Vertebrate Evolution

Chapter #1 The Diversity, Classification, and Evolution of Vertebra

Chapter #2 Vertebrates and Basic Structure, pg. 19 45.

Chapter #3 Early Vertebrates, Jawless and Jawed Vertebrates, pg. 4

Amphibians

What is a (Liss)Amphibian?
Liss = smooth
Amphibia = double life

(greek referring to skin)

(greek referring to metamorphosis)

Jurassic

Long hind limbs

Short, stiff bodi

(before break-up of Pangea)

HOP

Equal-sized limb

Long, slender bod

WADDLE

No limbs

Long, slender bod

SLITHER

Figure 6.1 Vertebrate Biology

Amphibians

Shared Derived Characteristics of Amphibians:

1) Smooth, Moist Skin:
Absence of bony / keratinized scales (exception = caecilians)
Absence of keratinized, true claws
Mucous glands (prevent desiccation / slippery / disease-free)
Other Methods for Controlling Water Loss:
1) Behavioral Modifications:
Example:
Puerto Rican Coqui
Cool, wet nights

Warm, dry nights

3) Permeable Bladder:
2) Pelvic Patch:
Store dilute urine;
Highly vascularized skin;
absorbs majority of water Canteen away from water

Amphibians

Shared Derived Characteristics of Amphibians:

1) Smooth, Moist Skin:

Absence of bony / keratinized scales (exception = caecilians)

Absence of keratinized, true claws
Mucous glands (prevent desiccation / slippery / disease-free)
Tarichatoxin
Poison glands:

(newts)
Water-soluble alkaloid

Concentrated on
dorsal surface

(blocks Na+ channels)

Produce diverse array

of chemicals
Epibatidine
(dart-poison frogs)
Batrachotoxin
Lipid-soluble Alkaloid (dart-poison frogs)
(Blocks Ach receptors)

Non-addictive
painkiller

Lipid-soluble alkaloid

(Blocks closing of Na+ channels)

Sequestered via diet

Amphibians

Shared Derived Characteristics of Amphibians:

2) Multiple Methods of Respiration:

Cutaneous Gas exchange occurs across moist skin

Buccopharyngeal Gas exchange occurs in buccal cavity / pharynx
Pulmonary Gas exchange occurs in paired lungs (too small for all gas ex
Gills Larval forms and neotenics (neoteny = adults that retain juvenile ch

3) Pedicellate Teeth:

Crown and base of tooth separated by uncalcified dentine / fibrous tiss

May allow for flex of tooth for prey handling (all carnivores)

4) Green Rods:
Distinct type of retinal cell

Hue discrimination in
dim light?

Teeth break easily continuously replaced

Amphibians

Shared Derived Characteristics of Amphibians:

5) Unique Hearing Adaptations:

Operculum Columella Complex:

Two bones in middle ear that transmit sound to inner ear (via oval wi
Dual high / low frequency system
High frequency (> 1000 Hz) = vibration of columella alone (air)
Frogs (needs tympanum for amplification); Vocalization
Low frequency (< 1000 Hz) = vibration of operculum-columella u
Vibration via leg bones / scapula (activated via opercularis mu
Frogs & Salamanders; Predator / prey detection
Inner Ear:
Basilar papillae:
Hair cell system
detects high frequencies
Amphibian papillae:
Hair cell system detects
low frequencies

Amphibians

Shared Derived Characteristics of Amphibians:

6) Levator bulbi muscle:

Bulges eye outward; increases buccal cavity

Amphibians

Salamanders (10 families; ~ 515 species):

Diversity:
First appearance in fossil record = Jurassic
Evolution associated w/ cool, wet forests
Order: Caudata (Tailed Ones)

(~ 245 sp.)

Primarily holarctic distribution:

North American = 9 of 10 families
(5 endemic)

Neotropic = greatest # of specie

All from single family
(Plethodontidae ~ 145 sp.)

Amphibians

Salamanders (10 families; ~ 515 species):

Morphology / Locomotion:

Presence of tail (larvae / juveniles / adults)

4 equally-sized limbs (extend at right angle from body)
Size range = 3 cm to >1 m
Paedomorphosis widespread (aquatic life)

Chinese Giant Salamander

Respiration:
Aquatic = primarily gills (some cutan
Cryptobrachidae = large skin fo

Terrestrial = lungs (50%) / skin (50%

Plethodontidae = lungless
skin (10%) & buccopharynx
(90%)
Hypothesis:

Adults retain:
laterally compressed tail / flattened head
functional lateral line system
external gills (not always)
Evolved in fast, cold water ( O2)
Anguilliform locomotion (walking-trot /Lungs
swim) disadvantageous (buoyancy)

Amphibians

Salamanders (10 families; ~ 515 species):

Diet / Prey Capture:
Generalist carnivores (primarily invertebrates)
Aquatic = mouth gape
Cryptobrachus alleganiensis
Suction pulls prey into oral cavity
movie
Works will with gills / gill slits (one-way flow)
Tongue = broad, flat, immobile
Terrestrial = Thick, sticky tongue grabs prey
Plethodontidae: Projective
tongueWhy Plethodontids?
Buccal pump not necessary for breathing

Taricha torosa
movie

(no lungs)

Hyoid bone specialized for projecting

the tongue (elongated / lightened)
Hydromantes platycephalus
movie
http://autodax.net/feedingmovieindex.html

Amphibians

Salamanders (10 families; ~ 515 species):

Sensory Systems:
Aquatic:
Taste chemicals in water
Vibrations Retain lateral line system
Terrestrial:
Smell Olfactory epithelium (volatile)
Smell Vomeronasal organ (Non-volatile)

Sexually dimorphic (larger in males)

Courtship / Identification (species / sex / individual)
Highly advanced in Plethodontidae
Nasolabial Grooves = Non-ciliated grooves; upper lip to
Aids in collection / delivery of chemical cues (capillary
Sexually dimorphic (esp. during reproductive season)

Hearing limited high frequency (no ear drum); primarily ground vi

Vision acute; especially in plethodontidae

Amphibians

Sirenidae

(sirens):

Nocturnal
Vocalization

Morphology:
Eel-like; lack hind limbs (lack pelvic girdle)
Paedomorphic; gills present
Habitat:
Swamps / lakes / marshes (slow-moving water)

Aquatic:

Reproduction:
SE United States / NE Mexico
External fertilization; female nest guarding

Cryptobranchidae

largest
individuals
(Hellbenders)
:
live 50+ years

Morphology:
Dorsal-ventral compressed body (paedomorphic)
Multiple folds in skin (cutaneous respiration)
Habitat:
Clear, cold mountain streams / lakes
Severely reduced ranges (loss of habitat)
Reproduction:
External fertilization; Males = den masters

Aquatic:

Central China / Eastern United States

Amphibians

Amphiumidae

: terrestrial egg-laying
(Amphiums)
aestivate

Morphology:
Eel-like; maintain four reduced limbs
Paedomorphic; lack gills (lungs present)
Habitat:
Sluggish streams / rivers; swamps

Reproduction:
Internal fertilization; female nest guarding

Proteidae

(Waterdogs):

Aquatic:
SE United States

dissection specimen
nocturnal

Morphology:
Paedomorphic; feathery gills / caudal fins
Habitat:
Lakes / streams
Limestone caves (drastic reduction in numbers)
Reproduction:
Internal fertilization; / nest guarding

Aquatic:
Eastern United States / SE Europe

Amphibians

Plethodontidae

(example = Ensatina escholtzii):

Anatomy:
Constriction at tail base = tail autotomy
Very costly (tail = large fat reserve)
~ 10 12% tail regeneration (~ 2 years)

Terrestrial:
Habitat:
Prefers cool forests with litter / dead trees North / Central / South America & Europ
Tolerates logging better than most plethodons

Reproduction:
Mating begins (Nov.) and ends (March) with rains
use hedonic glands / pheromones to attract
Methods of pheromone administration:
Slap nares with hedonic gland (video)
Innoculate with sharp teeth

Pheromone:
Elaborately patterned courtship dance
Chemical cue that affects the
Tail straddle-walking (unique to plethodons)
behavior and / or physiology
repeats dance (~ 1 5 hours)
of a conspecific
Pathway / movement unique to each species

Amphibians

Plethodontidae

(example = Ensatina escholtzii):

Reproduction:
lays down spermatophore; picks it up (video)
lay 8 12 eggs (~ 5 mm)
Terrestrial in logs; under logs; in burrows
Terrestrial:

North / Central / South America & Europ

Sperm Cap

guards eggs ( predation / fungal infection)

Direct development of eggs (~60 120 days)
Growth / Maturation:
Sexual Maturation ~ 4 years
Life Span ~ 10 years (best guess)

Gelatinous Base

Amphibians

Rhyacotritonidae:

(example = Rhyacotriton olympicus)

Anatomy:
Full metamorphosis
No operculum / opercular muscle ( hearing)
Highly reduced lungs (cutaneous respiration)
Habitat:
Cold, clear streams / seepages / waterfalls
Very desiccation intolerant
Low heat tolerance (susceptible to logging)
Reproduction:
Internal fertilization
Spermatophore deposition
Tail-wagging display
Aquatic egg-laying
No nest guarding
Anti-predator Display
Growth / Maturation:
Sexual Maturation ~ 4 years
Life Span ~ 10 years (best guess)

Terrestrial:
NW United States

Amphibians

Larval Anatomical Specializations:

Pond Larvae

Stream Larvae

Ambystomatidae
Salamandridae

Dicamptodontidae
Rhyacotritonidae
Plethodontidae

Depressed body profile

High body profile
broad tail fin extending up to head Narrow tail fin (not onto trunk)
Short, thread-like / curly gills
long, plume-like gills
All four legs at hatching
Only front legs at hatching

Amphibians

Salamandridae

(example = Taricha granulosa):

Habitat:
Prefer older growth forests during newt phase
Warning Coloration:
Flashes tail & brightly colored stomach

Terrestrial:
Chemical Defense:
Tarichatoxin (non-protein VERY poisonous) North America / Europe / NW Africa / Asi
Neurotoxin blocks NA+ channels (paralysis)
Small dose lethal to birds / mammals
Large dose lethal to humans

Red-spotted Garter Snake

The Arms Race

Assignment:

http://evolution.berkeley.edu/evolibrary/article/biowarfar
Questions Posted on Class Website

Amphibians

Salamandridae

(example = Taricha granulosa):

Reproduction:
Adults migrate to ponds / lakes (1 3 weeks prior)
Breeding from April July (annual / biannual)
release pheromone (hedonic gland)

Amplex with (ward against rivals)

Terrestrial:
North
America / Europe / NW Africa / Asi
signals willingness to mate by raising
head
lays down spermatophore
lay 200 375 eggs
Incubation ~ 20 35 days
Growth / Maturation:
Metamorphosis ~ 4 5 months
Sexual Maturation ~ 4 5 years
Life Span ~ 20 25 years

Amphibians

Dicamptodontidae:

(example = Dicamptodon tenebronsus)

Anatomy:
Full metamorphosis / facultative paedomorphosis
Large (up to 30 cm SVL)
Well-developed lungs / blade-like teeth
Habitat:
Terrestrial:
Damp coniferous forests (near streams / seeps)
NW United States / SW Canada
Paedomorphic = moving water
Diet: Opportunistic feeders (sit-and-wait)
Reproduction:
Internal fertilization
Breeding occurs in water
Will eat small mammals
Eggs (~ 200) deposited singly
wood / rocks in water (female nest-guarding)

Growth / Maturation:
Metamorphosis ~ 2 years
Sexual Maturation ~ 2 years
Life Span = long-lived

Amphibians

Ambystomatidae:

(example = Ambystoma macrodactylum

Anatomy:
Full metamorphosis / paedomorphosis (rare)
Stout-bodied; thick-tailed
Habitat:
Wide range of habitats (grasslands forests)
Subterranean (mole salamanders)
Close to water (~ within 100 m)
Enter torpor during winter (below frost line)

Terrestrial:
Western North America

Reproduction:
Earliest amphibian breeders (Pacific Northwest)
Migrate to breeding ponds (fidelity - arrive first)
Eggs (~ 100 400) deposited in clusters
Growth / Maturation:
Metamorphosis ~ 2 years
Sexual Maturation ~ 3 5 years
Life Span ~ 10 years
Cannibalistic
larvae

Amphibians

Caecilians (6 families; 180 species):

Long, slender body; small dermal scales
Legless; no post-anal tail
Most species blind as adults
Specialized sensory tentacles (snout)
Diet = small insects / earthworms

Order: Gymnophiona (Naked snak

Tropics (S. America / Africa / S.E. Asia

History:
Fossorial / Aquatic
Very little known about courtship
Internal fertilization (via intromittent organ)
75% viviparous (live young - matrotrophic)
Initial nutrition = yolk
Subsequent nutrition = uterine milk
Breath via fetal gills
Young caecilians 30 60% adult BL!
25% oviparous (lay eggs may brood eggs)

Life

Amphibians

Frogs (29 families; ~ 4800 species):

Diversity:

First appearance in
fossil record = Jurassic

Order: Anura (Without Tail)

(~ 1100 sp.)

Ubiquitous (except Antarctica):

North American = 10 or 29 families

Neotropic = greatest # of specie

Most Diverse Family
Leptodactylidae ~ 1100 sp.

Amphibians

Frogs (29 families; ~ 4800 species):

Morphology:
Body specialized for jumping:
Hind legs elongated; tibia / fibula fused
Large pelvis; short vertebral column
Urostyle (Fused posterior vertebra)
Size variable (1/2 1 )

Toe Pads:
Viscous mucus = sticky

Locomotion:

Jumping Frogs
Long hind limbs

Hopping Frogs
Semi-aquatic Frogs
Arboreal Frog
Short limbs / Robust bodyWebbed feet / stream-lined Slim waist / long legs

Amphibians

Pacific Tree Frog

Frogs (29 families; ~ 4800 species):

Bullfrog

Spadefoot Toad
Vocalization:
Associated with reproduction (advertisement calls)
Identify species / sex of caller
Can function in mate choice (e.g., male size)

Vocalization is a tradeoff:
Costs:
1) Vocalization energetically expensive
2) Vocalization announces location
Benefits:

Tungara Frogs:
Whine cluck
Females prefer / bats prefer
Whine
Females / bats less preference
Whine cluck only observed
when large choruses present

Amphibians

Frogs (29 families; ~ 4800 species):

Reproduction:
Mating systems:
1) Explosive Breeders (season = few days)
Temporary aquatic habitats
Large aggregations; limited mate choice
2) Prolonged Breeders (season = months)
Males defend territories
Males out-number females; strong mate choice
Fertilization primarily external
Amplexus: Embrace of male and female frog
Brings cloacae in close proximity (fertilization)

Inguinal Amplexus

May last hours to several days

Fertilization can
occur internally
Axillary Amplexus

Amphibians

Frogs (29 families; ~ 4800 species):

Reproduction:
Egg Development:

Directly in water Vegetation over water

(egg protection)

Parental Care

Carry Eggs

(Mid-wife Toad )

Direct Development
(egg protection)
(tadpole protection)

Guard Eggs

Foam Nest
(egg protection)

Bromeliad Nest
(egg protection)
(tadpole protection)

(common):

Carry Eggs / tadpoles Internal Brooding

(Sirinam Toad )

( Gastric Brooders)

Viviparous
(Morogoro Tree Toad)

Amphibians

Frogs (29 families; ~ 4800 species):

Tadpoles:
Aquatic larval stage of frogs

Still Water:
Ovoid bodies
Tails with large fins

Advantages:
Fast Water:
1) Exploit non-adult resources
Stream-lined bodies
Tadpoles = aquatic / herbivores
Small tail fins
Adults = terrestrial / carnivores
2) Efficient eating machines (rapid growth)
Metamorphosis:

Triggered by thyroid hormone

(3 stages)

Premetamorphosis

Prometamorphosis

Metamorphic Climax

Increase in size
Little change in form

Hind legs appear

Rapid decline in growth

Fore legs appear

Tail regresses

Mouth broadens; long short gut; degeneration of gills; dermal gland fo

Amphibians

Rhinophrynidae:

Single species
(Burrowing Toad)
Unique tongue

Morphology:
Short, powerful limbs; tubercle present
Robust body; pointed head (cornified tip)

Habitat:
Sub-humid areas; surface after heavy rains
Reproduction:
Explosive breeder; aquatic tadpole

Leptodactylidae

Fossorial
North America / Central America

Among
(Neotropical frogs)
:
largest
Many nocturnal

Morphology:
Highly variable

(10 mm 250 mm)

Habitat:
Variable:
Leaf litter layer
Fully aquatic

Arid regions
High plateaus

Reproduction:
External / internal fertilization
Aquatic tadpoles / direct development

Terrestrial
North / Central / South America

Amphibians

Microhylidae:

Commensal assoc.
(Microhylid frogs)
with spiders

Morphology:
Highly variable

(10 mm 100 mm)

Habitat:
Variable (Arid deserts Wet rain forests)
Reproduction:
Terrestrial / Fossorial / Arboreal
Explosive breeders / prolonged breeders
Ubiquitous
Aquatic tadpoles / direct development

Dendrobatidae

(Poison-dart frogs):

Morphology:
Relatively small; highly aposematic
Habitat:
Tropical rainforests

Poisonous

Introduced to
Reproduction:
Hawaii
Most lack amplexus
Small clutches; parental care

Terrestrial
Central / South America

Amphibians

Pipidae:

(clawed frogs)

Tongueless
Model lab organism

Morphology:
Dorso-ventrally compressed body
Limbs splayed laterally; large, webbed feet
Habitat:
Almost every type of body of water
Reproduction:
Aquatic tadpoles / direct development

Cane Toad

Aquatic
South America / Africa

(Bufo marinus)

one of the 100 worst invasive

Invasive Species
species worldwide.
Specialist Group
1)
2)
3)
4)

Prolific breeders (~ 33,000 eggs)

Tolerance for temperature / salinity
All stages of life poisonous (bufotoxin)
Generalist diet (including cat food)
No native toad species

Trap / hand catch frogs

Biocontrol (virus)

1935
~ 100 Marine toad
released

Amphibians

Ascaphidae

(example = Ascaphus truei):

Anatomy:
Males have tail-like intromittent organ
Highly vascularized cloacal extension
Tympana absent; do not call
Aquatic:

Habitat:
Northwest United States
Swift mountain streams with cobbled substrate
Primarily aquatic; terrestrial after heavy rains
Reproduction:
Internal fertilization (only frog species)
Inguinal amplexus; tail inserted into cloaca
Small clutches of large eggs (place under rocks)
Tadpoles have suction disks / reduced fins
Growth / Maturation:
Metamorphosis ~ 2 3 years
Sexual Maturation ~ 8 9 years
Life Span ~ 12 20 years

Amphibians

Pelobatidae

(example = Scaphiopus intermontana):

Anatomy:
Keratinous, spade-like metatarsal tubercle
Rotund body; short legs; large eyes
Glandular, tuberculate skin
Terrestrial:

Habitat:
North America / Europe / Asia / Africa
Found in sagebrush country (shallow burrows)
Over-winter in self-made burrows (~ 1 m deep)
Survive osmotic stress via urea in body fluids
Reproduction:
Explosive breeding; permanent / ephemeral waters
300 500 eggs / female; hatch = 2 4 days
Growth / Maturation:
Metamorphosis ~ 1 month
Sexual Maturation ~ 1 2 years
Life Span ~ 10 13 years

Amphibians

Bufonidae

(example = Bufo boreas):

Anatomy:
Bidders Organ: Rudimentary ovary (on male testis)
Rotund body; dull coloration
Prominent skin glands (toxic - peptides)
Habitat:
Various elevations; Close water contact
Over-winter in existing burrows near / in water
Reproduction:
Prolonged breeders; actively search out
Males lack call; chirp = release call
~ 6000 eggs / clutch; laid in double row strings
Growth / Maturation:
Metamorphosis ~ 1 2 months
Sexual Maturation ~ 4 6 years
Life Span ~ 10 15 years

Terrestrial:
Cosmopolitan

Amphibians

Hylidae

(example = Hyla regilla):

Indicator
Species

Anatomy:
Relatively small: 1 2 inches in length
Toe discs present
Polymorphic colors; change with temp / humidity
Primarily Arboreal:

Habitat:
North / Central / South America
Low shrubbery during wet season
Over-winter in moist, cool retreats during winter
Reproduction:
Migration to breeding ponds triggered by warm, winter rains
enter pond, begin calling (attract ; warn )
~ 500 - 750 eggs / clutch; ~ 3 clutches / season
Growth / Maturation:
Metamorphosis ~ 2 months
Sexual Maturation ~ 1 2 years
Life Span ~ ???

Amphibians

Ranidae

Red located on lower

abdomen / hind limbs

(example = Rana Aurora ):

Anatomy:
Mid-sized body: 2 - 3 inches in length
Long rear legs; exceptional leaping ability
Presence of vocal sacs dependent on sub-species
Terrestrial:

Habitat:
Cosmopolitan
Require riparian vegetation near ponds / streams
Move into upland forest outside of breeding season
Reproduction:
Reproduction occurs November April (assures cold water)
Extensive migrations to breeding sites not uncommon

~ 200 - 1000 eggs / clutch; attached to aquatic vegetation

Growth / Maturation:
Metamorphosis ~ 3 4 months
Sexual Maturation ~ 2 3 years
Believed to be inspiration for Twains
Life Span ~ 10 years
"The Notorious Jumping Frog
of Calveras Country"

Amphibians

Amphibian Decline:
Nearly 1/3 of amphibian species
are threatened (~ 2000 species)
~ 150 species believed extinct over
the past several years
43% of all species declining in
population (< 1% show increases)
Regions:
Latin America (largest #)
Caribbean (largest %)
Madagascar
Eastern Australia
Central / Southern China
West Africa

Global Distribution of Amphibians

Why are populations

declining?
(7 major hypotheses)

Global Distribution of Threatened Amphibian

Amphibians

Amphibian Decline - Reasons:

red1) Habitat Changes: California
legged frog

Habitat Destruction:
Complete elimination of ecosystem
leading to total loss of biological function

Habitat Fragmentation:
Populations isolated due to
removal of linking habitat

Habitat Alteration:
Changes to ecosystem that
adversely affect function

Amphibians

Amphibian Decline - Reasons:

red1) Habitat Changes: California
legged frog

Habitat Destruction:
Complete elimination of ecosystem
leading to total loss of biological function

Habitat Fragmentation:
Populations isolated due to
removal of linking habitat

Habitat Alteration:
Changes to ecosystem that
adversely affect function

2) Introduced Species:

Sunfish vs. Red-legged Frog / Pacific Tree Frog

Bullfrog vs. Anything that moves

Amphibians

Amphibian Decline - Reasons:

3) Overexploitation:
Education (e.g., dissection)
Research
Bait
Red-legged Frog:
Prized for legs
(~ 80,000 harvested / year)

Illicit Pet Trade

(Goliath Frog ~ $3000)

4) Climate Change:

Wood Frog New York Great-crested Newt

Breeding earlier than usual

Drought

Amphibians

Amphibian Decline - Reasons:

5) UV Radiation:

6) Chemical Contamination:
Atrazine (herbicide):
Disrupts steroidogenesis resulting in
demasculanization and hermaphroditism
Acidification:
Reduced hatching success

Pacific Tree Frog:

Developmental and physiological
abnormalities and reduces larval
survival

Nitrogen Pollution:
Reduced feeding activity, swim less vigorously,
Display disequilibrium, development malformations

7) Disease:

Common Frog
Virus

Yellow-legged Frog
Bacteria

Western Toad
Water Mold

Pacific Tree Frog

Trematodes

Amphibians

Amphibian Decline - Reasons:

5) UV Radiation:

6) Chemical Contamination:
Atrazine (herbicide):
Disrupts steroidogenesis resulting in
demasculanization and hermaphroditism
Acidification:
Reduced hatching success

Pacific Tree Frog:

Developmental and physiological
abnormalities and reduces larval
survival

7) Disease:

Common Frog
Virus

Yellow-legged Frog
Bacteria

Nitrogen Pollution:
Reduced feeding activity, swim less vigorously,
Display disequilibrium, development malformations

Synergisms most likely at work

Western Toad
Water Mold

Pacific Tree Frog

Trematodes