PROTOTHERIA

The subclass Prototheria contains the egg-laying mammals, which are the most ancestral forms
in the class Mammalia. There are only three living species grouped into two families which
comes under a single order, the Monotremata. Monotremata means single opening which refers
to the cloaca, a common opening for the fecal, urinary, and reproductive tracts. Prototherians are
endothermic, but they have unusually low metabolic rates and maintain a body temperature that
is lower than that of most other mammals. All prototherians are carnivorous, with their diets
consisting of various invertebrates. Platypuses searched food in the benthos of lakes and streams,
using their sensitive bills to find prey. They are generalist predators, whereas echidnas specialize
on either ants and termites (Tachyglossus) or worms (Zaglossus). Both species of echidna are
powerful diggers and use their claws and snouts to root through the earth to find food.
Classification:
Living monotremes are classified in two families :
1. Ornithorhynchidae (platypus family)
2. Tachyglossidae (echidnas)
Distribution :
Monotremes are restricted to Australia and New Guinea. Their fossil record is very poor; the
earliest fossil attributed to this group is from the early Cretaceous.
General Characters :
External Features:
Body is small , covered by hairs and spines. Snout produced into beaks. External ears
inconspicuous or absent. Tail present or absent . Mammary glands are without teats or nipples.
Males carry a hollow and horny ,tarsal spur on each hindleg , connected internally to a poison
gland. A temporary mammary pouch , equivalent to teats , develops during breeding season on
the abdomen of female.
Endoskeleton:
1. The skulls of monotremes are almost birdlike in appearance, with a long rostrum and
smooth external appearance.
2. Modern monotremes lack teeth as adults with no sutures ,the rostrum is elongate, beak-
like, and covered by a leathery sheath and lacrimal bones are absent.
3. Dentary bone is reduced and adults are edentate.
4. The jugals are reduced or absent, the dentary is a slender bone with only a vestige of a
coronoid process, the angle of the dentary is not inflected medially (unlike that of
marsupials), auditory bullae are missing (part of the middle ear is enclosed by tympanic
rings), and much of the wall of the braincase is made up by the petrosal rather than the
alisphenoid (unlike all other modern mammals).
5. Monotremes have no corpus callosum, a bundle of nerve fibers that integrate the 2
hemispheres of the brain.

Postcranial skeleton :
1. Their shoulder girdles are complex, including the standard components of modern
mammals (scapula and clavicle), but also additional elements including coracoid,
epicoracoid, and interclavicle.
2. The scapula, however, is simplified, lacking a supraspinous fossa. The shoulder girdle is
much more rigidly attached to the axillary skeleton than in other mammals.
3. Femur and humerus are held roughly parallel to the ground when the animal walks, more
in the fashion of therapsids and most modern reptiles than like modern mammals
4. Ribs are found on the neck (cervical) vertebrae as well as the chest (thoracic) vertebrae;
in all other modern mammals, they are restricted to the thoracic region.
 5. Another interesting skeletal characteristic of monotremes is the large epipubic bones in
the pelvic region. Epipubic bones were originally thought to be related to having a
pouch, but they are found in both males and females. They also occur in all species of
marsupials, whether a pouch is present or not (not all marsupials have a pouch). It is now
thought that epipubic bones are a vestige of the skeleton of therapsids, providing
members of that group with extra attachments for abdominal muscles to support the
weight of the hindquarters.
6. All male monotremes have spurs on their ankles that are presumed to be used in fighting
and in defense. In the Ornithorhynchidae, a groove along the spur carries poison secreted
by adjacent glands
Reproduction
1. The eggs layed by monotremes are small (13-15 mm diameter) and covered by a leathery
shell.
2. The number of eggs laid is small, usually 1-3, and they are placed in the mother's pouch,
except in the platypus, which does not have a pouch.
3. They contain a large yolk, which is concentrated at 1 end of the egg very much like the
yolk of a bird's egg
4. Only the left ovary is functional in the platypus, but both varies produce eggs in the
echidnas.
5. After an egg is shed into the infundibulum, it passes to the Fallopian tube, where
fertilization occurs. The shell is deposited in the oviduct over a period of about 2 weeks.
Nutrients are absorbed through the shell; thus, the eggs are permeable (not impermeable,
as is the case for birds).
6. Like the eggs of birds, monotreme eggs are incubated and hatched outside the body of the
mother. Incubation lasts about 12 days.
7. The young, which are tiny and at a very early stage of development when they hatch,
break out of the eggs using a "milk tooth.
8. They are fed milk produced by mammary glands; the milk is secreted onto the skin
within the pouch and sucked or lapped up by the babies.
9. Weaning takes place when the young are 16-20 weeks old.
10. After the eggs hatch the young are nursed via mammary glands without teats. Milk is
secreted from pores on the belly of the platypus and from paired glandular lobes in the
pouch of echidnas.
11. Males have a baculum, permanently abdominal testicles, and no scrotum.
Affinities of Monotremes :
Prototheria resemble the reptiles and birds with some advanced characters over them,
establishing mammalian ancestry.
Reptilian Affinities:
1. Presence of cloaca.
2. Presence of ectopterygoid in skull.
3. Vertebrae without epiphysis and with cervical ribs.
4. Ribs are single headed.
5. Cervical ribs are present.
6. Thoracic ribs are single headed.
7. A median T-shaped interclavicle present.
 8. Acetabulum in echidna is perforated.
9. Body temperature is not constant.
10. Cochlea of internal ear with lagina.
11. Ureters lead into a urinogenital sinus.
12. Corpus callosum is absent and anterior commissure is well developed.
13. Testis abdominal.
14. Oviparous and meroblastic segmentation.
Presence of strong reptilian features in Monotremata speaks of its primitiveness. These primitive
mammals have failed to cope up with many of the evolutionary transformations which
culminated in the establishment of better characteristics in higher mammals.

Avian affinity:
1. Beak of the platypus resembles that of birds.
2. Teeth in adults are absent.
3. Presence of webbed feet.
4. Oil gland is present.
The relationship between monotremes and birds does not stand on a solid ground. The
converging characters seen in them are due more to the fact that both possess common reptilian
ancestry.

Mammalian affinity:
The prototherians are essentially mammals as they possess the following typical mammalian
characters:
1. Presence of hair, mammary glands, oil gland and sweat glands.
2. Double occipital condyles.
3. Presence of palate.
4. A typical mammalian diaphragm is present in the body cavity.
5. Skull is dicondylic.
6. Sternum is segmented.
7. Lobes of liver typically mammalian.
8. Heart 4-chambered.
9. Only left aortic arch present.
10. Circulatory system is typically mammalian.
11. Presence of large ear ossicles.
12. Cochlea is slightly coiled.
13. Fertilization is internal.
14. A slender caecum demarcates two intestines.
15. Lobes of liver typically mammalian.
16. R.B.C. small, circular and non-nucleated.
17. Presence of 4 optic lobes (corpora quadrigemina).
18. Presence of milk glands secreting milk
Though monotremes show affinity with non-mammalian groups, the above mentioned characters
strongly speak of close and firm affinity with mammal

Phylogenetic status of Monotremes

There are two theories which explain the phylogeny of Monotremes. One theory explains that
Monotremes evolved independently from early mammal- like reptiles and continued to survive in
isolation as basically primitive mammals with certain specializations. Another theory advocates
that Monotremes have been derived from very Marsupials.
Among the mammals the Monotremes are very much controversial. They possess primitive,
degenerated and specialized features. It is reasonable to conclude that Monotremes originated as
a side line from the main line of mammalian evolution and have retained the characters through
which ancestors of higher mammals have passed.