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Main article: Neanderthal

Neanderthalization

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A human skull on the left facing a reconstructed Neanderthal skull on the right, emphasising the difference in braincase shape (more cranial length in Neanderthal), shorter forehead ratio, more defined brow ridge, larger nasal bone projection, pinned-back cheekbone angulation, straighter angled chin, and an occipital bun
Comparisons of a modern Eurasian male example (left) and a Neanderthal (right) skull reconstruction at the Cleveland Museum of Natural History
Front and side view diagram of Neanderthal skull reconstruction emphasising large circular orbits, straightened chin, projecting nasal bridge, large brow ridge, receded forehead, long topped braincase, occipital bun, fossa, and a large gap behind the third molar
Neanderthal skull features

Cranial vault

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Neanderthals had less developed chins, sloping foreheads, and longer, broader, more projecting noses. The Neanderthal skull is typically more elongated, but also wider, and less globular than that of most modern humans, and features much more of an occipital bun,[1] or "chignon", a protrusion on the back of the skull, although it is within the range of variation for modern humans who have it. It is caused by the cranial base and temporal bones being placed higher and more towards the front of the skull, and a flatter skullcap.[2]

Face

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The Neanderthal face is characterised by subnasal[3] as well as mid-facial prognathism, where the zygomatic arches are positioned in a rearward location relative to modern humans, while their maxillary bones and nasal bones are positioned in a more forward direction, by comparison.[4] Neanderthal eyeballs are larger than those of modern humans. One study proposed that this was due to Neanderthals having enhanced visual abilities, at the expense of neocortical and social development.[5] However, this study was rejected by other researchers who concluded that eyeball size does not offer any evidence for the cognitive abilities of Neanderthal or modern humans.[6]

The projected Neanderthal nose and paranasal sinuses have generally been explained as having warmed air as it entered the lungs and retained moisture ("nasal radiator" hypothesis);[7] if their noses were wider, it would differ to the generally narrowed shape in cold-adapted creatures, and that it would have been caused instead by genetic drift. Also, the sinuses reconstructed wide are not grossly large, being comparable in size to those of modern humans. However, if sinus size is not an important factor for breathing cold air, then the actual function would be unclear, so they may not be a good indicator of evolutionary pressures to evolve such a nose.[8] Further, a computer reconstruction of the Neanderthal nose and predicted soft tissue patterns shows some similarities to those of modern Arctic peoples, potentially meaning the noses of both populations convergently evolved for breathing cold, dry air.[9]

Teeth

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Neanderthals featured a rather large jaw which was once cited as a response to a large bite force evidenced by heavy wearing of Neanderthal front teeth (the "anterior dental loading" hypothesis), but similar wearing trends are seen in contemporary humans. It could also have evolved to fit larger teeth in the jaw, which would better resist wear and abrasion,[7][10] and the increased wear on the front teeth compared to the back teeth probably stems from repetitive use. Neanderthal dental wear patterns are most similar to those of modern Inuit.[7]

The incisors are large and shovel-shaped, and, compared to modern humans, there was an unusually high frequency of taurodontism, a condition where the molars are bulkier due to an enlarged pulp (tooth core). Taurodontism was once thought to have been a distinguishing characteristic of Neanderthals which lent some mechanical advantage or stemmed from repetitive use, but was more likely simply a product of genetic drift.[11] The bite force of Neanderthals and modern humans is now thought to be about the same,[7] about 285 N (64 lbf) and 255 N (57 lbf) in modern human males and females, respectively.[12]

Reconstruction of an elderly Neanderthal man

Brain

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The Neanderthal braincase averages 1,640 cm3 (100 cu in) for males and 1,460 cm3 (89 cu in) for females,[13][14] which is significantly larger than the averages for all groups of extant humans;[15] for example, modern European males average 1,362 cm3 (83.1 cu in) and females 1,201 cm3 (73.3 cu in).[16] For 28 modern human specimens from 190,000 to 25,000 years ago, the average was about 1,478 cm3 (90.2 cu in) disregarding sex, and modern human brain size is suggested to have decreased since the Upper Palaeolithic.[17] The largest Neanderthal brain, Amud 1, was calculated to be 1,736 cm3 (105.9 cu in), one of the largest ever recorded in hominids.[14] Both Neanderthal and human infants measure about 400 cm3 (24 cu in).[18]

When viewed from the rear, the Neanderthal braincase has lower, wider, rounder appearance than in anatomically modern humans. This characteristic shape is referred to as "en bombe" (bomb-like), and is unique to Neanderthals, with all other hominid species (including most modern humans) generally having narrow and relatively upright cranial vaults, when viewed from behind.[19][20][21][22] The Neanderthal brain would have been characterised by relatively smaller parietal lobes[23] and a larger cerebellum.[23][24] Neanderthal brains also have larger occipital lobes (relating to the classic occurrence of an occipital bun in Neanderthal skull anatomy, as well as the greater width of their skulls), which implies internal differences in the proportionality of brain-internal regions, relative to Homo sapiens, consistent with external measurements obtained with fossil skulls.[5][25] Their brains also have larger temporal lobe poles,[24] wider orbitofrontal cortex,[26] and larger olfactory bulbs,[27] suggesting potential differences in language comprehension and associations with emotions (temporal functions), decision making (the orbitofrontal cortex) and sense of smell (olfactory bulbs). Their brains also show different rates of brain growth and development.[28] Such differences, while slight, would have been visible to natural selection and may underlie and explain differences in the material record in things like social behaviours, technological innovation and artistic output.[29][30]

Build

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Neanderthals had more robust and stockier builds than typical modern humans,[31] wider and barrel-shaped rib cages; wider pelvises;[32][33] and proportionally shorter forearms and forelegs.[34][35]

Size

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Based on 45 Neanderthal long bones from 14 men and 7 women, the average height was 164 to 168 cm (5 ft 5 in to 5 ft 6 in) for males and 152 to 156 cm (5 ft 0 in to 5 ft 1 in) for females.[31] For comparison, the average height of 20 males and 10 females Upper Palaeolithic humans is, respectively, 176.2 cm (5 ft 9 in) and 162.9 cm (5 ft 4 in), although this decreases by 10 cm (4 in) nearer the end of the period based on 21 males and 15 females.[36] The average in the year 1900 was 163 cm (5 ft 4 in) and 152.7 cm (5 ft 0 in), respectively.[37]

The fossil record shows that adult Neanderthals varied from about 147.5 to 177 cm (4 ft 10 in to 5 ft 10 in) in height. A set of footprints from Le Rozel, France, may have been made by a 73.8 to 184.8 cm (2 ft 5 in to 6 ft 1 in) tall Neanderthal based on footprint length, or a 65.8 to 189.3 cm (2 ft 2 in to 6 ft 3 in) tall Neanderthal based on footprint width.[38]

For Neanderthal weight, samples of 26 specimens found an average of 77.6 kg (171 lb) for males and 66.4 kg (146 lb) for females.[39] Using 76 kg (168 lb), the body mass index for Neanderthal males was calculated to be 26.9–28.2, which in modern humans correlates to being overweight. This indicates a very robust build.[31] The Neanderthal LEPR gene concerned with storing fat and body heat production is similar to that of the woolly mammoth, and so was likely an adaptation for cold climate.[40]

Neanderthal hunters depicted in the Gallo-Roman Museum, Tongeren

Postcranium

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The neck vertebrae of Neanderthals are thicker from the front to the rear and transversely than those of (most) modern humans, leading to stability, possibly to accommodate a different head shape and size.[41]

Although the Neanderthal thorax (where the ribcage is) was similar in size to modern humans, the longer and straighter ribs would have equated to a widened mid-lower thorax and stronger breathing in the lower thorax, which are indicative of a larger diaphragm and possibly greater lung capacity.[33][42][43] The lung capacity of Kebara 2 was estimated to have been 9.04 L (2.39 US gal), compared to the average human capacity of 6 L (1.6 US gal) for males and 4.7 L (1.2 US gal) for females. The Neanderthal chest was also more pronounced (expanded front-to-back, or antero-posteriorly). The sacrum (where the pelvis connects to the spine) was more vertically inclined, and was placed lower in relation to the pelvis, causing the spine to be less curved (exhibit less lordosis) and to fold in on itself somewhat (to be invaginated). This condition of a lumbarised sacrum is rare in modern humans.[44] Such modifications to the spine would have enhanced side-to-side (mediolateral) flexion, better supporting the wider lower thorax. It is claimed by some that this feature would be normal for all Homo, even tropically adapted Homo ergaster or erectus, with the condition of a narrower thorax in most modern humans being a unique characteristic.[33]

Body proportions are usually cited as being "hyperarctic" as adaptations to the cold, because they are similar to those of human populations which developed in cold climates[45]—the Neanderthal build is most similar to that of Inuit and Siberian Yupiks among modern humans[46]—and shorter limbs result in higher retention of body heat.[35][45][47] Nonetheless, Neanderthals from more temperate climates—such as Iberia—still retain the "hyperarctic" physique.[48] In 2019, English anthropologist John Stewart and colleagues suggested Neanderthals instead were adapted for sprinting, because of evidence of Neanderthals preferring warmer wooded areas over the colder mammoth steppe, and DNA analysis indicating a higher proportion of fast-twitch muscle fibres in Neanderthals than in modern humans. He explained their body proportions and greater muscle mass as adaptations to sprinting as opposed to the endurance-oriented modern human physique,[34] as persistence hunting may only be effective in hot climates where the hunter can run prey to the point of heat exhaustion (hyperthermia). They had longer heel bones,[49] reducing their ability for endurance running, and their shorter limbs would have reduced moment arm at the limbs, allowing for greater net rotational force at the wrists and ankles, causing faster acceleration.[34] In 1981, American palaeoanthropologist Erik Trinkaus made note of this alternate explanation, but considered it less likely.[35][50]

Hair and skin colour

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The lack of sunlight most likely led to the proliferation of lighter skin in Neanderthals;[51] however, it has been recently claimed that light skin in modern Europeans was not particularly prolific until perhaps the Bronze Age.[52] Genetically, BNC2 was present in Neanderthals, which is associated with light skin colour; however, a second variation of BNC2 was also present, which in modern populations is associated with darker skin colour in the UK Biobank.[51] DNA analysis of three Neanderthal females from southeastern Europe indicates that they had brown eyes, dark skin colour and brown hair, with one having red hair.[53][54]

In modern humans, skin and hair colour is regulated by the melanocyte-stimulating hormone—which increases the proportion of eumelanin (black pigment) to phaeomelanin (red pigment)—which is encoded by the MC1R gene. There are five known variants in modern humans of the gene which cause loss-of-function and are associated with light skin and hair colour, and another unknown variant in Neanderthals (the R307G variant) which could be associated with pale skin and red hair. The R307G variant was identified in a Neanderthal from Monti Lessini, Italy, and possibly Cueva del Sidrón, Spain.[55] However, as in modern humans, red was probably not a very common hair colour because the variant is not present in many other sequenced Neanderthals.[51]

Metabolism

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Maximum natural lifespan and the timing of adulthood, menopause and gestation were most likely very similar to modern humans.[56] However, it has been hypothesised, based on the growth rates of teeth and tooth enamel,[57][58] that Neanderthals matured faster than modern humans, although this is not backed up by age biomarkers.[59] The main differences in maturation are the atlas bone in the neck as well as the middle thoracic vertebrae fused about 2 years later in Neanderthals than in modern humans, but this was more likely caused by a difference in anatomy rather than growth rate.[60][61]

Generally, models on Neanderthal caloric requirements report significantly higher intakes than those of modern humans because they typically assume Neanderthals had higher basal metabolic rates (BMRs) due to higher muscle mass, faster growth rate and greater body heat production against the cold;[62][63][64] and higher daily physical activity levels (PALs) due to greater daily travelling distances while foraging.[63][64] However, using a high BMR and PAL, American archaeologist Bryan Hockett estimated that a pregnant Neanderthal would have consumed 5,500 calories per day, which would have necessitated a heavy reliance on big game meat; such a diet would have caused numerous deficiencies or nutrient poisonings, so he concluded that these are poorly warranted assumptions to make.[64]

Neanderthals may have been more active during dimmer light conditions rather than broad daylight because they lived in regions with reduced daytime hours in the winter, hunted large game (such predators typically hunt at night to enhance ambush tactics), and had large eyes and visual processing neural centres. Genetically, colour blindness (which may enhance mesopic vision) is typically correlated with northern-latitude populations, and the Neanderthals from Vindija Cave, Croatia, had some substitutions in the Opsin genes which could have influenced colour vision. However, the functional implications of these substitutions are inconclusive.[65] Neanderthal-derived alleles near ASB1 and EXOC6 are associated with being an evening person, narcolepsy and day-time napping.[51]

Pathology

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Injuries

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Neanderthals suffered a high rate of traumatic injury, with an estimated 79–94% of specimens showing evidence of healed major trauma, of which 37–52% were severely injured, and 13–19% injured before reaching adulthood.[66] One extreme example is Shanidar 1, who shows signs of an amputation of the right arm likely due to a nonunion after breaking a bone in adolescence, osteomyelitis (a bone infection) on the left clavicle, an abnormal gait, vision problems in the left eye, and possible hearing loss[67] (perhaps swimmer's ear).[68]

In 1995, Trinkaus estimated that about 80% succumbed to their injuries and died before reaching 40, and thus theorised that Neanderthals employed a risky hunting strategy ("rodeo rider" hypothesis).[59] However, rates of cranial trauma are not significantly different between Neanderthals and Middle Palaeolithic modern humans (although Neanderthals seem to have had a higher mortality risk),[69] there are few specimens of both Upper Palaeolithic modern humans and Neanderthals who died after the age of 40,[70] and there are overall similar injury patterns between them. In 2012, Trinkaus concluded that Neanderthals instead injured themselves in the same way as contemporary humans, such as by interpersonal violence.[71]

A 2016 study looking at 124 Neanderthal specimens argued that high trauma rates were instead caused by animal attacks, and found that about 36% of the sample were victims of bear attacks, 21% big cat attacks, and 17% wolf attacks (totalling 92 positive cases, 74%). There were no cases of hyena attacks, although hyenas still nonetheless probably attacked Neanderthals, at least opportunistically.[72] Such intense predation probably stemmed from common confrontations due to competition over food and cave space, and from Neanderthals hunting these carnivores.[72]

Congenital defects

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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 unknown how this affected a single Neanderthal's genetic burden and, thus, if this caused a higher rate of birth defects than in modern humans.[73]

The 13 inhabitants of Sidrón Cave collectively exhibited 17 different birth defects likely due to inbreeding or recessive disorders.[74] Likely due to advanced age (60s or 70s), La Chapelle-aux-Saints 1 had signs of Baastrup's disease, affecting the spine, and osteoarthritis.[75] Shanidar 1, who likely died at about 30 or 40, was diagnosed with the most ancient case of diffuse idiopathic skeletal hyperostosis (DISH), a degenerative disease which can restrict movement, which, if correct, would indicate a moderately high incidence rate for older Neanderthals.[76]

Diseases

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A mostly complete skeleton laid out against a black background horizontally
La Ferrassie 1 (above) may have had periostitis in his femora and tibiae.[77]

Neanderthals were subject to several infectious diseases and parasites. Modern humans likely transmitted diseases to them; one possible candidate is the stomach bacteria Helicobacter pylori.[78] The modern human papillomavirus variant 16A may descend from Neanderthal introgression.[79] A Neanderthal at Cueva del Sidrón, Spain, shows evidence of a gastrointestinal Enterocytozoon bieneusi infection.[80] The leg bones of the French La Ferrassie 1 feature lesions that are consistent with periostitis—inflammation of the tissue enveloping the bone—likely a result of hypertrophic osteoarthropathy, which is primarily caused by a chest infection or lung cancer.[77]

Neanderthals had a lower cavity rate than modern humans, despite some populations consuming typically cavity-causing foods in great quantity, which could indicate a lack of cavity-causing oral bacteria, namely Streptococcus mutans.[81]

In Neanderthals, the Eustachian tubes (which connect the ear to the throat) are flat. In modern humans, as an infant grows, the Eustachian tubes become angled to permit drainage of the ear and prevent bacterial infection. The flatness of the Eustachian tubes throughout life may have made Neanderthals more prone to developing ear infections.[82] A 2019 study found that 48% of their 77 Neanderthal skull sample size presented bony growths consistent with swimmer's ear (inflammation of the ear canal).[68]

Poisoning

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Two 250,000-year-old Neanderthaloid children from Payré, France, present the earliest known cases of lead exposure. They were exposed on two distinct occasions either by eating or drinking contaminated food or water, or inhaling lead-laced smoke from a fire. There are two lead mines within 25 km (16 mi) of the site.[83]

Collapse

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Extended content
Cryptodira – 11 families, 74 genera, over 200 species
Family[84] Genera[85]
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
A beakless turtle with a snout shaped like that of a pigs
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
World's oceans excluding the polar regions
A white turtle with a beak, black eye-spots, and a dark-brown carapace
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
The Gulf of Mexico and the eastern coast of the United States
A green turtle with a white underside and beak
Olive ridley sea turtle L. olivacea
von Eschscholtz, 1829 		VU IUCN
Coasts of Oceania, South China Sea, Sea of Japan, the Indian Ocean, Africa (excluding the Mediterranean), and the Americas (excluding the eastern coast of North America and southern South America)
alt=A green turtle with a white underside and beak
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
Tropical and temperate oceans of the world
A dark brown turtle with a beak, a green carapace, and a white underside
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
Tropical and subtropical oceans of the world
A turtle with a beak, a black head, flippers, and carapace, a white neck and underside, and serrated scutes on the carapace
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
Around the coast of Australia, stretching to New Guinea and Java, excluding the southern coast
A grey turtle with a white beak
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
United States east of the Rocky Mountains
An orange-brown tortoise with a grey carapace
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
A green turtle with webbed feet
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
Oceans of the world
A large black turtle without a bony shell
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
A black tortoise with yellow speckles across the body and carapace
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
Mediterranean Europe, and around the Caspian Sea
A black tortoise with yellow speckles across the body and carapace
Sicilian pond turtle E. trinacris
Fritz, Fattizzo, Guicking, Tripepi, Pennisi, Lenk, Joger and Wink, 2005 		DD IUCN
Sicily, an island off the coast of Italy
A grey tortoise
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
Great Lakes region in the United States
A black tortoise with yellow spots covering the body
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
Western coast of the contiguous United States
A dark brown turtle with webbed feet, and a yellow-speckled head and neck
Genus Glyptemys Agassiz, 1857 – two species
Common name Scientific name IUCN Red List Status Range Picture
Bog turtle G. muhlenbergii
Schoepff, 1801 		CR IUCN
One population in New England and another population in Virginia, United States
A black turtle with an orange patch on its neck
Wood turtle G. insculpta
Le Conte, 1830 		EN IUCN
New England, Newfoundland, and the Greats Lakes
A black tortoise with a yellow plastron and spots on the head and neck
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 A dark grey tortoise
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 Brown tortoises
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
United States spilling over into Canada excluding the Mojave desert
Eastern painted turtle
C. p. picta 		Midland painted turtle
C. p. marginata 		Southern painted turtle
C. p. dorsalis 		Western painted turtle
C. p. bellii
A grey tortoise with square patterns on the carapace A brown tortoise A grey tortoise with a single thin, orange line running down the carapace from head to tail and white marks on the head and neck A grey tortoise with yellow stripes running down the neck from the head
Underside view, showing a tan plastron Underside view, showing a tan plastron Underside view, showing a tan plastron and webbed feet Underside view, the plastron is bright red with black and white Rorshach-like patterns
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

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  2. ^ Gunz, P.; Harvati, K. (2007). "The Neanderthal "chignon": variation, integration, and homology". Journal of Human Evolution. 52 (3): 262–274. Bibcode:2007JHumE..52..262G. doi:10.1016/j.jhevol.2006.08.010. PMID 17097133.
  3. ^ Lesciotto, K.M.; Cabo, L.L.; Garvin, H.M. (Received August 31, 2015, Accepted April 17, 2016). "A morphometric analysis of prognathism and evaluation of the gnathic index in modern humans" (PDF). "When discussing prognathism, it is important to distinguish it from midfacial projection. Midfacial, or facial, projection has been defined as the 'degree to which [the] face projects in front of [the] cranial base.' In contrast, prognathism, which may be referred to as lower facial, subnasal, or maxillary alveolar prognathism, is defined as the anterior projection of the lower face relative to the upper face. This is also distinct from mandibular prognathism, which may be referred to as basal prognathism, and is more often discussed in clinical literature. For the purposes of this paper, 'prognathism' will refer strictly to subnasal maxillary alveolar projection (i.e., subnasal prognathism)."
  4. ^ Romagnoli, Rivals & Benazzi 2022, p. 75
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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.
Extended content

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.[1] 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.[2][3]

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.[2][3]

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.[2][3]

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.[4] 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.[5] 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.[6]

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.[7] 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.[8] To prevent stretching the mouth too far, rorquals have a sensory organ located in the middle of the jaw to regulate these functions.[9] 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.[10]

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.[11]

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.[12] 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.[13] Toothed whales can retract and protrude its eyes thanks to a 2-cm-thick retractor muscle attached around the eye at the equator.[14]

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.[15]

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.[16]

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

Nervous system Central nervous system Cerebellum Identified neurons Immune system

See also

References

  1. ^ Prosser, C. Ladd (1991). Comparative Animal Physiology, Environmental and Metabolic Animal Physiology (4th ed.). Hoboken, NJ: Wiley-Liss. pp. 1–12. ISBN 0-471-85767-X.
  2. ^ a b c Bruno Cozzi; Sandro Mazzariol; Michela Podestà; Alessandro Zott (2009). "Diving Adaptations of the Cetacean Skeleton" (PDF). The Open Zoology Journal. 2: 24–32. doi:10.2174/1874336600902010024. Retrieved 5 September 2015.
  3. ^ a b c A. Thomas, J. (1916). Outlines of Zoology (5 ed.). pp. 766–771.
  4. ^ The Institute for Marine Mammal Studies. "Frequently asked questions". IMMS. Retrieved 17 February 2016.
  5. ^ Webster, D.; Fay, R.; Popper, A. (1992). "The Marine Mammal Ear: Specializations for aquatic audition and echolocation". In Ketten, D.R. (ed.). The Evolutionary Biology of Hearing. Springer-Verlag. pp. 717–750. ISBN 978-1-4612-7668-5.
  6. ^ Au, W.; Fay, R.; Popper, A. (2000). "Cetacean Ears". Hearing by Whales and Dolphins. SHAR Series for Auditory Research. Springer-Verlag. pp. 43–108. doi:10.1007/978-1-4612-1150-1. ISBN 978-0-387-94906-2.
  7. ^ W. Vogle, A.; A. Lillie, Margo; A. Piscitelli, Marina; A. Goldbogen, Jeremy; D. Pyenson, Nicholas; E. Shadwick, Robert (2015). "Stretchy nerves are an essential component of the extreme feeding mechanism of rorqual whales". Current Biology. 25 (9): 360–361. doi:10.1016/j.cub.2015.03.007.
  8. ^ A. Goldbogen, Jeremy (2010). "The Ultimate Mouthful: Lunge Feeding in Rorqual Whales". American Scientist. 98 (2): 124. doi:10.1511/2010.83.124.
  9. ^ Welsh, Jennifer (2012). "Whale's Big Gulp Aided by Newfound Organ". Retrieved 23 January 2016.
  10. ^ Kenney, Robert D. (2002). "North Atlantic, North Pacific and Southern Right Whales". In William F. Perrin, Bernd Wursig and J. G. M. Thewissen (ed.). The Encyclopedia of Marine Mammals. Academic Press. pp. 806–813. ISBN 0-12-551340-2.
  11. ^ Rommel, S. A.; Costidis, A. M.; Fernandez, A.; Jepson, P. D.; Pabst, D. A.; McLellan, W. A.; Houser, D. S.; Cranford, T. W.; van Helden, A. L.; Allen, D. M.; Barros, N. B. (2006). "Elements of beaked whale anatomy and diving physiology and some hypothetical causes of sonar-related stranding". Journal of Cetacean Research and Management. 7 (3): 189–209.
  12. ^ Mass et al. 2007, pp. 701–715.
  13. ^ Reidenberg, Joy S. (2007). "Anatomical adaptations of aquatic mammals". The Anatomical Record. 290 (6): 507–513. doi:10.1002/ar.20541.
  14. ^ Bjerager, P.; Heegaard, S.; Tougaar, J. (2003). "Anatomy of the eye of the sperm whale (Physeter macrocephalus L.)". Aquatic Mammals. 29 (1): 31–36. doi:10.1578/016754203101024059.
  15. ^ Tinker 1988, The Respiratory System, pp.65–68.
  16. ^ Cavendish, Marshall (2010). "Gray whale". Mammal Anatomy: An Illustrated Guide. ISBN 978-0-7614-7882-9.

Further reading

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