Amphibia

Paper Bios (Z): 301 / Unit – III

Origin of tetrapods,
Parental care,
Paedomorphosis
Position of Tetrapoda

Dr. J. K. Barot 2
Origin and Ancestory of Amphibia (Tetrapoda)
• Looking into the evolutionary aspect
– Earliest Amphibia.
– Piscine ancestry.
– Devonian fishes.
– Crossopterygian ancestry.
– Possible polyphyletic origin.
– Causes for transition from water to land.
– Adaptations to amphibious life.

Dr. J. K. Barot 3
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 Earliest Amphibia…
• The earliest fossils of
Amphibia (Tetrapoda)
are known as
Labyrinthodontia
because of the folded
nature of dentine of
their teeth.
• They probably originated
300 million years ago,
during Devonian period
and flourished through
Carboniferous and
Permian periods before
extinction in Triassic.
Dr. J. K. Barot 5
 Piscine ancestry…
• It is universally accepted that amphibians have originated
from fishes.
• Resemblance of Amphibia to fish is seen in most systems
of the body.
• Both are cold-blooded.
• Both respire by gills early in life, in some cases throughout
life.
• Both have air bladders serving as lungs.
• Like fishes, aquatic amphibians have a canal system.
• To prevent desiccation in air, both usually lay eggs in water
where larval development usually continues.
Dr. J. K. Barot 6
 Devonian fishes…
• Three major groups of advanced bony fishes were contemporaries of
labyrinthodontia during Devonian period— Actinopterygii, Dipnoi
and Crossopterygii.
• Which of these groups gave rise to the first amphibians ?
• Actinopterygii are far off the main course of tetrapod evolution
because of many specializations.
• Dipnoi have several features of close resemblance to Amphibia in
their anatomy and embryology.
• But they have also been too specialized to be considered as ancestral
to Amphibia. Particularly, the skeleton of their fins is not reminiscent
of that of a primitive tetrapod.
• Thus, the older view that Dipnoi, gave rise to amphibians is no longer
stand.
• Their similarity probably resulted due to convergent evolution.
• On the other hand, it is now believed
Dr. J. K. Barot
that the earliest amphibians 7
(labyrinthodonts) arose from primitive crossopterygian fishes.
 Crossopterygian ancestry…
• Origin of Amphibia can be traced back to the
freshwater carnivorous rhipidistian
crossopterygians of Devonian period,
represented by the genera Osteolepis and
Eusthenopteron.
• They show extensive morphological
similarities with early amphibians or
labyrinthodonts, which are evidence of their
close phylogenetic relationship.

Dr. J. K. Barot 8
Dr. J. K. Barot 9
(1) Pentadactyl limb of amphibians is supposed to have
arisen from the crossopterygian fin lobe. There are clear
homologies between the limb bones of ostelepid
crossopterygians and early tetrapods (labyrinthodonts).
(2) Their skulls and jaws are similar and can be equated bone
by bone.
(3) Both rhipidistians and early tetrapods were predators,
armed with sharp, strong teeth, with a peculiar
labyrinthine infolding of the enamel.
(4) Position of external and internal nares is similar.
(5) Air bladders were used as lungs during short migrations
from pond to pond.
(6) Ichthyostega, a primitive labyrinthodont fossil found in
Green land, has both crossopterygian and amphibian
characteristics and represents a transitional stage.
Dr. J. K. Barot 10
Dr. J. K. Barot 11
Possible polyphyletic origin…
• The three orders of living Amphibia
-Apoda,
-Urodela and
-Anura—are so different as to suggest a polyphyletic origin of
amphibia.
• Eric Jarvik supports a diphyletic origin.
• He suggests origin of Urodela and Apoda from porolepid
rhipidistians through lepospondyls (a group of small
Carboniferous and Permian forms), while origin of Anura from
osteolepid rhipidistians through labyrinthodonts.
• His views are NOT accepted by all.
• Another view traces the ancestry of all 3 orders from
lepospondyls.
Dr. J. K. Barot 12
Causes for transition from water to land…
• What was the need for early crossopterygians to leave their
aquatic habitat to come on land with unknown possibilities?
• According to one view, Devonian was a period of seasonal
droughts.
• Certain rhipidistian crossopterygians had sufficiently developed
lobe fins to move over land to ponds containing water.
• Periodic escapes resulted in conversion of their lobe fins into
pentadactyl tetrapod limbs.
• Another view contrasts between aquatic and terrestrial habitats
during Devonian. Compared to water, land had more 02 in air,
food, cover, shelter and breeding places.
• On the other hand, land had less impurities and less predators.

Dr. J. K. Barot 13
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 Adaptations to amphibious life…
• The transition from water to land was gradual.
• Amphibia, being the lowest and earliest
tetrapods are not fully adapted to a terrestrial
life like reptiles.
• Transition from water to land involved
important changes in almost every system of
body.
• Some striking modifications acquired by
amphibious Amphibia over the ancestral
aquatic fishes may be summarized as follows :
Dr. J. K. Barot 16
1. Locomotion. Streamlined body retained for
swimming. Limbs developed in place of paired fins.
2. Skeleton. Vertebral column becomes more rigid.
Pectoral girdle lost connection with skull. Pelvic
girdle attached to vertebral column and two halves
firmly united.
3. Muscles. Stronger muscles to lift body above
ground.
4. Respiration. Gills replaced by lungs for gaseous
exchange in air. Cutaneous respiration supplements
pulmonary respiration.
5. Circulatory system. Changes occurred to provide for
respiration by lungs and skin.
Dr. J. K. Barot 17
6. Eggs and development. Eggs small and still
laid in water. Developmental stages fish-like.
7. Sensory system. Addition of middle ear
containing a bone (columella auris) for
transmitting sound vibrations from tympanum
to inner ear.
8. Skin. Skin transformed to support terrestrial
life to resist desiccation.

Dr. J. K. Barot 18
 Parental care in amphibia…
• Looking after the eggs or the young until they are independent,
to defend from predators, is known as parental care.
• It is a very important factor for survival.
• Animals exhibit a great diversity in caring for their eggs and young
during their development.
• Perhaps, no group shows more diversity than to the amphibians,
especially those living in tropical regions.
• Anurans show much greater diversity than urodeles and apodans.
• The methods of caring by Amphibians generally fall under two
broad categories :
– (i) Protection by nests, nurseries or shelters and
– (ii) Direct caring by parents

Dr. J. K. Barot 19
[1] Protection by nests, nurseries or shelters
– 1. Selection of site.
– 2. Defending eggs or territories.
– 3. Direct development
– 4. Foam nests.
– 5. Mud nests.
– 6. Tree nests.
– 7. Gelatinous bags.

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Dr. J. K. Barot 21
 1. Selection of site.
• Many amphibians lay eggs in protected, moist microhabitats on land.
• Many tropical frogs and toads lay eggs on land near water.
• Many tree frogs lay their eggs not on land but on leaves and
branches overhanging water.
• Species of Phyllomedusa, Rhachophorus, Hylodes, etc. glue their
eggs to foliage hanging over water.
• Rhacophorus malabaricus in India and Chiromantis of Africa also
deposit their spawn on trees.
• Many tree frogs deposit eggs in water that accumulates in epiphytic
tropical plants.
• Free from reach of aquatic egg predators in their microhabitats, the
tadpoles on hatching drop into water beneath to complete the it
metamorphosis.
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2. Defending eggs or territories.
• Males of green frog Rana clamitans and other
species maintain territories and attack small
intruders to defend eggs.
• Male or female even guards the eggs.
• In Mantophryne robusta, the male actually sits over
and holds with hands the elastic gelatinous
envelope containing eggs numbering 17.
• Some tree frogs laying eggs above water may sit
beside the eggs or rest on top of them. Removal of
guarding frogs may result in desiccation or death of
eggs.
Dr. J. K. Barot 25
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3. Direct development
• In some terrestrial or tree
frogs, such as
Eleutherodactylus,
Arthmleptis, Hylodes and
Hyla nebulosa, the eggs
hatch directly into little
frogs thus avoiding larval
mortality.
• In the red backed
salamander, Plethodon
cinemas, the hatchlings
are miniatures of the
adults.
Dr. J. K. Barot 27
 4. Foam nests.
• Many amphibians convert copious
mucous secretions into nests for
their young.
• In the Japanes tree frog,
Rhacophorus schlegeli, the
mating couple digs a hole or
tunnel into which eggs are left in a
frothy mass to avoid desiccation.
• During rains, hatching tadpoles
are washed down the slopping
tunnel into pond or river water for
further development.
Dr. J. K. Barot 28
• The female of frog,
Leptodactylus mystacinus, stirs
up a frothy mass of mucus, fills
it in holes near water and lays
eggs in them.
• The tadpoles developing in
these improvised nests can
readily enter water.
• Some anurans lay eggs in nests
of foam floating on water.
• The female emits huge mucus
that she beats into a foam with
her hind-legs to lay eggs.
• When tadpoles hatch they drop
from foam into water.
Dr. J. K. Barot 29
 5. Mud nests.
• In Brazilian tree frog, Hyla
fabre, the male digs a little
crater-like hole or nursery in
mud in shallow water, in
which the female lays her
eggs.
• The nest is 30 cm in diameter
and 5 to 8 cm deep.
• Tadpoles hatch within this
relatively safer barrier and
develop until they are large
enough to defend
themselves.
Dr. J. K. Barot 30
6. Tree nests.
• The South American tree frog, Phyllomedusa
hypochondrales, lays eggs in a folded leaf nest with
margins glued together by cloacal secretion.
• The tadpoles when formed fall straight into water
below.
• Another tree frog, Hyla resinfictrix, lines a shallow tree
cavity with bees wax obtained from the hives of certain
stingless bees.
• Female lays eggs when this cavity is filled with rain
water.
• Here, the young develop relatively free from predators.
Dr. J. K. Barot 31
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 7. Gelatinous bags.
• In Phrynixalus biroi large
eggs are enclosed in a
sausage-shaped transparent
gelatinous membranous
bag, secreted by female and
left in mountain streams.
• Salanmndrella keyserlingi a
small aquatic salamander
also deposit 50 to 60 small
eggs in a gelatinous bag.

Dr. J. K. Barot 33
[II] Direct carrying by parents
– 1. Coiling around eggs.
– 2. Transferring tadpoles to water.
– 3. Eggs glued to body.
– 4. Eggs in back pouches.
– 5. Organs as brooding pouches.
– 6. Viviparity.

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1. Coiling around eggs.
• In Congo eel, Amphiuma and certain caecilians
like Ichthyophis and Hvpogeophis, the female
lays large eggs in burrows in damp soil and
carefully guards them by coiling her body
around them until they hatch.
• The female of Salamander, Plethodon also coils
round the eggs which are laid in small packages
in the hollow of a rotton log or beneath a rock.
• In Megalobactracus maximus, it is the male
who coils around the eggs.
Dr. J. K. Barot 36
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 2. Transferring tadpoles to water.
• Some species of small
frogs (e.g. Phyllobates,
Anthroleptis, Pelobates,
Dendrobates) in both
tropical Africa and South
America, deposit their
eggs on ground.
• The tadpoles hatching
out, fasten themselves to
the back of one of the
parents with their sucker-
like mouth and
transported to water. Dr. J. K. Barot 38
Dr. J. K. Barot 39
3. Eggs glued to body.
• Many amphibians, instead of remaining with the eggs, carry the
eggs glued to their body.
• In the dusky salamander, Desmognathus fuscus, female carries the
string of eggs coiled around her neck, until they have hatched.
• In Srilankan tree frog, Rhacophorus reticulatus, the eggs are glued
to the belly of female.
• An interesting case is that of the European midwife toad, Aiytes
obstetricans.
• When the female lays eggs, the male entangles them around his
hind-legs.
• He carries them with him until they are ready to hatch.
• At that time he releases the tadpoles into nearest water.

Dr. J. K. Barot 40
Dr. J. K. Barot 41
4. Eggs in back pouches.
• In one group of tree frogs called marsupial frogs
or toads, the female carries the eggs on her back,
either in an open oval depression, a closed pouch
or in individual pockets.
• The eggs develop into miniature frogs before
they leave their mother's back.
• In the Brazilian tree toad, Hyla goeldii or
Cryptobatrachus evansi, the posterior part of the
back of female forms a sort of incipient brood
pouch in which the eggs remain exposed.
Dr. J. K. Barot 42
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• In Nototrema (Gastrotheca), the eggs are covered by skin
forming a single large brood pouch which opens
posteriorly in front of the cloacal aperture.
• A similar adaptation is seen in the completely aquatic
Surinam toad, Pipa.
• In breeding season, skin of female's back becomes thick,
vascular, soft and gelatinous.
• The male presses fertilized eggs against female's back,
where they sink into individual pits.
• A hinged cover forms over each egg enclosing it in a small
capsule.
• Complete metamorphosis occurs with capsules.
• The tiny toads leaving mother are tailless and do not
enter water.
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 5. Organs as brooding pouches.
• Male of the terrestrial South American Darwin's frog,
Rhinoderma darwinii, pushes at least two fertilized eggs into his
relatively large vocal sacs.
• Here, they undergo complete development to emerge out as
fully formed froglets.
• In West African tree frog, Hylambates breviceps, the female
carries eggs in her buccal cavity.
• In Arthroleptis, it is the male who keeps the larvae in his mouth.
• The only known case of gastric incubation in vertebrates is found
in the Australian frog, Rheobatrachus silus.
• The female keeps the eggs in her stomach.
• The tadpoles are expelled through mouth after metamorphosis.
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6. Viviparity.
• Some anurans are ovoviviparous.
• They retain eggs in the oviducts and the females give birth to
living young.
• African toads, Nectophrynoids and Pseudophryne give birth
to little frogs.
• The European Salamander, Salamandra salamandra produces
20 or more small young while the Apline salamander, S. atra
gives birth to one or two fully developed young.
• Viviparity is widespread in order Gymnophiona, and 3 out of 4
families have this mode of reproduction.
• Common examples are Typhlonectes, Geotrypetes,
Dermophis, etc.

Dr. J. K. Barot 49
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 Neoteny and Paedomorphosis…
• The two terms, paedogenesis and neoteny (coined by Kollwann) are
often used as synonyms which is not correct.
• Neoteny refers to the retention of a larval or embryonic trait in the
adult body.
• Familiar examples are retention of embryonic cartilaginous skeleton
in adult in Chondrichthyes; and the larval gills in some adult
salamanders.
• Paedogenesis or paedomorphosis refers to development of gonads
and/or production of young ones by an otherwise immature, larval
or pre-adult animal.
• The examples are scattered in several groups or animals (e.g. gall fly,
liver fluke, salamanders).
• Thus, whereas neoteny emphasizes the retention of embryonic or
larval features in the adult body, paedogenesis stresses precocious
development of gonads in larval body.
Dr. J. K. Barot 51
 Amphibian examples…
• Some aquatic larval urodeles delay or fail to
metamorphose, yet become sexually mature, mate and
produce fertile eggs.
• Do they represent neoteny or paedogenesis ?
• The distinction between these two processes becomes
blurred or overlapped in these amphibians.
• These animals may either be looked upon as adults,
which retain certain larval characteristics not having
metamorphosed. This is neoteny.
• Or, they may be considered larvae in which reproductive
organs develop precociously. This is paedogenesis.
Dr. J. K. Barot 52
Axolotl.
• Classical and most informative examples of neoteny and
paedogenesis among Amphibia (vertebrates) are furnished by
Ambystoma (or Amblystoma).
• A. maxicanum lives in Lake Xochimilco in the highlands of
Mexico and the closely related. A. tigrinum (tiger salamander)
in high altitudes of Colorado (North America).
• Ordinarily they go through typical gilled aquatic larval stages,
then metamorphosis, to transform into adult air-breathing
land forms.
• However, under certain circumstances, the larvae do not
metamorphose, retain their gills and aquatic habitat but
become sexually mature.
• This sexually mature but morphologically immature, larval
stage with external gills is called an axolotl.
Dr. J. K. Barot 53
• The name was given by natives of Mexico who captured
them for food.
• Axolotl means "servant of water".
• At one time axolotls were considered a separate genus,
Siredon, S. axoiotl, S. mexicanum, S. pisciformis etc.
(Wagler, 1830).
• However, in 1865, some axolotls brought to Paris in
France, discarded their gills and fins and underwent
metamorphosis to become adult terrestrial
salamanders.
• Axolotls have been important laboratory experimental
animals for a century, for most of them now used
worldwide have descended directly from those sent to
Paris in 1864 and 1868
Dr. J. K. Barot 54
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Environmental factors affecting neoteny.
• The significance and causes of neoteny are not
properly understood.
• Environmental factors affect metamorphosis in
several ways.
• Abundance of food, cold temperature or insufficient
iodine (a component of thyroxin hormone that
induces amphibian metamorphosis) may cause failure
of metamorphosis and retention of larval features.
• This is indicated by the fact that drying up of swamps,
lack of food and rise in temperature in surrounding
water induce axolotls to metamorphose.
Dr. J. K. Barot 56
• As already mentioned, axolotls breed in Mexico and
south-western parts of U.S.A. When transferred to the
eastern states or experimentally treated with thyroxine
or TSH, these axolotls lose their gills, assume lungs and
become adult air-breathing native tiger salamanders.
• Response of larval tissues to thyroid hormones is
reduced as temperature drops and disappears entirely
below 5°C.
• The genetic basis for metamorphosis seems to be
multifactorial, variable and subject to selective
pressure.
• Genes for transformation have become suppressed in
neotenic populations but not entirely absent for there is
occasional appearance of metamorphosed individuals.
Dr. J. K. Barot 57
Dr. J. K. Barot 58
 Extrinsic factors
(1) Abundance of food and other favourable requisites in the
aquatic life is the cause of retention of larval features (Gadow,
1903).
(2) Deepwater and coldness inhibits the secretion of thyroxin
(Shufeldt).
(3) Saline nature of water is responsible for neoteny (Weismann).
(4) Low temperature is responsible for the arrest of
metamorphosis (Huxley, 1929).

• Despite of extensive reaserches on the role of extrinsic factors

on metamorphosis it is still not clear that whether extrinsic
factors are exclusively responsible for arrest of metamorphosis.
• So the existence of other factors, internal and physiological
becomes apparent. Dr. J. K. Barot 59
 Intrinsic factors

• Many experimental evidences have been advanced by

different investigators.
• Gressner (1928) was of the opinion that insulin
hormone inhibits metamorphosis.
• But recent researches incline to reveal that
metamorphosis is primarily influenced by
– (i) varying threshold levels of thyroxin and its analogues and
– (ii) by the degree of responsiveness of the larval tissues to
the hormones.
• During early pre-metamorphic stage in amphibian
development, the level of thyroxin is kept very low in
the body by genetic mechanism (Etkin, 1968).
Dr. J. K. Barot 60
• Etkin and his coworkers have also established
the role of prolactin on metamorphosis.
• They have shown that the level of prolactin
which acts as an inhibitor in the overall control
of metamorphosis remains high at this time.
• In the light of modern genetics it may be
suggested that the structural genes guiding
the synthesis of thyroxin are switched off* by
some operator genes, whereas, the genes
guiding the formation of prolactin are
"switched on".
Dr. J. K. Barot 61
• In such condition, hypothalamus becomes sensitive to the
available level of thyroid hormone in the blood stream.
• The neurosecretory apparatus of the hypothalamus
produces a substance called thyrotropin releasing factor
(TRF).
• TRF stimulates the anterior lobe of pituitary to produce
thyroid stimulating hormone (TSH), which in turn
enhances the rate of thyroid secretion.
• As the level of TSH rises during protnetamorphosis, the
level of prolactin suddenly falls, so the metamorphosis
starts.
• Poor secretion of thyroid glands and the
irresponsiveness of the larval tissues to the hormone are
responsible for neoteny.
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Dr. J. K. Barot 63
• In amphibian development the tadpole larva undergoes
progressive metamorphosis and transforms into an adult.
• This is a normal occurrence in amphibians.
• But deviation from the normal pathway of development is found
in the life cycle of many urodeles.
• Such deviated pathways of development in axolotls due to
extrinsic as well as intrinsic environmental factors may be
regarded as "canalisation", i.e., buffering of development
against environmental change.
• Neoteny is looked upon as a consequence of adaptations to
neighbouring environments where retention of larval gills and
other larval features may be advantageous.
• G.K. Noble (1954) regarded that the retention of larval features
during sexual maturity has nothing to do in the phytogeny of the
amphibians.
Dr. J. K. Barot 64
 Types of neoteny…
– Partial neoteny,
– Intermediate neoteny,
– Total neoteny
• Neoteny is- partial when metamorphosis is delayed due to
temporary ecological or physiological changes in environment.
It is shown by tadpoles and larvae tiding over winter.
• Intermediate neoteny is shown by axolotls which also reproduce
sexually but undergo metamorphosis in suitable conditions.
Under experimental conditions in laboratories, it is possible to
produce either axolotls or transformed individuals.
• The extreme or total neoteny is shown by several
perennibranchiate salamanders such as Necturus, Siren and
Proteus. They remain larval throughout. Even treatment with
thyroxine fails to induce metamorphosis; the tissue response is
Dr. J. K. Barot 65
absent.
Significance of neoteny…

• Weismann (1875) thought neoteny to be a case

of retarded evolution or atavism, that is,
reversion to ancestral condition.
• However, this is now regarded to be of secondary
specialization, a physiological adaptation of
advantage.
• This is also proved by the great heterogeneity of
all neotenous perennibranchiate forms.

Dr. J. K. Barot 66