Bird" where the golden eagle was a symbol of power. There is evidence from Krapina, Croatia, from wear use and even remnants of string, that suggests that raptor talons were worn as personal ornaments.

Bibliography

• Broodbank, Cyprian (2013). The Making of the Middle Sea: A History of the Mediterranean from the Beginning to the Emergence of the Classical World. London, UK: Thames & Hudson. ISBN 978-0-500-29208-2.
• Papagianni, D.; Morse, M. A. (2013). "Still with us?". Neanderthals rediscovered: how modern science is rewriting their story. Thames and Hudson. ISBN 978-0-500-77311-6.
• Pettitt, Paul; White, Mark (2012). The British Palaeolithic: hominin societies at the edge of the Pleistocene world. London: Routledge. ISBN 978-0415674546.
• Tattersall, I. (2015). "Neanderthals, DNA, and creativity". The strange case of the Rickety Cossack: and other cautionary tales from human evolution. St. Martin's Publishing Group. ISBN 978-1-4668-7943-0.
• Shipman, P. (2015). "How humans and their dogs drove Neanderthals to extinction". The invaders: how humans and their dogs drove Neanderthals to extinction. Harvard University Press. doi:10.2307/j.ctvjf9zbs. ISBN 978-0-674-42538-5. JSTOR j.ctvjf9zbs.
• Finlayson, C. (2019). The smart Neanderthal: bird catching, cave art, and the cognitive revolution. Oxford University Press. ISBN 978-0-19-251812-5.
• Romagnoli, Francesca; Rivals, Florent; Benazzi, Stefano (2022). Updating Neanderthals: Understanding Behavioural Complexity in the Late Middle Palaeolithic. Academic Press. ISBN 978-0-12-821429-9.</ref>

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Cryptodira – 11 families, 74 genera, over 200 species Family Genera Carettochelyidae Boulenger, 1887 (1 genus) Genus Carettochelys Ramsay, 1886 – one species Common name Scientific name IUCN Red List Status Range Picture Pig-nosed turtle C. insculpta Ramsay, 1886 VU IUCN Southern New Guinea and northern Northern Territory Cheloniidae (sea turtles) Oppel, 1811 (5 genera) Genus Caretta Rafinesque, 1814 – one species Common name Scientific name IUCN Red List Status Range Picture Loggerhead sea turtle C. caretta Linnaeus, 1758 VU IUCN Genus Lepidochelys (Ridley sea turtles) Fitzinger, 1843 – two species Common name Scientific name IUCN Red List Status Range Picture Kemp's ridley sea turtle L. kempii Garman, 1880 CR IUCN Olive ridley sea turtle L. olivacea von Eschscholtz, 1829 VU IUCN Genus Chelonia Brongniart, 1800 – one species Common name Scientific name IUCN Red List Status Range Picture Green sea turtle C. mydas Linnaeus, 1758 EN IUCN Genus Eretmochelys Fitzinger, 1843 – one species Common name Scientific name IUCN Red List Status Range Picture Hawksbill sea turtle E. imbricata Linnaeus, 1758 CR IUCN Genus Natator McCulloch, 1908 – one species Common name Scientific name IUCN Red List Status Range Picture Flatback sea turtle N. depressus Garman, 1880 DD IUCN Chelydridae Gray, 1831(2 genera) Genus Chelydra (snapping turtles) Schweigger, 1812 – three species Common name Scientific name IUCN Red List Status Range Picture Common snapping turtle C. serpentina Linnaeus, 1758 LC IUCN Central American snapping turtle C. rossignonii Bocourt, 1868 VU IUCN Southeastern Mexico, southern Belize, central Guatemala, and northwestern Honduras Dermatemydidae Gray, 1870 (1 genus) Genus Dermatemys Gray, 1847 – one species Common name Scientific name IUCN Red List Status Range Picture Central American river turtle D. mawii Gray, 1847 CR IUCN Eastern Mexico, Guatemala, Honduras, and Belize Dermochelyidae Fitzinger, 1843 (1 genus) Genus Dermochelys de Blainville, 1816 – one species Common name Scientific name IUCN Red List Status Range Picture Leatherback sea turtle D. coriacea Vandelli, 1761 VU IUCN Emydidae Rafinesque, 1815 (12 genera) Genus Clemmys von Ritgen, 1828 – one species Common name Scientific name IUCN Red List Status Range Picture Spotted turtle C. guttata Schneider, 1792 EN IUCN Great Lakes region Genus Emys Duméril, 1805 – two species Common name Scientific name IUCN Red List Status Range Picture European pond turtle E. orbicularis Linnaeus, 1758 EN IUCN Sicilian pond turtle E. trinacris Fritz, Fattizzo, Guicking, Tripepi, Pennisi, Lenk, Joger and Wink, 2005 DD IUCN Genus Emydoidea Holbrook, 1838 – one species Common name Scientific name IUCN Red List Status Range Picture Blanding's turtle E. blandingii Holbrook, 1838 EN IUCN Genus Actinemys Baird and Girard, 1852 – one species Common name Scientific name IUCN Red List Status Range Picture Western pond turtle A. marmorata Baird and Girard, 1852 VU IUCN Genus Glyptemys Agassiz, 1857 – two species Common name Scientific name IUCN Red List Status Range Picture Bog turtle G. muhlenbergii Schoepff, 1801 CR IUCN Wood turtle G. insculpta Le Conte, 1830 EN IUCN Genus Terrapene (box turtles) Merrem, 1820 – four species Common name Scientific name IUCN Red List Status Range Picture Common box turtle T. carolina Linnaeus, 1758 VU IUCN Eastern coast of North America, and the Gulf of Mexico Eastern box turtle T. c. carolina Florida box turtle T. c. bauri Gulf Coast box turtle T. c. major Three-toed box turtle T. c. triunguis Mexican box turtle T. c. mexicana Yucatán box turtle T. c. yucatana Coahuilan box turtle T. coahuila Schmidt and Owens, 1944 EN IUCN Cuatro Ciénegas, Coahuila, Mexico Spotted box turtle T. nelsoni Stejneger, 1925 DD IUCN Sierra Madre Occidental, Mexico Terrapene ornata] Terrapene ornata Agassiz, 1857 NT IUCN Central United States, including the Mojave desert and the Midwest region Ornate box turtle left, Desert box turtle right Genus Chrysemys Gray, 1844 – one species Common name Scientific name IUCN Red List Status Range Picture Painted turtle C. picta Schneider, 1783 LC IUCN Eastern painted turtle C. p. picta Midland painted turtle C. p. marginata Southern painted turtle C. p. dorsalis Western painted turtle C. p. bellii Geoemydidae Theobald, 1868 24 Kinosternidae Agassiz, 1857 4 Platysternidae Gray, 1869 1 Testudinidae Batsch, 1788 12 Trionychidae Fitzinger, 1826 14 Pleurodira – 3 families, 16 genera, over 60 species Family Genera Chelidae Gray, 1831 15 Pelomedusidae Cope, 1868 2 Podocnemididae Gray, 1869 3 References Further reading David T. Kirkpatrick (November–December 1995). "Platysternon megacephalum". Reptile & Amphibian Magazine. pp. 40–47. Retrieved 26 June 2010. Cogger, H.G.; R.G. Zweifel; D. Kirschner (2004). Encyclopedia of Reptiles & Amphibians Second Edition. Fog City Press. ISBN 1-877019-69-0. External links John B. Iverson; A. Jon Kimerling; A. Ross Kiester. "EMYSystems". Terra Cognita Laboratory, Geosciences Department of Oregon State University. Retrieved 26 June 2010.

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Cetacean anatomy is the study of the form or morphology of cetaceans (whales, dolphins and porpoises). It can be contrasted with cetacean physiology, which is the study of how the component parts of cetaceans function together in these living marine mammals. In practice, cetacean anatomy and cetacean physiology complement each other, the former dealing with the structure of a cetacean, its organs or component parts and how they are put together, such as might be observed on the dissecting table or under the microscope, and the latter dealing with how those components function together in the living marine mammal. The anatomy of cetaceans have common characteristics with other terrestrial mammals, and, in addition, is often shaped by the physical characteristics of aquatic living, the medium in which these mammals live. Water is much denser than air, holds a relatively small amount of dissolved oxygen, and absorbs more light than air does. Body Skeleton Skull Unlike toothed whales (left), baleen whales (right) do not have a melon The skull of all cetaceans is extended, which can be clearly seen in baleen whales. The nostrils are located on top of the head above the eyes. The back of the skull is significantly shortened and deformed. By shifting the nostrils to the top of the head, the nasal passages extend perpendicularly through the skull. The teeth or baleen in the upper jaw sit exclusively on the maxilla. The braincase is concentrated through the nasal passage to the front and is correspondingly higher, with individual cranial bones that overlap. The bony otic capsule, the petrosal, is only cartilaginous when connected to the skull, so that it can swing independently. Vertebrae The number of vertebrae that make up the spine varies between species, anywhere between 40 and 93 individual vertebrae. The cervical spine, found in all mammals, consists of seven vertebrae which, however, are greatly reduced or fused together. This gives stability during swimming at the expense of mobility. The fins are carried by the thoracic vertebrae, ranging from 9 to 17 individual vertebrae. The sternum is only cartilaginous, but nonetheless strong. The last two to three pairs of ribs are not connected at all and hang freely in the body wall. Behind it is the stable lumbar and tail part of the spine which includes all other vertebrae. Below the caudal vertebrae is the chevron bone; the vortex developed provides additional attachment points for the tail musculature. Limbs The front limbs are paddle-shaped with shortened arms and elongated finger bones, to support the movement. They are united by cartilage. It also leads to a proliferation of the finger members, a so-called hyperphalangy, on the second and third fingers. The only functional joint is the shoulder joint in all cetaceans except for the Amazon river dolphin. The collarbone is completely absent. The movement of cetaceans on land is no longer necessary nor possible, due to the great body weight and the atrophied hindlimbs. In fact the rear limbs have become a rudimentary internal appendage without connections to the spine. External organs Jaw The jaws of toothed whales are designed for catching swift prey. Porpoises have spade-shaped teeth, but dolphins have conical teeth. Cetaceans are monophydonts, meaning they have one set of teeth their entire life. Toothed whales use their jaw to recieve pulses for echolocation. Echoes are received using complex fatty structures around the lower jaw as the primary reception path, from where they are transmitted to the middle ear via a continuous fat body. Lateral sound may be received though fatty lobes surrounding the ears with a similar density to water. Some researchers believe that when they approach the object of interest, they protect themselves against the louder echo by quieting the emitted sound. This is known to happen in bats, but here the hearing sensitivity is also reduced close to a target. As opposed to toothed whales, baleen whales have different jaw designs depending on their feeding behavior. Lunge-feeders, like rorquals, have to expand their jaw to a volume that can be bigger than the whale itself; to do this, the oral cavity inflates to expand the mouth. The inflation of the oral cavity causes the cavum ventrale, the folds (throat pleats) on the throat stretching to the naval, to expand, increasing the amount of water that the mouth can store. The mandible is connected to the skull by dense fibers and cartilage, allowing the jaw to swing open at almost a 90° angle. The mandibular symphysis is also fibrocartilaginous, allowing the jaw to bend which lets in more water. To prevent stretching the mouth too far, rorquals have a sensory organ located in the middle of the jaw to regulate these functions. Gulp-feeders, like right whales, on the other hand swim with an open mouth, filling it with water and prey. This makes their head, which can make up a third of their body weight, huge in order to feed effectively. Not able to expand their mouth like rorquals, right whales must have a head that is large enough to take in enough water and food to feed effectively, carrying their bulk all the time. Beaked whales have a somewhat similar jaw anatomy as rorquals. The throats of beaked whales have a bilaterally paired set of grooves that are associated with their unique feeding mechanism, suction feeding. Instead of capturing prey with their teeth, beaked whales suck it into their oral cavity. Suction is aided by the throat grooves, which stretch and expand to accommodate food. Their tongue can move very freely. By suddenly retracting the tongue and distending the gular (throat) floor, pressure immediately drops within the mouth sucking the prey in with the water. Eyes The whale eye is relatively small for its size, yet they do retain a good degree of eyesight. As well as this, the eyes of a whale are placed on the sides of its head, so their vision consists of two fields, rather than a binocular view like humans have. When belugas surface, their lens and cornea correct the nearsightedness that results from the refraction of light; they contain both rod and cone cells, meaning they can see in both dim and bright light, but they have far more rod cells than they do cone cells. Whales do, however, lack short wavelength sensitive visual pigments in their cone cells indicating a more limited capacity for colour vision than most mammals. Most whales have slightly flattened eyeballs, enlarged pupils (which shrink as they surface to prevent damage), slightly flattened corneas and a tapetum lucidum; these adaptations allow for large amounts of light to pass through the eye and, therefore, a very clear image of the surrounding area. In water, a whale can see around 10.7 metres (35 ft) ahead of itself, but, of course, they have a smaller range above water. They also have glands on the eyelids and outer corneal layer that act as protection for the cornea. Toothed whales can retract and protrude its eyes thanks to a 2-cm-thick retractor muscle attached around the eye at the equator. Blowhole Main article: Blowhole (anatomy) The blowhole is the hole at the top of a whale's head through which the animal breathes air. When a whale reaches the water surface to breathe, they will forcefully expel air through the blowhole. Mucus and carbon dioxide from the animal's metabolism, which have been stored in the whale while diving, are also expelled. The exhalation is released into the comparably lower-pressure and colder atmosphere, so any water vapor condenses. This spray, known as the blow, is often visible from far away as a white splash, which can also be caused by water resting on top of the blowhole. Baleen whales have two blowholes, causing a V-shaped blow, while toothed whales have only one blowhole. The trachea only connects to the blowhole and there is no connection to the esophagus as with humans and most other mammals. Because of this, there is no risk of food accidentally ending up in the animal's lungs, and likewise the animal cannot breathe through its mouth. Consequently, whales have no pharyngeal reflex. Skin Fins Internal organs Intestines The small intestines is divided into three sections: the duodenum, the jejunum, and the ileum. The mesentery is thin in baleen whales. The caecum is present in all whales with the exception of the Amazon river dolphins and the right whales, however it is relatively short in baleen whales. The appendix is absent in all cetaceans. Stomach In most whales, food is swallowed and travels down through the esophagus where it meets a three-chambered-stomach. The first compartment is known as the fore-stomach; this is where food gets ground up into an acidic liquid, which is then squirted into the main stomach. Like in humans, the food is mixed with hydrochloric acid and protein-digesting enzymes. Then, the partly digested food is moved into the third stomach, in which fat-digesting enzymes, and then mixed with an alkaline liquid to neutralize the acid from the first stomach to prevent damage to the intestinal tract. Once the solution is safe, it is moved into the intestinal tract. Kidneys Whale kidneys are specially designed for excreting excess salt content. Water is typically gained by the food they eat, however, the invertebrates they consume have the same salt content as seawater. As in other vertebrates, whale salt levels are three times less than that of seawater. However, the kidneys are inefficient at retaining water, and expel much of it while excreting salt. Spleen Liver The liver in whales is bilobed, as opposed to the five-lobed liver in humans, and they lack a gall bladder. Toothed whales have one bile duct and baleen whales have two. Like other mammals, the liver is located in the right side of the body, just below the diaphragm. Heart Swim bladder Weberian apparatus Reproductive organs Testes Ovaries La peinture de l'Embryon d'un Marsouins (Painting of the embryo of a Porpoise) - from The natural history of marine fish to strange with real peincture & description Daulphin, and several others of his species observed by Pierre Belon du Mans. Nervous system Central nervous system Cerebellum Identified neurons Immune system See also Anatomical terms of location Mammal anatomy References Further reading W. Tinker, Spencer (1988). Whales of the World. ISBN 978-0935848472.