Ungulates and Subungulates. Ungulata Subungulates –Proboscidea elephants –Hyracoidea hyraxes...
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Transcript of Ungulates and Subungulates. Ungulata Subungulates –Proboscidea elephants –Hyracoidea hyraxes...
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Ungulates and Subungulates
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Ungulata
• Subungulates– Proboscidea
• elephants
– Hyracoidea• hyraxes
– Sirenia• dugongs and manatees
• Ungulates– Perrisodactyla
• odd toed ungulates
• horses, tapirs, and rhinos
– Artiodactyla• even toed ungulates
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Subungulates
• 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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Subungulates
• 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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Subungualta• 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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Subungulata
• Proboscideans and Sirenians were much more diverse during the Oligocene and Miocene.
• Their future does not look promising.
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Proboscidea
• 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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Proboscidea
• 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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Proboscideans
• 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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Proboscideans
• 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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Proboscideans
• 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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Proboscideans
• 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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Proboscideans
• 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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Proboscideans
• 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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Proboscideans
• 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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Proboscideans
• 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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Proboscideans
• 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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Proboscideans
• 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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Hyracoidea
• 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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Hyraxes
• 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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Hyraxes
• 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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Hyraxes
• 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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Sirenians
• 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
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Manatees
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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) Deer
F) CamelG) Pronghorn
Calcaneum is shaded and
articulates w/ Astragalus (H)
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Perissodactyla• 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 Pleistocene
Note the fore-limb
dominance.
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Tapirs
• 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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Tapirs
• 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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Tapirs
• Compare the teeth of Tapirus with those of the rhino
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Black Rhino
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Tapirs
• 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• 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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Hyracodontids
• 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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Hyracodontids
• 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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Hyracodontids
• 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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Amynodontids
• 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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Rhinocerotidae
• 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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Rhinocerotidae
• 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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Rhinocerotidae
• 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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Chalicotheres
• 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 Pleistocene
Note 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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Horses
• 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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Horses
• 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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Horses.
• 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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Horses.
• 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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Horses.
• 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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Horses
• 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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Horses
• 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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Artiodactyla
• 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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Artiodactyls
• 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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Artiodactyls
• 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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Artiodactyls
• 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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Pig
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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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Peccaries
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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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Hippos
• 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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Hippos
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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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Dromedary
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Dromedary
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Dromedary
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Llama
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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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Hydropotes
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Hydropotes
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Hydropotes
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Cervidae
• 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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Caribou
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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
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Pronghorn nasals
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Pronghorn
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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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Caribou
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Eland
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Greater Kudu
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Nyala