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THE ORIGIN OF MAMMALS

What traits have mammals got that make them unique?
A. Glands
Of all the vertebrates, mammals have the greatest variety of integument (=skin) glands
All are derivatives of the epidermis
1. Sweat Glands
Tubular coiled glands that occur over much of the body
Secrete a watery sweat that when evaporated on the skin's surface draws heat from the
skin and cools it
Much reduced or absent in rodents and whales
2. Scent Glands
Present in nearly all mammals
Used in communication with members of the same species
3. Sebaceous glands
These are oil-secreting glands
Function to soften and lubricate the skin and hair and to kill some kinds of bacteria
In humans, they are highly concentrated on the face
4. Mammary Glands
Probably modified apocrine glands
Occur on all female mammals and in a rudimentary form on all male mammals
In females these glands produce milk; all mammalian females nourish their young with
milk, a balanced diet rich in fats, sugars, proteins, minerals and vitamins
B. Hair
All mammals possess hair, which helps to insulate the animal and maintain a constant
body temperature.
In some mammals hair has been substantially reduced, e.g. in whales it is reduced to a
few hair bristles around the snout and eyelids
Mammals characteristically have 2 kinds of hair:
Underhair - dense, soft hair; important as insulation
Guard Hair - course and longer: functions in protection and provides coloration
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C. Diaphragm
Mammals have a diaphragm, a sheet of muscle and tendon that separates the body cavity
into two sections.
No other animal has a diaphragm.
D. High metabolic rate/high body temperature
Though this trait is not unique to mammals, they are one of just two important living
groups of vertebrates that possess the trait
Note:
All of these traits involve "soft stuff"; it's hard to judge their appearance in the fossil
record.
Instead, we look at novel hard parts in mammals that correlate with this soft stuff.
E. Big brains with modified skulls
F. Three ear bones
G. Complex teeth
H. Reduced number of bones in the lower jaw
These hard parts arose in a mosaic fashion over millions of years of synapsid evolution,
from pelycosaurs to therapsids, and from therapsids to mammals.
These transformations were in no sense a straight line "march" from pelycosaur to
mammal.
Rather, "important" mammalian traits arose on some lines, whereas other traits that
weren't ultimately passed on to long-lasting descendents arose on other lines.
Mammal teeth are different from teeth of other vertebrates
Mammal teeth have more complex shapes that the teeth of other vertebrates.
Many more cusps, cones, and ridges.
They have teeth that differ from front-to-back in the jaw: Incisors, canines, premolars,
molars.
Complex occlusion between teeth.
When chewing, mammal teeth lock together in a very complex and rigid fashion.
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When did this happen?
Pelycosaurs have simple, cone-shaped teeth that don't lock together tightly during biting.
Some do have big "canine" teeth, but little other front back differentiation.
Some therapsids have much more complex shapes and occlusion and substantial front-
back differentiation.
Extremely complex teeth and occlusion, and full front-back differentiation developed
after the first appearance of true mammals.
Why innovations in teeth?
As therapsids and early mammals began to eat a greater diversity of foods, particularly
plants, they needed to develop specialized chewing systems.
Consequences?
Mammals have only one set of replacement teeth (most vertebrates have many).
They can't continuously replace their teeth and maintain complex occlusion.
Mammal teeth have a thick coating of enamel to make sure they last a lifetime.
Mammal skulls are different from skulls of other vertebrates
They greatly increase their brain size.
They have elaborate musculature on their head to control chewing.
Many have a big crest on the top of their heads for muscle attachment.
They enlarge the synapsid opening greatly to accommodate added muscles and swelling
brain.
Mammals have new solutions to breathing/walking problems.
Secondary palate: allows simultaneous chewing and breathing.
This structure first began to develop in therapsids.
Diaphragm: pumps air in and out without ribs
Upright posture and a spine that flexes in the vertical, not horizontal, plane.
Pumps air in-and-out, rather than from side-to-side, when the animal moves
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Mammals have bizarre ears and jaws
Mammals have three bones in their ears that transmit sound from the eardrum to the inner
ear.
Eardrum - malleus - incus - stapes - inner ear
A Brief Overview on Ear-Jaw Evolution among Mammals
The oldest reptiles having mammal-like features, the synapsids, occur in rocks of
Pennsylvanian age formed about 305 mya.
However, the first mammals do not appear in the fossil record until Late Triassic time,
about 210 mya.
Hopson (1994) noted, "Of all the great transitions between major structural grades within
vertebrates, the transition from basal amniotes [egg-laying tetrapods except amphibians]
to basal mammals is represented by the most complete and continuous fossil record....
Structural evolution of particular functional systems has been well investigated, notably
the feeding mechanism... and middle ear, and these studies have demonstrated the gradual
nature of these major adaptive modifications."
A widely used definition of mammals is based on the articulation or joining of the lower
and upper jaws.
In mammals, each half of the lower jaw is a single bone called the dentary; whereas in
reptiles, each half of the lower jaw is made up of three bones.
The dentary of mammals is joined with the squamosal bone of the skull.
This condition evolved between Pennsylvanian and Late Triassic times.
Evolution of this jaw articulation can be traced from primitive synapsids (pelycosaurs), to
advanced synapsids (therapsids), to cynodonts, to mammals.
In mammals, two of the extra lower jawbones of synapsid reptiles (the quadrate and
articular bones) became two of the middle-ear bones, the incus (anvil) and malleus
(hammer).
Thus, mammals acquired a hearing function as part of the small chain of bones that
transmit air vibrations from the eardrum to the inner ear.
Where did these extra ear bones come from?
Pelycousaurs (like lizards and amphibians) transmitted sound from the eardrum to the
inner ear with just the staples, which had evolved earlier via transformation of an old gill
support bone.
The joint between the head and lower jaw was between the quadrate (on the skull) and
the articular (on the lower jaw).
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In many therapsids, the quadrate and articular reduce in size and shift towards the back
of the head.
Sound waves were transmitted via the eardrum (located on the lower jaw), through the
articular-quadrate jaw joint, and on to the stapes and inner ear.
Some sound waves also moved up from the feet, into the shoulder, through the jaw joint,
and on into the head.
In true mammals, the articular-quadrate joint lost all function in jaw opening and closing.
Instead, the articular became the malleus, the quadrate transformed in the incus, and both
bones were transferred to the lower jaw and used solely for hearing.As a consequence,
the jaw joint had to move forward, and was now between the major bone in the lower jaw
(the dentary) and a different bone on the skull.
Summary
This shift to hearing with three shaking bones, and moving the jaw joint forward, is the
shared novelty that unites all true mammals.
Ultimately, mammals would keep just one bone in their lower jaw, and the ear bones
would shift to the skull, but this was not the case early in mammal evolution.

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Who are the early mammals?
Morganucodon: late Triassic:
The first mammal
Small, shrew-like insectivore, agile climber and jumper
Good mammalian ear, but it is attached to lower jaw, not the skull.
Upright posture for hind limb.
Primitive in many other respects (more than one bone in lower jaw, sprawling front
limbs).
Monotremes: Cretaceous-Recent
Egg-laying, aquatic predators on arthropods and worms
Milk oozes from the skin (no breasts).
Hair
Ear bones shift from lower jaw to skull during embryonic development.
Electroreception (receptors on front edge of rostrum)
Multituberculates: Jurassic-Eocene
Small rodent-like animals (rat- to rabbit-sized)
Important small herbivore in Cretaceous and early Cenozoic.
Incisor (grasp/puncture), splicing blade-shaped premolar, grinding molars.
Single bone in lower jaw.
Fossil record shows that they had hair.
Some may have had live birth.
Marsupials: Cretaceous-Recent
Pouched mammals.
Born as gross little embryos.
Crawl into pouch, attach to nipple, and develop.
Cretaceous ones were fairly opposum-like in their ecology.Later ones are more diverse.
Today, they are most diverse on Australia and South America.
They share complex type of molar tooth shape with Placental mammals.
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Placentals: Cretaceous-Recent
Nourish their young internally with a placenta, a complex intergrowth of maternal tissue
with the embryonic membranes of the amniotic eggs.neonate tissue
Placentals give birth to offspring that are more "adult" like and independent.
Cretaceous ones were shrew-like in their ecology.
Later placentals are spectacularly diverse.

How do Placentals and Marsupials differ?

Marsupials Placentals
Slower growth and maturation. Faster growth and maturation.
Smaller brains.Biggers brains.
Slower metabolic rates. Higher metabolic rates.
The first two differences probably relate to the more efficient feeding of developing
placental mammals relative to marsupials.
The reason for the latter difference is not clear.
What is one advantage of having young develop in a pouch?
Flexibility in inconstant environment.