Early Tetrapods and Modern Amphibians

25-1

Movement Onto Land

Adaptations
Movement from water to land is perhaps the most
dramatic event in animal evolution
Animals are composed mostly of water
Land represents a relatively dangerous habitat

Vascular plants, pulmonate snails, and tracheate

arthropods made transition earlier
Amphibians only living vertebrates that have a
transition from water to land in both their ontogeny
and phylogeny
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Movement Onto Land

Quasiterrestrial
Hover between aquatic and land environments

Physical differences that must be addressed when

moving from water to land
Oxygen is 20 times more abundant in air
Diffuses much more rapidly through air

Air is 1000 times less dense

Provides less buoyancy than water
Limbs and skeleton must support more weight

Air fluctuates in temperature more rapidly than water

Animals must adjust to these extremes

Variety of terrestrial habitats allows greater

opportunities for adaptation
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Early Evolution of Terrestrial

Vertebrates
400 million years ago, bony fishes diversified to
include many freshwater forms
Combinations of characteristics that evolved in
aquatic habitats, made it possible to explore
terrestrial habitats
Two structures connected to pharynx
Air-filled cavity functioned as a swim bladder
Paired internal nares functioned in chemoreception
On land, combination would be used to draw in oxygen-rich air
through nares into the air-filled cavity

Bony elements of paired fins

Modified for support and movement underwater
On land, would provide same function
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25-5

Early Evolution of Terrestrial

Vertebrates
Freshwater habitats are inherently unstable
Multiple fish groups, given a combination of structures
that could be coopted for terrestrial breathing and
locomotion, evolved some degree of terrestriality
Ex: Mudskippers and lungfish

One transition in late Devonian period provided the

ancestral lineage of all tetrapod vertebrates
Evolved adaptations to air breathing
Increased vascularization of the air-filled cavity
Double circulation to direct deoxygenated blood into the
lungs and oxygenated blood out of the lungs to other
body tissues
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Early Evolution of Terrestrial

Vertebrates
Bony elements of fins of
lobe-finned fishes
resemble homologous
structures of
amphibians
Eusthenopteron
Could paddle itself
through bottom mud
Had both lungs and
walking fins

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25-9

Early Evolution of Terrestrial

Vertebrates
Tiktaalik
Recently discovered
fossil genus
Intermediate between
lobe-finned fishes and
tetrapods
Probably used limbs to
support body while
placing snout above
water to breathe air in
shallow water
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Early Evolution of Terrestrial

Vertebrates
Acanthostega
Had clearly
formed digits on
both forelimbs
and hindlimbs
Body drug on the
ground

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Early Evolution of Terrestrial

Vertebrates
Ichthyostega
Bulkier limb muscles to walk onto land, but
not with great efficiency

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Early Evolution of Terrestrial

Vertebrates

Lobe-finned fishes are the sister group to tetrapods

Adaptations for life on land include skull, teeth,
pectoral girdle and jointed limbs
Tetrapods

Have stronger backbone

Muscles to support the body in air
Muscles to elevate the head
Stronger shoulder and hip girdles
More protective rib cage
Modified ear structure to detect airborne sounds
Foreshortening of skull
Longer snout
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Early Evolution of Terrestrial

Vertebrates

Ichthyostega still retained aquatic features including

fin rays and opercular bones
Several extinct lineages plus the Lissamphibia with
modern amphibians formed the temnospondyls
Temnospondyls generally had four rather than five
digits on the forelimb
During the Carboniferous, amphibians developed
additional adaptations for living in water
Lepospondyls and anthracosaurs are, based on skull
structure, closer to amniotes than temnospondyl

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Modern Amphibians
Diversity

Over 60 living species are known in the three

amphibian orders
Metamorphosed adults use a redesigned olfactory
epithelium to sense airborne odors and the ear
detects sounds
Remain tied to water
Eggs deposited in water or must be kept moist
Larvae depend on gills for respiration

Some salamanders have retained aquatic

morphology throughout life
Others lack the larval phase

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Modern Amphibians
Thin skin loses water rapidly
Restricts terrestrial forms to moist habitats

Ectotherms
Body temperature depends on environment and restricts
range

Eggs easily dessicate and must be shed into water or

kept moist
A few brood young

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Los anfibios en Cifras

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Modern Amphibians
Caecilians: Order Gymnophiona (Apoda)
Approximately 173 living species
Elongate, limbless, burrowing animals
Inhabit tropical forests in South America, Africa,
India, and Southeast Asia
Bodies have many vertebrae, long ribs, no limbs, and
terminal anus
Feed primarily on worms and small underground
invertebrates
Fertilization is internal
Male has a protrusible copulatory organ
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Modern Amphibians

Eggs deposited in moist ground near water

Some species have aquatic larvae
In others, larval development occurs within egg
Eggs are guarded and develop in folds of the body in
some
In other species, viviparity allows embryos to obtain
nourishment by eating wall of oviduct

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Gymnophiona en Colombia

Familia Caeciliidae (21 spp.)

Familia Dermophiidae (1 sp.)
Familia Rhinatrematidae (3 spp.)
Familia Siphonopidae(4 spp.)
Familia Typhlonectidae (3 spp.)

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Orden: Urodela (Caudata)

Familia Plethodontidae (24 spp.)
Dos gneros en Colombia
Bolitoglossa
Oedipina

Familia Abystomatidae (1
sp.) INTRODUCIDA
Ambystoma mexicanum (Shaw &
Nodder, 1798)
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 Bolitoglossa (22 spp.)

El Gnero Bolitoglossa est constituido por las Salamandras terrestres de desarrollo directo, lo cual genera
importantes restricciones en su distribucin que abarca las reas de bosque natural hasta los pramos. En la
actualidad son descritas 22 especies en el territorio colombiano.

Bolitoglossa adspersa (Peters, 1863)

Bolitoglossa altamazonica (Cope, 1874)
Bolitoglossa biseriata Tanner, 1962
Bolitoglossa capitana Brame & Wake, 1963
Bolitoglossa guaneae Acosta & Gutierrez, 2012
Bolitoglossa hiemalis Lynch, 2001
Bolitoglossa hypacra (Brame & Wake, 1962)
Bolitoglossa leandrae Acevedo,Wake,Mrquez,Silva,Franco & Amzquita 2013.
Bolitoglossa lozanoi Acosta & Restrepo, 2001
Bolitoglossa medemi Brame & Wake, 1972
Bolitoglossa nicefori Brame & Wake, 1963
Bolitoglossa palmata (Werner, 1897)
Bolitoglossa pandi Brame & Wake, 1963
Bolitoglossa phalarosoma Wake & Brame, 1962
Bolitoglossa ramosi Brame & Wake, 1972
Bolitoglossa savagei Brame & Wake, 1963
Bolitoglossa silverstonei Brame & Wake, 1972
Bolitoglossa sima (Vaillant, 1911)
Bolitoglossa tamaense Acevedo,Wake,Mrquez, Silva,& Amzquita 2013.
Bolitoglossa tatamae Acosta & Hoyos, 2006
Bolitoglossa vallecula Brame & Wake, 1963
Bolitoglossa walkeri Brame & Wake, 1972

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Oedipina (2 spp.)

Las salamandras cripticas de Gnero Oedipina estn

pobremente representadas en nuestro pas al igual que las del
gnero Bolitoglossa su estrategia reproductiva involucra el
desarrollo directo; su biologa que est ligada a los ambientes
selvticos y en particular en microambientes asociados la
hojarasca; estos aspectos han contribuido a generar
restricciones sobre el conocimiento de su historia natural y
distribucin en nuestro pas. En la actualidad estn
pobremente representados en las colecciones nacionales.

Oedipina complex (Dunn, 1924)

Oedipina parvipes (Peters, 1879)

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Modern Amphibians
Salamanders: Order Caudata (Urodela)
Approximately 553 living species are found primarily
in northern temperate regions
Most are small, under 15 cm long
Japanese giant salamander is 1.5 meters long

Limbs usually at right angles to trunk

Forelimbs and hindlimbs about equal in length
Burrowing species and some aquatic forms may have
lost their limbs
Carnivorous as both larvae and adults
Feed on worms, small arthropods, and molluscs

Ectotherms with a low metabolic rate

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Modern Amphibians
Breeding Behavior

Some aquatic throughout life cycle

Most have aquatic larvae and terrestrial adults
Internal fertilization in most
Female recovers in cloaca a spermatophore deposited on a
leaf or stick
Aquatic species lay eggs in clusters or stringy masses
Completely terrestrial species deposit eggs in small, grapelike clusters under logs or in soft earth
Terrestrial species undergo direct development
Hatch as miniature adults

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Modern Amphibians
Some North American newts have aquatic larvae that
metamorphose into terrestrial juveniles that again
metamorphose into secondarily aquatic, breeding adults
Some newt populations skip the terrestrial red eft stage
and remain entirely aquatic

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Modern Amphibians
Respiration
Wide array of respiratory mechanisms
Extensive vascular nets in skin that exchange both oxygen
and carbon dioxide
At various stages, may also have external gills, lungs, both
gills and lungs, or neither
Salamanders with aquatic stage hatch with gills and which
are lost at metamorphosis
Several diverse lineages fail to undergo metamorphosis
and retain gills and a fin-like tail
In species with lungs, lungs are present from birth and
become functional following metamorphosis

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Modern Amphibians
Aquatic amphiumas lose gills and respire by lung
Hold nostrils above water surface
Many species of Plethodontidae are lungless and strictly
terrestrial
Respiratory gases may also be exchanged across
vascularized lining of mouth cavity
Lungless salamanders likely evolved in cold streams where
lungs would have been too buoyant

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Modern Amphibians
Paedomorphosis
Descendants retain into adulthood features that were
present only in pre-adult stages of ancestors
Eliminating ancestral adult morphology is a trend found in
salamander evolution
Non-metamorphic species that retain gills, etc. are
perennibranchiate
Obligate perennibranchiates, like the mudpuppy, have
never been observed to metamorphose
Others reach sexual maturity with larval morphology but
can change depending on the conditions

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Modern Amphibians
In Mexico and the U.S.
Ambystoma tigrinum may stay in a gilled stage as an axolotl
When ponds dry up, may metamorphose into a terrestrial
form and migrate to a new pond
Axolotls treated with thyroid hormone will metamorphose
artificially
Pituitary gland appears to be the controlling factor in natural
populations

Paedomorphosis may be partial

Amphiumas shift to lungs but otherwise remains larval in
body form

In some species, larval appendages have been

maintained to preserve a climbing ability
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Anfibios en Colombia

25-43

Modern Amphibians
Frogs and Toads: Order Anura (Salientia)
Approximately 5283 species
Dates from Triassic period, 250 million years ago
Must live near water source
Reproduction mode requires water
Skin is water-permeable skin

Ectothermy prevents anurans from inhabiting polar

and subarctic habitats
All have a tailed larval stage and tailless, jumping
adults (except for 1 species)

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Diversidad por departamentos

En relacin a la distribucin de las familias en Colombia, se tiene que las ranas de

desarrollo directo de la familia Craugastoridae contiene la mayor riqueza.

Acosta-Galvis A.R. 2013, Bolitoglossa adspersa en Lista de los Anfibios de Colombia. V.02.2013.0. www.batrachia.com

25-49

Diversidad por departamentos

Acosta-Galvis A.R. 2013, Bolitoglossa adspersa en Lista de los Anfibios de Colombia. V.02.2013.0. www.batrachia.com

25-50

Modern Amphibians
44 families of frogs and toads
Family Ranidae
Contains the common larger frogs in North America

Family Hylidae
Includes the tree frogs

Family Bufonidae (true toads)

Contains toads with thicker skins and prominent warts

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Modern Amphibians
Habitats and Distribution
Genus Rana are common in temperate and tropical regions
Rana sylvatica, wood frog, spends most of its time on damp
forest floors, returns to pools for breeding
Bullfrogs and green frogs occur in or near permanent water
and swamps
Leopard frog is widespread and commonly studied in
laboratories
Pickerel frog is restricted to certain localities
Anuran are declining worldwide and becoming
geographically fragmented
Cause is not known
Malformed limbs are often associated with infection by
trematodes
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Modern Amphibians
Life Cycle
Most larger frogs are solitary until breeding season
During breeding season, males are especially noisy when
trying to attract a female
Hold forelimbs near body when swimming with powerful
hindlimbs
Surface to breathe with only the head and foreparts
exposed
During winter in temperate climates
Hibernate in soft mud in bottom of pools
During hibernation period, the little energy they use is
provided from stored glycogen and fat
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Modern Amphibians
Frost-tolerant frogs
Prepare for freezing by accumulating glucose and glycerol in body
fluids
Protection from damaging effects of ice-crystal formation

Many are easy prey

Defend themselves by aggression, concealment, and poison
glands
Many species have suffered from changes in the
environment and climate brought about by humans
Climatic changes that reduce water depth at oviposition
sites
Increases ultraviolet exposure of embryos
Also, makes them more susceptible to fungal infection

Decline of some amphibians may be caused by other

amphibians such as Bufo marinus
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Modern Amphibians

Integument and Coloration

Frog skin is thin, moist and attached loosely to the body at

a few points
Skin is composed of an outer stratified epidermis and an
inner spongy dermis
Epidermal layer is shed periodically
Contains deposits of keratin

More terrestrial amphibians have heavier deposits of

keratin
Amphibian keratin is soft

Epidermis has two types of integumentary glands

Mucous glands secrete protective waterproofing
Large serous glands produce a whitish, watery poison

Dendrobatid frogs of South America secrete highly toxic

skin poisons
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Modern Amphibians
Specialized pigment cells, chromatophores, produce
skin color in frogs
Primarily in dermis
Xanthophores (upper region)
Yellow, orange or red pigments

Iridophores (middle region)

Silvery light-reflecting pigment that acts like a mirror

Melanophores (lower region)

Black or brown melanin

Green hue is an interaction of xanthophores

containing yellow pigment and underlying iridophores
Many can adjust color to match with background and
camouflage themselves
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Modern Amphibians

Skeletal and Muscular Systems

Well-developed endoskeleton of bone and cartilage provides

for body support and muscular movements
Movement to land posed new mechanical stress problems
Dramatic changes in the musculoskeletal system for jumping
and swimming
Vertebral column lost much flexibility in order to transmit
force from limbs to body
Extremely shortened body
Only nine trunk vertebrae and a urostyle

Caecilians have not moved toward tetrapod locomotion and

have as many as 285 vertebrae

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Modern Amphibians
Front of the skull, containing the brain, eyes, and nose is
lightweight and flattened
Back of the skull, which contained the gill apparatus in
fishes, is reduced
Posterior limbs have three main joints
Hip, knee and ankle

Foot generally has five rays

Hand is four-rayed
Both have several joints in the digits
System derived from the pattern in rhipidistian lobe-finned fish

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Modern Amphibians
Limb musculature
Anterior, ventral group pulls the limb forward and toward the
midline
Posterior, dorsal group draws the limb backward and away from
body

Myomeres highly modified to support head and brace the

vertebral column

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Modern Amphibians
Respiration and Vocalization
Amphibians use 3 respiratory surfaces for gas exchange in
air
Skin provides cutaneous breathing
Mouth provides buccal breathing
Lungs are usually present in adults

Frogs and toads depend on lung breathing more than

salamanders
Skin is critical during winter hibernation
Carbon dioxide is mostly lost across skin while oxygen is
absorbed across the lungs
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Modern Amphibians
Lungs supplied by pulmonary arteries and return blood to
the left atrium
Frog lungs
Ovoid, elastic sacs
Inner surfaces divide into a network of smaller
chambers
Absorptive surface is 20 cm2/cc of air compared to 300 cm2
for humans
Positive Pressure Breathing
Air moved into lungs by force
Rhythmic throat movements gulp air and force it
backward
Rib cage does not expand to draw air into the lung
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Modern Amphibians
Vocal cords are located in the larynx
Much more developed in males than females
Air is passed back and forth over vocal cords between the
lungs to a large pair of vocal sacs
Most species have unique sound patterns

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Modern Amphibians
Circulation
Circulation is closed with a single pressure pump moving
blood through the peripheral network
Main difference in circuitry is the shift from gill to lung
breathing
Elimination of gills reduced one obstacle to blood flow in
the arterial circuit
Conversion of sixth aortic arch into a pulmonary artery
provided a blood circuit to lungs
Separation of oxygenated blood from deoxygenated blood
circuit is only partial

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Modern Amphibians
Frog Heart
Single undivided ventricle and two separate atria
Blood from body enters through the sinus venosus and right
atrium
Blood from the lung enters left atrium
Both atria contract at the same time
Pumps blood into ventricle
Ventricle contracts
Moving blood moves to the lungs or body
Although there is no septum
Deoxygenated blood goes primarily to the lungs
Oxygenated blood goes mostly to the body

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Modern Amphibians
Right and left atria contract asynchronously
Oxygenated and deoxygenated blood remains mostly
separated when entering ventricle
Blood separation aided by
Spiral valve which divides systemic and pulmonary flows in the conus
arteriosus
Different blood pressure in the pulmonary and systemic blood vessels
leaving the conus arteriosus

Exact mechanism and precision of separation of oxygenated

and deoxygenated blood in the conus arteriosus remain
unclear

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Modern Amphibians
Feeding and Digestion
Most adult are carnivorous
Feed on insects, spiders, worms, slugs, etc.

Catch prey with a tongue that is attached at the front of

mouth
Free end of tongue is glandular
Sticky secretion adheres to prey

Any teeth that are present function to hold prey

Do not bite or chew

Short digestive tract produces enzymes for digesting fats,

carbohydrates and proteins
Larval stages of tadpoles are usually herbivorous
Digestive tract is relatively long
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Modern Amphibians
Nervous System and Special Senses
Nervous System
Brain has 3 fundamental parts
Forebrain or telencephalon
Interprets smells

Midbrain or mesencephalon
Perceives vision

Hindbrain or rhombencephalon
Perceives hearing and balance

Brain is gradually assuming more information processing

independent of spinal cord
However, a headless frog still has highly coordinated
behavior based on spinal cord alone
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Modern Amphibians
Forebrain contains the olfactory center
Remainder of forebrain has little importance
Complex integrative activities occur in the midbrain optic
lobes
Hindbrain is divided into an anterior cerebellum and a
posterior medulla
Cerebellum, which coordinates equilibrium and
movement, is not well developed in frogs
All sensory neurons except those for vision and smell pass
through the medulla which is on the anterior end of the
spinal cord

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Modern Amphibians
Medulla has centers for auditory reflexes, respiration,
swallowing, and vasomotor control
Special Senses
Pressure-sensitive lateral line found only in amphibian
larvae and aquatic adults
Ear specialized for detecting airborne sounds
Large tympanic membrane or eardrum passes vibrations to
inner ear via the columella
Inner ear has a utricle with three semicircular canals and a
saccule with a lagena
Lagena is covered with a tectorial membrane that is similar
to the mammalian cochlea
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Modern Amphibians
Frogs are sensitive to low-frequency sound energy under
4000 Hz (cycles per second)
Except for blind caecilians, vision is dominant sense in
many amphibians
Lachrymal glands and eyelids
Keep eye moist, free of dust, and protected
Cornea and lens
Bend light rays to focus an image on the retina

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Modern Amphibians
At rest, the fish eye focuses on near objects and the frog
eye focuses on distant objects
Amphibian retina contains rods and cones
Cones provide frogs with color vision

Iris can rapidly change aperture to adjust to light levels

Upper eyelid is fixed
Lower eyelid is folded into a transparent nictitating
membrane
Other sensory receptors include chemical receptors in skin,
taste buds on tongue, and olfactory epithelium in nasal
cavity

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Modern Amphibians
Reproduction and Development

Frogs and toads are ectothermic

Breed, feed, and grow during warm seasons
In spring, males call to attract females
When eggs are mature, females enter the water and the
males clasp them in amplexus
After fertilization, jelly layers of egg absorb water and swell
Eggs usually laid in large masses

Development begins immediately

Tadpole may hatch in 69 days

Tadpole head has horny jaws for feeding and a ventral

adhesive disc for clinging to objects
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Modern Amphibians
Eggs of most hatch into tadpoles with a long, finned
tail, no legs, internal and external gills and
specialized mouthparts for (usually) herbivorous
feeding
Tadpoles look and act entirely different from adult
frogs
Perennibranchiate condition never occurs in frogs
and toads

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Modern Amphibians
3 pairs of external gills develop into internal gills covered with
a flap of skin
On right side of a tadpole, operculum fuses with body wall
On left side, a spiracle remains
Water enters mouth, flows past gills, and then out spiracle
Metamorphosis
Hindlegs are first to appear
Forelegs temporarily hidden in folds of operculum
Tail is resorbed
Intestine becomes shorter
Mouth transforms to the adult condition
Lungs develop and gills are resorbed

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Modern Amphibians
Males migrate back to breeding ponds or streams
Tropical anurans have different reproductive strategies
Some lay eggs in foam masses that float on surface of
water
Some deposit eggs on leaves over-hanging ponds and
streams into which tadpoles drop
Other place eggs in water trapped in tree cavities or
water-filled chambers of bromeliads
Poison-dart frog tend to their eggs
Tadpoles hatch on their back and can be carried for
varying lengths of time

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Modern Amphibians
Marsupial frogs carry eggs in a pouch on the back
Eleutherodactylus mate on land and eggs hatch directly
into froglets
Some salamanders have a strong homing instinct,
returning to the same pool to breed
Stimulation to migrate depends on hormone changes and
sensitivity to temperature and humidity

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Modern Amphibians
Clasificacin Taxonmica
Phylum: Chordata
Subfilo: Vertebrata
Superclase: Gnathostomata
Class Amphibia
Order Gymnophiona Apodos
Order Caudata Urodelos
Order Anura Salientia

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