Species within a Genus.

1
It is easier to study the evolutionary
relationships among related

• Species within a Genus.
• Families within an Order.
• Phyla within a Kingdom.

2
It is easier to study the evolutionary
relationships among related

• Species within a Genus.
• Families within an Order.
• Phyla within a Kingdom.

WHY? Deep phylogenetic splits are usually
heavily impacted by convergence and
heterochrony. At this scale how valuable are
outgroups?
3
WHY NOT NEWLY DIVERGED TAXA? Recent splits are
difficult to study because of a lack of
informative characters? Also, this scale
suffers from parallel evolution and vicariance.
4
Origin and Relationships of Modern Amphibians
Things I actually know something about (Im no
herpetologist)
Modern Reptiles Birds/Crocs (By Craggy)
Modern Amphibians
Lungfish
Coelocanths
Ray-finned Fish
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This is like a detective novel where we are
trying to find out what happened 400 MYA.
Dont forget there are plenty of false cluesmany
groups evolved in a short period of time, and
the available fossils are often side roads of
evolution.
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SarcopterygiiLobe Fin Fishes

• Appendages are Similar in Structure to Amphians
• Most Modern Forms Use Them for Locomotion
• Walk on Bottom
• Swim
• Two Major Groups
• Coelacanths2 Living Fossils from Indian Ocean
• Lungfish
• 1 Species each in Australia, Africa, and South
  America
• Sister to Terrestrial Vertebrates
• Australian Species is Very Different
• Walks on Bottom of Ponds/Streams
• Very similar to Paleozoic and Mesozoic forms
• South American African Forms
• Gills Mostly Remove Carbon Dioxide
• Drown Without Air
• African Form Aestivate
• Both Originally Considered Salamanders

(Images from Pough et al., Vertebrate Life)
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Pugh et al., 2002. Vertebrate Life, Fig 7-5
Late Carboniferous of Europe--Any place to look
for this around here?
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2 Competing Hypotheses for Terrestrial Vertebrates
1. Drying pond hypothesis a. lobe-finned fish
in drying ponds b. fish crawl to deeper
ponds c. problems 1) fish go back to
water? 2) African lung fish aestivate 3)
Fossilized remains in burrows 4) Oldest
fossils are in brackish areas 2. Edge of the
sea hypothesis a. lobe-finned fish at the edge
of the water b. feed on prey that falls in c.
become more agile at darting out of the surf d.
dry land is free of competitors e.
Problems 1. Some modern fish do jump
ponds 2. Why are they gone?
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Here is One Recent Idea of Relationships
Lissamphibia
Lepospondyls
Temnospondyls
Stegacephalians
Modern Lungfish
(Images from Pough et al., Vertebrate Life, and
Herpetology)
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Here is Competing Idea of Relationships
Why do we see these differences
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The traditional view has always assumed a close
relationship between temnospondyls and modern
Amphibians. Thus outgroup and character
selection may be biased
How do these biases impact the results of the
study?
12
LETS PLAY CLUE
Maybe it was a Branchiosaur (a temnospondyl
in the Permian.. via paedomorphosis.
Explains Small size, Combination of Primitive
Derived Characters, Difficulty Pinning Things
Down, Absence of Last Bones to Develop in
Branchiosaurs (Supratemporals,
PrePostfrontals, Jugals, Parietal, Lacrimals),
Large Orbits, Tooth Structure. Matches Fossil
Developing Branchiosaurs BUTTesting requires
complete phylogenetic information.
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SO WHAT ABOUT LISSAMPHIBAINS
Urodela
Gymnophonia
Anura
Can We Do Any Better? Are they Monophyletic?
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Synapomorphies Supporting Lissamphibian Monophyly
1. Teeth are Pedicilate and Bicuspid
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• Ears are unique having a stapes and operculum
• (not present in caecillians) and a
• unique patch of sensory epithelial cells
• (papilla amphibiorum--detects low frequency
  sounds)

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• Stapes is directed dorsolaterally vs
  ventrolaterally
• (shared by some probable relatives)
• 4.Fat bodies develop from different germ layer
  (germinal ridge)
• Skin contains Mucus and poison glands

17

• Green Rods Present in Frogs and Salamanders
• Muscle beneath Eye Allows Elevation
• (used in breathing Swallowing)
• Buccopharyngeal Pump for Respiration
• (use throat like a billows). Means air is
  pushed
• into the lungs
• Short Strait Ribs
• 2 Occipital Condyles (Shared with Mammals)

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11. Reduction of Many Cranial Elements
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SO WHAT ABOUT LISSAMPHIBAINS
Urodela
Gymnophonia
Anura
This arrangement supported by Soft
Anatomy (Presence of a tail, costal groves)
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SO WHAT ABOUT LISSAMPHIBAINS
Urodela
Gymnophonia
Anura
This arrangement supported by Skeletal Anatomy
Combined Morphological and Genetic Data (What
does this tell you about the of soft characters
used?) (What would a combined morphological
genetic analysis say?) Frogs
SalamandersBatrachia
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ORDER URODELLA

• Characters
• Long Tail
• Four Limbs
• Costal Grooves
• D. Paedomorphosis is Common (aquatic salamanders)
• Fossil Record Dates to Middle Jurassic (170-159
  MYA)
• Use of Caudata
• Difficulties of Metamorphosis
• Biodiversity
• 515 Species
• Many Cryptic Species
• Mainly Holoarctic (recent invasion of Neotropics)

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SirenidaeSirens

• 2 Genera 4 Species
• Primarily Indiana south and East
• Eel-Like and Aquatic
• Keratinous Beak
• No Hind Limbs
• Some of the Largest Salamanders
• Prey Primarily on Inverts
• Siren intermedia

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CryptobranchidaeHellbenders

• 2 Genera 3 Species
• Eastern NA, China, Japan
• Mostly Aquatic
• Incomplete Metamorphosis
• Largest Indiana Salamander
• Prey Primarily on Crayfish
• Cryptobranchus alleganiensis

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AmphiumidaeAmphiumas

• 1 Genus 3 Species
• Mississippi River System
• Eel-Like and Aquatic
• Reduced Limbs
• Nasty Biters
• Prey Primarily on Aquatic Animals
• Amphiuma

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PlethodontidaeLungless Salamanders

• 27 Genera 360 Species
• Mostly NA with some in SA and Europe
• Most specious group of salamanders
• Full Metamorphosis (Mostly)
• Typically Small
• Prey Primarily on Small Inverts
• Plethodon, Eurycea, Psudotriton, Eurycea,
  Desmognathus

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ProteidaeMudpuppies and Kin

• 2 Genera 6 Species
• Mostly Central Europe Necturus
• Aquatic Salamanders with Well-developed Gills 4
  Legs
• Prey Primarily on Inverts (occasionally caught on
  worms)
• Necturus maculosus

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SalamadridaeNewts

• 15 Genera 62 Species
• Holoarctic
• Eft Stage and Return to Water
• Complete Metamorphosis
• Highly Toxic
• Bad News for Frog Eggs
• Notophothalmus viridescens

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AmbystomidaeMole Salamanders

• 1 Genus 30 Species
• Mostly US
• Robust Bodies made for Digging
• Most Have Full Metamorphosis
• (Axotyls)
• Some Paedomorphic Forms
• Prey Primarily on Inverts
• Ambystoma jeffersonianum, tigrinum, and texanum

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ORDER GYMNOPHONIA

• Characters
• Short Post-Anal Tail
• No Limbs
• Costal Grooves?
• D. Paedomorphic
• Fossil Record Dates to Early Jurassic (190 MYA)
• Fossils from NA (What does this mean)
• Had Legs in Fossil Records
• Very Similar to Salamander Legs
• Snakes Before Snakes
• Biodiversity
• 170
• Pantropical Except Central Africa
• Systematics are a Mess

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ORDER ANURA

• Characters
• No Post-Anal Tail
• No Limbs
• No Costal Grooves
• Caudal Vertebra Fused Urostyle (rod)
• Vertebral Column Reduced
• Elongate Hind Limbs
• Triadobatrachus from Lower Triassic (230 MYA)
• Madagascar
• Poland
• Same Arizona Deposit as a Caecilian
• Biodiversity
• 4300 Species
• Cosmopolitan Except Deserts and Tundra
• Very Diverse In Tropics

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Triadobatrachus massinoti
Modern Leptodactylus
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AscaphidaeTailed Frogs

• Monotypic
• Ascaphus truei
• Tail is an Intermittent (Penile) Organ
• Highly Aquatic
• Fast Flowing Streams
• Pacific Northwest
• Tadpoles have oral suckers, which are sometimes
  used to climb

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PaleobatidaeSpadefoot Toads

• 3 Genera 11 Species
• NA, Europe, and North Africa
• Keratin Spade on Foot
• Used to Dig
• Animals Emerge During Heavy Rains
• Explosive Breeders
• Rapid Metamorphosis
• Scapheopus holbrooki, Spea bombifrons

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BufonidaeToads

• 33 Genera 455 Species
• Once a Trashcan
• Nearly Cosmopolitan
• Many are Terrestrial as Adults
• Highly Toxic Skin Glands (Parotid)
• Maintain Bidders Organ (Rudimentary Ovary
• No Teeth
• Bufo woodhousi and americanus and maritimus

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PipidaeClawed Frogs

• 5 Genera 30 Species
• Sub-Saharan Africa, Panama, Tropical South
  America
• Aquatic as Adults
• Odd Morphology
• Lack Tongues
• Claw-like Tips to Toes
• Dorsoventral Compression
• Well-developed Lateral Lines
• Official Lab Rat of Developmental Biology
• Xenopus

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Myobatrachidae

• 23 Genera 122 Species
• Australia, New Guinea, Tasmania
• Highly Variable Reproduction
• Young in Inguinal Pouches of Males
• Young in the Stomach
• Many falling victim to chytid fungal infections

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HylidaeTree Frogs

• 40 Genera 835 Species
• Paraphyletic with Psuedidae excluded
• Mostly Arboreal
• Toe Pads and claw-like terminal phalanges
• 2 Hyla, 2 Psuedacris, Acris

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MicrohylidaeNarrowmouth Toads

• 69 Genera 350 Species
• Eastern US, South America, Southern Africa
• Small Palatal Folds--Head Vail
• Very Small Frogs
• Local Versions are Fossorial
• Other places get Terrestrial and Arboreal Forms
• 2 Form Commensal Relationships with Spiders
• Often Found Under Cow Poop
• Gastrophryne

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DendrobatidaePoison Dart Frogs

• 8 Genera 205 Species
• Central and South America
• Terrestrial or Arboreal
• Parents Cary Tadpoles on Backs
• Highly Toxic Skin Glands
• Used to Poison Blow-gun Darts
• Probably Diet Related
• Common in Pet Trade
• Official Lab Rat of Developmental Biology
• Dendrobates

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MantelliidaeMadagascar Poison Frogs

• 5 Genera 140 Species
• Madagascar
• Have Aposematic Coloration and Poison in
  Mantellus
• Some Mantellus also resembles Rana and Bufo
• 2 Genera Converge on Spea
• Boophis resembles Hyla
• Why do we see these patterns?
• What is the difference between convergence and
  parallelism?

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RanidaePond Frogs

• 38 Genera 600 Species
• Cosmopolitan
• Highly Variable and Para/Polyphyletic
• Trashcan taxon
• Economically Important
• Rana catesbienna widely Introduced
• 8 Local Species

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ILLUSTATION OF DIVERSITY OTHER FAMILIES

• Racophoridae
• Leptodactylidae
• Rhinodermatidae
• Pseudidae
• Allophrynidae
• Hemisotidae
• Artholeptidae
• Petropedetidae
• Hyperoliidae

• Leiopelmatidae
• Bombinatoridae
• Discoglossidae
• Megaphyridae
• Pelodytidae
• Rhinophrynidae
• Brachycephalidae
• Heleophrynidae

Mostly in African, SA, and Asiatic Tropics