Dr. J. SWAMY, Assistant Professor of Zoology B.

Sc ZOOLOGY Semester – I
3.4 Amphibia: General characteristics and classification up to orders: Parental
care, Neoteny and Paedogenesis in Amphibians

GENERAL CHARACTERS OF AMPHIBIA

Amphibia: Amphi – Double; bios – Life (Greek)

The amphibians are the first anamniotic tetrapods. They have evolved from
crossopterygian “Osteolepid fishes” during “Devonian period”. The golden age of
amphibians was the “Carboniferous period”. The term 'Amphibia’ coined by Linnaeus.
The study of Amphibians is known as “Batrachology”. The largest living amphibian is
“Andrias davidianus” (Chinese giant salamander) and the smallest one is “Paedophryne
amanuensis” (7.7 m.m.). According to Romer, osteolepids modified their appandages
to reach water not to leave water. Bucher, Professor of Yale university observed that
the foot prints of Thynopus were intermediary to fins of fishes and limbs of
amphibians.

1. Amphibians live partly in water and partly on land. They breed in water. Usually
larvae are aquatic. Some adults are also aquatic.

2. Rana cancrivora is the only marine amphibian.

3. Poikilotherms. Exhibit hibernation or aestivation.

4. Head distinct, trunk elongated. Neck and tail may be present or absent.
5. Limbs usually two pairs (tetrapod), some limbless. Toes 4-5 (pentadactylus) or less.
Paired fins absent. Median fins if present, without fin rays.
6. Skin soft, moist and glandular. Pigment cells (chromatophores) present.
7. Exoskeleton absent. Digits clawless. Some with concealed dermal scales.
8. Coelome consists pericardial and pleuroperitoneal cavities
9. Endoskeleton mostly bony. Notochord does not persist. Skull with two occipital
condyles (Dicondylic skull). Autostylic jaw suspension is seen.
10. Vertebrae are Procoelous or Ophisthocoelous or Amphicoelous.
11. Two pairs if zygopophyses strengthen the intervertebral joints.
12. Single sacral vertebra is present.
13. Sternum appeared for the first time in amphibians.

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 Dr. J. SWAMY, Assistant Professor of Zoology B. Sc ZOOLOGY Semester – I
14. Mouth large. Upper or both jaws with small teeth. Homodont, acrodont and
polyphyodonty type of dentition
15. . Tongue often protrusible. Alimentary canal terminate into cloaca.
16. Respiration by lungs, skin and mouth lining. Larvae with external gills which may
persist in some aquatic adults.
17. Larynx appeared for the first time, hence these are the directors of sound
production. Vocal cords are present in anurans.
18. Heart 3-chambered (2-auricle and 1-ventricle). Sinus venosus and truncus
arteriosus are present. Aortic arches 1-3 pairs. Renal and hepatic portal systems
well developed. Erythrocytes large, oval and nucleated.
19. Kidneys mesonephric. Urinary bladder large. Urinary ducts open into cloaca.
Larvae are ammonotelic and adults are ureotelic organisms.
20. Brain is well developed, protected by two mwninges namely outer dura matter and
inner pia matter. Cerebral hemispheres are smooth. Cerebellum is small. Cranial
nerves are 10 pairs.
21. One pair of eyes with movable eye lids, nictitating membrane are present.
22. Nostrils connected to buccal cavity.
23. External ear is absent. Middle ear with a single root like ossicle, columella auris,
formed by the modification of hyomandibular arch.
24. Larval forms and some aquatic adults with lateral line sensory system.
25. Sexes separate. Male without copulatory organ. Gonoducts opens into cloaca.
26. Fertilization mostly external. Females mostly oviparous.
27. Anamniotes.
28. Development is indirect.
29. Mesolecithal eggs. Cleavage is holoblastic but unequal.
30. Larva a tadpole which metamorphosis into adult. Parental care is seen
CLASSIFICATION OF AMPHIBIA

The living amphibians are represented by about 2500 species. They dominated
the world during carboniferous, but most of them have become extinct since long.
G. Kingsley Noble (1924) recognized 3 orders of extinct and 3 orders of living
amphibians.

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 Dr. J. SWAMY, Assistant Professor of Zoology B. Sc ZOOLOGY Semester – I
Subclass I: Stegocephalia (Extinct)

 Limbs pentadactylus.
 Skin with scales and bony plates.
 Skull with a solid bony roof, living openings for eyes and nostrils.
Order 1: Labyrinthodontia

 Oldest known tetrapod called stem Amphibia.

 Fresh water or land forms.
 Salamander or crocodile like.
 Teeth large with characteristically much folded dentine similar to their
crossopterygian ancestors.
Example: Eryops, Eogyrinus, Paleogyrinus, Cyclotosaurus

Order 2: Phyllospodyli

 Small salamander-like.
 Head large, flat.
 Vertebrae tubular.
 Notochord and spinal cord housed in common cavity.
 Believed to be ancestors of modern salientia and urodela.
 Carboniferous to Permian.
Example: Branchiosaurus (Ichthyostega).

Order 3: Lepospondyli

 Small salamander or eel-like.

 Vertebrae cylindrical, each made of a single piece.
 Neural arch and centrum continuous.
 Ribs articulating intervertebrally.
 Regarded ancestral to modern caecilians (Gymnophiona).
 Carboniferous to Permian.
Example: Diplocaulus, Lysorophus.

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 Dr. J. SWAMY, Assistant Professor of Zoology B. Sc ZOOLOGY Semester – I

Subclass II: Lissamphibia (living)

 Modern Amphibia lacking dermal bony skeleton.

 Teeth small simple.
Order 1: Gymnophiona or Apoda
(gymnos-naked; ophioneos-serpent-like Greek) (a-without; podos-foot Greek)

 Limbless, blind, elongated worm like, burrowing (fussorial) tropical forms known
as caecilians (Blind worms).
 Tail short or absent, cloaca terminal.
 In some dermal scales embedded in skin which is transversally wrinkled.
 Skull compact, roofed with bone.
 Limb girdle absent.
 Eyes are small, non-functional, present below the bones of the skull.
 Amphicoelous vertebrae, long ribs are present.
 Tooth found on both the jaws.
 Lungs and kidneys are asymmetrical.
 Gills are absent in adults.
 A small blood vessel that connects systemic and pulmonary arches is known as
ductus botali is present.
 In males, cloacal wall is eversible and acts as copulatory organs.
Example: Ichthyophis, Gegenophis, Uroaeotyphlus, Typhlonectes.

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 Dr. J. SWAMY, Assistant Professor of Zoology B. Sc ZOOLOGY Semester – I

Order 2: Urodela or Caudata

(Ura= tail + delos= visible Greek) or (cauda= tail Latin)

 Lizard-like amphibians with a distinct tail.

 Limbs 2 pairs, usually weak, almost equal.
 Skin devoid of scales and tympanum.
 Gills permanent or lost in adults.
 Males without copulatory organs.
 Larvae aquatic, adult-like, with teeth.
 About 300 species in 5 suborders.
Suborder 1: Cryptobranchoidea

 Most primitive. Permanently aquatic.

 Adults without eyelids and gills.
 Angular and prearticular separate.
 Premaxillary spine is short.
 Fertilization external.
Example: Cryptobranchus, Megalobranchus.

Suborder 2: Ambystomatoidea

 Adult terrestrial with two eyelids.

 Angular fuse with prearticular.
 Premaxillary spine short.
 Vertebrae amphicoelous.

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 Dr. J. SWAMY, Assistant Professor of Zoology B. Sc ZOOLOGY Semester – I
 Fertilization internal.
Example: Amblystoma (Tiger salamander).

Suborder 3: Salamandroidea

 Vertebrae opisthocoelous.
 Teeth on palate and pre-vomers.
 Three sets of cloacal glands.
 Fertilization internal.
Example: Triton and Triturus (newts), Salamandra (Salamander), Desmognathus,
Amphiuma (Congo eel), Plethodon.

Suborder 4: Proteidae

 Aquatic bottom dwellers representing permanent larval forms, without eyelids.

 Adults with 3 pairs of external gills and 2 pairs of gill slits.
 Skull cartilaginous, without maxillae.
 Jaws with teeth.
Example: Proteus (Blind salamander), Necturus (Mud-puppy)

Suborder 5: Meantes

 Aquatic representing permanent larvae.

 Fore limbs small, hind limbs absent.
 Three pairs of external gills.
 Jaws with horny covering.
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 Dr. J. SWAMY, Assistant Professor of Zoology B. Sc ZOOLOGY Semester – I
Example: Siren (mud eel), Pseudobranchus.

Order 3: Salientia or Anura

(saliens-leaping Latin) or (an-without; aura-tail Greek)

 Specialized Amphibia without tail in adults.

 Hind limbs usually adapted for leaping and swimming.
 Adults without gills or gill openings.
 Eyelids well-formed. Tympanum present.
 Skin loosely-fitting, scaleless.
 Mandible toothless.
 Pectoral girdle bony. Ribs absent of reduced. Vertebral column very small of 5-9
presacral vertebrae and a slender urostyle.
 Fertilization always external.
 Fully metamorphosed without neotenic forms.
 About 2200 species of frogs and toads in 5 suborders.

Suborder 1: Amphicoela

 Vertebrae amphicoelous. Presacral 9.

 Free ribs and 2 relict tail muscles.
 Fertilization internal.
Example: Leopelma, Ascaphus.

Suborder 2: Opisthocoela

 Vertebrae opisthocoelous.
 Scapula small.
 Ribs free in adult or larva.
Example: Alytes (midwife toad), Bombinator, Discogiosus, Pipa, Xenopus.

Suborder 3: Anomocoela

 Vertebrae procoelous or amphicoelous.

 Free ossified ribs absent.
 Upper jaw with teeth.

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 Dr. J. SWAMY, Assistant Professor of Zoology B. Sc ZOOLOGY Semester – I
Example: Pleobates, Scaphiopus.

Suborder 4: Procoela

 Vertebrae procoelous. Presacral 5-8.

 Urostyle with two condyles. No free ribs.
Example: Bufo (common toad), Rhinoderma, Dendrobates, Hyla (tree toad),
Gastrotheca (marsupial frog).

Suborder 5: Diplasiocoela

 First 7 vertebrae procoelous, 8th vertebrae amphicoelous sacral or 9th vertebrae

convex anteriorly and bears 2 condyles.
 Pectoral girdle usually fuses to sternum
 Ribs absent.
Example: Rana (common frog), Polypedates or Rhacophorus (tree frog)

PARENTAL CARE IN AMPHIBIA

Parental care is the care of the eggs or the young until they become able to
protect themselves from the predators.
Looking after the eggs or young until they are independent, to defend from

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predators, is known as parental care. In amphibians there are many devices for the
protection of the eggs during the early stages of development.
The methods of caring by Amphibia generally fall under two broad categories:
(A) Protection by means of nests, nurseries, or shelters and
(B) Direct caring by parents.
The different modes of protection are given below in the three important orders of
class Amphibia.
A- Protection by Means of Nests, Nurseries and Shelters:
Amphibians have evolved countless interesting methods to give protection to their
defenceless eggs and larvae from predators. Different species of frogs and toads
construct nests or shelters in which the eggs are deposited and the young are
developed.
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, Rhacophorus 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 accumulate in epiphytic tropical
plants. The tadpoles on hatching drop into water to complete their metamorphosis.
Defending Eggs or Territories: Males of green frog Rana clamitans and other species
maintain territories and attack small intruders to defend 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 are rest on top of them.
Direct Development- In some terrestrial or tree frogs such as Arthroleptis, Hylodes
and Hyla nebulosa, the eggs hatch directly into little frogs thus avoiding larval
mortality.
Foam Nests: Many amphibians convert copious mucus secretion into nests for their
young. In the Japanese 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 sloping tunnel into pond or river water for
further development. The female of South American tree frog Leptodactylus stirs up a
frothy mass of mucus, fills in holes near water and lay eggs in them. Some anurans lay
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 Dr. J. SWAMY, Assistant Professor of Zoology B. Sc ZOOLOGY Semester – I
eggs nests of foam floating on water.
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.
Tree Nests: The South American tree frog Phyllomedusa hypochondrales lay 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 lay eggs when this cavity is filled with rainwater. Here the young develop
relatively free from predators.
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. Salamandrella keyserlingi also deposits 50 to 60 small eggs in a gelatinous
bag which is fastened to aquatic plants.
(B) Direct caring by parents:
Coiling Around Eggs: In Congo eel, Amphiuma (urodele) and certain caecilians like
Ichthyophis and Hypogeophis the female large eggs in burrows in damp soil and carefully
guards them by coiling her body around them until they hatch. The female Plethodon
(Salamander) also coil round the eggs which are laid in small packages beneath the stones or in
the hollow of rotten log. In Megalobatrachus maximus, (urodele) the male coils round the eggs.

Organs as Brooding Pouches: In Rhinoderma darwini, (South American Darwin’s

frog) the eggs (few and large) are transferred by the male to the relatively large vocal
sacs that extend over its ventral surface, where eggs develop. In Arthroleptis, it is the
male frog who keeps the larvae in his mouth. Australian frog Rheobatrachus silus keeps
the eggs in her stomach. The tadpoles are expelled through mouth after metamorphosis.
Tadpoles Transported from One Place to Another: Small South American frogs
Phyllobates and Dendrobates and tropical African frogs Arthroleptis and Pelobates lay
their eggs on ground. The hatched tadpoles adhere by their sucker-like lips and
flattened abdomen to the back of one of their parents and, thus, they are carried from
one place to the other and in this way they can even go from one pool to the other and
this is particularly when one pond is to dry up.

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 Dr. J. SWAMY, Assistant Professor of Zoology B. Sc ZOOLOGY Semester – I

Eggs Carried by the Parents: In Obstetric toad (Alytes obstetricans) of Europe, the
male entangles the eggs by adhesion of their gelatinous secretion round his hindlegs.
Here they are retained until the tadpoles are ready to be hatched. Female Rhacophorus
reticulatus (Sri Lankan tree frog) carries the eggs glued to her belly. In Desmognathus
fusca (urodele) the string of eggs is bound around the neck until they hatched.
Eggs in Back Pouches: In one group of tree frogs called marsupial frogs or toads, the
female carries the eggs on her back. In a Brazilian tree-frog, Hyla goeldii, the female
carries the eggs on the back within an incipient brood pouch in which the eggs remain
exposed. How they reached there is not known but probably male does it. In Nototrema
also the eggs are placed over the back in a single large brood pouch covered by the skin
and opened posteriorly in front of cloacal aperture. In Pipa americana (Surinam toad) the
eggs are carried on the back of the mother. In breeding season the back skin of female
becomes thick, vascular, soft and gelatinous. The male places and spaces the eggs.
Each egg sinks into a small pouch, over which develops an operculum, which comes
from a remnant of the egg envelope, reinforced by integumental secretions.

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 Dr. J. SWAMY, Assistant Professor of Zoology B. Sc ZOOLOGY Semester – I

Viviparous or Viviparity: Some anurans are ovoviviparous. They retain eggs in the
oviducts and the females give birth to living young. Two small East African toads,
Pseudophryne vivipara and Nectophryne tornieri, are known to be viviparous, but no
observations have yet been made on them beyond the fact that larvae are found in the
uteri. Caecilians like Typhlonectes, Geotrypetes, Schistometopum, Chthonerpeton,
Gymnopis are ovoviviparous.
NEOTENY AND PAEDOGENESIS IN AMPHIBIANS

The two terms neoteny and paedogenesis coined by Kollwann. The term
neoteny is derived from Latin neotenia means larval life is extended. Neoteny also
called juvenilization, is the retention, by adults in a species, of traits previously seen
only in juveniles.
Retention of larval or embryonic features in the adult body. e.g. retention of embryonic
cartilaginous skeleton in adult Chondrichthyes and the larval gills in some adult
salamanders.

Paedogenesis is the act of reproduction by an organism that has not achieved

physical maturity. In other words, paedogenesis is the production of offspring by an
organism in its larval or juvenile form and elimination of the adult phase of the life
cycle. It is associated with progenesis, where sexual maturity is achieved in the
juvenile form and further physical maturity is not achieved. Paedogenesis is found in

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gall fly, liver fluke, salamanders, insects in which the larval stage reproduces without
achieving maturity. It occurs in the females of certain beetles, Strepsiptera, bagworms,
scale insects and gall midges.

EXAMPLES OF NEOTENY
Classical and most informative examples of neoteny and paedogenesis among
Amphibia are furnished by Ambystoma. Amblystoma maxicanum lives in Lake
Xochimilco in the highlands of Mexico and A. tigrinum in high altitudes of Colorado
(North America). Normally they go through typically gilled aquatic larval stages,
then metamorphose to become 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.

Axolotl larva of Ambystoma tigrinum (tiger salamander) possesses three pairs

of delicate bushy external gills, four pairs of gill-slits, a flat long tail with prominent
tail-fin and a dorsal fin merged with tail-fin. The axolotl larvae possess the power of
retention.
The axolotl larvae become sexually mature when they attain the age of 6
months only. The new-born larvae are hatched out within 15 days. The optimum
temperature for the development of the eggs is 18-24°C. At the age of 6 months, the
axolotls attain a length of 2.0-2.5 cm and become sexually mature to start breeding. A
sexually mature axolotl, at the age of 18–24 months, ranges in length from 15–45
centimeters possess features typical of salamander larvae, including external gills and
a caudal fin. Three pairs of external gills are used for respiration, although buccal
pumping (gulping air from the surface) may also be used in order to provide oxygen
to their lungs.

Types of Neoteny: Kollmann (1882) classified neoteny into two categories, viz.,
Partial neoteny and Total neoteny. But it can be divided into three categories-

 Partial neoteny: Partial neoteny involves the simple postponement of

metamorphosis beyond the normal period due to temporary change in ecological
condition or due to sudden physiological abnormality. Wintering of the tadpoles of
Pelobates fuscus, Pelodytes punctatus, Alytes obstetricans, Hyla arborea, Bufo

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 Dr. J. SWAMY, Assistant Professor of Zoology B. Sc ZOOLOGY Semester – I
vulgaris, Bufo viridis, Rana temporaria, Rana esculenta, Bombinator pachypus and
many others furnish the typical examples of partial neoteny. In Alytes the broods
usually complete their development within autumn. But the larvae which hatched
later in the months of July or August usually hibernate and retain their larval
features up to the next autumn.
 Intermediate Neoteny: Intermediate stages between partial and total neoteny are
also recorded where the larvae become sexually functional and may metamorphose
into adults with the advent of favourable conditions. The sexually mature axolotl
larvae come under this category.
 Total Neoteny: In this category the specimens become sexually mature at the
larval stage but retain larval characters, like (i) external gills, (ii) tail-fin, (iii) ill-
developed eyes, (iv) ill-developed fin on the back and (v) very weak limbs. Total
neotenic animals are paedo-genic. Because paedogenesis involves the capability of
reproduction at the larval stage. In this case of total neoteny the sexually functional
larvae cease to metamorphosis. In extreme cases of total neoteny (e.g., Necturus,
Proteus, etc.) the larvae attain sexual maturity and they remain in that stage without
undergoing metamorphosis. Total neoteny is seen in many urodele (Ex: Necturus,
Amphiuma, Triturus vulgaris, Triton cristatus, Triton waltli, Ambystoma, Triturus
alpestris, Siren, Proteus, etc.).
Factors affecting neoteny

In the axolotl, metamorphic failure is caused by a lack of thyroid stimulating

hormone (TSH), which is used to induce the thyroid to produce thyroxine in
transforming salamanders. Axolotls can be induced to metamorphose by an injection of
iodine (used in the production of thyroid hormones) or by shots of thyroxine hormone.
Another method for inducing metamorphosis is to keep them in shallow water tanks.
They will then, over a period of weeks, slowly metamorphose into adult salamanders.
However, most attempts at inducing metamorphosis led to death.

Extrinsic factors:
● Gadow (1901) advanced the idea that the cause of retention of larval features in
axolotl is the abundance of food and other favourable requisites in aquatic life.
● Shufeldt holds that deep water and coldness inhibit thyroxine secretion which
retards metamorphosis.

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 Dr. J. SWAMY, Assistant Professor of Zoology B. Sc ZOOLOGY Semester – I
● Drying up of swamps, lack of adequate food supply and rise in temperature in
surrounding water induce metamorphosis.
● Weissmann again claimed that the retardation of metamorphosis of the axolotls is
possibly due to the saline nature of the water of the lakes where they live.

Intrinsic factors influencing neoteny:

Zondeck and Leiter (1923) established that calcium delays metamorphosis in axolotls.
Gressner (1928) also advanced that insulin hormone inhibits metamorphosis.
● But recent researches incline to reveal that the metamorphosis is primarily influenced
(i) by varying threshold levels of thyroxine and its analogs and (ii) by the degree of
responsiveness of the larval tissue to hormones.
● During early pre-metamorphic stage in amphibian development, the level of
thyroxine (T4) is kept very low in the body by genetic mechanism. Etkin (1968) and his
collaborators 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 thyroxine are
‘switched off’ by some operator genes whereas the genes guiding the formation of
prolactin are ‘switched on’.
● In such condition the 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 the pituitary to produce thyroid-stimulating hormone
(TSH) which in turn enhances the rate of thyroid secretion.
● As the level of TSH rises during pro-metamorphosis, the level of prolactin suddenly
falls. So the metamorphosis starts. The time of shift in hormone balance is possibly
determined by the initiation of positive thyroid feed-back to the hypothalamus. Poor
secretion of thyroid glands and the irresponsiveness of the larval tissues to the hormone
are responsible for neoteny.
Genetics of Neoteny: The role of thyroxine in urodele metamorphosis has been
revealed by studies in genetics. The thyroid hormone binds to nuclear receptors that are
in immediate contact with DNA. The hormone helps to change the transcription of
genes that also influence to develop the larval characteristics to one, these gradually

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 Dr. J. SWAMY, Assistant Professor of Zoology B. Sc ZOOLOGY Semester – I
change into juvenile and adult characteristics. But the exact role of genetics in
paedomorphosis or neoteny has not clearly understood. However, a single gene
hypothesis is thought to control the axolotl’s life cycle. If this theory is correct, then
there are two alternate alleles at a major locus primarily responsible for determining the
expression of metamorphosis or paedomorphosis.

Significance of Neoteny:
Neoteny is looked upon as a consequence of adaptation to neighbouring environments
where retention of larval gills and other larval features may be advantageous.
● Weissmann (1875) regarded neoteny as a case of reversion to atavistic ancestral
conditions by assuming that all amphibia were originally gill-breathing aquatic
creatures and that every feature seen in a larva represented a phylogenetic stage and the
axolotl as such is a case of reversion to an ancestral stage.
● External gills of urodeles have been evolved as an adaptation to aquatic life. The
external gills actually developed first in the embryos as additional respiratory organs.
The external gills were first initially restricted to embryonic life, which may be
prolonged in aquatic larval life.
● Possession of long external gills in the viviparous embryos of Salamandra altra
supports the contention that the external gills are embryonic but not larval features. So
existence of such gills in neotenous larvae is a secondary but not an atavistic feature.
Besides external gills, the tail with tail-fin and epidermal sense organs of the neotenous
larvae are secondary acquisitions rather than ancestral reminiscences.
● G. K. Noble (1931) regarded that the retention of larval features during sexual
maturity has nothing to do in the phylogeny of the amphibians. This is quite evident
from the heterogeneous characters of the Perenni branchiata where all the neotenous
species are included. Neoteny as such may have some importance in the individual
groups.

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 Dr. J. SWAMY, Assistant Professor of Zoology B. Sc ZOOLOGY Semester – I

PAEDOGENESIS IN AMPHIBIANS
Paedogenesis is the retention of those characters which occurred in younger
stages of the ancestors of a sexually mature descendant. It is considered that
paedogenesis (reproduction in a pre-adult form) has been a powerful influence in the
evolution of at least some animal types (Garstang). A classical case of paedogenesis is
provided by the Mexican Axolotl, which frequently breeds in the gilled or larval state.
Paedogenesis is of two types, namely
1. Obligatory paedogenesis
2. Facultative paedogenesis
Obligatory paedogenesis:
Some species of urodeles or larval stages always remain mature permanent

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 Dr. J. SWAMY, Assistant Professor of Zoology B. Sc ZOOLOGY Semester – I
larval stages. They do not metamorphose at any time. The developing tissues fail to
respond to the thyroid hormone (T 4), when in other forms the tissues respond to the
thyroid hormone and metamorphose into adult. In obligatory paedomorphosis, the
genes for transformation during metamorphosis have become suppressed.
Necturus maculosus live permanently in
ponds and lakes of U.S.A. They have external
gills, mid-dorsal caudal fin and the lung is highly
reduced. Proteus, a blind urodele, lives in
European caves with external gills and no
pigments in the body. Siren of U.S.A. shows
almost larval features with external gills and caudal mid-fin.
Amphiuma of United States have no
external gills but larval gill-slits. They have
vestigial limbs but without eyelids. They are all
obligatory paedomorphic individuals. They do not
metamorphose and cannot be induced by T4 and
are genetically fixed. The Mexican axolotl of
Amblystoma mexicanum which lives in Lakes
Xochimilco and Chalco of Mexico, representing a typical example of paedomorphosis.
Facultative paedogenesis:
The axolotl larva of some urodeles, e.g., some species of Amblystoma of north
western part of U.S.A. and Triturus exhibit paedomorphosis. The larvae become
sexually mature and breed but they can metamorphose and change into adults when the
available conditions are changed. They are not the permanent larvae like Necturus or
Proteus.
In this case the hormone of thyroid gland, thyroxine (T 4) induces the
metamorphosis. When they remain in aquatic condition, they possess external gills,
gill-slits, median dorsal fin and are able to reproduce but when water is dried up or
changes the environmental condition, they metamorphose into adult individuals losing
its larval features. Thus, this type of paedomorphosis is called facultative
paedomorphosis.

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 Dr. J. SWAMY, Assistant Professor of Zoology B. Sc ZOOLOGY Semester – I
3.5 Evolutionary Trends in Early Vertebrates: Transition from water to land;
Adaptive features in early tetrapod

EVOLUTIONARY TRENDS IN EARLY VERTEBRATES

Soft-bodied multicellular animals evolved in the late Precambrian, some of these

were primitive ancestors of modern invertebrates such as corals, invertebrate animals
with hard skeletal parts (molluscs) evolved about 543 million years ago. Essentially all
of the ancestors of modern animal phyla evolved during the Cambrian period and many
forms that became extinct.
Evolution of Vertebrates - The Fishes
Vertebrates belong to phylum chordata, subphylum vertebrata. Oldest chordates
were pikaia of the middle Cambrian. Oldest vertebrate fossils (scales of jawless fish)
are from the upper Cambrian vertebrates have backbone and braincase. Jawless fish
(agnathans) had cartilage skeletons, some had bony plates on the head. Lampreys and
hagfish are about the only remaining jawless fish.
Placoderms probably evolved in the Silurian (common in Devonian) - now
extinct had jaws but no teeth. Jaws modified into tooth-like shapes cartilage skeletons
and bony plates armoring their head some very large predators. Acanthodians probably
evolved by the Silurian - now extinct cartilage skeletons more streamlined form than
placoderms probably ancestors of the bony, ray-finned fish Sharks, rays & skates
probably evolved by the Silurian, common in Devonian cartilage skeletons produce
thousands of enamel covered teeth in their lifetime. Sharks, rays, and skates still survive
very successfully.
Bony Fish - Ray-fins evolved in the Devonian period. Bony skeleton fine bones
support fins. Ray-fins are the most diverse of all fish Bony Fish. Lobe fins evolved in
the Devonian period. Bony skeleton muscular fleshy lobes support finer bones of the
fins. The modern coelacanth and a few others still survive.
Evolutionary Trends in Early Vertebrates

The evolution of vertebrates is a fascinating journey marked by a series of significant

innovations that allowed these animals to diversify and conquer various environments,
from ancient oceans to terrestrial landscapes. The earliest vertebrates emerged during
the Cambrian period, over 500 million years ago, resembling simple, jawless fish. From

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 Dr. J. SWAMY, Assistant Professor of Zoology B. Sc ZOOLOGY Semester – I
these humble beginnings, a remarkable array of anatomical and physiological changes
unfolded.

Here are the key evolutionary trends observed in early vertebrates:

1. Development of a Cranium and Vertebral Column

 From Notochord to Vertebrae: The earliest chordates possessed a notochord, a

flexible rod providing axial support. Early vertebrates evolved a cranium (skull) to
protect the brain, followed by the development of a vertebral column (backbone)
that gradually replaced or supplemented the notochord. Initially, this backbone was
cartilaginous, as seen in early fish and modern sharks.

 Mineralization of Skeleton: A crucial step was the development of mineralized

tissues, first appearing as dentine and enamel-like tissues in conodonts and later
evolving into true bone. A bony endoskeleton provided stronger support, allowing
for larger body sizes and more powerful muscle attachments.

2. Emergence of Neural Crest Cells and Placodes

 Complex Head and Sensory Organs: The evolution of neural crest cells, a unique
embryonic cell population, was a defining vertebrate synapomorphy. These cells
contributed to the formation of a more complex head, including sensory capsules
(eyes, ears, nose) and a protective cranium. Placodes, another set of embryonic
structures, further enabled the development of sophisticated sensory systems like
paired eyes with complex retinas and olfactory organs.

3. Evolution of Jaws

 Revolution in Feeding: One of the most significant breakthroughs was the

development of jaws, which are believed to have evolved from modified gill arches.
This innovation, appearing around 450 million years ago (Late Ordovician to Early
Silurian), dramatically improved feeding efficiency, allowing early vertebrates to
exploit new food sources and adopt active predatory lifestyles, leading to a rapid
diversification of jawed vertebrates (gnathostomes).

4. Paired Appendages (Fins)

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 Dr. J. SWAMY, Assistant Professor of Zoology B. Sc ZOOLOGY Semester – I
 Enhanced Maneuverability: While the earliest vertebrates had only median fins,

the evolution of paired fins (pectoral and pelvic fins) provided enhanced
maneuverability, stability, and control in aquatic environments. These paired fins
were later modified into the limbs of tetrapods.

5. Specialized Sensory Organs and Enlarged Brain

 Advanced Environmental Interaction: Along with the development of complex

eyes and otic capsules for balance and hearing, early vertebrates evolved an
elaborate lateral line system for detecting water movements. These sensory
advancements were accompanied by the evolution of an enlarged brain with distinct
regions (forebrain, midbrain, hindbrain), allowing for more sophisticated processing
of environmental information and complex behavioral responses.

6. Efficient Respiratory and Circulatory Systems

 Supporting Active Lifestyles: The evolution of more efficient gills with increased
surface area and a complex circulatory system, including a muscular heart and
closed blood vessels, enabled better oxygen delivery to tissues. This supported the
increasingly active and predatory lifestyles of early vertebrates.

7. Transition to Land (Tetrapods)

 Lungs and Limbs: Around 365 million years ago, lobe-finned fish ancestors
evolved into the first tetrapods (four-limbed vertebrates). This transition involved
the adaptation of swim bladders into lungs for air breathing and the transformation
of muscular paired fins into walking legs capable of supporting the body on land.

 Amniotic Egg and Waterproof Skin: Reptiles, evolving from amphibians,

developed the amniotic egg, which contained specialized membranes and a shell,
allowing reproduction on land without dependence on water. They also evolved
dry, scaly skin to prevent desiccation.

 Mobile Neck: The development of a mobile neck, allowing the head to turn
independently of the body, was crucial for terrestrial life, enabling better scanning
of the environment for prey and predators.

8. Endothermy

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 Dr. J. SWAMY, Assistant Professor of Zoology B. Sc ZOOLOGY Semester – I
 Internal Temperature Regulation: A later but highly significant trend, observed

in the evolution of mammals and birds from reptile-like ancestors, was the
development of endothermy (warm-bloodedness). This ability to maintain a
constant internal body temperature through metabolic processes allowed these
vertebrates to be active in a wider range of environmental conditions.

These evolutionary trends, occurring over millions of years, represent a series of

interconnected adaptations that collectively contributed to the remarkable diversity and
ecological success of vertebrates across the planet.

TRANSITION FROM WATER TO LAND

The water-to-land transition is one of the most important and inspiring

major transitions in vertebrate evolution. And the question of how and when
tetrapods transitioned from water to land has long been a source of wonder and
scientific debate.

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 Dr. J. SWAMY, Assistant Professor of Zoology B. Sc ZOOLOGY Semester – I
Early ideas posited that drying-up-pools of water stranded fish on land and
that being out of water provided the selective pressure to evolve more limb-like
appendages to walk back to water. In the 1990s newly discovered specimens
suggested that the first tetrapods retained many aquatic features, like gills and a tail
fin, and that limbs may have evolved in the water before tetrapods adapted to life on
land. There is, however, still uncertainty about when the water-to-land transition
took place and how terrestrial early tetrapods really were. High-resolution fossil
data shows that these early tetrapods were still tied to water and had aquatic features,
they also had adaptations that indicate some ability to move on land. Although, they
may not have been very good at doing it, at least by today's standards.
The Remarkable Transition: From Water to Land
The move from aquatic to terrestrial life, primarily by early vertebrates, was
a watershed moment in evolution, occurring roughly 390 to 360 million years ago
during the Devonian period. This complex transition involved overcoming
numerous challenges posed by the new terrestrial environment and led to the
diversification of all land-dwelling vertebrates, including amphibians, reptiles,
birds, and mammals.
Environmental Pressures and Opportunities made them to move to land:
The Devonian period saw fluctuating environmental conditions, particularly in
freshwater habitats.
Drying Pools and Hypoxia: Periods of drought could cause shallow pools and
streams to dry up, leading to increasingly foul water and a drastic reduction in
dissolved oxygen. Fish that could breathe atmospheric oxygen would have a
significant survival advantage.
New Niches and Resources: Land offered untapped resources. Early plants and
invertebrates had already colonized terrestrial environments, providing potential
food sources without the intense competition and predation found in the crowded
aquatic realms.
Favorable Temperatures: While water temperatures could fluctuate, certain
terrestrial environments might have offered more stable or advantageous thermal
conditions for some species.

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 Dr. J. SWAMY, Assistant Professor of Zoology B. Sc ZOOLOGY Semester – I

ADAPTIVE FEATURES IN EARLY TETRAPOD

The transition required profound changes across almost all bodily systems.
1. Respiration: From Gills to Lungs
 Swim Bladder to Lungs: Fish possessed a swim bladder, an air-filled sac used
for buoyancy control. In the lineage leading to tetrapods, this organ became
increasingly vascularized and adapted for gas exchange, effectively functioning
as a primitive lung.
 Loss of Gills: While early transitional forms like Acanthostega still retained
gills, true tetrapods eventually lost them as lungs became the primary means of
oxygen uptake.
 Nostrils and Choanae: The development of nostrils connected to the lungs
(choanae) allowed for breathing air with the mouth closed, a more efficient

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 Dr. J. SWAMY, Assistant Professor of Zoology B. Sc ZOOLOGY Semester – I
method on land.
2. Locomotion: From Fins to Limbs
 Lobe-Fins as Precursors: Lobe-finned fish had fleshy, muscular fins with
bones homologous to the limb bones of tetrapods (e.g., humerus, radius, ulna).
These robust fins were initially used for "walking" along the bottom of shallow
waters.
 The humerus bone because it is not only abundant and well preserved in the
fossil record, but it is also present in all sarcopterygians, a group of animals
which includes coelacanth fish, lungfish, and all tetrapods, including all of their
fossil representatives. According to Pierce "We expected the humerus would
carry a strong functional signal as the animals transitioned from being a fully
functional fish to being fully terrestrial tetrapods and that we could use that to
predict when tetrapods started to move on land".
 Strengthened Girdles and Vertebral Column: To support the body against
the full force of gravity (unlike the buoyant support of water), the vertebral
column became stronger, and the pectoral (shoulder) and pelvic (hip) girdles
became more robust and connected to the axial skeleton.
 Digits: The evolution of distinct digits at the end of the limbs, as seen in fossils
like Acanthostega (which had more than five digits), provided better traction
and articulation for movement on uneven terrestrial surfaces.
3. Support and Skeleton
 Rib Cage: A strong rib cage evolved to support the internal organs against
gravity and to aid in lung ventilation.
 Skull Changes: The skull became flatter in some early forms, with eyes
positioned more dorsally, allowing them to peer above the water surface. Later,
a mobile neck evolved, allowing the head to turn independently of the body,
crucial for surveying the terrestrial environment.
4. Sensory Systems
 Vision: Light behaves differently in air than in water. Terrestrial animals
experienced a greatly enlarged visual range. Eyes adapted to focus in air, and
the development of a mobile neck further enhanced visual perception.
 Hearing: The lateral line system, vital for detecting water movements, became

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 Dr. J. SWAMY, Assistant Professor of Zoology B. Sc ZOOLOGY Semester – I
obsolete on land. New adaptations for detecting airborne sounds, such as the
evolution of a tympanum (eardrum) in an otic notch, emerged.
 Olfaction: The sense of smell became more important for detecting airborne
chemical cues, such as food or predators.
5. Reproduction and Water Conservation
 Amphibian Dependence: Early amphibians remained tied to water for
reproduction, as their eggs lacked a protective shell and required a moist
environment. Their skin was also permeable, necessitating moist habitats.
 Amniotic Egg: The evolution of the amniotic egg in reptiles was a monumental
step, providing a self-contained aquatic environment for the embryo, thus
freeing reproduction from external water bodies. This allowed for full terrestrial
colonization.
 Kidneys and Waste Excretion: Terrestrial life required more efficient water
conservation. Kidneys adapted to excrete nitrogenous waste with less water loss
(e.g., urea or uric acid instead of ammonia). Skin also became less permeable to
water.
Transitional Fossils
“Department of Organismic and Evolutionary Biology and curator of vertebrate
paleontology in the Museum of Comparative Zoology at Harvard University,
examined 40 three-dimensional models of fossil humeri (upper arm bone) from
extinct animals that bridge the water-to- land transition”. According to Dickson
"Because the fossil record of the transition to land in tetrapods is so poor we went to
a source of fossils that could better represent the entirety of the transition all the
way from being a completely aquatic fish to a fully terrestrial tetrapod".
The fossil record provides compelling evidence for this stepwise transition:
 Eusthenopteron: A lobe-finned fish with a skull and fin bones resembling early
tetrapods, suggesting it could "flop" along the bottom.
 Tiktaalik: Often called a "fishapod," this fossil shows a blend of fish and
tetrapod features, including a fish-like body, scales, and fins, but also a
flattened skull, eyes on top of its head, a mobile neck, and robust, limb-like fins
with wrist bones.
 Acanthostega: One of the earliest known tetrapods, it had well-formed limbs

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 Dr. J. SWAMY, Assistant Professor of Zoology B. Sc ZOOLOGY Semester – I
with digits (though likely more than five), but its limbs were still too weak for
effective terrestrial walking, suggesting it was primarily aquatic. It also retained
gills and a tail fin.
 Ichthyostega: A contemporary of Acanthostega, Ichthyostega had more robust
limbs and a stronger shoulder girdle, indicating a greater ability to pull itself
onto land, though it likely still moved with a sprawling, "crutching" motion
rather than true walking.
This incredible journey from water to land highlights the power of natural selection
to shape life in response to new environmental challenges and opportunities,
ultimately leading to the diverse array of vertebrates that inhabit our planet today.

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