The origin of tetrapods and movement onto land

• The term "tetrapod" (meaning four-limbed or four-

footed) has historically been applied to the land
vertebrates (amphibians, reptiles, dinosaurs, birds,
and mammals).
• but this group includes many animals that don't have
four feet.
• All these animals are tetrapods because
they are descendents of the first animal with four
legs.
 The origin of tetrapods and movement onto land
• There are a suite of challenges associated with
terrestrial living that the first tetrapods had to
overcome.

• These included the fact that water supports a fish’s

body but
air does not,
so stronger skeletal structures were needed
to support the body and limbs to allow movement.
 Cont…
• In addition, gills collapse out of water so
an alternative breathing system (i.e. lungs) was
needed.

• In addition, there was a constant threat of water loss

from the skin or through eggs,
which limited the first tetrapods to a close
connection with water
until hard-shelled amniotic eggs evolved.
Transition from aquatic to terrestrial through the evolution of:
• Limbs instead of fins,
• Lungs instead of gills,
• Kidney advancement,
• Axial skeleton, for support and movement
• Cornified skin – to resist drying,
• Modified eye, ear and nose,
• Oral glands to moisten food,
• Eggs and delicate larva – egg with shell and
fetal membranes.
The origin of tetrapods and their fish ancestor

• The first tetrapods were Acanthostega and

Ichthyostega, and were closely related to a
group of fish known as lobe-finned fish e.g.
Eusthenopteron.

• Eusthenopteron had a number of expatiation

that pre-adapted it to life on land:
 Cont…

it had limbs (with digits) that allowed it to move

around on the bottom of pools,
lungs - which meant it could gulp air at the
surface, and
the beginnings of a neck.

• This last is important as a terrestrial predator

• cannot rely on water current to bring food into
its mouth,
• but must move its head to catch prey.
 Cont…
• And the bones in Eusthenopteron's fins are
almost identical to those in the limbs of the
earliest amphibians, an example of homology

Ichthyostega's skull was almost identical to that of

the lobe-finned fish Eusthenopteron,
a definite neck separated its body from its head,
and
it retained a deep tail with fins.
Cont…
Figure 25.01a

17.1a Eusthenopteron
 Panderichthys
• The next fossils are a series of more tetrapod-like
lobe-finned fish including
Panderichthys from the late Devonian.

• Panderichthys is a “fishapod” intermediate between

fish and tetrapods with
a tetrapod-like skull and body,
braincase, and lungs,
but still retaining true fins.
Panderichthys
 Tiktaalik
• The gap between Panderichthys and the early tetrapods
Acanthostega and Ichthyostega has recently been neatly bridged
by the discovery of a classic intermediate form, Tiktaalik
roseae.

• Discovered in 2004 on Ellesmere Island in the Canadian Arctic

Tiktaalik roseae is an extremely important fossil link in the
origin of tetrapods.

• Another “fishapod” Tiktaalik like Panderichthys has a mixture of

fish and tetrapod characteristics, but has several tetrapod
characteristics than Panderichthys lacks.
 Tiktaalik roseae
Unlike a fish however it has a mobile neck and
the ear is structured so that it can hear both in and
out of water.
• Most strikingly, in addition to the other tetrapod limb bones
Tiktaalik has a wrist.
• Tiktaalik’s elbow could bend like ours and
• the wrist could bend too, which allowed the animal to
make its “palm” lie flat.
Tiktaalik roseae 375 mya
Brief Evolutionary history of amphibians

• Amphibian is derived from the Ancient Greek term

amphíbios which means both kinds of life, amphi
meaning “both” and bio meaning life.

• The term was used to refer to animals that live both

in the water and on land.
 General characteristics of modern
amphibians
• It include frog, toads, salamanders & worm like
caecilians.

• Skeleton mostly bony with varying number of vertebrate;

• Ribs present in some (e.g Apoda), absent or fused to

vertebrae in other;

• Notochord does not persist;

• Exoskeleton absent
 Cont..
• Body forms vary greatly: from an elongated trunk with distinct
head, neck and tail to a compact depressed body with fused head
and trunk and no intervening neck.

• Limbs usually four (tetrapod- although some are legless, (e.g

caecilians).

• Forelimbs of much smaller than hind limbs (frogs),

• in others all limbs small & inadequate (e.g in some salamanders).

• Webbed feet often present, no true nails or claws.

• Forelimb is usually with four digits but sometimes five, which is

vestigial and in male its function for reproduction and sometimes
fewer.
 Cont…
• Skin smooth and moist with many glands;

• Some of which may be poison glands to avoid their enemy;

• Pigment cells (chromatophores) are common;

• Have no scales, except concealed dermal ones in some e.g.

caecilians (Apoda).

• Mouth usually large with small teeth in upper or both

jaws; two nostrils open into anterior part of mouth cavity;
also they have well developed two eyes, two ear
 cont…
• Respiration by lungs (absent in some salamander),

• Skin and gills in some, either separately or in combination;

• External gills in larval forms & may persist throughout life in

some.

• Heart with a sinus venousus, two atria, one ventricle, a conus

arteriosus and a double circulation through the heart;

• Skin abundantly supplied with blood vessels.

• They are exothermal

 cont…
• They have excretory system of paired mesonephric
kidneys

• Urea is main nitrogenous waste products

• They have ten pairs of cranial nerves

• Separate sexes; Fertilization mostly internal via

spermatophore in salamanders & caecilians;

• mostly external in frogs & toads;

• predominantly oviparous, some ovoviviparous or

viviparous; Metamorphosis usually present.
Classification of living amphibians
(Lissamphibia)
The amphibians are represented by about 4800
hundred species divided into three orders.

Urodela: “tailed ones”: salamanders

Anura: “tailless ones” frogs, toads.

Gymnophiona (“naked snake”; previously

Apoda “legless ones”) caecilians.
Eastern Mud salamander
Caecilian
Red-eyed Tree frog
 1. Order Gymnophiona (Apoda)
The order Apodas contains approximately 160 species of elongated,

limbless,

burrowing creatures commonly called caecilians (like earthworms).

There is small scale in the skin of some;

many vertebrae,

long ribs (why?)& terminal anus.

Eyes are small, and most species are totally blind as adults (because
the eyes covered with skin or bone).
 Order Gymnophiona

• Special sensory protrusible tentacles occur on their snout

(probably helped to find their food).

• Fertilization is internal and males have a protrusible

copulatory organ.

• Viviparity also is common in some, with embryos

obtaining nourishment by eating the wall of the oviduct.

• Tail very short or absent.

• They have 5 families, 33 genera, and 160 species.

Yellow-striped Caecilian
Caecilian: note prominent annuli.
 2. Order Urodela (Caudata), e.g
salamanders
Order caudata consists of tailed amphibians
with head and trunk.

They have no scales;

usually two pairs of equal limbs.

They are carnivores both as larvae & adults.

 Order Urodela
• Some species of order caudata practice internal fertilization;

• very little deference between the larvae & adult;

• the adult is viviparous.

• They show paedomorphosis (the retention of ancestral

juvenile characteristics by descendant adult).

• These can include a lack of eyelids, a lateral line system,

larval tooth and bone patterns and, in some instances,
external gills.
 Order Urodela
• Some characteristics of ancestral adult morphology
consequently eliminated.

• This condition is paedomorphosis; larva became sexually

matured without assuming adult appearance.

• They have 10 families, 61 genera, 500 species.

• Several groups of salamanders have become specialized

for living in caves.
Japanese Giant Salamander
Texas Blind Salamander
3. Order Anura (Salientia), e.g forgs
& toads
The frogs and toads are the largest group of
amphibians
They have no neck region, head & trunk fused;
no tail, no scales.
They have large mouth & lungs;

smooth and moist skin (e.g frog).

They are cosmopolitan, predominantly tropical;
some tropical species live on trees.
 Order Anura
• They are specialized for jumping

• All are carnivores.

• The larva is tadpole.

• They show metamorphosis.

• Toads have dry skin, short legs and well adapted to

dry environment.

• They have 29 families, 352 genera, approximately

4840 species
 Leaping Frog

Woodhouse Toad
 Arboreal frogs
• There are many species of frogs specialized for
climbing in trees and shrubs.

• These are usually slim and long legged with large

heads and eyes.

• Some of the most specialized arboreal species are

called tree frogs and possess specially modified toe
pads that allow them to adhere to surfaces.
Common Tree Frog
White-lipped Tree frog: note toe pads
 Difference between frog and toad
• The frog has relatively thin, moist and smooth skin.

• It acts as a respiratory organ and has many mucous glands.

• They are mostly aquatic and hardly come on land.

• While the skin of toad is much thicker, dry and rough.

• It has no role to play in respiration and has very few

mucus glands.

• They are mostly terrestrial and come to water for

copulation.
 Difference between frog and toad
• Frogs are almost invariably associated with aquatic habitats;

• toads are found much more in every habitat, due to in part

their integument.

• The frog has a triangular head with maxillary teeth (upper

jaw);

• toad not have

• Frogs usually diurnal and lay their eggs in clusters; toads

usually nocturnal and lay eggs in string (lines).
Morphology and Physiology of Amphibians
1. Skin and Coloration of Amphibia
• Amphibians have amazing skin.

• They can drink water and also breathe through their

skin.

• The skin is composed of two layers:

• An outer stratified epidermis which is molt and

contains deposits of keratin.
 Skin and Coloration of Amphibia
• Terrestrial; amphibians have heavy deposits of keratin; e.g. toads

• The inner layer of epidermis gives rise to two types of

integumentary glands that grow in to the loose dermal tissues
below.

1. Small mucous glands secrete protective mucous water proofing

on to the skin surface

2. Large granular serous glands produce a whitish watery poison

highly irritating to would be predators.
Poison Arrow Frog
The skin of some amphibians, such as this poison arrow frog, contains poisons that help keep
predators from eating the amphibian.
Green and Black Poison Dart Frog
Skin and Coloration of Amphibia
Skin color in amphibians is produced by special pigment cell
chromatophores, located mainly in the dermis. Most
amphibians have three types of chromatophores.

-Uppermost- Xanthophores containing yellow, orange or red

pigment

-Beneath- iridophores containing silvery, light reflecting

pigment

-Lower most- melanophores containing black or brown melanin

Skin and Coloration of Amphibia
• There is no green pigment, the green color result
from a combined effect of light reflected from
underlying iridophores and the yellow pigment
through which the light passes.

• The color change brought about primarily by the

hormones intermedin, produced by pituitary gland.

• Light is the main stimulus for color change.

 Skin and Coloration of Amphibia
• Intermedin causes a dispersion of pigment in the
chromatophores.

• Another cause is internal and external condition. E.g.

Temperature (To), When To increases the pigment
concentrated, skin change to light color, while To decreases
black color obtained due to expansion of pigment.

• Nervous system takes in part on color change; this color

change important for camouflages to void enemies.
 Medical applications of skin secretions
• There have been a variety of investigations into the
medical benefits of various amphibian skin secretions.

• For example a powerful painkiller called epibatidine

has been isolated from a poison dart frog.

• a bacteria-killing antibiotic peptide called magainin

has been isolated from the skin of the African clawed
frog.
 2. Skeletal and Muscular systems
• They have well development endoskeleton of bone
and cartilage provides a frame work for muscles.

• They have one pair of forelimbs, which are short,

and important to landing on ground, hand with four
digits, the 5th digit is vestigial and its function in
male for reproduction.
 Skeletal and Muscular systems
• They have two or a pair of hind limbs, which are
elaborate, long and strong and specialized for
jumping and swimming.

• Skeletal system modified for land life.

• Common frogs have only 9 trunk vertebrae and a rod

like urostyle, which represents several fused caudal
vertebrae (coccyx).
 Skeletal and Muscular systems
• The skull of amphibians much lighter in weight, flattened
in profile, has fewer bones and less ossification.

• They have immovable upper jaws and movable lower

jaws.

• The muscular system of amphibians well developed and

organized, spindle shaped and their muscle attached to
the bone by Tendon.
 3. Feeding and Digestion
• Most adult amphibians are carnivorous; they feed on
insects, worms, and mollusks; usually capture of their
prey by the specialized extensile or protrusible tongue,
which produces sticky secretion.

• Most amphibians have small, numerous, identical teeth

(homodont dentition).

• The function of the teeth is to grasp and hold prey not to

chew it.
Hydromantes salamander shooting its tongue to catch a housefly
 Feeding and Digestion
• Stomachs of amphibians are simple, with enzyme,
and ends with a valve and

• their intestines are short and produces a variety of

enzymes for digesting of proteins, carbohydrates and
fats.
 4. Respiration and vocalization
• Amphibians use three respiratory surfaces for gas
exchange in water and air.

• These are: Skin (cutaneous breathing) - on land and

swimming, mouth (buccal breathing), lungs- for
terrestrial group.

• They use also gills - in aquatic and larval stage.

• Frogs and toads are more dependent on lung

breathing than are salamanders.
• To enhance gas exchange the skin in many species
(e.g. Eastern hellbenders) is highly folded and heavily
vascularized.
 Gills
• Larval amphibians breathe using external gills.

• In anuran tadpoles the gills are concealed behind a flap of

tissue and water flows through the mouth across the gills and
out of a spiracle.

• Most species lose their gills when they mature but some
retain them into adulthood (neotony).

• For example, the aquatic salamanders Necturus and

Crytobranchus possess both gills and lungs. These species
usually air breathe only when oxygen levels in the water are
low
In larval salamanders and caecilians the gills are exposed and project
from the sides of the head.

Larval Tiger Salamander

 Respiration and vocalization
• Both male and female frogs have vocal cords but these of
males are much better developed.

• Vocal cords located in the larynx or voice box.

• A frog produces sound by passing air back and forth

over the vocal cords between lungs and large pair of sacs
(vocal pouches) in the floor of mouth.

• Male use this voice to attract females.

 5. Circulation
• The principle changes in circulation involve the shift from
gill to lung breathing.

• They have double circulation consisting of separate

pulmonary and systemic circulation;

• Double circulation" has an interior circuit within the

heart--blood enters the heart, leaves the heart and gets
oxygenated, enters the heart again, and then gets pumped
out to the body.
 Circulation
• Because "Double circulation" allows oxygenated blood to
be pumped back into the heart before going out to the
body, it pumps blood with much more pressure and much
more vigorously than "single circulation".

• Amphibians have a three-chambered heart, with two

separate atria and a single undivided ventricle.
 Circulation
• The right and left atria contract asynchronously so that
although the ventricle is undivided, blood remains mostly
separated when it enters this chamber.

• When the ventricle contracts, oxygenated pulmonary

blood enters the systemic circulation and deoxygenated
systemic blood enters the pulmonary circulation.
 Circulation

• This separation is aided by a spiral valve, which divides

the systemic and pulmonary flows in the conus arteriosus
and

• by different blood pressure in the pulmonary and systemic

blood vessels leaving the conus arteriosus.
Amphibian circulatory system
 6. Nervous systems and special senses
• The frog has a highly developed nervous system.

• It consists of a brain, a spinal cord, and nerves.

• Amphibians have three fundamental parts of the brain;

• Forebrain (telencephalon), which is concerned with the

sense of smell, that contains olfactory center important
for detection of dilute air borne odor son land.
 Nervous systems …
• The sense of smell is one of the dominant special senses
in frogs.

• Midbrain (mesencephalon) is concerned with vision.

• Vision is the dominant special sense in many amphibians;

adjusting focus for near and distant objects is
accomplished by moving the lens.

• A retina contains both rods and cones, the latter

providing frogs with color vision.

• Frogs and toads generally possess good vision.

 Nervous systems …
• Hind brain (rhombenaephalon), which is concerned with
hearing and balance;

• a middle ear closed externally by a large tympanic

membrane (ear drum) and containing a columella
(stapes) that transmits vibrations to the inner ear.

• Other sensory receptors are tactile and chemical

receptors in skin, taste buds on the tongue.
 7. Reproduction/Breeding habit
• They are ectothermic; they breed, feed and grow only
during warm seasons.

• One of the first drives after the dormant period is

breeding.

• Most of them are oviparous.

• Migration of frogs and toads depends up on their breeding

habits.

• Eggs of most salamanders are fertilized internally;

 Reproduction
• Amphibian eggs do not have a hard shell and dry out
quickly if not kept in a moist environment.

• Many species lay their eggs directly in water or on the

undersides of leaves over water so the larvae fall in when
they hatch.

• Others show more parental care and brood eggs in

(depending on the species) the mouth, stomach or pressed
into soft skin on their backs.
 A female, American pygmy marsupial frog, Flectonotus
pygmaeus, carries developing larvae in a dorsal pouch.

• Female Surinam frog carries eggs embedded in

specialized brooding pouches on the dorsum.

• Male poison arrow frog, Phyllobates bicolor carries

tadpoles adhering to its back.

• Tadpoles of male Darwin frog, Rhinoderma darwins

develop in to frog lets in its vocal pouch.

• One species of Australian frog even brood it’s young in

the stomach while discontinuing digestion.
Figure 25.04
• What is the difference between Male
and Female Frogs?
• • Males are smaller than females of the
same species.
• • Males are more vocal than females,
especially during the mating season.
• • The female cloaca is more visible to the
exterior than the male cloaca.
• • Males are usually brighter than females
in colourations.
 Reproduction/Breeding habit
• In most anurans fertilization is external.

• When frogs eggs are matured females enter water and are
clasped by male in a process called amplexus in which
eggs are fertilized externally.

• As female lays eggs, a male discharges sperm over the

eggs to fertilize them; a fertilized egg (zygote) begins
development almost immediately.
 Reproduction/Breeding habit
• By repeat division (cleavage) an egg is converted in to a hollow ball
of cell.

• The blastula undergoes gastrulation and then continues to differentiate

to form an embryo with a tail bud.

• At 2 to 21 days, depending on To, a tadpole hatches from the

protective jelly coats.

• In caecilians fertilization is internal.

• However, the majority of caecilians are viviparous (about 75%) and

matrotrophic (young obtain nutrition from the mother).

• Most terrestrial species have direct developments.

Figure 25.26
Figure 25.05

Many caecilians lay their eggs in the mud near water,

but some caecilians brood their eggs in burrows.
Caecilian with young