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Ungulates and Subungulates. Subungulates Proboscidea elephants Hyracoidea hyraxes Sirenia dugongs and manatees. Ungulates Perrisodactyla odd toed ungulates horses, tapirs, and rhinos Artiodactyla even toed ungulates. Ungulata. Subungulates.

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Ungulates and Subungulates

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Ungulates and subungulates l.jpg

Ungulates and Subungulates

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dugongs and manatees



odd toed ungulates

horses, tapirs, and rhinos


even toed ungulates


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  • This seems to be an odd grouping of organisms. However, it is not by accident, and does not represent another ‘garbage’ group.

  • Proboscideans, Hyraxes, and Sirenians are all derived from Condylarthrans, that evolved in the Paleocene about 65mya.

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  • The Paenungulata was one group within the Condylarthra, and by the Eocene of Africa, they gave rise to the Proboscidea, Sirenia, and Hyracoidea.

  • If that is the case, you would expect some morphological similarities between the groups.

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  • They all lack clavicles.

  • They all have short nails on their digits.

  • Females have 2 pectoral mammae (Hyraxes have 2 inguinal pairs as well).

  • All females have a bicornuate uterus.

  • All males have abdominal testes and have no baculum.

  • All are non-ruminating, hind gut fermenting, herbivores.

  • All have a cecum.

  • Elephants and Sirenia have horizontal molariform tooth replacement.

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  • Proboscideans and Sirenians were much more diverse during the Oligocene and Miocene.

  • Their future does not look promising.

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  • There is but 1 family (Elephantidae) and 2 species: African elephant - Loxodonta africana, and Asian elephant - Elaphas maximus.

  • African elephants are much larger than Indian (Asian) elephants. The teeth differ, Africans have higher shoulders, larger ears, and a more complex trunk.

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African vs. Asian Elephant

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  • Reproduction is not easy

    • Females are sexually mature by 9 to 12 years, with peak reproductive value between 25 and 45 years. Gestation is 22 months, but estrus lasts only 2 to 4 days, with about 4 years between estrus events.

    • Copulation is no simple deal either.

    • There is sexual dimorphism, and young small males generally do not reproduce.

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  • Size of the males tusks seems to be an important character for reproduction. African elephant females look for a minimum tusk length, and will not mate with ‘short’ males even if no ‘long’ males are available. This has some implications for the ivory industry.

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  • African elephant males weigh up to 7500kg, while Indian elephants weigh about 4500kg.

  • They exhibit indeterminant growth.

  • They have graviportal limbs, and are capable of one gait only.

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  • Feldhammer claims that large size in elephants is a consequence of ‘competition’ with other herbivores.

    • Is this the most parsimonious explanation?

    • Does it reduce the importance of predation?

    • What about the cost of transport?

  • What does large size mean for an endotherm?

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  • Elephants are inefficient herbivores, and require large home ranges. They are usually found in groups. Thus, as they move long distances each day, they are capable of significant habitat modification.

  • Consider what it means to be so large. How is it possible that 50% of what passes through the gut of an elephant is undigested?

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  • The trunk of elephants is actually part of the upper lip and the nostrils.

  • It is prehensile, and is essential since the animal can not reach the ground with its mouth.

  • It is used to manipulate food, suck up water (and then spray water into the mouth), and suck up dust and mud as well.

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  • Dental formula is 1/0, 0/0, 3/3, 3/3 = 28.

  • Tusks are dentine (with only the tip covered in enamel).

  • Tooth replacement is horizontal, they are worn and replaced from the rear. Note: although they have 6 molariform teeth in each jaw, only one is functional at any time.

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  • Elephants were once much more diverse than they are today.

  • In the Pleistocene they were in Europe and North America. In fact, until just recently, there were 2 species in N. America at the same time, mastadons (Mammut americanus), and Mammoths.

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  • Oldest fossils are from the Eocene of Africa

  • We have fossil evidence from Asia, Europe, Africa, and N. America.

  • Moeritheriids were relatively small (1m) in Africa during the Eocene and Oligocene, while Deinotheriids were in Asia and Europe from the Miocene to the Pliocene.

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  • The Deinotheriids had weird tusks, based on the lower incisors rather than upper.

  • Gomphotheriidae were contemporaries, and had tusks in upper and lower jaws.

  • Mammutidae were the mastodons from the early Miocene.

  • Stegodontidae were from the mid-Miocene.

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A) Moeritherium, B) Deinotherium, C) Gomphotherium, D) Wooly

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  • Only the Elephantidae persist today.

  • The genus Primelephas from the late Miocene/early Pliocene is probably ancestral to modern elephants as well as the Wooly Mammoths.

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  • There are 5 species of rock hyraxes, and 3 species of bush hyraxes, all inhabiting rocky habitats in Africa and the middle east.

  • Were first thought to be rodents, but are clearly subungulates.

  • They are not ruminants, but have a large cecum as well as a smaller paired cecum.

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  • Have a mid-dorsal gland surrounded by light hair.

  • They have unique pads on the feet, which function as suction cups on rocky surfaces. Glands on the feet provide moisture for ‘suction’

  • Toes have hoof-like nails (except 2nd on rear, which has a grooming claw).

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  • They have no canines, and have a diastemma, hence the early confusion with rodents.

  • Upper incisors are pointed and triangular with no enamel on posterior.

  • Unlike elephants and sirenians, dentition is not replaced horizontally.

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Modern Hyrax vs. Megalohyrax from the Oligocene. Note the diastemma in the modern form.

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Rock Hyrax: Procavia capensis

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Rock Hyrax: Procavia capensis

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Rock Hyrax: Procavia capensis

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  • Fossils are known from the Eocene of Europe and Africa.

  • There is always the speculations that the diversity of Hyraxes suffered as a consequence of competition with ungulates. More about this later.

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  • These are the dugongs and manatees.

  • 2 families: monotypic Dugongidae from western Pacific, and Trichechidae (3 species) form the Atlantic.

  • Essentially tropical, feeding on aquatic vegetation.

  • Poor thermoregulatory abilities and low metabolic rates - hence warm waters.

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Sirenian Morphology

  • Large fusiform bodies - valvular nostrils, no pinnae, horizontal tail, no external hind-limbs, and flipper-like fore-limbs.

  • Dense bone to facilitate negative bouyancy.

  • Lungs run nearly length of body to even out bouyant forces.

  • Teeth replaced horizontally.

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Stellar’s Sea Cow (Extinct), Manatee, and Dugong

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Note position of lungs in the Manatee.

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Dugongs vs. Manatees

  • Dugongs eat aquatic vegetation which is much softer than that consumed by manatees.

  • Feldhammer uses competition to explain distribution of species.

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Dugong vs Manatee

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Sirenian Fossil History

  • There were once at least 20 genera of Sirenians.

  • There are Eocene sirenians from india, Europe, and N. America (Protosiren).

  • Eocene sirenians are unique in that thay have a fifth premolar.

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Dusisiren: Miocene sea cow.

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Dugong vs Manatee: Deflected rostrum in Dugong is ‘adaptation’ to bottom feeding.

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Perissodactyla and Artiodactyla

  • Both forms of modern ungulates are digitigrade.

  • Teeth are usually hypsodont.

  • Limbs operate in a single plane, and are designed for cursorial locomotion.

  • Calcaneum usually does not articulate with the fibula.

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A) TapirB) RhinoC) HorseD) PigE) DeerF) CamelG) PronghornCalcaneum is shaded and articulates w/ Astragalus (H)

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  • Horses, Tapirs, and Rhinos.

  • Odd toed ungulates, with the 3rd digit bearing most of the weight (Mesaxonic).

  • Teeth are usually hypsodont and lophodont.

  • Horses and tapirs have upper incisors, rhinos generally do not.

  • Stomach is simple, but they have a cecum. Gut retention times are half that of ruminating artiodactyls. Thus, only about 70% as efficient.

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Malayan tapir

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Indian Rhino

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Perissodactyla: Fossil History

  • The Condylarthra are ancestral to the Perissodactyla, as well as the Artiodactyla, Proboscidea, Sirenia, and Cetaceans.

  • It is not necessarily true that the Perissodactyla and Artiodactyla are monophyletic.

  • Based on 67 hard and soft morphological characters, we can propose the following:

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Ungulate Evolution

  • Note the implications:

    • Closest relatives of the perissodactyls are the cetaceans.

    • Mammals invaded wate completely at least twice independently (Cetceans and Sirens).

    • Note the close relationship between hyraxes, elephants, and sirenians.

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Perissodactyl Evoltuion

  • Originally 14 families at their peak in the Eocene.

  • By the end of the Oligocene there were only 4 families.

  • They were the dominant ‘medium to large’ herbivores of the Tertiary.

  • Both the Brontotheres and Chalicotheres went extinct.

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Chalico-there: went extinct in the PleistoceneNote the fore-limb dominance.

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  • Origin and early differentiation in the Paleocene

  • Heptodon is one of the earliest, and comes from the Eocene of Wyoming.

  • Modern Tapirus is remarkably similar to Heptodon, but bigger. Both have 4 toes in front and 3 in rear, both have ulna and fibula complete and unfused.

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  • Both Heptodon and Taprius have complete dentition w/ a small diastemma; upper canine is reduced and lateral incisor is caniniform.

  • Upper molars have 3 lophs, the lower molear have 2 transverse lophs (as in Rhinos)

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Malayan tapir

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Tapir teeth - sort of

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  • Compare the teeth of Tapirus with those of the rhino

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Black Rhino

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  • This similarity in structure is one of the reasons why tapirs and rhinos are generally considered to share a common ancestor.

  • The cladogram for Perissodactyla lumps tapirs and rhinos, with horses as the outlying sister group.

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  • Tapirs persist in S. America and Central America, and in Southeast Asia. Can you explain this distribution?

  • Hyrachyus (Family Hyrachyidae) may be transitional between tapirs and rhinos. It was abundant in the eocene of n. America and Europe.

  • Hyracodontids and Amynodontids were abundant in the Eocene and oligocene of N. America and Asia.

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  • About 12 genera

  • Moderate size

  • Slender limbs like horses w/ light builds.

  • Cursorial

  • incisors were spatulate (primitive) and equal sized.

  • Canines were moderate size.

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  • In the Oligocene, from Hyracodont lineage, came a series of gigantic hornless rhinos in the subfamily Indricotherinae

  • Indricotherinae ranged from central Asia to China.

  • Indricotherium was the largest land mammal to ever live.

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  • Indricotherium was 5.4m tall at the shoulders, had a long neck and a skull which was 1.3m long.

  • Could reach vegetation 8m above the ground.

  • Had a probable weight of 30 tons, 4.5 times greater than Loxodonta, and about twice as great as the largest Mammoth.

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  • About 10 genera

  • Large heavy bodies

  • Short stocky limbs

  • Short faces

  • Prominent canine tusks.

  • The bulk of the Amynodontid radiation was over by the close of the Oligocene.

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  • Hyracodontids during the Oligocene obtained a unique dental variation: chisel like I1 and tusk like I2. This formed the basis of a 2nd radiation… the Rhinocerotidae.

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  • About 50 genera

  • N. America, Eurasia, and Africa from the Miocene to Pleistocene.

  • Rhinocerotids included wooly rhinos and rhinos w/ horns (Elasmotherium) as long as 2m.

  • Wooly rhinos show up as cave paintings by palaeolithic man.

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  • Elasmotherium’s horn was not nasal like most, but originated on the forehead. It had no incisors.

  • Today, rhinos occur only in India, Java, Sumatra, and Africa.

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Black Rhino

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Black Rhino

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Black Rhino

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Black Rhino

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  • From the Eocene on in N. America, Eurasia, and Africa.

  • Simple premolars and bunolophodont molars.

  • Probably a bipedal browser.

  • Had long forearms and hooked claws - very un-ungulate like.

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Chalico-there: went extinct in the PleistoceneNote the fore-limb dominance.

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Titanotheres (=Brontotheres)

  • From the early Eocene to early Oligocene of N. America and eastern Asia.

  • Medium to very large size.

  • Probably succeeded in Asia by Indricothere Rhinos.

  • Had graviportal limbs and nasal horns which were probably covered by skin.

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  • Evolution of horses has been used as best example of gradualism.

  • Over 55 million years, the progression from Eohippus to Equus has involved:

    • Increase in size from small lamb size to present size.

    • Reduction of toes from 3 to 1.

    • Increased complexity of enamel pattern on molars.

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  • Eohippus (= Hyracotherium)

  • Eocene of N. America, W. Europe, and E. Asia.

  • 4 toes fromt, 3 rear.

  • Horses died out (Together w/ horse-like Tapirs) in W. Eeurope by the Oligocene. Also died out in Asia by this time.

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  • In Oligocene, N. America horses are Mesohippus and Miohippus.

  • Sheep size, 3 toes w/ middle digit largest.

  • Snout elongating.

  • Premolars beginning to look like molars w/ lophs and lophids.

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  • By Miocene, Anchitherium had split off from other N. American horses, and migrated through Europe and Asia.

  • By the end of the Miocene, forest-dwelling Hypohippus migrated into China.

  • From Oligocene Anchitheres came the Miocene Parahippus, a precursor to mid-miocene Merychippus.

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  • Merychippus is first grazer horse.

  • True hypsodont cheek teeth, elaborately lophed and had cementum.

  • Had fused ulna/radius and tibia/fibula to improve gallop and minimize twisting of legs.

  • All later horses evolved from Merychippus.

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  • First successful descendent of Merychippus were the Hipparionines, which included as many as 6 lineages. They invaded the old world several times and were finally extinct by the late Pleistocene.

  • In the late Miocene, Merychippus was replaced by Pliohippus, the 1st one-toed horse.

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  • Pliohippus gave rise to Equus during the Pleistocene of N. America, from where it radiated to the old world.

  • Equus became extinct in the N. American recent. Why?

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  • 1/3 of all mammalian genera are Herbivores. Of these, 50% are Artiodactyla or Perissodactyla.

  • Origin is probably I the Palaeocene.

  • Today, there are 6 genera of Perissodactyls vs. about 80 genera of Artiodactyls.

  • Whereas perissodactyls were once most diverse, artiodactyls now have significant edge. Why?

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  • Currently there are 12 famillies of herbivores, there are 24 extinct families.

  • Origin is probably in northern continents with movements into southern ones (except Australia).

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Horns, Antlers, Ossicones.

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Diacodexis: early Eocene artiodactyl.

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  • Primary axis of support is between 3rd and 4th toes (paraxonic).

  • 2nd and 5th digits are absent or non-functional.

  • Pigs (Suiformes) are plantigrade, while ruminants are digitigrade (Unguligrade).

  • Dentition varies from bunodont and brachydont to solenodont and hypsodont.

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  • Upper incisors and canines are reduced or absent.

  • Suids and Tayasuids have non-ruminating stomachs while more derived families have 4 chambered ruminating stomachs.

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Suiformes: Suidae

  • 5 genera and 16 species.

  • Simple stomachs and bunodont teeth, large ever-growing canines.

  • Cartilaginous disk on snout.

  • Endemic to Europe, Africa, and Asia. Introduced almost everywhere else.

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Warthog, Babirusa, and Wild Boar.

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Suiformes: Tayassuidae

  • Least specialized of the suiformes.

  • Peccaries - legs are thin and feet end in hooves. Upper canines point downward rather than upward as in pigs.

  • Restricted to the New World, from the desert southwest to Argentina.

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Suiformes: Hippopotamidae

  • 2 species only.

  • Little or no hair, also lack sweat glands for thermoregulation.

  • They do have glandular skin that produces pigmented secretions to protect against sunlight.

  • Bunodont cheek-teeth, ever-growing tusk-like lower canines and incisors, with alveoli for canines anterior to those for incisors.

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  • Not ruminants, but septa in stomach increase gut retention times.

  • H. amphibius grazes on land at night fo rup to 6 hrs.

  • Hexaprotodon liberiensis is less aquatic.

  • Both are African.

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Hippo: note elevated eyes and nares.

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Pigmy hippo

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Tylopoda: Camelidae

  • North American origin in Eocene, extinct here by the Pleistocene.

  • 3 genera and 6 species

  • Dromedaries, Bactracians, Quanaco, Llama, Alpaca, and Vicugna.

  • Small head, long snout, cleft upper lip, long thin neck, long legs w/ canon bone.

  • Upper and lower canines, and selenodont cheek teeth.

  • Toes spread out under load.

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Tylopoda: Camelidae

  • Outer spatulate upper incisor is retained in adults.

  • 3-chambered stomachs and a cecum.

  • Dromedary was once throughout the Middle East, but now exists only in domestication.

  • Bactracians were once throughout Asia, but are now restricted to the Gobi.

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Tylopoda: Camelidae

  • Vicunas and Llamas are restricted to S. America.

  • Camelids consume plants w/ high salt content, foods avoided by other grazers.

  • Unique gaits in Camels.

  • Heat and water strategies - the hump is not what you think.

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Lama glama

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Lama glama

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Lama glama

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Lama glama

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Ruminantia: Tragulidae

  • 3 genera and 4 species of Chevrotains in Africa and Asia.

  • Most underived of all ruminants, once had a worldwide distribution.

  • Mouse deer is smallest artiodactyl at 2.5kg.

  • No antlers, but curved upper canines.

  • 3-chambered ruminating stomach.

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Tragulus napu

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Ruminantia: Giraffidae

  • 2 genera and 2 species: Giraffa camelopardalis and Okapia johnstoni.

  • Small brachydont teeth, prehensile tongues, ossicones.

  • Consider circulatory problems of great height.

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Giraffa camelopardalis incisors

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Ruminantia: Moschidae

  • 4 species of musk deer.

  • Lack antlers, but have curved canines.

  • Distributed from Siberia to the Himalayas.

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Musk deer

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Musk deer

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Hydropotes inermis: water deer F

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Hydropotes inermis: water deer M

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  • 16 genera and 42 extant species, ranging in size from the pudu at 8kg to Alces alces at 800kg.

  • Absent only from sub-Saharan Africa and Antarctica, were introduced to Australia and New Zealand.

  • Sexually dimorphic - males have antlers, females (except caribou) do not. Why?

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Moose cow and calf

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Moose bull

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Mule deer

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White tail

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White tail nasals

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Ruminantia: Antilocapridae

  • 1 genus, 1 species.

  • Restricted to N. America and Mexico.

  • Unique horns.

  • Forage on Artemisia tridenta.

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Pronghorn nasals

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Ruminantia: Bovidae

  • 45 genera and 137 species.

  • 4 chambered ruminating stomachs.

  • All have 2 horns except the the four-horned antelope.

  • Worldwide distributin except S. America and Australia. Why?

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African buffalo

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Bush Buck

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Greater Kudu

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