Origin of Mammals
The mammals of today are but one branch of the Synapsida, a great vertebrate group with a 300 million year history. Pre-mammalian synapsids -- the Pelycosaurs and Therapsids -- dominated the land vertebrate fauna of the Permian before losing ground during the end-of-the-Permian extinction and then to the diversifying dinosaurs and other archosaurs in the Mesozoic.


Characteristics of Mammals

1. Warm blooded - along with adaptations such as hair to control heat loss and sweat glands to augment evaporation and cooling.

2. Mammary Glands - modified sweat glands that provide nutrients for young. Most mammals are viviparous (give birth to live young), but the monotremes are oviparous, and most, if not all, non-mammalian synapsids were probably oviparous.

3. Diaphragm - to increase the inspiration of oxygen and expiration of carbon dioxide (necessary for a high metabolic rate).

4. A four chambered heart that separates oxygenated from unoxygenated blood:


5. Expansion of the neocortex of the brain - resulting in greater intelligence.

6. An ear more sensitive to a range of vibrations because the bone connected to the ear drum has detached from the rigid jaw, joined three new bones and become the bones of the middle ear where they are free to vibrate more easily.



7. A mandible composed of only one bone (the dentary) that articulates with the squamosal bone of the skull.

8. A bony secondary palate separating the nasal passage from the mouth.

9. Greater differentiation of the teeth characterized by multi-rooted teeth with multicusped crowns.



Transition From the Other Synapsids to Mammals

Mammals first appeared around 220 million years ago (in the Jurassic). They differ in many ways from these ancestors, and almost all of the differences appear to be reflections of a more active life, supported by a relatively high, constant metabolic rate. The transformation of the therapsids into true mammals is gradual and the new features (see list above) do not all appear at once.

Major Changes in the Transition

Locomotion

Mammalian locomotion involved a change in limb posture from sprawling (in Pelycosaurs) to erect, with elbows pointing out and backward, and knees pointing out and facing forward. As discussed earlier, erect body posture places the hands and feet closer to the body's center of gravity, resulting in less need for large ventral limb muscles just to hold the body up off the ground. All of the movement of the limb goes into rotation back and forward (i.e., moving forward, not into lifting the body off the ground). The long bones of mammalian limbs became more slender in conjunction with the shift to more erect body posture.

Another feature that goes along with the erect body posture: the vertebral column becomes more rigid. To compensate for the loss of flexibility, more neck bones are added and a more rounded occipital condyle (joint at the base of the cranium) develops which allow more flexible movement of the head.

Feeding
Major changes in the teeth, jaws, and jaw musculature modified the feeding system.

Teeth - the mammalian dentition differentiated into small incisors, large canines, and increasingly complex premolars and molars (called the cheek teeth). Increased complexity in the cheek teeth included the growth of additional cusps on the crowns with precise interlocking of cusps and ridges between upper and lower teeth.

The more elaborate cheek crowns provided mammals with shearing and crushing capability during chewing rather than the simple puncturing of single cusped reptile teeth.

As crown complexity increased, the cheek teeth developed multiple roots for firm anchorage.

One of the features of all mammals is the presence of interlocking teeth which results in improved chewing (the upper and lower teeth in the jaw work together to cut or crush and grind food). BUT, the interlocking cusps of upper and lower teeth cannot be maintained if the teeth are continuously lost and replaced. This led to evolution of just two sets of teeth one set replacing the other in the juvenile (milk teeth and adult teeth -- I think the tooth fairy must have also evolved at about this time). This altered design means the early egg-laying mammals must have evolved milk glands and lactation. Once milk is available, the young can be born with few or no teeth that can later appear when the jaw is larger and closer to the adult size. Without milk, a newborn mammal would need a full set of teeth to eat and survive.


Sensory
Brain becomes larger; hearing better
(1) reptiles have at least four bones in the lower jaw (e.g. the dentary, articular, angular, surangular, and coronoid), while mammals have only one (the dentary), and
(2) reptiles have only one middle ear bone (the stapes), while mammals have three (the hammer, anvil, and stapes)





Major Groups of Mammals:

The first mammals appear in the Jurassic. These first mammals were small shrew-sized animals (about 25 gms or 1 oz in size). This small mammal may have been successful because warm-bloodedness allowed it to live as a nocturnal insectivore.

By the late Cretaceous, 3 main types of mammals had evolved:

1. Multituberculates (now all extinct)
2. Monotremes
3. Therians (which include the Marsupial and Placental mammals)


Multituberculates - The "Lost Tribe" of Mammals

Multituberculates are the only major branch of mammals to have become completely extinct, and have no living descendants. Multituberculates get their name from their teeth, which have many cusps, or tubercles arranged in rows.

Although not known to many people, they have a 100 million-year fossil history (the longest of any mammalian lineage) and were distributed throughout the world.


Multituberculates first appeared in the Jurassic, and went extinct in the early Oligocene, with the appearance of true rodents. Over 200 species are known, some as small as the tiniest of mice, the largest the size of beavers. Some, such as Lambdopsalis from China, lived in burrows like prairie dogs, while others, such as the North American Ptilodus (skull and reconstruction pictured above), climbed trees as squirrels do today. The narrow shape of their pelvis suggests that multituberculates gave birth to tiny, undeveloped pups that were dependent on their mother for a long time before they matured.



Monotremes - Monotremes are mammals that lay eggs, and have only one external opening, called a cloaca, through which all waste matter and reproductive substances pass. (The word monotreme means "one opening.") The modern duck-bill platypus and the spiny anteater (or echidna) that live in Australia and New Guinea are the only living monotremes.

The platypus incubates its eggs in a nest, but the anteater egg is incubated in a temporary pouch that forms on the abdomen of the female.

Monotremes today produce milk from modified seat glands - but do not have nipples. The young feed by licking the milk off the mother's fur.

The echidnas and platypus have a sixth sense which all other mammals lack. In their snouts are electro-receptors, allowing echidnas to detect small electric currents. The purpose of this sense is unclear but scientists have calculated that it is sensitive enough to detect the natural electrical activity of underground grubs and worms, though probably not ants and termites.



Fossil Record:

The time and place of monotreme origin is still largely unknown. Most fossil monotremes have been found in Australia, though a platypus tooth has been recovered from Argentina, suggesting they were once distributed across southern Gondwana. There were never many different kinds of monotremes (most of the fossils can be categorized as either a platypus or an echidna).

Echidnas
:
The echidna is a medium sized animal, about a foot long and weighing around 7 kilograms (15.4 pounds, females are somewhat smaller) with a smallish head attached to a stocky body, and a long cylindrical snout. The echidna is covered on its back and sides with stout spines mixed with bristly hair, and superficially resembles a large hedgehog. It is common throughout Australia and New Guinea.


Platypus:
The duckbill platypus is one of the oddest looking mammals alive today. It lives in burrows on the banks of streams and has webbed feet to aid in paddling through the water. Its fleshy nose is used to search out earthworms in the bottom of stream beds.




Therian Mammals - In the early Cretaceous, a new group of mammals appeared which have an improved inner ear for detecting and analyzing sound. This was achieved by the coiling and enlarging of the cochlea bone of the inner ear (see figure of ear above).

There are two types of therians: the marsupial (or pouched mammals) and the placental mammals. Both marsupials and placentals give birth to live young, but the marsupials have very immature newborns that they usually place in a pouch, while the placentals carry their young in their bodies until a later stage of growth and give birth to relatively more mature newborns.

(1) Marsupial Mammals - Though marsupials today do not have as many species as do the placental mammals, they are quite structurally diverse. They are characterized by premature birth and continued development of the newborn while attached to the nipples on the lower belly of the mother.



The pouch, or marsupium, from which the group takes its name, is a flap of skin covering the nipples. Although prominent in many species, it is not a universal feature among marsupials. Where a pouch occurs, it tends to open anteriorly in upright and climbing forms, and posteriorly in quadrupedal, ground-dwelling species. The young remain firmly attached to the milk-giving teats for a period corresponding roughly to the latter part of development of the fetus in the womb of a placental mammal.

In many marsupials, the hind legs are noticeably larger than the forelegs (this is most obvious in the kangaroos); and they have a unique bone associated with the pelvic girdle which helps support the pouch.

The largest and most varied assortment of marsupials--more than 100 species--is found in Australia alone: kangaroos, wallabies, wombats, the koala, and a bewildering assemblage of smaller rodent-like forms. They also include the marsupial wolf, which may have recently gone extinct. About 70 more species are distributed more widely, in Australia, New Guinea, and a cluster of nearby islands. The wide array of Australian marsupials is reflected in the extensive popular vocabulary of names, many of which are derived from descriptive Aboriginal words. Only a few marsupials are found in the Americas, the most "famous" of which is the oppossum.

As marsupials evolved into diverse forms, they came to occupy various habitats. The result is that several have converged on placental mammal body forms :

The Tasmanian Wolf is convergent with the placental dog family:


The Koala resembles a small bear:



and the Wombat is similar to a groundhog:


In general, marsupials have a lower metabolic rate and smaller brain size than placentals. This led to a speculation that marsupials were biologically inferior to placentals, and have only managed to survive in the isolation of Australia and New Guinea because placentals were absent. There was some evidence for that position - when placentals were introduced to Australia with aboriginal man (e.g., the dog or dingo), native marsupials did not fair well and many carnivorous marsupials went extinct except on the isolated island of Tasmania (e.g., the Tasmanian Devil). Then again, when European settlers introduced other placentals (rabbits, mice, foxes, sheep, cats, etc.) marsupials declined again. Furthermore, South America supported a huge fauna of marsupials (some were spectacular like the saber-toothed "cat"). When South America collided with North America, placental mammals from the North invaded and wiped out the marsupials (except for the possum - which, of course, migrated the other direction and invaded North America).

But recently, Mike Archer (U. of Sydney) has discovered fossil placental mammal teeth in Australia. So at least once in history, marsupials apparently outcompeted placentals and took over the continent of Australia.

Placental Mammals
Placental mammals (nearly 4000 species) include such diverse forms as whales, bats, elephants, shrews, and armadillos. They are also some of the most familiar organisms such as dogs and cats, as well as many farm and work animals, such as sheep, cattle, and horses. And humans, of course, are also placental mammals.

Placental mammals all bear live young, which are nourished before birth in the mother's uterus through a specialized embryonic organ attached to the uterus wall, the placenta. The placenta is derived from the same membranes that surround the embryos in the amniote eggs of reptiles, birds, and monotreme mammals. The term "placental mammals" is somewhat of a misnomer because marsupials also have placentae. The difference is that the placenta of marsupials is temporary and does not make as much of a contribution to fetal nourishment as it does in placental mammals.



The Cenozoic is commonly called the age of mammals. During this era, the modern groups of mammals arose, diversified and came to dominate the vertebrate world.

Cenozoic Time-Line:

The first part of the Cenozoic is called the Tertiary. It began about 65 mya with the Paleocene Epoch.

Paleocene

During the Paleocene a tropical to subtropical climate stretched to the polar regions.

It seems that the placental mammals took several million years to evolve into even moderately large body sizes. One reason may have been the dense forests - large animals would have difficulty moving about, whereas small tree-dwelling animals would have been favored.

Eocene

Began 55 million years ago and lasted for 16 million years. Climate grew noticeably warmer - with the tropics reaching from Britain to the equator and temperate climates further north. South America, Africa, North America, Greenland, New Zealand and Eurasia were all distinct. Australia, and Antarctica remained connected.

The most marked feature of the Eocene is the origin and radiation of the modern mammalian orders.