TETRAPODS.

Tetrapods’ origin and early evolution are the result of the terrestrialization process of
vertebrates. Before them, two other groups of organisms had already made the "great
transition": in the Silurian we already have tracheophytes and arthropods populating the
earth. Vertebrates followed plants and arthropods and their process of terrestrialization
began in the late Devonian.

The fish that gave rise to tetrapods had two important preadaptations to terrestrial life:
functional lungs (which they used to breathe out of water) and muscular fins
(sarcopterygii) capable of providing a type of locomotion similar to the tetrapod gait
and, in any case, alternative to that typical of fish (based in undulating movements of
the trunk operated by the lateral muscles of the trunk. However, sarcopterygian fins had
to undergo many modifications in order to be efficient on land.

The transformation from fish to tetrapod was a gradual process and there are now quite
a few fossils known that illustrate that transition. Historically Ichthyostega is one of the
first fossils to be placed in the tetrapod lineage (Ichthyostega is now considered a
tetrapod stem group). Ichthyostega still shows characters from his aquatic ancestors
(especially in the skull and tail) and probably led an amphibious (and perhaps more
aquatic than previously thought) life. He lived on the banks of rivers and coastal
lagoons. It seems that its limbs were adapted to walk on shallow water bottoms rather
than on land. Ichthyostega is almost a metre long and was a carnivore, probably feeding
on fish and invertebrates. Acanthostega is another well-known fossil similar to
Ichthyostega. Both Acanthostega and Ichthyostega had more than 5 fingers on their
extremities.

LISSAMPHIBIANS
All modern amphibians share a common ancestor, so they are a monophyletic group
(known as LISSAMPHIBIA). Modern amphibians or lissamphibians (prefix "liss"
means soft and refers to the soft skin of lissamphibians): Anura (frogs and toads),
Urodela (salamanders and newts) and Gymnophiona (caecilians) have an incomplete
fossil record but seem to have originated in the early Mesozoic.
The early tetrapods had an amphibian physiology, but many of these primitive
"amphibians" were very different from today's amphibians. These first tetrapods
radiated extensively during the Carboniferous (period with abundant humid forests) and
the Lower Permian, some were small semi-aquatic animals like many of today's
amphibians, but many were large piscivorous and carnivorous animals with a markedly
terrestrial way of life (although they had an amphibian physiology).

AMNIOTES
The first fossil amniotes are Carboniferous. They are small tetrapods, the size and
appearance of a lizard. Small head but with more powerful jaws than amphibians: they
were probably insectivorous.
The amniotes complete the process of terrestrialization started by the amphibians, they
are independent from the water even during the reproductive period:
-internal fertilization.
-amniote egg: semi-permeable shell that allows certain gases, but not water, to
pass.
-The development of the embryo is completed inside the egg.

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Historically, amniotes were divided into three groups based on an anatomical feature of
the skull: the temporal fenestrae (cranial openings; their origin is attributed to a
tendency to reduce the weight of the skull, although it has also been proposed that they
could provide sites for muscle insertion):
-anapsids: no temporal fenestrae. It is the plesiomorphic condition in amniotes, but can
also occur secondarily (as seems to be the case in turtles).
-synapsids: one temporal fenestra; the mammal lineage
-diapsids: lineage that gave rise to modern lizards, snakes and crocodiles, and also to
birds.
The first amniotes are anapsids, but in the upper Carboniferous there are already
representatives of diapsids and synapsids.

First diapsid: small reptile (40 cm), slender, with a small head and neck and long legs.
Agile insectivores similar to modern lizards. Thus the ancestor of snakes, lizards,
crocodiles and birds resembled a small lizard.

First synapsid: Large reptile, up to 3 m. in length. Very large skull and massive limb
bones. It was a carnivore that would probably feed on fish and tetrapods rather than
insects. The first synapsids are known as pelycosaurs.

BIRDS:
Some characters of birds already present in their latest common ancestor:
-bipedalism, feathers, nesting behaviour, incubation of eggs (brooding behaviour),
homeothermy, flight.

Archaeopteryx has often been called "the first bird". There are 10 known specimens of
Archaeopteryx (9 skeletons and a loose feather). All from the lithographic limestones of
Solnhofen (Bavaria, Germany). The feather was found in 1860 and the first skeleton in
1861 (two years after Darwin published "The Origin of the Species"). It is the size of a
magpie. It shows intermediate traits between reptiles and birds. Among the reptilian
characters are the small, sharp, spaced teeth, a long and straight tail with 22 or 23 caudal
vertebrae and claws at the end of fingers. Archaeopteryx lived in the Upper Jurassic.
There has been much discussion as to whether or not Archaeopteryx could fly, and to
which degree; the question continues being controversial.
The crown group "Birds" begins with the latest common ancestor of all modern birds;
Archaeopteryx is placed below, in the stem group birds.
However (perhaps because of tradition) there are scientists who prefer to give the name
"Birds" to the clade that begins with Archaeopteryx, and have coined the name
"Neornithes" for crown group birds.

During the 19th and 20th centuries there was much speculation about the origin of birds,
and they were derived from different groups of reptiles. In recent years a great deal of
evidence has accumulated that tells us without a doubt that birds are descended from a
group of dinosaurs, the maniraptorans.

The first known dinosaurs date from the middle/upper Triassic. They are biped
carnivores of small to moderate size. One of them is Herrerasaurus (possible example
of a basal dinosaur, but its phylogenetic position is controversial: basal dinosaur, basal
saurischian or even basal theropod). It was biped, carnivorous, 3-4 m long. It comes
from the Ischigualasto Formation, Argentina.

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Traditionally dinosaurs have been divided into two large groups, the
ORNITHISCHIANS (bird-hipped) and the SAURISCHIANS (lizard-hipped). The
plesiomorphic condition for dinosaurs is a saurischian pelvis, which is also present in
basal archosaurs.

ORNITHISCHIANS: group of herbivorous dinosaurs: includes the hadrosaurs or duck-

billed dinosaurs, Triceratops and Protoceratops, and the armoured quadruped
dinosaurs: ankylosaurs, stegosaurs.

SAURISCHIANS: includes the sauropods (large quadruped herbivores) and the

theropods (agile carnivores). Among the theropods are large dinosaurs such as
Tyrannosaurus, small carnivores such as Compsognathus and the maniraptorans.

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ANATOMICAL EVIDENCE of skeletons indicates that birds descend from a group of

theropods known as maniraptorans → small and medium-sized biped predators that
used their forelimbs to hunt (instead of using the jaws like Tyrannosaurus).

BEHAVIOURAL EVIDENCE → Nesting behaviour. Crocodiles build nests that they

protect with plant material, and take care of → perhaps this behaviour already existed in
the common ancestor of birds and crocodiles.
There is evidence that some dinosaurs had a nesting behaviour similar to that of modern
birds. There is evidence of nesting colonies in duck-billed dinosaurs (hadrosaurs) where
the little ones remained in the nest after hatching. In maniraptorans → there is evidence
they sat on their eggs exactly as modern birds do, suggesting incubation of eggs
(brooding behaviour).

PLUMAS: since the 1990s, a number of fossil maniraptorans (and other more basal
theropods such as compsognathids) with feathers have been described. These feathered
dinosaurs are from the early Cretaceous of China. Some have fine feathers, much like
down, while others have vaned feathers very similar to those of modern birds. This is
yet another argument that birds descended from maniraptoran dinosaurs, and it also
proves that feathers evolved before flight.
It is also known, both from fossil evidence and from biological data (ontogenetic
development of feathers), that the first feathers were down-like and that vened feather
occur later. The original function of the feathers is not known. Several functions have
been suggested (thermal insulators, communication and display, water insulators).

Pterosaurs also had feather-like structures covering their bodies, which complicates the
scenario of feather evolution. If these structures are homologous to the feathers of
theropods then it is possible that feathers evolved at the base of the birds' stem group
and that either all dinosaurs had them or that some had lost them secondarily. Perhaps
the same could be said about homeothermy.

Acquisition of avian characters:

(1) Bipedalism and nesting behaviour, (2???) homeothermy, feathers and incubation of
eggs, (3) flight.

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Bipedalism → the last common ancestor of dinosaurs was already biped. It seems that
many archosaurs were also biped, or facultative biped.
Homeothermy → it is not clear when this character evolved but it is likely that the small
active theropods were homeotherms and of course the maniraptorans.
Feathers → In the group of theropods? It is possible, however, that feather precursors,
in the form of down-like structures, evolved at the base of the bird stem group.
Flight → flight arose at the end of the bird stem group, before the common ancestor of
modern birds.

SYNAPSIDS → TOWARDS THE MAMMALS.

The oldest crown group mammals date from the middle Jurassic, but the first synapsids
appeared at the end of the Carboniferous. Therefore the "stem group" of mammals is
very long (more than 100,000 Ma).

The fossil record of synapsids is relatively complete, and documents the gradual
evolution from the first synapsids (pelycosaurs) to the mammals. The synapsids
experienced a series of consecutive radiations, and each one brings more and more
mammal innovations.
The fossil record documents the gradual transition from organisms with ectothermic
metabolism to fully homeothermic mammals. Synapsids form the mammalian stem and
crown group. The mammal stem group consists of pelycosaurs, therapsids and, in the
uppermost part, animals that are already very similar to modern mammals and that have
been grouped under the term "mammaliaformes" (all three groups are paraphyletic).
Traditionally, pelycosaurs and therapsids are known as "mammal-like reptiles".

At the basal end of the stem group:

-pelycosaurs: mostly large amniotes. Many are characterised by the presence of a sail →
dorsal crest supported by vertebral extensions → It seems that the sail (structure with a
large surface and a small volume) could have a thermoregulatory function (it would
have a highly vascularised skin). Pelycosaurs were supposed to have been ectothermic
animals, and the possession of a sail could have been important for such large animals,
with so much volume to heat and so little surface to capture heat. Pelycosaurs originated
at the end of the Carboniferous. The first pelycosaurs were large amniotes, up to 3 m.
long, with very large skull and massive limb bones. They were carnivores that probably
feed on fish and tetrapods. Pelycosaurs were the dominant and most diverse tetrapods
during the lower Permian, there were herbivores and carnivores. Pelycosaurs gave rise
to therapsids.

At the other extreme, "mammaliaformes" and "crown group" mammals:

endothermic animals with a high metabolism. They have hair, have large brains, feed
their young with milk produced by mammary glands and the care of the young is
prolonged.

Therapsids, the transition:

The first therapsids appear in the Middle Permian and continue until the Triassic. They
become the dominant tetrapods during the late Permian and constitute a diverse
association of herbivores and carnivores:

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Three classic changes of the pelycosaur/mammal transition (they are not the only ones,
of course, but they are very significant).

1- MASTICATION:
Pelycosaurs had a dentition adapted for capturing and holding the prey, but not to
triturate it into small pieces.
Mammals are capable of tearing apart and triturate food into small pieces: this facilitates
the action of digestive enzymes by giving them more surface on which to act →
accelerates the obtaining of energy from the ingestion of food. This change implies:
a- specialized dentition
b- changes in the jaw articulation and its musculature allow chewing: mammals
can move their lower jaw from front to back and from one side to the other →
these movements are necessary for mammal mastication.
c- occlusion of teeth: teeth fit perfectly (this feature is related to the fact that
mammals have a reduced number of tooth replacements instead of replacing
teeth as they wear out).

2- RESPIRATION
In pelycosaurs, nostrils opened into the oral cavity, so the air has to pass through it
before reaching the lungs. When food blocked the passage of air, the animal had to stop
breathing. Mammal nostrils do not open into the oral cavity; the air passage is separated
from the oral cavity by a bony plate, the secondary palate. Breathing does not stop
during feeding.

3- HEARING:
The change of the jaw articulation is linked to the development of the bones of the
middle ear and the development of hearing.
The bones that form the mandibular articulation in reptiles are different from
those in mammals. The change of articulation is documented in the fossil record and
went through a stage in which there were two points of articulation (a case of
redundancy in evolution).
All modern-day amniotes, except mammals, have only one small bone in the middle
ear: the stapes or stirrup, while mammals have three: stapes (stirrup), malleus (hammer)
and incus (anvil).
The two bones involved in the pelycosaurian mandibular articulation (quadrate
and articular) became the two extra bones of the mammalian middle ear, and one of the
largest bones of the pelycosaurian lower mandible (angular) became the ectotympanic
bone (the eardrum is held by this curved bone located behind the modern mandibular
articulation).

EVIDENCE OF HAIR AND MAMMARY GLANDS IN THE STEM GROUP OF

MAMMALS:
some Triassic therapsids have small pores in the snout skeleton very similar to those
that represent the passages for the nerves and blood vessels that go towards the sensitive
whiskers of modern mammals. If these advanced therapsids had whiskers, they must
also have had hair. This evidence relates to the presence of mammary glands: mammary
glands are thought to have their origin in modified sweat glands and heat-dissipating
sweat glands are functionally related to the presence of hair that maintains heat.

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THE FIRST MAMMALS:
Mammals are the clade that begins with the last common ancestor of modern
monotremes (echidna and platypus), marsupials and placentals. The immediate
ancestors of "crown group" mammals (the "mammaliaformes"), are animals very similar
to modern mammals:
-Agile insectivores of small size, perhaps similar to a weasel. Locomotion entirely
mammal-like.
-They were homeotherms and had hair and mammary glands, but a monotreme
reproductive physiology.
-They still have two mandibular articulations, the old pelycosaur and the new mammal
articulations.

HOMININ EVOLUTION
Fossil hominins are generally rare and often very fragmentary, making it difficult to
accurately assess the number of different species that existed (as well as their
intraspecific variety ranges, etc.). Only a few species of hominins, such as A. afarensis,
are well known.

The first hominin (stem + crown group Homo), dates from around 4.5 Ma and was
found in Ethiopia (East Africa) in 1992. The fossil was identified as a new species of
Australopithecus, Australopithecus ramidus (although it was later assigned to the new
genus Ardipithecus). The remains are incomplete but suggest that it was an animal of
about 40 kg, with a posture more upright than that of chimpanzees. The teeth suggest a
diet similar to that of the chimpanzees and its burial context suggests that it lived in the
forest. Neither the size of its brain nor its mode of locomotion are known.
The following fossil hominins (from approx. 4 Ma) belong to Australopithecus
afarensis and have also been found in East Africa. There is plenty of paleontological
data on A. afarensis, which is a much better known species.
A. afarensis was a species with a high sexual dimorphism, with females of around 25 kg
and males of up to 50 kg. This range is quite high for such a small animal which has led
scientists to question whether the remains of A. afarensis could actually include more
than one species. The cranial volume of A. afarensis is similar to that of chimpanzees.
A. afarensis was bipedal and between 1 and 1.5 m high. Habitat reconstructions of A.
afarensis suggest that they lived in "open forest" areas.

Among the Australopithecines there are species with a more robust morphology
(generally grouped in the genus Paranthropus), with more massive skulls and,
particularly, lower jaws, and more developed molars. They were hominids adapted to a
diet of tougher vegetation.

The first Homo (Homo habilis) have a significantly larger brain than the
Australopithecines. Some fossils of H. habilis have been found associated with stone
tools and are considered the first hominids to work the stone to produce recognizable
tools.