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Amphibians
Amphibians
Amphibians
īŽ First vertebrates
adapted to land
environment
īŽ Preceded onto land
by plants, insects,
snails
īŽ ~350 million years
ago
Amphibians
īŽ Not completely land
adapted
īŽ Structurally between
fish & reptiles
īŽ Most cannot be far
removed from water
Amphibians
īŽ Things to be
considered in water to
land movement:
īŽ 1) O2 content of air
greater than water
(less effort to take up
O2 from air)
īŽ 2) air less dense (lack
of body support)
Amphibians
īŽ 3) greater
temperature
fluctuations
(unpredictable
temperature
extremes)
īŽ 4) greater habitat
variety (cover, shelter,
breeding areas)
Earliest Amphibians
īŽ Appeared to have arisen from lobe-finned fishes
īŽ Were abundant during period when amphibians
first appeared
īŽ Possessed characteristics that benefited first
amphibians
Earliest Amphibians
īŽ Strong, mobile fins - used as stabilizers during
swimming
īŽ Used as paddles to pull themselves across land
in search of water
Earliest Amphibians
īŽ Lungs - lived in freshwater lakes & streams
during periods of alternating floods, droughts
īŽ Gills useless in low dissolved O2 areas or dry
areas
īŽ Surviving fishes of this period developed simple
lung
Earliest Amphibians
īŽ Salamander-like
īŽ Retained many characteristics of fish
īŽ Skull, tooth structure similar - snout more elongated
īŽ Fish tail with fin rays
īŽ Opercular bones present
Earliest Amphibians
īŽ Several differences
īŽ Jointed limbs with 5 digits (pentadactyl)
īŽ Ears for picking up airborne sounds (replace lateral
line)
īŽ Improved olfactory powers - for airborne odors
Amphibian Diversity
īŽ Wet, warm conditions
followed alternating
dry-wet
īŽ Perfect conditions for
amphibians - radiated
off into many lines
īŽ Little pressure to
become better land-
adapted because of
water everywhere
Amphibian Diversity
īŽ Became better
adapted to water
īŽ Bodies became more
flattened
īŽ Some developed
webbed feet
īŽ Some developed
weaker legs &
stronger tails for more
efficient swimming
Amphibian Diversity
īŽ Produced 3 basic
groups of amphibians
present today
īŽ Order Anura (without
tail) - frogs, toads
īŽ Order Caudata (with
tail, Urodela) -
salamanders & newts
īŽ Order Gymnophiona
(naked snake) -
caecilians
Lungs & Circulation
īŽ Improved on
efficiency of simple
lung
īŽ Supplied it with
capillary network to
improve O2 uptake
īŽ Double circulation
īŽ Systemic
īŽ Pulmonary
Jointed Limbs
īŽ Strengthening of limbs - 5 digits, joints
īŽ Improvements in pelvic, pectoral girdles
īŽ Developed for continued survival in aquatic
environments, but pre-adapted others for life on
land
Amphibian Characteristics
īŽ Aquatic ectotherms - body temperature varies
with the environment
īŽ Difficult to withstand temperature extremes of
terrestrial environment
Amphibian Characteristics
īŽ Skin must be kept moist - obtain much of their O2
through the skin
īŽ Thin, unprotected from desiccation
Amphibian Characteristics
īŽ Mode of reproduction requires water or moisture
īŽ Eggs fertilized externally
īŽ Larvae must pass through an aquatic tadpole
stage
Amphibian Characteristics
īŽ Skelton mostly bony
īŽ Varying number of
vertebrae
īŽ Some may have ribs
īŽ Notochord does not
persist
Amphibian Characteristics
īŽ Great variety of body
forms
īŽ Elongated trunk with
distinct head, neck,
tail, 4 legs (tetrapods)
īŽ Frogs with compact
depressed body,
fused head & trunk,
no neck
Amphibian Characteristics
īŽ Some legless forms
without distinct body
regions (caecilians)
Amphibian Characteristics
īŽ Webbed feet common
īŽ No “true” claws or
nails on digits
Amphibian Characteristics
īŽ Skin smooth and
moist
īŽ No ectodermal scales
(some have
concealed dermal
scales)
īŽ Many glands - some
poisonous
Amphibian Characteristics
īŽ Two nostrils open into
mouth cavity
īŽ Breathe with mouth
closed
Amphibian Characteristics
īŽ Respiration via lungs,
skin, gills
īŽ Some salamanders
lack lungs
īŽ Skin well vascularized
īŽ External gills in larvae
may persist
throughout life in
some
Order Gymnophiona
īŽ Caecilians
īŽ Body long, slender,
wormlike
īŽ Limbs and limb
girdles absent
īŽ Many vertebrae, long
ribs
īŽ Tail short or absent
Order Gymnophiona
īŽ Burrowers, tropical
forests of S. Amer.,
Africa, SE Asia
īŽ Small, mostly
degenerate eyes
(mostly blind as
adults)
īŽ Sensory tentacles on
snout
īŽ Tail short or absent,
terminal anus
Order Gymnophiona
īŽ Feed on worms, small
invertebrates
īŽ Fertilization internal,
eggs deposited in
moist soil near water
īŽ Some species guard
eggs (body folds)
Order Gymnophiona
īŽ May or may not have
aquatic larval stage
īŽ Complete larval
development may
occur in egg
īŽ Viviparity common in
more advanced forms
īŽ Embryos eat oviduct
wall
Order Caudata
īŽ Salamanders and
newts
īŽ Least specialized
amphibians
īŽ Found in almost all
tropical and northern
temperate regions of
world
īŽ Most species in N.
America
Order Caudata
īŽ Typically small, most
<15 cm long
īŽ Aquatic species may
be much larger
īŽ Carnivorous
Japanese giant
salamander may be
>1.5 m long
Order Caudata
īŽ No scales
īŽ 2 pairs of equal-sized limbs
īŽ Primitive limbs set at right angles to body
Order Caudata
īŽ Some species have
rudimentary limbs
īŽ Amphiuma has tiny
limbs
īŽ Sirens have minute
forelimbs, no
hindlimbs
īŽ Different enough for
separate order?
Order Caudata
īŽ Amphiuma
īŽ Siren
Order Caudata
īŽ Carnivorous - prey on
worms, small
arthropods, small
molluscs
īŽ Most eat only things
that are moving
Order Caudata
īŽ Food is protein-rich -
do not store great
quantities of fat or
glycogen
īŽ Ectotherms, low
metabolic rate - low
food demand
Order Caudata
īŽ Most salamanders
are terrestrial
īŽ Live in moist places
under stones, rotten
logs
īŽ Seldom far from water
Order Caudata
īŽ Some salamanders
wholly aquatic
throughout life cycle
Order Caudata - reproduction
īŽ Little diversity of breeding habits
īŽ Eggs of most salamanders fertilized internally
īŽ Female picks up packet of sperm
(spermatophore) previously deposited by male
onto some substrate
Order Caudata - reproduction
īŽ Ritualistic behaviors help ensure female
receptivity
īŽ Additional behaviors ensure recovery of
sperm mass by female
Order Caudata - reproduction
īŽ Aquatic species lay
eggs in clusters or
stringy masses in
water
īŽ Often attached to
submerged objects
Order Caudata - reproduction
īŽ Terrestrial species lay
eggs in tiny, grape-
like clusters under
logs, in vegetation, or
in excavations in soft
earth
Order Caudata - development
īŽ Some species remain
(female) and guard
eggs until hatching
Order Caudata - development
īŽ Larvae which hatch
are not tadpole-like
īŽ Resemble parents
Order Caudata - development
īŽ Still must undergo
metamorphosis to
attain adult form
īŽ Gills generally lost
and fin-like tail
modified during
metamorphosis
Order Caudata - development
īŽ Size and type of gills
dependent on larval
habitat
īŽ Larger in lake, pond,
smaller in stream
habitat
īŽ Lungs usually replace
gills in adults (except
in aquatic forms)
Order Caudata - development
īŽ Development
variation: American
newts
īŽ Have a terrestrial
stage between
aquatic larva and
aquatic breeding adult
īŽ Red-spotted newt -
red eft stage - 1-3
years
Order Caudata - no lungs
īŽ Some salamanders
don’t bother
developing lungs
when they lose gills
īŽ Family
Plethodontidae -
lungless salamanders
- most of familiar N.
Amer. salamanders
Order Caudata - no lungs
īŽ Only adult vertebrates
with neither lungs nor
gills
īŽ Skin highly
vascularized -
vascular net
īŽ Extension of
capillaries into
epidermis OR
thinning of epidermis
over dermal
capillaries
Order Caudata - no lungs
īŽ Provides 90-95%
of gas exchange
īŽ Other 5-10% from
capillaries inside
mouth
īŽ Buccopharyngeal
breathing
Order Caudata - no lungs
īŽ Originated in swift,
cool, highly
oxygenated streams
in Appalachia
īŽ Cutaneous respiration
could provide enough
O2
īŽ Lungs would provide
too much buoyancy in
this environment
Order Caudata - metamorphosis
īŽ Most salamanders
undergo
metamorphosis, but
some retain gills or
other larval
characteristics after
becoming sexually
mature
īŽ Neoteny (young, to
extend)
Order Caudata - metamorphosis
īŽ Some retain
characters for entire
lifetime - permanent
larvae
īŽ Obligatory neoteny
īŽ Necturus - mudpuppy
īŽ Live on bottom of
ponds, lakes, large
rivers
Order Caudata - metamorphosis
īŽ Either developing
tissues fail to respond
to thyroxin
(metamorphosis
hormone) OR pituitary
gland doesn’t develop
fully to produce
thyrotropin that
stimulates thyroid
gland to produce
thyroxin
Order Caudata - metamorphosis
īŽ Others retain
characteristics, but
can change if
conditions are
favorable
īŽ Facultative neoteny
īŽ Achieved by delaying
somatic development,
but not sexual
maturation
Order Caudata - metamorphosis
īŽ Example of facultative
neoteny: Ambystoma
species
īŽ Tiger salamander
subspecies called
American axolotl
īŽ Mexico and SW US
Order Caudata - metamorphosis
īŽ Remains aquatic, gill-
breathing, fully
reproductive larval
form unless water
begins to dry up
īŽ Metamorphoses to
adult, loses gills,
develops lungs,
assumes appearance
of ordinary
salamander
Order Caudata - metamorphosis
īŽ Can be artificially
induced to
metamorphose by
treating then with
thyroxin
īŽ Essential for normal
metamorphosis in all
amphibians
Order Caudata - territoriality
īŽ Some salamanders
display territorial
behavior
īŽ Use scent to mark
territories -
Plethodontidae
īŽ Males of some
species defend
territories against
conspecifics
Order Caudata - territoriality
īŽ Attack intruders with
foreign scent, but not
neighbors with
familiar scent
īŽ Nasolabial grooves -
chemosensory
system
Order Caudata - territoriality
īŽ Territory defended
against intruders at
expense of reduced
energy intake
īŽ Attack nose of
intruders - ruin
chemosensory
system?
īŽ Less able to feed,
smaller
Order Caudata - toxicity
īŽ Many newts contain
mucous glands in skin
that secrete irritating
or toxic materials
īŽ E.g., red eft
(terrestrial red-spotted
newt) produces a
potent neurotoxin
Order Caudata - toxicity
īŽ Produces unpleasant
numbness or tingling
sensation in animals
touching it
īŽ Secretions have been
known to kill animals
that ingested them
(including humans)
Order Caudata - toxicity
īŽ Species may
advertise their toxicity
with conspicuous
aposematic, or
warning, colors and
behaviors
īŽ Red eft is brilliant red-
orange
Order Caudata - toxicity
īŽ Other species that are
more plainly often flip
when attacked to
display brightly
colored bellies
Order Caudata - toxicity
īŽ Some species have
adapted to mimic toxic
forms
īŽ E.g., red-backed
salamander has a red
color morph (normally
only red on back) that
resembles red eft
īŽ Studies: red morph
suffers less bird
predation than normal
color morph
Order Anura
īŽ Frogs & toads
īŽ >3400 species
īŽ Most familiar,
successful
amphibians
īŽ Obvious, common
characteristic - lack of
tail in adult
īŽ All pass through tailed
larval stage
Order Anura
īŽ Specialized for
jumping mode of
locomotion
Order Anura
īŽ Further
distinguished from
Caudata by:
īŽ Larvae - tadpole or
polliwog stage
īŽ Metamorphosis
during
development
Order Anura
īŽ Tadpole
īŽ Long, finned tail
īŽ Both internal, external
gills
īŽ No legs
īŽ Specialized
mouthparts for
herbivory (salamander
larvae are
carnivorous)
īŽ Highly specialized
internal anatomy
Order Anura
īŽ Striking
metamorphosis from
tadpole to adult
īŽ Neoteny never
exhibited in frogs or
toads
Order Anura
īŽ Represent
specialized side
branch of
amphibian
evolution
īŽ Not good
representatives of
vertebrate body
plan
Order Anura
īŽ No visible neck
īŽ Caudal vertebrae
fused (to form
urostyle)
īŽ Ribs generally
absent
īŽ Hindlegs much
enlarged
Order Anura
īŽ 21 different
families
īŽ Common North
American frogs
īŽ Ranidae - grass
frogs
īŽ Hylidae - tree frogs
Order Anura
īŽ Bufonidae - true toads
īŽ Short legs, stout
bodies, and thick
skins, usually with
prominent warts
īŽ “toad” often used to
refer to more
terrestrial members of
many families
Order Anura
īŽ Largest anuran -
West African goliath
frog
īŽ >30 cm from nose to
anus, ~1 m with
outstretched legs
īŽ 3 kg+ (up to 15 lbs.)
īŽ Eats animals as large
as rats, ducks
Order Anura
īŽ Smallest anuran -
Cuban leafclimber
frog
īŽ ~1 cm - covered by a
dime
Order Anura
īŽ Bullfrog - up to 20 cm
long
Order Anura
īŽ Most abundant,
successful frogs belong
to genus Rana
īŽ 260 species
īŽ Temperate, tropical
regions (not New
Zealand, oceanic
islands, southern S.
Amer.)
īŽ Most often near water
Order Anura
īŽ Most larger frogs are
solitary except
during breeding
season
īŽ Males call noisily for
females
Order Anura
īŽ Winter - hibernate in
soft mud at bottom
of pools, streams
īŽ Low energy demand
īŽ Glycogen, fat stored
during spring,
summer to provide
energy during winter
Order Anura
īŽ Tree frogs winter
under humus on
forest floor
īŽ Tolerant of low
temperatures
Order Anura
īŽ May survive prolonged
freezing of all
extracellular fluid (35%
of body water)
īŽ Accumulate glucose,
glycerol in body fluids
to protect tissue from
damage from ice
crystals
Order Anura - enemies
īŽ Natural enemies -
snakes, aquatic
birds, turtles,
racoons, humans
īŽ Usually
defenseless
Order Anura - enemies
īŽ Some tropical,
subtropical forms
are aggressive,
jump at and bite
attackers
īŽ Others play dead
Order Anura - enemies
īŽ Others blow up
lungs - difficult to
swallow
īŽ Jumping is best
defense
īŽ Some species
have poison
glands
Order Anura - skin
īŽ Thin and moist
īŽ Attached loosely to
body only at
certain points
Order Anura - skin
īŽ Stratified epidermis,
inner dermis
īŽ Outer layer of
epidermis shed
periodically
īŽ Deposits of soft
keratin - protection
against water loss,
some abrasion
Order Anura - skin
īŽ Dermis, lower
epidermis have two
types of glands
īŽ Mucous - secrete
protective
waterproofing mucous
īŽ Serous - secrete
whitish, watery poison
Order Anura - skin
īŽ Poisons of
dendrobatid frogs
(small, S. Amer.
frogs) are most lethal
animal secretions
known
īŽ More poisonous than
venoms of sea
snakes, any
arachnids
Order Anura - skin
īŽ Dermis also contains
several types of
pigment cells -
chromatophores
īŽ Many can adjust color
to blend in with
background
Order Anura
īŽ Entire
musculoskeletal
system is specialized
for jumping and
swimming, via thrusts
of the hindlimbs
Order Anura - skeleton
īŽ Vertebral column - rigid structure - lost much of
flexibility found in fishes
īŽ Transmits forces from limbs to body
Order Anura - skeleton
īŽ Body much shorter - only 9 trunk vertebrae,
several fused caudal vertebrae
īŽ Caecilians have 285 vertebrae
Order Anura - skeleton
īŽ Skull much lighter than in fishes
īŽ Changed for improved special senses, feeding,
breathing
Order Anura - muscles
īŽ Limb muscles -
probably derived from
radial muscles that
move fish fins up and
down
īŽ Arranged in
antagonistic groups
(actions of one
oppose another)
Order Anura - muscles
īŽ Abductor - away from
median axis
īŽ Adductor - toward
median axis
īŽ Flexor - bend part
īŽ Extensor - straighten
part
īŽ Rotators - rotate
Order Anura - muscles
īŽ Myotome muscles
modified to provide
support to various
portions of body
(head, belly)
Order Anura - lungs
īŽ Frogs and toads are
more highly
dependent on lung
breathing than are the
salamanders
īŽ Several
disadvantages of
cutaneous respiration
Order Anura - lungs
īŽ 1) skin must be kept
thin, moist - too
delicate for wholly
terrestrial life
īŽ 2) gas exchange rate
across skin nearly
constant - cannot vary
with increased activity
Order Anura - lungs
īŽ Cutaneous respiration
important to anurans
during winter
hibernation
īŽ Also, under normal
conditions most CO2
is lost across skin
surface,most O2 taken
up by lungs
Order Anura - lungs
īŽ Lungs are elastic
sacs with inner areas
subdivided into many
chambers
īŽ Not as much internal
surface area relative
to higher vertebrates
Order Anura - lungs
īŽ Frogs are positive-
pressure breathers
īŽ Breathe by
swallowing air -
forcing it into their
lungs
īŽ All higher vertebrates
use negative-
pressure system
īŽ Expansion of lungs to
pull air inward
Order Anura - lungs
īŽ Both male and
female anurans
have vocal cords
īŽ Males are better
developed
īŽ Located in larynx -
voicebox
Order Anura - lungs
īŽ Sound produced by
passing air back and
forth between lungs
and large pair of air
sacs (vocal pouches)
in floor of mouth
īŽ Pouches also serve
as resonators in
males
īŽ Function to attract
mates
Order Anura - circulation
īŽ Shift from gill to
lung breathing
solved some
circulatory
problems, but
caused others
īŽ Eliminated major
obstacle to arterial
blood flow - gills
Order Anura - circulation
īŽ Needed to develop
new circuitry to serve
lungs - pulmonary
circulation
īŽ 6th aortic arch
developed into
pulmonary artery
īŽ New pulmonary veins
developed for return
Order Anura - circulation
īŽ More difficult
evolutionary problem
- separate
oxygenated from
deoxygenated blood
īŽ Amphibians have
made it halfway
īŽ Two atria, but single
ventricle - potential
mixing
Order Anura - circulation
īŽ Actually little mixing of
oxygenated,
deoxygenated blood
īŽ Spiral valve in conus
arteriosus (leading
from ventricle) may
help maintain
selective distribution
īŽ Controversial - defied
complete analysis
Order Anura - feeding
īŽ Adult frogs are
carnivorous
īŽ Feed on insects,
spiders, worms,
slugs, snails,
millipedes, anything
moving and small
enough to eat
Order Anura - feeding
īŽ Snap at prey with protrusible tongue
īŽ Attach at front, free behind
īŽ Free end very glandular, produces sticky
secretion, adheres to prey
Order Anura - feeding
īŽ Teeth on premaxillae,
maxillae, vomers
īŽ Not biting or chewing
īŽ Used to prevent prey
escape
Order Anura - digestion
īŽ Digestive tract
relatively short in
adults, longer in
tadpoles
īŽ Carnivore adaptation
vs. herbivore
adaptation
īŽ Variety of enzymes
produced
Order Anura - nervous system
īŽ Dramatic changes
occurred in nervous
system as vertebrates
moved onto land
īŽ Increased
cephalization to cope
with increased need
for information
processing
Order Anura - nervous system
īŽ Increased senses of
smell, vision, hearing,
increased demand for
balance
īŽ Has resulted in loss of
independence of the
spinal ganglia
Order Anura - nervous system
īŽ Spinal ganglia can
serve as “mini brains”
in lower vertebrates
īŽ Coordinate actions
without involving
brains
īŽ But even with some
loss of
independenceâ€Ļâ€Ļ
Order Anura - nervous system
īŽ Headless frog with
only spinal cord intact
still maintains high
degree of purposeful,
highly coordinated
behavior
īŽ Normal posture, raise
leg to remove
disturbing object
(other if one is
restrained)
Order Anura - nervous system
īŽ Two senses highly
advanced from fishes
and most other
amphibians
īŽ Hearing and sight
Order Anura - hearing
īŽ Lateral line system
remains in aquatic
larvae, but not most
adults
īŽ Role is replaced by
ear in most adult
frogs
īŽ Obvious structure -
tympanum - serves as
eardrum
Order Anura - hearing
īŽ Picks up vibrations
of airborne sounds
and transmits them
through middle ear
to inner ear
Order Anura - hearing
īŽ Contains
semicircular canals
and structures
similar, but simpler,
than mammalian
cochlea
Order Anura - hearing
īŽ Hearing of species
is most sensitive to
frequency range of
their mating calls
īŽ Peeper - higher
frequency
īŽ Bullfrog - lower
frequency
Order Anura - sight
īŽ Eye is dominant
special sense in
most amphibians
īŽ Especially good in
frogs
īŽ Fish eye had to be
modified for land
use
Order Anura - sight
īŽ Lachrymal glands,
eyelids evolved to
keep eye moist,
dust-free, shielded
from injury
Order Anura - sight
īŽ Upper and lower
eyelids
īŽ Upper fixed
īŽ Lower with folded
nictitating
membrane that
moves across eye
surface
Order Anura - sight
īŽ Cornea exposed to
air became an
important refractive
surface
īŽ Removed some of
light bending and
focusing burden
from lens
Order Anura - sight
īŽ Lens normally
focused on distant
objects
īŽ Moved to focus on
nearby objects
īŽ Opposite of
arrangement in
fishes
Order Anura - reproduction
īŽ Breeding is first
desire after coming
out of hibernation
īŽ Males migrate to
water first -
olfactory cues
īŽ Croak, call to
attract females
Order Anura - reproduction
īŽ Females enter
water when eggs
mature and are
clasped by males -
amplexus
īŽ Females lay eggs
and males fertilize
them
Order Anura - reproduction
īŽ Eggs laid in large
masses, anchored
to vegetation
īŽ Jelly layer absorbs
water and swells
after fertilization
īŽ Development
occurs quickly
Order Anura - reproduction
īŽ Tadpole hatches from
protective jelly coat
within 6-9 days
(temperature-
dependent)
īŽ Head and body, tail,
ventral mouth with
horny jaws, ventral
adhesive disc, gills,
etc.
Order Anura - reproduction
īŽ Length of tadpole
stage is species-
dependent
īŽ Leopard frog
completes
metamorphosis within
3 months
īŽ Bullfrog takes 2-3
years to complete
process

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Amphibians.

  • 3. Amphibians īŽ First vertebrates adapted to land environment īŽ Preceded onto land by plants, insects, snails īŽ ~350 million years ago
  • 4. Amphibians īŽ Not completely land adapted īŽ Structurally between fish & reptiles īŽ Most cannot be far removed from water
  • 5. Amphibians īŽ Things to be considered in water to land movement: īŽ 1) O2 content of air greater than water (less effort to take up O2 from air) īŽ 2) air less dense (lack of body support)
  • 7. Earliest Amphibians īŽ Appeared to have arisen from lobe-finned fishes īŽ Were abundant during period when amphibians first appeared īŽ Possessed characteristics that benefited first amphibians
  • 8. Earliest Amphibians īŽ Strong, mobile fins - used as stabilizers during swimming īŽ Used as paddles to pull themselves across land in search of water
  • 9. Earliest Amphibians īŽ Lungs - lived in freshwater lakes & streams during periods of alternating floods, droughts īŽ Gills useless in low dissolved O2 areas or dry areas īŽ Surviving fishes of this period developed simple lung
  • 10. Earliest Amphibians īŽ Salamander-like īŽ Retained many characteristics of fish īŽ Skull, tooth structure similar - snout more elongated īŽ Fish tail with fin rays īŽ Opercular bones present
  • 11. Earliest Amphibians īŽ Several differences īŽ Jointed limbs with 5 digits (pentadactyl) īŽ Ears for picking up airborne sounds (replace lateral line) īŽ Improved olfactory powers - for airborne odors
  • 12. Amphibian Diversity īŽ Wet, warm conditions followed alternating dry-wet īŽ Perfect conditions for amphibians - radiated off into many lines īŽ Little pressure to become better land- adapted because of water everywhere
  • 13. Amphibian Diversity īŽ Became better adapted to water īŽ Bodies became more flattened īŽ Some developed webbed feet īŽ Some developed weaker legs & stronger tails for more efficient swimming
  • 14. Amphibian Diversity īŽ Produced 3 basic groups of amphibians present today īŽ Order Anura (without tail) - frogs, toads īŽ Order Caudata (with tail, Urodela) - salamanders & newts īŽ Order Gymnophiona (naked snake) - caecilians
  • 15. Lungs & Circulation īŽ Improved on efficiency of simple lung īŽ Supplied it with capillary network to improve O2 uptake īŽ Double circulation īŽ Systemic īŽ Pulmonary
  • 16. Jointed Limbs īŽ Strengthening of limbs - 5 digits, joints īŽ Improvements in pelvic, pectoral girdles īŽ Developed for continued survival in aquatic environments, but pre-adapted others for life on land
  • 17. Amphibian Characteristics īŽ Aquatic ectotherms - body temperature varies with the environment īŽ Difficult to withstand temperature extremes of terrestrial environment
  • 18. Amphibian Characteristics īŽ Skin must be kept moist - obtain much of their O2 through the skin īŽ Thin, unprotected from desiccation
  • 19. Amphibian Characteristics īŽ Mode of reproduction requires water or moisture īŽ Eggs fertilized externally īŽ Larvae must pass through an aquatic tadpole stage
  • 20. Amphibian Characteristics īŽ Skelton mostly bony īŽ Varying number of vertebrae īŽ Some may have ribs īŽ Notochord does not persist
  • 21. Amphibian Characteristics īŽ Great variety of body forms īŽ Elongated trunk with distinct head, neck, tail, 4 legs (tetrapods) īŽ Frogs with compact depressed body, fused head & trunk, no neck
  • 22. Amphibian Characteristics īŽ Some legless forms without distinct body regions (caecilians)
  • 23. Amphibian Characteristics īŽ Webbed feet common īŽ No “true” claws or nails on digits
  • 24. Amphibian Characteristics īŽ Skin smooth and moist īŽ No ectodermal scales (some have concealed dermal scales) īŽ Many glands - some poisonous
  • 25. Amphibian Characteristics īŽ Two nostrils open into mouth cavity īŽ Breathe with mouth closed
  • 26. Amphibian Characteristics īŽ Respiration via lungs, skin, gills īŽ Some salamanders lack lungs īŽ Skin well vascularized īŽ External gills in larvae may persist throughout life in some
  • 27. Order Gymnophiona īŽ Caecilians īŽ Body long, slender, wormlike īŽ Limbs and limb girdles absent īŽ Many vertebrae, long ribs īŽ Tail short or absent
  • 28. Order Gymnophiona īŽ Burrowers, tropical forests of S. Amer., Africa, SE Asia īŽ Small, mostly degenerate eyes (mostly blind as adults) īŽ Sensory tentacles on snout īŽ Tail short or absent, terminal anus
  • 29. Order Gymnophiona īŽ Feed on worms, small invertebrates īŽ Fertilization internal, eggs deposited in moist soil near water īŽ Some species guard eggs (body folds)
  • 30. Order Gymnophiona īŽ May or may not have aquatic larval stage īŽ Complete larval development may occur in egg īŽ Viviparity common in more advanced forms īŽ Embryos eat oviduct wall
  • 31. Order Caudata īŽ Salamanders and newts īŽ Least specialized amphibians īŽ Found in almost all tropical and northern temperate regions of world īŽ Most species in N. America
  • 32. Order Caudata īŽ Typically small, most <15 cm long īŽ Aquatic species may be much larger īŽ Carnivorous Japanese giant salamander may be >1.5 m long
  • 33. Order Caudata īŽ No scales īŽ 2 pairs of equal-sized limbs īŽ Primitive limbs set at right angles to body
  • 34. Order Caudata īŽ Some species have rudimentary limbs īŽ Amphiuma has tiny limbs īŽ Sirens have minute forelimbs, no hindlimbs īŽ Different enough for separate order?
  • 36. Order Caudata īŽ Carnivorous - prey on worms, small arthropods, small molluscs īŽ Most eat only things that are moving
  • 37. Order Caudata īŽ Food is protein-rich - do not store great quantities of fat or glycogen īŽ Ectotherms, low metabolic rate - low food demand
  • 38. Order Caudata īŽ Most salamanders are terrestrial īŽ Live in moist places under stones, rotten logs īŽ Seldom far from water
  • 39. Order Caudata īŽ Some salamanders wholly aquatic throughout life cycle
  • 40. Order Caudata - reproduction īŽ Little diversity of breeding habits īŽ Eggs of most salamanders fertilized internally īŽ Female picks up packet of sperm (spermatophore) previously deposited by male onto some substrate
  • 41. Order Caudata - reproduction īŽ Ritualistic behaviors help ensure female receptivity īŽ Additional behaviors ensure recovery of sperm mass by female
  • 42. Order Caudata - reproduction īŽ Aquatic species lay eggs in clusters or stringy masses in water īŽ Often attached to submerged objects
  • 43. Order Caudata - reproduction īŽ Terrestrial species lay eggs in tiny, grape- like clusters under logs, in vegetation, or in excavations in soft earth
  • 44. Order Caudata - development īŽ Some species remain (female) and guard eggs until hatching
  • 45. Order Caudata - development īŽ Larvae which hatch are not tadpole-like īŽ Resemble parents
  • 46. Order Caudata - development īŽ Still must undergo metamorphosis to attain adult form īŽ Gills generally lost and fin-like tail modified during metamorphosis
  • 47. Order Caudata - development īŽ Size and type of gills dependent on larval habitat īŽ Larger in lake, pond, smaller in stream habitat īŽ Lungs usually replace gills in adults (except in aquatic forms)
  • 48. Order Caudata - development īŽ Development variation: American newts īŽ Have a terrestrial stage between aquatic larva and aquatic breeding adult īŽ Red-spotted newt - red eft stage - 1-3 years
  • 49. Order Caudata - no lungs īŽ Some salamanders don’t bother developing lungs when they lose gills īŽ Family Plethodontidae - lungless salamanders - most of familiar N. Amer. salamanders
  • 50. Order Caudata - no lungs īŽ Only adult vertebrates with neither lungs nor gills īŽ Skin highly vascularized - vascular net īŽ Extension of capillaries into epidermis OR thinning of epidermis over dermal capillaries
  • 51. Order Caudata - no lungs īŽ Provides 90-95% of gas exchange īŽ Other 5-10% from capillaries inside mouth īŽ Buccopharyngeal breathing
  • 52. Order Caudata - no lungs īŽ Originated in swift, cool, highly oxygenated streams in Appalachia īŽ Cutaneous respiration could provide enough O2 īŽ Lungs would provide too much buoyancy in this environment
  • 53. Order Caudata - metamorphosis īŽ Most salamanders undergo metamorphosis, but some retain gills or other larval characteristics after becoming sexually mature īŽ Neoteny (young, to extend)
  • 54. Order Caudata - metamorphosis īŽ Some retain characters for entire lifetime - permanent larvae īŽ Obligatory neoteny īŽ Necturus - mudpuppy īŽ Live on bottom of ponds, lakes, large rivers
  • 55. Order Caudata - metamorphosis īŽ Either developing tissues fail to respond to thyroxin (metamorphosis hormone) OR pituitary gland doesn’t develop fully to produce thyrotropin that stimulates thyroid gland to produce thyroxin
  • 56. Order Caudata - metamorphosis īŽ Others retain characteristics, but can change if conditions are favorable īŽ Facultative neoteny īŽ Achieved by delaying somatic development, but not sexual maturation
  • 57. Order Caudata - metamorphosis īŽ Example of facultative neoteny: Ambystoma species īŽ Tiger salamander subspecies called American axolotl īŽ Mexico and SW US
  • 58. Order Caudata - metamorphosis īŽ Remains aquatic, gill- breathing, fully reproductive larval form unless water begins to dry up īŽ Metamorphoses to adult, loses gills, develops lungs, assumes appearance of ordinary salamander
  • 59. Order Caudata - metamorphosis īŽ Can be artificially induced to metamorphose by treating then with thyroxin īŽ Essential for normal metamorphosis in all amphibians
  • 60. Order Caudata - territoriality īŽ Some salamanders display territorial behavior īŽ Use scent to mark territories - Plethodontidae īŽ Males of some species defend territories against conspecifics
  • 61. Order Caudata - territoriality īŽ Attack intruders with foreign scent, but not neighbors with familiar scent īŽ Nasolabial grooves - chemosensory system
  • 62. Order Caudata - territoriality īŽ Territory defended against intruders at expense of reduced energy intake īŽ Attack nose of intruders - ruin chemosensory system? īŽ Less able to feed, smaller
  • 63. Order Caudata - toxicity īŽ Many newts contain mucous glands in skin that secrete irritating or toxic materials īŽ E.g., red eft (terrestrial red-spotted newt) produces a potent neurotoxin
  • 64. Order Caudata - toxicity īŽ Produces unpleasant numbness or tingling sensation in animals touching it īŽ Secretions have been known to kill animals that ingested them (including humans)
  • 65. Order Caudata - toxicity īŽ Species may advertise their toxicity with conspicuous aposematic, or warning, colors and behaviors īŽ Red eft is brilliant red- orange
  • 66. Order Caudata - toxicity īŽ Other species that are more plainly often flip when attacked to display brightly colored bellies
  • 67. Order Caudata - toxicity īŽ Some species have adapted to mimic toxic forms īŽ E.g., red-backed salamander has a red color morph (normally only red on back) that resembles red eft īŽ Studies: red morph suffers less bird predation than normal color morph
  • 68. Order Anura īŽ Frogs & toads īŽ >3400 species īŽ Most familiar, successful amphibians īŽ Obvious, common characteristic - lack of tail in adult īŽ All pass through tailed larval stage
  • 69. Order Anura īŽ Specialized for jumping mode of locomotion
  • 70. Order Anura īŽ Further distinguished from Caudata by: īŽ Larvae - tadpole or polliwog stage īŽ Metamorphosis during development
  • 71. Order Anura īŽ Tadpole īŽ Long, finned tail īŽ Both internal, external gills īŽ No legs īŽ Specialized mouthparts for herbivory (salamander larvae are carnivorous) īŽ Highly specialized internal anatomy
  • 72. Order Anura īŽ Striking metamorphosis from tadpole to adult īŽ Neoteny never exhibited in frogs or toads
  • 73. Order Anura īŽ Represent specialized side branch of amphibian evolution īŽ Not good representatives of vertebrate body plan
  • 74. Order Anura īŽ No visible neck īŽ Caudal vertebrae fused (to form urostyle) īŽ Ribs generally absent īŽ Hindlegs much enlarged
  • 75. Order Anura īŽ 21 different families īŽ Common North American frogs īŽ Ranidae - grass frogs īŽ Hylidae - tree frogs
  • 76. Order Anura īŽ Bufonidae - true toads īŽ Short legs, stout bodies, and thick skins, usually with prominent warts īŽ “toad” often used to refer to more terrestrial members of many families
  • 77. Order Anura īŽ Largest anuran - West African goliath frog īŽ >30 cm from nose to anus, ~1 m with outstretched legs īŽ 3 kg+ (up to 15 lbs.) īŽ Eats animals as large as rats, ducks
  • 78. Order Anura īŽ Smallest anuran - Cuban leafclimber frog īŽ ~1 cm - covered by a dime
  • 79. Order Anura īŽ Bullfrog - up to 20 cm long
  • 80. Order Anura īŽ Most abundant, successful frogs belong to genus Rana īŽ 260 species īŽ Temperate, tropical regions (not New Zealand, oceanic islands, southern S. Amer.) īŽ Most often near water
  • 81. Order Anura īŽ Most larger frogs are solitary except during breeding season īŽ Males call noisily for females
  • 82. Order Anura īŽ Winter - hibernate in soft mud at bottom of pools, streams īŽ Low energy demand īŽ Glycogen, fat stored during spring, summer to provide energy during winter
  • 83. Order Anura īŽ Tree frogs winter under humus on forest floor īŽ Tolerant of low temperatures
  • 84. Order Anura īŽ May survive prolonged freezing of all extracellular fluid (35% of body water) īŽ Accumulate glucose, glycerol in body fluids to protect tissue from damage from ice crystals
  • 85. Order Anura - enemies īŽ Natural enemies - snakes, aquatic birds, turtles, racoons, humans īŽ Usually defenseless
  • 86. Order Anura - enemies īŽ Some tropical, subtropical forms are aggressive, jump at and bite attackers īŽ Others play dead
  • 87. Order Anura - enemies īŽ Others blow up lungs - difficult to swallow īŽ Jumping is best defense īŽ Some species have poison glands
  • 88. Order Anura - skin īŽ Thin and moist īŽ Attached loosely to body only at certain points
  • 89. Order Anura - skin īŽ Stratified epidermis, inner dermis īŽ Outer layer of epidermis shed periodically īŽ Deposits of soft keratin - protection against water loss, some abrasion
  • 90. Order Anura - skin īŽ Dermis, lower epidermis have two types of glands īŽ Mucous - secrete protective waterproofing mucous īŽ Serous - secrete whitish, watery poison
  • 91. Order Anura - skin īŽ Poisons of dendrobatid frogs (small, S. Amer. frogs) are most lethal animal secretions known īŽ More poisonous than venoms of sea snakes, any arachnids
  • 92. Order Anura - skin īŽ Dermis also contains several types of pigment cells - chromatophores īŽ Many can adjust color to blend in with background
  • 93. Order Anura īŽ Entire musculoskeletal system is specialized for jumping and swimming, via thrusts of the hindlimbs
  • 94. Order Anura - skeleton īŽ Vertebral column - rigid structure - lost much of flexibility found in fishes īŽ Transmits forces from limbs to body
  • 95. Order Anura - skeleton īŽ Body much shorter - only 9 trunk vertebrae, several fused caudal vertebrae īŽ Caecilians have 285 vertebrae
  • 96. Order Anura - skeleton īŽ Skull much lighter than in fishes īŽ Changed for improved special senses, feeding, breathing
  • 97. Order Anura - muscles īŽ Limb muscles - probably derived from radial muscles that move fish fins up and down īŽ Arranged in antagonistic groups (actions of one oppose another)
  • 98. Order Anura - muscles īŽ Abductor - away from median axis īŽ Adductor - toward median axis īŽ Flexor - bend part īŽ Extensor - straighten part īŽ Rotators - rotate
  • 99. Order Anura - muscles īŽ Myotome muscles modified to provide support to various portions of body (head, belly)
  • 100. Order Anura - lungs īŽ Frogs and toads are more highly dependent on lung breathing than are the salamanders īŽ Several disadvantages of cutaneous respiration
  • 101. Order Anura - lungs īŽ 1) skin must be kept thin, moist - too delicate for wholly terrestrial life īŽ 2) gas exchange rate across skin nearly constant - cannot vary with increased activity
  • 102. Order Anura - lungs īŽ Cutaneous respiration important to anurans during winter hibernation īŽ Also, under normal conditions most CO2 is lost across skin surface,most O2 taken up by lungs
  • 103. Order Anura - lungs īŽ Lungs are elastic sacs with inner areas subdivided into many chambers īŽ Not as much internal surface area relative to higher vertebrates
  • 104. Order Anura - lungs īŽ Frogs are positive- pressure breathers īŽ Breathe by swallowing air - forcing it into their lungs īŽ All higher vertebrates use negative- pressure system īŽ Expansion of lungs to pull air inward
  • 105. Order Anura - lungs īŽ Both male and female anurans have vocal cords īŽ Males are better developed īŽ Located in larynx - voicebox
  • 106. Order Anura - lungs īŽ Sound produced by passing air back and forth between lungs and large pair of air sacs (vocal pouches) in floor of mouth īŽ Pouches also serve as resonators in males īŽ Function to attract mates
  • 107. Order Anura - circulation īŽ Shift from gill to lung breathing solved some circulatory problems, but caused others īŽ Eliminated major obstacle to arterial blood flow - gills
  • 108. Order Anura - circulation īŽ Needed to develop new circuitry to serve lungs - pulmonary circulation īŽ 6th aortic arch developed into pulmonary artery īŽ New pulmonary veins developed for return
  • 109. Order Anura - circulation īŽ More difficult evolutionary problem - separate oxygenated from deoxygenated blood īŽ Amphibians have made it halfway īŽ Two atria, but single ventricle - potential mixing
  • 110. Order Anura - circulation īŽ Actually little mixing of oxygenated, deoxygenated blood īŽ Spiral valve in conus arteriosus (leading from ventricle) may help maintain selective distribution īŽ Controversial - defied complete analysis
  • 111. Order Anura - feeding īŽ Adult frogs are carnivorous īŽ Feed on insects, spiders, worms, slugs, snails, millipedes, anything moving and small enough to eat
  • 112. Order Anura - feeding īŽ Snap at prey with protrusible tongue īŽ Attach at front, free behind īŽ Free end very glandular, produces sticky secretion, adheres to prey
  • 113. Order Anura - feeding īŽ Teeth on premaxillae, maxillae, vomers īŽ Not biting or chewing īŽ Used to prevent prey escape
  • 114. Order Anura - digestion īŽ Digestive tract relatively short in adults, longer in tadpoles īŽ Carnivore adaptation vs. herbivore adaptation īŽ Variety of enzymes produced
  • 115. Order Anura - nervous system īŽ Dramatic changes occurred in nervous system as vertebrates moved onto land īŽ Increased cephalization to cope with increased need for information processing
  • 116. Order Anura - nervous system īŽ Increased senses of smell, vision, hearing, increased demand for balance īŽ Has resulted in loss of independence of the spinal ganglia
  • 117. Order Anura - nervous system īŽ Spinal ganglia can serve as “mini brains” in lower vertebrates īŽ Coordinate actions without involving brains īŽ But even with some loss of independenceâ€Ļâ€Ļ
  • 118. Order Anura - nervous system īŽ Headless frog with only spinal cord intact still maintains high degree of purposeful, highly coordinated behavior īŽ Normal posture, raise leg to remove disturbing object (other if one is restrained)
  • 119. Order Anura - nervous system īŽ Two senses highly advanced from fishes and most other amphibians īŽ Hearing and sight
  • 120. Order Anura - hearing īŽ Lateral line system remains in aquatic larvae, but not most adults īŽ Role is replaced by ear in most adult frogs īŽ Obvious structure - tympanum - serves as eardrum
  • 121. Order Anura - hearing īŽ Picks up vibrations of airborne sounds and transmits them through middle ear to inner ear
  • 122. Order Anura - hearing īŽ Contains semicircular canals and structures similar, but simpler, than mammalian cochlea
  • 123. Order Anura - hearing īŽ Hearing of species is most sensitive to frequency range of their mating calls īŽ Peeper - higher frequency īŽ Bullfrog - lower frequency
  • 124. Order Anura - sight īŽ Eye is dominant special sense in most amphibians īŽ Especially good in frogs īŽ Fish eye had to be modified for land use
  • 125. Order Anura - sight īŽ Lachrymal glands, eyelids evolved to keep eye moist, dust-free, shielded from injury
  • 126. Order Anura - sight īŽ Upper and lower eyelids īŽ Upper fixed īŽ Lower with folded nictitating membrane that moves across eye surface
  • 127. Order Anura - sight īŽ Cornea exposed to air became an important refractive surface īŽ Removed some of light bending and focusing burden from lens
  • 128. Order Anura - sight īŽ Lens normally focused on distant objects īŽ Moved to focus on nearby objects īŽ Opposite of arrangement in fishes
  • 129. Order Anura - reproduction īŽ Breeding is first desire after coming out of hibernation īŽ Males migrate to water first - olfactory cues īŽ Croak, call to attract females
  • 130. Order Anura - reproduction īŽ Females enter water when eggs mature and are clasped by males - amplexus īŽ Females lay eggs and males fertilize them
  • 131. Order Anura - reproduction īŽ Eggs laid in large masses, anchored to vegetation īŽ Jelly layer absorbs water and swells after fertilization īŽ Development occurs quickly
  • 132. Order Anura - reproduction īŽ Tadpole hatches from protective jelly coat within 6-9 days (temperature- dependent) īŽ Head and body, tail, ventral mouth with horny jaws, ventral adhesive disc, gills, etc.
  • 133. Order Anura - reproduction īŽ Length of tadpole stage is species- dependent īŽ Leopard frog completes metamorphosis within 3 months īŽ Bullfrog takes 2-3 years to complete process