SYSTEMATIC BIOLOGY

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CLASS MAMMALIA

Mammalia (mam-A-li-a) is derived from a single Latin root that means breast (mamma).  The name refers to the occurrence of milk glands, a synapomorphic character of this group.

The mammals are synapsids and a sister group to the "mammal-like reptiles", the Eosynapsida.  In general, the mammals have a suite of synapomorphic characters that define them as a monophyletic group.  They have hair, and a single lower tooth-bearing jaw bone (the dentary) that articulates with the squamosal. The teeth are replaced one time and show a differentiation between incisors, canines, and cheek teeth (premolars and molars).  The mammary glands are sweat glands that have been modified to deliver milk, a protein and lipid-rich substance, to newborn mammals before they can ingest food on their own. Though not unique to mammals, they are endothermic and capable of high levels of activity.  Mammals have diversified to occupy almost all terrestrial and aquatic environments.  

The approximately 4,700 extant mammals can be divided into three very unequal groups: the monotremes, the marsupials, and the placentals.  The monotremes are highly modified burrowing animals that are restricted to the continent of Australia and represented by only three extant species: the platypus, echidna, and spiny anteater.  All are toothless; furthermore, the platypus has an unusual horny beak (see Figure A).  They retain the primitive character of laying eggs rather than gestating the young within their bodies.

Marsupials are characterized by having a marsupium, a protective pouch in which the almost fetus-like young resides and feeds attached to the nipple from the time it is born until it can move about on its own.  Their teeth have a dental formula similar to other mammals, but they replace only the premolars and molars.  The southern continental distribution of marsupials is a consequence of their having evolved on Gondwanaland during the Cretaceous period.  The South American marsupials are called the Ameridelphids and are represented primarily by opossums (see Figure B), mainly omnivorous animals that have prehensile tails.  The exchange of animals between North and South America after the completion of the Central American land bridge, led to the mass extinction of many diverse South American marsupials during the past 5 million years.  Only the Virginia possum (Didelphis virginiana) of all the marsupials has made its way into North America where it has become very successful.

Australia also broke away from Gondwanaland, but it never became connected to a major land mass.  Thus, its marsupial fauna remained diverse and rich until the introduction of placental mammals by humans (who also are placental mammals).  Perhaps, the most renowned members of the australidelphid marsupials are the kangaroos and wallabies (Figure C).  Large kangaroos are the ecological equivalents of gregarious herbivores like the ungulates of North America (e.g. Bison) and Africa (e.g. antelope).  Aside from terrestrial mammals, the Australian marsupials include arboreal and fossorial species.  They also range from carnivores to omnivores as well as herbivores.  They also include species that specialize on eating nectar, insects, and eucalyptus leaves.

Placental, also called the eutherian, mammals comprise more than 90% of all extant mammals.  They are characterized by the occurrence of a placenta, a connection between the uterine wall and the developing young.  Placental mammals, because they allow the gestation of a fairly well-developed offspring before birth, also have a relatively large birth canal.  Though they appeared much earlier in the Cretaceous period in which much of their diversification occurred, the successful exploitation of almost all continents and oceans by the placental mammals has occurred over the last 65 million years.  Much of the organization of eutherians seems to be associated with plate tectonics and is reflected in the names of the four distinct clades: the Afrotheria, Xenarthra, Laurasiatheria, and Euarchontoglires.

Six orders form a monophyletic group of very different animals that evolved in Africa and, therefore, are called the Afrotheria.  The largest members of the group are the elephants (see Figure D) and dugongs.  Most are small to medium animals: elephant shrews, aardvarks, tenrecs, and hyrax.  Although molecular evidence suggests a strong, ancient relationship between these taxa, structural synapomorphies characteristic of the Afrotheria have not been identified.

Xenarthra is a monophyletic group that has been recognized for some time (formerly the order Edentata).  The extant taxa include armadillos and sloths (see Figure E), which, as the old ordinal name implies, no teeth.  However, this is not characteristic of the group.  The extinct giant sloth had more than 100 teeth, a very large number for a mammal.  Structural synapomorphies include extra joints between some of the trunk and tail vertebrae.  Also, the ischium and the ilium are fused to the first caudal vertebra.

Animals of the Laurasiatheria evolved on the northern supercontinent of Laurasia after the breakup of Pangea.  The diversity in form, habitat, and size in this group is astounding and contains the largest and among the smallest of all mammals.  The Laurasiatheria includes 7 or 8 orders depending on how the cetaceans and artiodactyls are lumped together into a group called the Cetartiodactyla.  This group (here treated as a superorder) is an unlikely collection of animals like pigs, cows, camels together with whales and dolphins.  As remarkable as the collection seems, the relationship has been confirmed by fossil, molecular, and anatomical evidence (Thewissen and Williams 2002).  Furthermore, the emerging story of whale evolution has become one of the most exciting demonstrations of macroevolution to have emerged in vertebrate paleohistory.

Other orders of the Laurasiatheria include Chiroptera (bats), Perissodactyla (horses, rhinoceroses, and tapirs), Carnivora (cats, dogs, mongooses, bears, weasels, seals, walruses, etc.), Eulipotyphla (contains moles and shrews, animals formerly included in a group called the Insectivora), and the Pholidota (scaly anteaters).  Though these animals appeared in Laurasia, they entered and became important members of the landscape in all continents and all oceans.  Perhaps, most notable are the Chiroptera, one of the few groups of vertebrates to achieve true flight.

The Euarchonotoglires, sometimes called the Supraprimates, are sisters to the Laurasiatheria and include the rodents, hares, primates, tree shrews and flying lemurs.  The rodents and hares are very speciose and comprise more than 40% of all living mammals.  The primates include lemurs, monkeys (see Figure H), and apes.  Our species, Homo sapiens, is a primate and one of the most successful species on earth occurring in almost all biomes and transforming the globe.  The Euarchonotoglires and Laurasiatheria together are in a clade called the Boreotheria (animals of the northern continents) and contain the crown taxa of the mammals.  

HIERARCHICAL TAXONOMY OF THE CLASS MAMMALIA. The following descriptions come from Benton (2005), Kemp (2005), and the Animal Diversity Web (from the University of Michigan Museum of Zoology).  The structure of the following system is also based, in part, on the phylogenetic treatment by Mikko's Phylogeny Web (from the Finnish Museum of Natural History) and Kielan-Jaworowska et al. (2004).     Taxa in red are extinct.

FIGURE A. Duck-billed Platypus, a monotreme from Australia.

FIGURE B. A Virginia possum, a member of the Ameridelphia branch of marsupials.

FIGURE C. A kangaroo, a large bipedal member of the Australidelphia branch of marsupials.

FIGURE D. An African elephant, an Afrotheria placental mammal.

FIGURE E. A sloth, a Xenarthra placental mammal.

FIGURE F. A horse (Equus), a Laurasiathera placental mammal.

FIGURE G. A blue whale, a member of the Laurasiatheria, is the largest animal that has ever lived.

FIGURE H. A spider monkey in the rainforest of Costa Rica.  It is a member of the Euchontoglires.

Image of Platypus from: http://www.epa.qld.gov.au/images/nature_conservation/platypus.jpg
Image of Virginia Possum (Didelphis) from: http://condor.depaul.edu/~gandrus/jpg/pics/possum.jpg 
Image of the African Elephant from: http://www.bioinquiry.vt.edu/SouthAfrica/species.html 
Image of the horse came from the Systematic Biology collection.
Image of the sloth from: http://arnica.csustan.edu/boty1050/Evolution/sloth_extant.jpg 
Image of the Blue Whale from: http://www.yoto98.noaa.gov/books/whales/whale3.htm 

 

CLASS MAMMALIA

In general, the mammals have a suite of synapomorphic characters that define them as a monophyletic group.  They have hair, and a single lower tooth-bearing jaw bone (the dentary) that articulates with the squamosal. The teeth are replaced one time and show a differentiation between incisors, canines, and cheek teeth (premolars and molars).  They are endothermic and capable of high levels of activity.  Mammals have diversified to occupy almost all terrestrial and aquatic environments.  One group, the bats, has achieved true flight.  

Living mammals can be divided into the following major groups:

MONOTREMATA

MARSUPIALIA

AMERIDELPHIA The didelphids and caenolestoids (mainly in South America).

AUSTRALIDELPHIA The marsupials that occur primarily on Australia.

PLACENTALIA

AFROTHERIA  includes Macroscelidea, Tubulidentata, Proboscidea, Hyracoidea, Syrenia, Tenreca, Chrysochlorida.

XENARTHRA includes Xenarthra.

LAURASIATHERIA  includes Cetartiodactyla (Cetacea and Artiodactyla), Perissodactyla, Carnivora, Chiroptera, Eulipotyphla.

EUARCHONTOGLIRES  includes Rodentia, Lagomorpha, Primates, Dermoptera, Scandentia.

 

SUBCLASS UNNAMED+

ORDER UNNAMED

FAMILY UNNAMED

Known from fragments of the braincase.  Elements present indicate a mammalian organization of the skull, including the enclosure of the expanded brain by sheets of bone. 

Upper Triassic

Adelobasileus

FAMILY SINOCONODONTIDAE

It had the same skull features as Adelobasilius.  In addition, it had a fully-developed dentary-squamosal jaw joint.  The middle ear was approaching the mammalian state.  The cheek teeth did not occlude well and seem to have been replaced through the individual’s life.  Such indeterminate growth of the teeth and skull was a primitive feature similar to that of the cynodonts. 

Lower Jurassic

Sinoconodon

SUBCLASS MAMMALIAFORMES

INFRACLASS UNNAMED+

ORDER UNNAMED

It retained the extradentary bones in the lower jaw which served in hearing.  The cheek teeth were differentiated between premolars and molars.  It seemed to have developed the mammalian diphydont or single tooth replacement.  All cheek teeth occlude.  The lower jaw was narrower than the upper jaw, a condition common to most mammals.  Ribs only in the thoracic region.  The forelimbs were still somewhat sprawling.  The hind legs, however, were erect.  They seem to have had determinate growth (animals were of one size).  Also, they seem to have been nocturnal (inferred from the architecture of the brain case which emphasized parts of the brain responsible for hearing and smell) and enthothermic (inferred from the small size and adaptations to rapid respiration).  Also, likely they were insectivorous.  Thus, they were like shrews.

FAMILY MORGANUCODONTIDAE

Morganucodon, Megazostrodon

INFRACLASS HOLOTHERIA

SUPERORDER UNNAMED

ORDER UNNAMED+

Known only from teeth and jaw fragments, it had clear mammalian features of the cheek teeth, primarily 3 cusps- a large central cusp and two smaller lateral cusps.

Lower Jurassic

FAMILY KUEHNEOTHERIIDAE

Kuehneotherium

ORDER DOCODONTA+

Also known from fragmentary skeletal evidence, mainly jaws and teeth.  These had the tricusp molars as well as differentiated incisors and canines. 

Middle and upper Jurassic

Docodon

ORDER UNNAMED+

These animals had cheek teeth that interlock in a mortar and pestle grinding action called a tribosphenic-like molar (others have an anterior and lingual shelf of bone rather than a full cupule in the lower molar).

Middle Jurassic to lower Cretaceous

FAMILY AUSKTRIBOSPHENIDAE

Ausktribosphenos, Bishops, Asfaltomylos, Ambondro

ORDER MONOTREMATA

This order includes two extant animals, the platypus and echidna.  Both animals reproduce by laying eggs, and have an interclavicle bone, both primitive characters.  However, both extant monotreme groups are toothless as adults and difficult to compare to the early mammalian fossils.  Fortunately, the platypus has teeth that do not erupt and are absorbed before the formation of the horny beak.  Those teeth and the earlier ones show similarities with the ausktribosphenid teeth. 

Lower Cretaceous to the present: Steropodon+

FAMILY ORNITHORHYNCHIDAE

Duck-Billed Platypus.  These are aquatic mammals of streams and lakes of Australia .  Its feet are webbed, and it has fur that repels water.  The ears have no pinnae.  The most distinctive aspect of the animal is it horny beak.  Platypus adults have no teeth and feed on aquatic invertebrates by crushing and chewing them with their horny plates.  Males have venomous spurs on their hind legs.

Platypus

FAMILY TACHYGLOSSIDAE

Echidnas and Spiny Anteaters.  These are medium-sized mammals of Australia and New Guinea .  They have a compact body that is covered with spines and a head with arelatively long, thin snout.  They dig for ants and termites which they lap with a mucous-covered tongue.  They crush the insects against their palate.

Echidna, Zaglossus..

ORDER TRICONODONTA+

Mainly known from isolated tricuspid molars, but some skeletal remains have been found, the most spectacular of which was Jeholodens, a burrowing anomal.  It was and insectivore with plantigrade stance (feet flat on the ground).  Both fore and hind limbs were sprawling. 

Middle Jurassic to Upper Cretaceous

Triconodont, Jeholodens

ORDER MULTITUBERCULATA+

The largest group of Mesozoic mammals.  The multituberculates were omnivorous and rodent-like.  As in rodents, the incisors are long and separated from the cheek teeth by a gap.  There were no canine teeth.  Unlike rodents, the last premolar forms a large shearing surface.  They had two small epipubic bones (also seen in the extant monotremes and other extinct Cretaceous groups). At least one appeared to have been arboreal with a prehensile tail and hind feet that can swivel backwards like a squirrel. 

Upper Jurassic to upper Eocene.

Kamptobaatar, Nemegtbaatar, Ptilodus

ORDER SYMMETRODONTA+

The middle cusp of the tricusp molars was offset toward the tongue.  Some had a very large coronoid process and extended angular process at the back angle of the jaw. 

Lower Cretaceous

Spalacotherium, Zhangeotherium, Cusafontia

ORDER DELTATHEROIDA+

These animals stood at the inflexion point of the holotherians and the therians.  They had cheek teeth (form and replacement pattern; see below) that shared characters with the marsupials.  Also, the pattern of cranial vascularization was marsupial in form. 

Upper Cretaceous

Deltatheridium

INFRACLASS THERIA (a clade of marsupial + placental mammals)

SUPERORDER MARSUPIALIA

Marsupials replace only the premolars and molars.  The anterior teeth are not replaced.  Marsupials have 3 premolars and 4 molars.

AMERIDELPHIA

ORDER DIDELPHIMORPHIA

Opossums.  Marsupials of Central and South America with one found through much of the US .  They occur in aquatic to forest to dry scrub habitats and are typically omnivorous though some are carnivores.  They have a full set of teeth (5 upper incisors and 4 lower incisors) on each side followed by a canine, 3 premolars, and 4 molars.  Extant taxa are typically small to medium-sized animals.  The first toe on the hindfoot is opposable and has a nail (all other toes have claws).  Tail is long, naked, and prehensile.

Upper Cretaceous to the present

FAMILY DIDELPHIDAE

Alphadon+

Subfamily Caluromyinae (Wooly Opossums): Caluromys, Calurmysiops, Glironia.

Subfamily Didelphinae (American Opossums): Chironectes, Didelphis, Gracilinanus, Lestodelphys, Lutreolina, Marmosa, Marmosops, Metachirus, Micoureus, Monodelphis, Philander, Thylamys.

ORDER PAUCITUBERCULATA

Shrew Possums.

FAMILY CAENOLESTIDAE

Shrew Possums.  These animals seem to be insectivores of western South America on the floor of cold and wet forests in the Andes .  Shrew-like, they have small eyes and a tick coat.  The upper lip has a distinctive flap of skin.  Females do not have pouches.  The lower jaw has a large incisor on each side that projects forward.  The upper canines are large.  The tail is long, but not prehensile.  Paleothentes+ had elongate lower incisor and blade-like cheek teeth. 

Upper Oligocene to present

Caenolestes, Lestoros, Rhyncholestes, Paleothentes+

FAMILY ARGYROLAGIDAE+

Argyrolagus was similar in form to a kangaroo rat; adapted to hopping and large cheek teeth. 

Argyrolagus

FAMILY CAROLOAMEGHINIDAE+

Small insectivores.

Early Cenozoic

Roberthoffestetteria

ORDER SPARASSODONTA+

Carnivores

FAMILY BORHYAENIDAE

They were somewhat dog-like with short limbs.

Paleocene to Pliocene

Prothylacynus

FAMILY THYLACOSMILIDAE

These animals developed long saber-like upper canines that fit into a sheath formed by the lower jaw.  The canine had indeterminate growth. 

Upper Miocene to Pliocene

Thylacosmilus

AUSTRALIDELPHIA

ORDER MICROBIOTHERIA

Oligocene to present

FAMILY MICROBIOTHERIDAE

Monitos del Monte.  They live in humid vegetation of the mountainous regions of souther Argentina and Chile .  A small arboreal animal with a prehensile tail, which might serve to store fat; short, rounded ears.  The animal has a well-developed marsupium.  They superficially resemble the didelphids, with which they were classified.  Though its dental formula is the same as the didelphids, the incisors and cheek teeth are quite unique (incisors spatulate and cheek teeth rounded).

Dromiciops, Microbiotherium+, Khasia+

ORDER DASYUROMORPHIA

Marsupial mice and rats, cat-like animals, and the Tasmanian wolf

FAMILY DASYURIDAE

Dasyurids.  Arboreal and terrestrial; New Guinea and Australia .  These animals have a long body with a pointed snout.  They are plantigrade and are not syndactylous (no fused toes); the tail is not prehensile.

Antechinus, Dasycercus, Dasykaluta, Dasyurus, Murexia, Myoictis, Neophascogale, Ningaui, Parantechnicus, Phascogale, Phascolosorex, Planigale, Pseudantechnicus, Sarcophilus, Sminthopsis.

FAMILY MYRMECOBIIDAE

Numbats. Ant-eating marsupials of southern Australia .  These animals do not have a pouch; and have dentition but the teeth appear to be reduced.  They have a very long tongue.

Myrmecobius.

FAMILY THYLACINIDAE +

Tasmanian Wolf.  Once common from New Guinea to Tasmania, now likely extinct.  The marsupial has many dog-like features, including its posture and head.

Thylacinus.

ORDER PERAMELEMORPHIA

Bandicoots and bilbis.

Early Miocene to present

FAMILY PERAMELIDAE

Bandicoots and Bilbies. Ground-dwelling animals of drier habitas of Australia .  They have flattened heads and long hind legs adapted for hopping and running (some adaptations also suggest digging – e.g. enlarged fibula).

Chaeropus, Isodon, Macrotis, Perameles.

FAMILY PERORYCTIDAE

Spiny Bandicoots and Mouse Bandicoots.  Humid tropical forests of New Guinea and Australia .  They have cylindrical heads, but otherwise resemble the peramelids.

Echymipera, Microperoryctes, Peroryctes, Rhynchomeles.

ORDER NOTORYCTEMORPHIA

Marsupial moles

FAMILY NOTORYCTIDAE

Marsupial Moles.  Fossorial animals of the deserts of Australia .  They are mole-like with reduced eyes, hevy digging fore feet, no external ears, and snouts covered by a shield.  They swim through the sandy soil with their powerful forelegs.  They feed on grubs.  Females have a marsupium.

Notoryctes.

ORDER DIPROTODONTIA

Very diverse order including: Australian possums, gliding phalangers (including the marsupial lion), wallabies, kangaroos, koalas, wombats.  Australia and New Guinea .

Lower Miocene to present

FAMILY ACROBATIDAE

Feathertail Gliders.  Small arboreal animals of Australia and New Guinea .  Although only one genus has a gliding membrane (it seems to have been lost in Distoechurus), both genera are united in that they feed on nectar, have stiff hairs on either side of the tail. 

Acrobates, Distoechurus.

FAMILY BURRAMYIDAE

Pygmy Possums.  Small arboreal animals of New Guinea and Australia .  They have a mouse-like head with a short muzzle.  The tail is long and prehensile, and they have an opposable halux on the hind feet..  Their pouches open anteriorly.  They eat insects, nectat, and sometimes small reptiles.

Burramys, Cercartetus.

FAMILY PETAURIDAE

Gliders, Leadbeater’s Possum, and Striped Possum.  Arboreal animals of New Guinea and Australia .  All have a distinctive stripe that runs from the head to the base of the tail.  The tail is prehensile and bushy.  Many have a gliding membrane.  They have opposable digits on the front feet and an opposable halux on the hind feet.  They have protruding incisors.  Their pouches open anteriorly. 

Dactylopsila, Gymnobelidius, Petaurus.

FAMILY POTOROIDAE

Bettongs, Potoroos, and Rat Kangaroos.  They are terrestrial mammals of open areas of Australia.  They have large hind legs and feet and hop like rabbits, but can come up to a bipedal hop at high speeds.  They have a tail that is somewhat prehensile.  They are omnivores and seem to have specialized on underground fungi, tubers, and insects.  Like the macropodids, their premolars are replaced by a single premolar with a shearing edge.

Aepyprymnus, Bettongia, Caloprymnus, Hypsiprymnodon, Potorous.

FAMILY MACROPODIDAE

Kangaroos and Wallabies. Animals are terrestrial or arboreal and tend to be herbivores (grazers and browsers) which occur over most of Australia and into New Guinea and surrounding islands.  They are bipedal and plantigrade.  The hind legs are powerful and allow for a hopping motion that can propel them at great speeds (to 50km/hr).  The large tail serves as a balancing organ.  Their lower incisors stick forward and close against a pad on the upper jaw (as in the bovids).  Premolars move forward during replacement and become shearing blades.The pouch opens anteriorly.

Dendrolagus, Dorcopsis, Dorcopsulus, Lagochestes, Lagotrophus, Macropus, Onychogalea, Petrogale, Setonix, Thylogale, Wallabia.

FAMILY PSEUDOCHEIRIDAE

Ringtailed Possums.  Small to medium-sized arboreal animals of New Guinea and Australia .  Some are gliders.  Most have prehensile tails.  They have opposable digits on the front feet and an opposable halux on the hind feet.  Mostly herbivores.

Hemibelideus, Petaluroides, Petropseudes, Pseudocheirus, Pseudochirops.

FAMILY TARSIPEDIDAE

Honey Possum.  They are small arboreal animals of southwestern Australia .  The have a long muzzle and a prehensile tail.  The syndactylous toes on the hind feet have nails.  Teeth are small and reduced.  The cheekteeth are peg-like.  They have bristle-tipped tongues with which they gather nectar.

Tarsipes.

FAMILY PHALANGERIDAE

Brushtail Possums, Cuscuses, and Phalangers.  They are semi-arboreal animals of Australia , New Guinea and some surrounding islands.  Although good climbers, they spend much  time on the ground.  They have long tails that generally are prehensile.  Digits 1 and 2 are opposable to the other three.  They are omnibvores that are nocturnal or crepuscular.  Pouch opens anteriorly.

Ailurops, Phalanger, Spilocuscus, Strigocuscus, Trichosurus, Wyulda.

FAMILY VOMBATIDAE (PHASCOLOMYIDAE)

Wombats. Stocky terrestrial animals of Australia .  They are badger-like, thick-bodied and tailess with short legs and strong digging claws.  Incisors resemble those of rodents.  Herbivores.  The pouch opens posteriorly.

Lasiorhinus, Vombatus.

FAMILY PHASCOLARCTIDAE

Koalas. Small (~60cm high) teddy-bear arboreal animals of Eastern Australia .  Digit 1 opposable.  Toes 2 and 3 united on hind feet.  Pouch opens backwards. They are unusual in that they form a placenta for a brief time.  They are herbivores that specialize on eucalyptus.

Phascolarctos.

SUPERORDER PLACENTALIA (EUTHERIA)

AFROTHERIA

ORDER TUBULIDENTATA

Aardvarks. These are terrestrial animals of Africa .  They are large (pig-sized) with a tubular snout and a reduced dentition.  They are adapted to eating termites with a long tongue, thick skin, and strong digging claws.

Miocene to present

FAMILY ORYOCTEROPIDAE

Orycteropus.

ORDER AFROSORISCIDA

FAMILY TENRECIDAE

Tenrecs.  These are insectivores of Madagascar and western Africa .  They range in size from very small shrew-like animals to cat-sized animals.  Some are spiny and resemble hedgehogs. They occur in habitats that range from aquatic to terrestrial, to fossorial, to arboreal.  They have no jugals and have incomplete zygomatic arches. 

Miocene to present.

Subfamily Geogalinae (Large-Eared Tenrecs): Geogale.

Subfamily Oryzorictinae (Aquatic Tenrecs, Rice Tenrecs, and Shrew Tenrecs): Limnogale, Microgale, Oryzoryctes.

Subfamily Tenrecinae (Tenrecs): Echinops, Hemicentetes, Setifer, Tenrecs.

Centetes, , Ericulus, , ,.

FAMILY POTAMOGALIDAE

Otter Shrews. Aquatic animals of western and central Africa .  They superficially resemble small otters with smooth fur.  They have long muzzels; reduced eyes and ears, and laterally-flattened tails.  These are sometimes placed within the Tenrecidae as a subfamily.

Micropotamogale, Potamogale.

FAMILY CHRYSOCHLORIDAE

Golden moles.  These are fossorial insectivores of south and east-central Africa   They are adapted to a burrowing lifestyle in that they have large claws on the forefeet, a leathery pad over the nose, and small ears.  They have a tabular bone and a single cloaca. 

Miocene to present

Ambltsomus, Calcochloris, Chlorotalpa, Chrysochloris, Chrysopalax, Erimitalpa.

ORDER MACROSCELIDEA

Elephant Shrews. The elephant shrews have dentition that resemble rodents (long incisors) and a body that resembles shrews.

Middle Eocene to present

FAMILY MACROSCELIDIDAE

Jumping or Elephant Shrews.  These terrestrial leaping mammals are from south and east-central Africa .  They have long snouts, large eyes and ears.  They are adapted for leaping.

Elephantulus, Macroscelides, Petrodromus, Rhynchodon.

PAENUNGULATA

ORDER HYRACOIDEA

Hyraxes, Dassies, and Conies.  They are herbivores from sub-Saharan Africa and the Middle East .  Hyraxes are rabbit sized animals with incisors like those of rodents.

Eocene to present

FAMILY PROCAVIIDAE

Hyraxes.  

Dendrohyrax, Heterohyrax, Procavia.

ORDER SIRENIA

Sea cows.  Early forms had short legs.  Later the forelegs became paddle-like and the hindlegs became reduced to a few bones that do not emerge from the animal and acquired a whale-like tail.  Animals of tropical and subtropical coastal waters where they eat water plants.  Their dentition is reduced to a few molars.  Also, they have a short elephant-like proboscis. 

Eocene to present

Pezosiren+, Dusisiren+

FAMILY DUGONGIDAE

Dugongs and Sea Cows.  They are fully marine animals of eastern Africa , southern Asia over to the Philippines and Australia .  They have no vestigial nails on their flippers, and their flukes are notched.  The rostrum of the skull is bent strongly downward.  They have a pair of upper incisors on each side of the jaw.  The second one is like a tusk in males.  Canines and other incisors are vestigial.  Cheek teeth are simple and columnar.  They feed on vegetation in shallow coastal waters.

Dugong, Hydrodamalis+

FAMILY TRICHECHIDAE

Manatees.  These are marine animals of the coastal regions of southeastern US, the West Indies , eastern South America , and western Africa .  They have a small head with small eyes and flippers that have vestigial nails.  The tail fin is not lobed.  Incisors are absent in adults.  Cheek teeth are numerous and migrate forward with age.  They feed on aquatic plants in shallow coastal waters.

Trichechus.

ORDER PROBOSCIDEA

The developed a reduced jugal and orbit that opens to the maxilla.  Enlarged second incisors; lower canines and first premolars absent.  Modifications in the limbs to support great weight.  Presumably, all had extensions of the nose that became the trunk in large forms that could not lower their heads to the ground.

SUBORDER UNNAMED+

FAMILY MOERITHERIIDAE

These were the earliest forms.  They had a deep skull with second incisors of both upper and lower jaws enlarged and projecting forward.  They were relatively small semi aquatic animals, much like small hippos.

Lower Eocene to Oligocene

Phosphatherium, Moeritherium

FAMILY DEINOTHERIIDAE

These animals had a pair of recurved lower tusks and no upper tusks. 

Eocene to middle Pleistocene

Deinotherium

SUBORDER ELEPHANTIFORMES

These animals lost more of their premolars

FAMILY MAMMUTIDAE+

The paleomastodontids.  A pair of elongate upper tusks; molars with rounded cusps.

Upper Eocene to Miocene

FAMILY GOMPHOTHERIIDAE+

Upper and lower incisors elongate as tusks.  Mastodon-type teeth with mounded cusps. 

Miocene

Gomphotherium

FAMILY STEGODONTIDAE+

Molars with more incised (higher) cusps.

Pliocene

Stegodon

FAMILY ELEPHANTIDAE

Elephants and Mammoths.  Large terrestrial animals of Africa and southern Asia .  They have large heads, a long prehensile proboscis and incisors that emerge as tusks (Asian Elephant females do not have tusks).  They have a limited number of molars that move forward to replace those that have worn out.  The legs are large and columnar, and the ears are relatively large.  Mammoths had ridged cusps on their molars which might be present as one or two at a time. Pleistocene (a dwarf mammoth survived on Wrangel Island until 4,000BCE) to present.

Elephas, Loxodonta, Mammuthus+.

XENARTHRA

These animals have a suite of unique characters including extra joints between some of the trunk and tail vertebrae.  Also, the ischium and the ilium are fused to the first caudal vertebra.  They have reduced teeth or no teeth at all. 

ORDER CINGULATA

FAMILY DASYPODIDAE

Armadillos.  They are terrestrial animals that range from the central US south through South America .  They are covered by bony scutes with horny plates on the outside.  The head and tail are similarly armored.  The eyes are small.  All have muscular forelimbs that terminate in strong digging claws.  They have a very simple dentition that lacks canines. 

Upper Paleocene to present

Subfamily Chlamyphorinae (Fairy Armidillos and Pichiciegos): Chlamyphorus.

Subfamily Dasypodinae (Armadillos): Cabassous, Chaetophractus, Dasypus, Euphractus, Priodontes, Tolypeutes, Zaedyus.

FAMILY GLYPTODONTIDAE+

The glyptodonts were similar to armadillos, but many grew to very large size and were much more heavily armored.  They had continuously growing cheek teeth.  Some had a spiked club at the end of the tail.

Pliocene to Pleistocene

Glyptodon

ORDER PILOSA

Sloths and South American Anteaters

Miocene to present

FAMILY MYRMECOPHAGIDAE

South American Anteaters.  These are remarkable animals of Central and South America .  They have very elongate snouts from which they can extend a very long tongue that is tipped with sticky saliva with which they gather ants and termites.  Tails are long and prehensile.  The front legs are strong with powerful claws that they use to tear into ant and termite nests.  They have no teeth and an opening that is small and tubular. 

Lower Miocene to the present.

Cyclopes, Myrmecophaga, Tamandua.

FAMILY BRADYPODIDAE

Three-Toed Sloths.  They are arboreal animals that range from Central America to southern Brazil .  They have hairs with grooves in which algae grow (adding to their cryptic coloration).  The forelegs are much larger than the hind legs.  Fore and hind legs terminate in three curved claws.  They have no incisors or canines.  They also have a reduced number of cheek teeth.  They have 9 cervical vertebrae.  They feed primarily on leaves.

Bradypus, Hapalops+

FAMILY MEGALONYCHIDAE

Two-Toed Sloths.  These are arboreal animals that range from Central America to the Amazon basin.  They have long shaggy fur on which algae grow and aid in cryptic coloration.  The forelimbs are slightly longer than the hind limbs and terminate in two large, curved claws (the hind limbs have three claws).  Incisors and canines are absent, but the cheek teeth are vary and are enlarged toward the front of the mouth (the forward most are canine-like).  Their body temperature is quite variable (24-33C).

Miocene to the present.

Choloepus.

FAMILY MEGATHERIIDAE+

Giant Ground Sloths.

Megatherium

FAMILY MYLODONTIDAE?

BOREOEUTHERIA

ORDER LEPTICTIDA+

Small shrew-like animals; long snouts and small sharp teeth (insectivores).  They retain the jugal. 

Paleocene to Oligocene

Leptictis

ORDER ANAGALIDA+

Herbivorous.  These animals had broad grinding molars.  Rabbit-like in appearance.  They retain a postorbital process.

Paleocene

Anagale

ORDER APATEMYIDA+

Insectivorous.  Their dentition seemed to be a mixture of rodent and shrew. The incisors were long and project forward.

Paleocene to Oligocene

Sinclairella

ORDER TAENIODONTA

Herbivores. The forelimbs and forefeet were larger than the hind limbs and hind feet.  Claws were narrow and recurved (for digging?). 

Paleocene to Eocene

Stylinodon

ORDER TILLIODONTIA

Herbivores.  Some were bear-sized.  Their teeth indicate that they fed on tough plants.

Paleocene to Eocene

ORDER PANTODONTA

Herbivores, relatively large (pig to hippo-sized) that resembled ground sloths.  They had massive limbs and plantigrade feet; some may have been semi-erect. 

Paleocene to Oligocene

Titanoides

ORDER PANTOLESTA

These resembled otters with flattened skulls.  Their enlarged molars suggest that they may have fed on shellfish.  Stomach contents also indicate that they fed on fish.

Paleocene to Oligocene

Buxolestes

ORDER DINOCERATA

Also known as the uintotheres, these animals were very large and built much like a rhinoceros.  They had bony protuberances on the skull and large canines that may have been used for ritual combat. 

Paleocene and lower Eocene

Uintatherium

LAURASIATHERIA

ORDER LIPOTYPHLA

This order includes most of the animals once regarded as Insectivora. 

SUBORDER ERINACEOMORPHA

The hedgehogs.  Early forms were dog-sized and covered with stiff hairs.  Later forms evolved the spines characteristic of today’s hedgehog. 

Eocene to present

FAMILY ADAPISORICIDAE+

Cretaceous.

FAMILY ERINACEIDAE

Hedgehogs and Rat Shrews. These are terrestrial mammals of Africa , Eurasia , over to Borneo .  They are plantigradeand some have enlarged incisors.  The hedgehogs have spines (rat shrews lack spines).  They are good diggers and are omnivorous.  Sme are good climbers and swimmers.

Subfamily Erinaceinae (Hedgehogs): Atelerix, Erinaceus, Hemiechinus, Mesechinus.

Subfamily Hylomyinae (Gymnures and Moonrats): Echinosorex, Hylomys, Podogymnura.

Deinogalerix+.

SUBORDER SORICOMORPHA

The shrews and moles.  W-shaped pattern of ridges on the upper molars is diagnostic for the group. 

Middle Paleocene to present

FAMILY SORICIDAE

Shrews. World-wide (except Australia and southern South America ).  Small insectivores with long muzzle.  There are two major groups: red-toothed shrews ( Europe , northern Asia and North America ) and whilte-toothed shrews ( Africa ans southern Asia ).

Subfamily Crocidura (White-Toothed Shrews): Congosorex, Crocidura, Diplomesodon, Feroculus, Myosorex, Paracrocidura, Ruwenzorisorex, Scutisorex, Solisorex, Suncus, Surdisorex, Sylvisorex.

Subfamily Soricinae (Red-Toothed Shrews): Anourosorex, Blarina, Blarinella, Chimarrogale, Cryptotis, Megasorex, Microsorex, Nectogale, Neomys, Notiosorex, Praesorex. Sorex, Soriculus.

Domnina+.

FAMILY TALPIDAE

Moles and Desmans. Fossorial and aquatic animals of North America and Eurasia .  The fossorial taxa have evolved the typical large forelegs that terminate in powerful claws and no external ears.  They have very small eyes.  Moles are adapted to burrowing with large digging claws and weak hind legs; small eyes and ears.  Desmans look like fat shrews (they are aquatic) with webbed feet.

Eocene to the present.

Subfamily Desmaninae (Desmans): Desmana, Galemys.

Subfamily Talpinae (Moles): Condylura, Euroscaptor, Mogera, Nesoscaptor, Neurotrichus, Parascalops, Parascaptor, Scalopus, Scapanulus, Scapanus, Scaptochirus, Scaptonyx, Talpa, Urotrichus.

Subfamily Uropsillinae (Chinese Shrew Moles): Uropsilus.

ORDER CHIROPTERA

The bats use fingers 2-5 as the frame for their wings.  The shoulder girdle is adapted to large flight muscles on the scapula, ribs, and sternum.  Hind feet can turn backwards. 

Upper Paleocene to present

SUBORDER MEGACHIROPTERA

Fruit bats.  This suborder has a single family.

FAMILY PTEROPODIDAE

Flying Foxes and Old World Fruit Bats.  They are large bats of the tropics and subtropics of Africa , southern and central Asia , Australia , and Pacific islands.  Some are quite large and have wingspans up to 1.7m.  Most are much smaller.  Pteropodids have a suite of unique characters that include: a second finger with a claw, elongated bony palate, no more than 4 upper and 4 lower incisors, relatively simple ears and small cochlea, tail absent or, if present, much reduced.  They eat fruit or nectar.

Oligocene to the present.

Subfamily Macroglossinae (Blossom Bats, Dawn Bats, Long-Tongued Bats): Eonycteris, Macroglossus, Megaloglossus, Melonycteris, Notopteris, Syconycteris.

Subfamily Pteropodinae (Old World Friut Bats): Acerodon, Aethalops, Alionycteris, Balionycteris, Boneia, Casinycteris, Chironax, Cynopterus, Dobsonia, Dyacopterus, Eidolon, Epomophorus, Epomops, Haplonycertis, Hypsignathus, Latidens, Megaerops, Micropteropus, Myonycteris, Nanonycteris, Neopteryx, Nyctimene, Otopteropus, Paranyctimene, Penthetor, Plerotes, Ptenochirus, Pteralopex, Pteropus, Rousettus, Scotonycteris, Sphaerias, Styloctenium, Thoopterus.

SUBORDER MICROCHIROPTERA

Insect-eating bats.  Mainly small with relatively large and elaborate ears, many with nose-leaves, both of which function in echolocation.  Index finger without a claw; tail usually long.  Found from the tropics through temperate latitudes.  Teeth broad with many cusps.  Most are sociable; some hibernate.

FAMILY CRASEONYCTERIDAE

Bumblebee Bat and Hog-Nosed Bat.  Tiny bats of the Kwai River basin in Thailand .  They have no tail.  They have a hog-like nose and large ears.

Craseonycteris.

FAMILY EMBALLONURIDAE

Sac-Winged Bats and Tomb Bats. Insectivores of tropical and subtropical areas world-wide.  Most notable is the glandular sac on the leading edge of the wing.  The tail is in a membranous sheath.  They use echlolocation to catch prey and for communication.

Upper Eocene to the present.

Balantiopteryx, Centronycteris, Coelura, Cormura, Cyttarops, Diclidurus, Emballonura, Mosia, Peropteryx, Rhynchonycteris, Saccolaimus, Saccopteryx, Taphozous.

FAMILY FURIPTERIDAE

Smoky Bats and Thumbless Bats.  Insectiverous bats of South America .  Aside from being very small, they have very reduced thumbs.  They have large ears that seem to cover the eyes

Amorphochilus, Furipterus.

FAMILY MEGADERMATIDAE

False Vampire Bats. India to the Philippines . They are  relatively large bats with large ears that are united at their bases.  Their nose leaves are large, and they have large canines.  They feed on small vertebrates, birds, small bats, lizards, frogs, mice, and fish.

Cardioderma, Lavia, Macroderma, Megaderma.

FAMILY MOLOSSIDAE

Free-Tailed Bats. They occur from Argentina northward to the central US and tropical and subtropical Asia and Australia .  They are gregarious bats in buildings or caves.  They have broad ears, united at the bases; and at least one-half of the tail free of the interfemoral membrane.

Chaerephon, Cheiromeles, Eumops, Molossops, Molossus, Mops, Mormopterus, Myopterus, Nyctinomops, Otomops, Promops, Tadarida.

FAMILY MORMOOPIDAE

Ghost-Faced Bats, Moustached bats, and Naked-Backed Bats.  They occur from Brazil to the southern US.  They do not have a noseleaf, but they have elaborate folded skin for lips.  They have moustache of stiff hairs.  Insectivores and usually near water.

Pleistocene to the present.

Mormoops, Pteronotus.

FAMILY MYSTACINIDAE

New Zealand Short-Tailed Bats.  These bats are restricted to New Zealand .  They can hide their wings within a leathery sheath.  Thus, they can easily walk on the forest floor where they spend much of their time searching for food.  The nose is pointed.  They eat insects as well as fruit, nectar, and pollen.

Mystacina.

FAMILY MYZOPODIDAE

Old-World Sucker-Footed Bats.  These bats are restricted to Madagascar .  Their wrists are modifeied into suction devices that allow them to roost inside rolled leaves.  They have relatively large ears and toes with only 2 phalanges.

Pleistocene to the present.

Myzopoda.

FAMILY NATALIDAE

Funnel-Eared Bats.  They are small insect eating bats of lowland areas from northern Mexico to Brazil .  These bats have relatively long legs and large ears.  They do not have a noseleaf, but males do have a natalid organ on the face (the function of it is unknown). 

Natalus.

FAMILY NOCTILIONIDAE

Bulldog Bat or Fish-Eating Bat.  Central America . Bulldog face and large mouth; long ears and nose leaf. Unusually long legs, toes, and claws.  They skim over water catching small fish and crustaceans.

Noctilio.

FAMILY NYCTERIDAE

Slit-Faced Bats.  Bats of tropical forests and arid regions from Africa and southeastern Asia .  Tey are characterized by having a peculiar slit that lies on top of the muzzle.  They have no noseleaf.  They have relatively large ears.  Most eat arthropods, but some eat small vertebrates.

Nycteris.

FAMILY PHYLLOSTOMIDAE

New World Leaf-Nosed Bats. Bats of the New World ( Mexico to the Amazon basin).  Mainly insectivores, but they also eat blood, fruit, birds, and small bats.  Nose leaf simple, erect and pointed; some with grotesque folds over the face. 

Miocene to the present.

Subfamily Brachyphyllinae (Antillian Fruit-Eating Bats): Brachyphylla.

Subfamily Carolliinae Little Bats and Short-Tailed Bats): Carollia, Rhynophylla.

Subfamily Desmodontinae (True Vampire Bats. Tropical South and Central America .  The most derived of all bats.  Its upper incisors are rodent-like but razor-sharp and can cut the skin of a living animal quickly and somewhat painlessly.  They lap the fresh blood.  Cheek teeth are reduced and vestigial.  Very agile on the ground.): Desmodus, Diaemus, Diphylla.

Subfamily Glossophaginae (Long-Nosed Bats, Long-Tongued Bats, Tailless Bats): Anoura, Choeroniscus, Glossophaga, Hylonycteris, Leptonycteris, Lychnonycteris, Monophyllus, Musonycteris, Scleronycteris.

Subfamily Lonchophyllinae (Nectar Bats): Lionycteris, Lonchophylla, Platalina.

Subfamily Phyllonycterinae (Flower Bats): Erophylla, Phyllonycteris.

Subfamily Phyllostomatinae (Round-Eared Bats, Spear-Nosed Bats, Sword-Nosed Bats): Chrotopteris, Lonchorhina, Macrophyllum, Macrotus, Micronycteris, Mimon, Phylloderma, Phyllostomus, Tonatia, Trachops, Vampyrum.

Subfamily Stenodermatinae (Fruit-Eating Bats, Tent-Making bats, Yellow-Shouldered Bats): Ametrida, Ardops, Ariteus, Artibeus, Centurio, Chiroderma, Ectophylla, Mesophylla, Phyllops, Platyrrhinus, Pygoderma, Sphaeronycteris, Stenoderma, Sturnira, Uroderma, Vampyressa, Vampyrodes.

FAMILY RHINOLOPHIDAE

Horseshoe Bats and Old-World Leaf-Nosed Bats. Bats of temperate and tropical areas of Eurasia , Africa , Australia , and many Pacific islands.  They have complex noseleaves that form a horshoe beneath the nostrils, a large sternum and anterior portion of the pelvic girdle.  They eat insects.

Subfamily Hipposiderinae (Leaf-Nosed bats): Anthops, Asellia, Anselliscus, Cloeotis, Coelops, Hipposideros, Paracoelops, Rhinonicteris, Triaenops.

Subfamily Rhinolophinae (Horshoe Bats): Rhinolophus.

FAMILY RHINOPOMATIDAE

Mouse-Tailed Bats. Bats of deserts and semi-deserts northeastern Africa and southern Asia .  They have a tail that is not enclosed or attached to a membrane.  They have large ears that are attached at the base.They have no noseleaf.  They are insectivores.

Rhinopoma.

FAMILY THRYOPTERIDAE

Disc-Winged Bats.  Bats of the neotropics.  They have suction structures on their wrists and ankles (like the myzopodids of Madagascar ) which they use for attachement inside unrolling banana leaves.  The ears are moderately large, and they have no noseleaf.

Thryoptera.

FAMILY VESPERTILIONIDAE

Common Bats. Almost global; Eurasia , northern Africa , North America to northern South America .

Subfamily Kerivoulinae (Wooly Bats):  Kerivoula.

Subfamily Miniopterinae (Bent-Winged Bats and Long-Fingered Bats): Miniopterus.

Subfamily Murininae (Hairy-Winged Bats and Tube-Nosed Bats): Harpiocephalus, Murina.

Subfamily Tomopeatinae (Blunt-Eared Bats): Tomopeas.

Subfamily Vespertillioninae (Evening Bats, Long-Eared Bats, Myotis Bats, Pipistrelles, Serotines): Antrozous, Barbastella, Chalinolobus, Eptesicus, Euderma, Eudiscopus, Glischropus, Hesperoptenus, Histiotus, Ia, Idionycteris, Laephotis, Lasionycteris, Lasiurus, Mimetillus, Myotis, Nyctalus, Nycticeius, Nyctophilus, Otonycteris, Pharotis, Philetor, Pipistrellus, Plecotus, Rhogeessa, Scotoecus, Scotomanes, Scotophilus, Tylonycteris, Vespertilio.

THE FERUNGULATA = THE CERTARTIODACTYLA

ORDER UNNAMED

FAMILY ZHELESTIDAE

ORDER ARCTOCYONIA+

Herbivore that was sheep-sized, but it looked like a dog.  Its molars, however, were broad and adapted to crushing rather than slicing flesh.  Some were adapted to climbing with muscular limbs and, possibly, a prehensile tail. 

Paleocene

Arctocyon

ORDER UNNAMED+

FAMILY MESONYCHIDAE

Carnivores. Early forms were dog-like.  Later, they were quite large.  This group likely was related to the whales. 

Paleocene to upper Eocene

Mesonyx, Andrewsarchus

ORDER ARTIODACTYLA

These are the even-toed ungulates.

SUBORDER UNNAMED+

FAMILY DICHOBUNIDAE

The basal artiodactyls.  These animals had multiple digits on the fore and hind feed (5 and 4, respectively), but they bore the weight on digits 3 and 4, each of which terminated in a small hoof.  They had the diagnostic double pulley astragalus.  They also showed artiodactyls features in the skull. 

Lower Eocene to Oligocene

Diacodexis

SUBORDER SUIFORMES (BUNODONTIA)

Generally, omnivores.  The pigs and hippos are characterized by having bulbous cusps on their molars.  Also, they have powerful canines. 

Upper Oligocene to the present

FAMILY ENTELODONTIDAE+

These pig-like animals had long skulls.  Omnvivores. 

Oligocene

Dinohyus

FAMILY SUIDAE

Pigs.  Formerly distributed through Eurasia , south of 48 degrees; to the Philippines and Africa . Now, feral animasl occur throughout the world.  Suids have stocky bodies; eyes are small and high on the skull; and ears are pointed.  They have have long canines (continuously growing) used in feeding and fighting.  Omnivores with a muscular, mobile snout.  They have a 2-chambered stomach that does not ruminate.

Upper Oligocene to present: Perchoerus+

Subfamily Babyrousinae (Babyrousa): Babyrousa.

Subfamily Phachoerinae (Warthogs): Phacochoerus.

Subfamily Suinae (Hogs and Pigs): Hylochoerus, Potamochoerus, Sus

FAMILY TAYASSUIDAE

Peccaries.  Omnivores of the southwestern US to Argentina .  These pig-like animals resemble suids in many ways from the overall appearance to their mobile, disc-like snout.  They have a scent gland on the rump and their feet are different.  Their tusks also are directed downward.  Their stomach has 3 chambers and is not ruminating.

Oligocene to present.

Catagonus, Pecari, Tayassu.

FAMILY ANTHRACOTHERIIDAE

These animals were sisters to the hippos.  Small at first but large later (the size of a pygmy hippo). 

Eocene to Pliocene

FAMILY HIPPOPOTAMIDAE

Hippopotamuses.  These animals ranged through all of sub-Saharan Africa and Madagascar .  The large hippopotamus is a semi-aquatic animal while the pygmy hippo is a forest grazer.  Their bodies are large and barrel-shaped with stubby legs. 

Middle Miocene to present

Hippopotamus, Hexaprotodon, Choeropsis+

SUBORDER SELENODONTIA

These animals have specialized cheek teeth with a selenodont pattern (pairs of crescent moon ridges).  Upper incisors reduced or missing (elongate as sabers in some for sexual display).  Compound stomach. 

TYLOPODA

FAMILY MERYCOIDODONTIDAE

Oreodonts.  These animals were pig-sized browsers that moved in large herds.  They had 4 toes on each foot.

Merycoidodon, Bathygenys

FAMILY PROTOCERATIDAE

They were deer-like in form, but with shorter legs.  They also evolved horns (as seen in the Ruminantia) above the eyes and on the nose.

Upper Eocene to Pliocene.

FAMILY CAMELIDAE

Camels, Llamas, Vicugna.  Early camels had two toes with hooves.  Later, they evolved pads.  The center of origin was North America .  The skull is elongate with a well-developed saggital crest.  They have a 3-chambered ruminating stomach.