AMPHIBIANS

THE FIRST TERRESTRIAL

VERTEBRATES
 PHYLOGENETIC RELATIONSHIPS
• Amphibians are tertrapods
• Origin of amphibians was from ancient
sarcopterygians
• No one knows, therefore, what animal was
the first amphibian, but the structure of limbs,
skulls, and teeth suggests that Ichthyostega is
probably similar to the earliest amphibians
Modern amphibians belong to three orders:
• Caudata, the salamanders;

• Gymnophiona, the caecilians

• Anura, the frogs and toads;

 ORDER CAUDATA (Salamanders)
• Possess a tail throughout life, and both pairs of legs, when
present, are relatively unspecialized
• Salamanders range in length from only a few centimeters to 1.5
m (the Japanese giant salamander, Andrias japonicus)
 Family Plethodontidae are the most fully terrestrial
salamanders. Their eggs are laid on land, and the young hatch as
miniatures of the adult.

 Members of the family Salamandridae are commonly called

newts. They spend most of their lives in water and frequently
retain caudal fins.
• Most salamanders have internal fertilization
• Male produce gelatinous structures called
spermatophores with sperm deposited at their tips. A receptive
female is induced to pass over the spermatophore and take the
packet of sperm into her cloaca.
• Sperm are stored in the spermatheca associated with
the oviduct until ovulation. The eggs are fertilized with stored
sperm as they are shed, often in the absence of males.
• Larvae are similar to adults but smaller. They often possess
external gills, a tail fin, larval dentition, and a rudimentary
tongue
• The aquatic larval stage usually metamorphoses into a
terrestrial adult.
• Many other salamanders undergo incomplete metamorphosis
and are paedomorphic (e.g., Necturus); that is, they become
sexually mature while still showing larval characteristics.
ORDER GYMNOPHIONA (caecilians)
• Wormlike burrowers that feed on worms and other
invertebrates in the soil.
• Appear segmented because of folds in the skin that overlie
separations between muscle bundles.
• Skin covers the eyes; thus, caecilians are probably nearly blind.
• Fertilization is internal in caecilians.
• Larval stages are often passed within the oviducts, where they
scrape the inner lining of the oviducts with fetal teeth to feed.
The young emerge from the female as miniature adults.
• Other caecilians lay eggs that develop into either aquatic larvae
or embryos that develop on land.
 ORDER ANURA (SALIENTIA)
• Includes species of frogs and toads.
• Adults lack tails, and caudal (tail) vertebrae fuse into a rodlike
structure called the urostyle.
• Hindlimbs are long and muscular and end in webbed feet.
• Fertilization is almost always external, and eggs and larvae are
typically aquatic.
• Larval stages, called tadpoles, have well-developed tails. Their
plump bodies lack limbs until near the end of their larval existence.
• Unlike adults, the larvae are herbivores and possess a
proteinaceous, beaklike structure used in feeding.
• Anuran larvae undergo a drastic and rapid metamorphosis from
the larval to the adult body form
https://www.yourdictionary.com/articles/frog-toad-differences
https://www.youtube.com/watch?v=K3oG6lyqZDI
EVOLUTIONARY PRESSURES
 EXTERNAL STRUCTURE AND LOCOMOTION
Importance of Skin
 Protects against infective microorganisms, ultraviolet light,
desiccation, and mechanical injury
 Functions in gas exchange, temperature regulation, and
absorption of water
 Skin lacks a covering of scales, feathers, or hair. It is, however,
highly glandular, and its secretions aid in protection. These glands
keep the skin moist to prevent drying.
 They also produce sticky secretions that help a male cling to a
female during mating
 Produce toxic chemicals that discourage potential predators
 Chromatophores are specialized cells in the epidermis and dermis
of the skin that are responsible for skin color and color changes
Support and Movement
• Skull is flattened, is relatively smaller, and has fewer bony elements
• Have a neck
• Jaws are modified for crushing prey held in mouth
• Vertebral Column acts like the arch of a suspension bridge by
supporting the weight of the body between anterior and posterior
paired appendages
• Supportive processes called zygapophyses on each vertebra
prevent twisting.
• The first vertebra is a cervical vertebra, which moves against the
back of the skull and allows the head to nod vertically.
• The last trunk vertebra is a sacral vertebra. This vertebra anchors
the pelvic girdle to the vertebral column to provide increased
support.
• The pelvic girdle of amphibians consists of three bones (the ilium,
ischium, and pubis) that firmly attach pelvic appendages to the
vertebral column.
• Tetrapods depend more on appendages than the
body wall for locomotion. Thus, body-wall
musculature is reduced, and appendicular
musculature predominates.
• Terrestrial salamanders move by a pattern of limb and body
movements in which the alternate movement of appendages
results from muscle contractions that throw the body into a curve
to advance the stride of a limb.

• Caecilians have an accordion-like movement in which adjacent

body parts push or pull forward at the same time.
• The long hindlimbs and the pelvic girdle of anurans are modified
for jumping
 NUTRITION AND THE DIGESTIVE SYSTEM
• Most adult amphibians are carnivores
• Diet depends on prey size and availability
• Most amphibians locate their prey by sight and simply wait for prey to pass
by.
• Olfaction plays an important role in prey detection by aquatic salamanders
and caecilians
• Many salamanders only their jaws to capture prey.
• Anurans and plethodontid salamanders, however, use their tongue and jaws
in a flip-and-grab feeding mechanism
• A true tongue is first seen in amphibians. Mucous and buccal glands on the
tip of the tongue exude sticky secretions
• The tip of the tongue entraps the prey, and the tongue and prey are flicked
back inside the mouth.
• The amphibian holds the prey by pressing it against teeth on the roof of the
mouth, and the tongue and other muscles of the mouth push food toward
the esophagus. The eyes sink downward during swallowing and help force
 CIRCULATION
• The separation of pulmonary and systemic circuits is less
efficient in amphibians than in lungfish.
• Gas exchange occurs across the skin of amphibians, as well as in
the lungs
• Blood entering the right side of the heart is nearly as well
oxygenated as blood entering the heart from the lungs.
• When an amphibian is completely submerged, all gas exchange
occurs across the skin and other moist surfaces; therefore, blood
coming into the right atrium has a higher oxygen concentration
than blood returning to the left atrium from the lungs
• Under these circumstances, blood vessels leading to the lungs
constrict, reducing blood flow to the lungs and conserving
energy
• The atrium is partially divided in salamenders and
completely divided in anurans. The ventricle has
no septum.
• A spiral valve in the conus arteriosus or ventral
aorta helps direct blood into pulmonary and
systemic circuits.
• Adult amphibians have fewer aortic arches than
fishes. After leaving the conus arteriosus, blood
may enter the carotid artery (aortic arch III), which
takes blood to the head; the systemic artery
(aortic arch IV), which takes blood to the body; or
the pulmonary artery (aortic arch VI)
 Gas Exchange
 Cutaneous Respiration
o Amphibian skin is moist and richly supplied with capillary beds.
These two factors permit the skin to function as a respiratory organ.
o Gas exchange across the skin is called cutaneous respiration and can
occur either in water or on land.
o This ability allows a frog to spend the winter in the mud at the
bottom of a pond. In salamanders, 30 to 90% of gas exchange occurs
across the skin
 Buccopharyngeal Respiration
o Gas exchange also occurs across the moist surfaces of the mouth
and pharynx.
o This buccopharyngeal respiration accounts for 1 to 7% of total gas
exchange.
 Pulmonary (lung) ventilation
o Most amphibians, except for plethodontid salamanders,
possess lungs.
o The lungs of salamanders are relatively simple sacs.
o The lungs of anurans are subdivided, increasing the
surface area for gas exchange.
o Pulmonary (lung) ventilation occurs by a buccal pump
mechanism. Muscles of the mouth and pharynx create
a positive pressure to force air into the lungs
(b) With the opening of the respiratory tract (the glottis) closed, the
floor of the mouth lowers, and air enters the mouth cavity.
(c) The glottis opens, and the elasticity of the lungs and contraction
of the body wall force air out of the lungs, over the top of air just
brought into the mouth.
(d) The mouth and nares close, and the floor of the mouth raises,
forcing air into the lungs.
(e) With the glottis closed, oscillations of the floor of the mouth
exchange air in the mouth cavity to facilitate buccopharyngeal
respiration.

Blue arrows show air movements. Red arrows

show movements of the lungs and body wall
 Temperature Regulation
• Amphibians are ectothermic.
• When amphibians are in water, they take on the temperature of their
environment to prevent heat loss. On land, however, their body temperatures
can differ from that of the environment.
• Temperature regulation is mainly behavioral.
• Some cooling results from evaporative heat loss. In addition, many
amphibians are nocturnal and remain in cooler burrows or under moist leaf
litter during the hottest part of the day.
• Amphibians may warm themselves by basking in the sun or on warm surfaces.
Basking after a meal is common, because increased body temperature
increases the rate of all metabolic reactions—including digestive functions,
growth, and the fat deposition necessary to survive periods of dormancy
• Amphibians have correspondingly wide temperature tolerances. Critical
temperature extremes for some salamanders lie between -2 and 27° C, and
for some anurans between 3 and 41° C.
 NERVOUS AND SENSORY FUNCTIONS
 Brain
o The forebrain contains olfactory centers and regions
that regulate color change and visceral functions.
o The midbrain contains a region called the optic tectum
that assimilates sensory information and initiates
motor responses. The midbrain also processes visual
sensory information.
o The hindbrain functions in motor coordination and in
regulating heart rate and the mechanics of respiration.
 Lateral line System
o Lateral-line organs are distributed singly or in small groups
along the lateral and dorsolateral surfaces of the body,
especially the head.
o These receptors respond to low-frequency vibrations in
the water and movements of the water relative to the
animal.

 Chemoreceptors
Chemoreceptors are in the nasal epithelium and the lining of
the mouth, on the tongue, and over the skin.
 Olfaction
Olfaction is used in mate recognition, as well as in detecting
noxious chemicals and in locating food
 Vision
o Eyes are on the front of the head, providing the
binocular vision and well-developed depth
perception necessary for capturing prey
o The lower eyelid is movable, and cleans and
protects the eye. Much of it is transparent and is
called the nictitating membrane.
o Orbital glands lubricate and wash the eye.
o The lens is large and nearly round. It is set back
from the cornea, and a fold of epithelium called the
iris surrounds it. The iris can dilate or constrict to
control the size of the pupil.
o Focusing, or accommodation, involves bending
(refracting) light rays to a focal point on the
retina
• Receptors called rods and cones are in the
retina. Cones are associated with color vision
 Auditory system
o It transmits both substrate borne vibrations and, in
anurans, airborne vibrations.
o The ears of anurans consist of a tympanic membrane, a
middle ear, and an inner ear.
o The sense of equilibrium and balance is similar to that
described for fishes.
o The inner ear of amphibians has semicircular canals that
help detect rotational movements and other sensory
patches that respond to gravity and detect linear
acceleration and deceleration
 EXCRETION AND OSMOREGULATION
• The kidneys of amphibians lie on either side of the
dorsal aorta on the dorsal wall of the body cavity
• The nitrogenous waste product that amphibians
excrete is either ammonia or urea.
• Amphibians that live in freshwater excrete
ammonia. It is the immediate end product of
protein metabolism; therefore, no energy is
expended converting it into other products. The
toxic effects of ammonia are avoided because it
rapidly diffuses into the surrounding water.
• Amphibians that spend more time on land excrete urea that is
produced from ammonia in the liver. Although urea is less toxic
than ammonia, it still requires relatively large quantities of
water for its excretion. Unlike ammonia, urea can be stored in
the urinary bladder
• In water, amphibians face the same osmoregulatory problems
as freshwater fishes. Amphibian kidneys produce large
quantities of hypotonic urine, and the skin and walls of the
urinary bladder transport Na , Cl and other ions into the blood.
• On land, amphibians must conserve water. They limit water
loss by behavior that reduces exposure to desiccating
conditions
 Many are nocturnal.
 During daylight hours, they retreat to areas of high humidity,
such as under stones, or in logs, leaf mulch, or burrows.
 Diurnal amphibians usually live in areas of high humidity and
rehydrate themselves by entering the water.
 Many amphibians reduce evaporative water loss by curling
their bodies and tails into tight coils and tuck their limbs
close to their bodies.
 Some amphibians prevent water loss by forming cocoons
that encase the body during long periods of dormancy.
 When an amphibian flattens its body on moist surfaces,
the skin, especially in the ventral pelvic region, absorbs
water.
 Amphibians can also temporarily store water. Water
accumulated in the urinary bladder and lymph sacs can be
selectively reabsorbed to replace evaporative water loss.
 REPRODUCTION & DEVELOPMENT
• Dioecious
• Gonads are located near the dorsal body wall
• Fertilization is usually external
• Developing eggs lack any resistant coverings so
development is tied to moist habitats, usually water
• 10% of all salamanders have external fertilization. All others
produce spermatophores, and fertilization is internal
• All caecilians have internal fertilization, and 75% have
internal development
• Amphibians larval stages are called tadpoles. Amphibian
tadpoles often differ from the adults in mode of respiration,
form of locomotion, and diet.
• Salamanders rely primarily on olfactory and visual cues
in courtship and mating,
• Male vocalizations and tactile cues are important for
anurans.
 Male vocalizations are usually species specific and
function in the initial attraction and contact between
mates.
 After that, tactile cues become more important. The
male grasps the female—his forelimbs around her waist
—so that they are oriented in the same direction, and
the male is dorsal to the female. This positioning is
called amplexus and may last from 1 to 24 hours.
During amplexus, the male releases sperm as the
female releases eggs.
 Vocalization
Sound production is primarily a reproductive function
of male anurans.
 Advertisement calls attract females to breeding
areas and announce to other males that a given
territory is occupied. Advertisement calls are
species specific. The calls may also help induce
psychological and physiological readiness to breed.
 Release calls inform a partner that a frog is
incapable of reproducing. Unresponsive females
give release calls if a male attempts amplexus, as do
males that have been mistakenly identified as
female by another male.
Distress calls are not associated with
reproduction; either sex produces these calls
in response to pain or being seized by a
predator. The calls may be loud enough to
cause a predator to release the frog.
• The sound-production apparatus of frogs
consists of the larynx and its vocal cords. This
laryngeal apparatus is well developed in
males, who also possess a vocal sac.
 Parental Care
The most common form of parental care in
amphibians is attendance of the egg clutch by either
parent.
Maternal care occurs in species with internal
fertilization (predominantly salamanders and
caecilians),
Paternal care may occur in species with external
fertilization (predominantly anurans). It may involve
aeration of aquatic eggs, cleaning and/or moistening
of terrestrial eggs, protection of eggs from predators,
or removal of dead and infected eggs.
• Females of the genus Pipa carry eggs on their backs.
• Rheobatrachus females broode tadpoles in their
stomachs, and the young emerged from the females’
mouths. During brooding, the female’s stomach
expanded to fill most of her body cavity, and the
stomach stopped producing digestive secretions.

• Viviparity and ovoviviparity occur primarily in

salamanders and caecilians
 Metamorphosis
o Series of abrupt structural, physiological, and
behavioral changes that transform a larva into an
adult.
o Metamorphosis is under the control of
neurosecretions of the hypothalamus, hormones
of the anterior lobe of the pituitary gland and the
thyroid gland
o In caecilians and salamenders, slight
morphological changes occur like reproductive
structures develop, gills are lost, and a caudal fin
(when present) is lost.
o In the Anura, however, changes from the
tadpole into the small frog are more dramatic.
Limbs and lungs develop, the tail is
reabsorbed, the skin thickens, and marked
changes in the head and digestive tract
(associated with a new mode of nutrition)
occur