11

The diversity and evolution of Amphibia:What are they and where did they come from?

Lecture goal

To familiarize students with characteristics of theClass Amphibia, the diversity of extant amphibians,

and the fossil record of amphibians.

Reading assignments:Wells: pp. 1-15, 41-58, 65-74, 77-80

Supplemental readings on amphibian taxa:Wells: pp. 16-41, 59-65, 75-77

Lecture roadmap

Extant amphibian orders

Amphibian fossil record and evolution

Characteristics of amphibian orders and diversity

Characteristics of amphibians

22

What are amphibians?

These foul and loathsome animals are abhorrent becauseof their cold body, pale color, cartilaginous skeleton, filthyskin, fierce aspect, calculating eye, offensive smell, harshvoice, squalid habitation, and terrible venom; and so theirCreator has not exerted his powers to make many of them.

Carl von Linne (Linnaeus)Systema Naturae (1758)

What are amphibians?

Ectothermic tetrapods that have a biphasic life cycleconsisting of anamniotic eggs (often aquatic) and a

terrestrial adult stage.

Orders:•Anura (frogs)•Caudata (salamanders)•Gymnophiona (caecilians)

Kingdom: Animalia

Class: Amphibia

Phylum: ChordataSubphylum: Vertebrata

Subclass: Lissamphibia(amphibios: “double life”)

Amphibia characteristics1) Cutaneous respiration

2) Skin glands

Oxygen and CO2 Transfer

Family Plethodontidae

Gills (larvae, few adult salamanders), 2 Lungs (adults)

Mucous glands

Granular glands

Plethodon dorsalis

33

Amphibia characteristics

3) Modifications of middle and inner ear

Middle ear consists of 2 elements

•Stapes (columella)

•Operculum

Inner ear consists of 2 sensory epithelial patches

•Papilla bassilaris (>1000 Hz)

•Papilla amphibiorum (≤1000 Hz)

Amphibia characteristics

6) Bicuspid pedicellate teeth

Crown (above gum), Pedicel (connected to jawbone)

5) Focus eye by changing position of lens

Levator bulbi underlying the eye controls elevation

7) Reductions in skull bones

General trend associated with paedomorphosis

4) Green rods in retina (excluding caecilians)Involved in hue discrimination (433 nm = blue light)

Modern Orders of Amphibia

3. Anura (frogs)

2. Caudata (salamanders)

1. Gymnophiona (caecilians)

Dendrobates tinctorius

Plethodon shermani

Ichthyophis kohtaoensis

176 species

576 species

5,695 species

88%Anurans

3%Caecilians

9%Caudates

280 Species in U.S.(86 Species in TN)

44

Gymnophiona (Caecilians)Characteristics:

•Degenerate Eyes (covered with skin or bone)•Internal Fertilization (phallodeum)

•Left lung reduced or absent•Tentacle between eye and nostril

Tropical Distribution

6 Families

•Limbless (pectoral & pelvic girdles absent)•Elongate and annulated bodies

•Some with dermal scales•Distinct skulls

•Stegokrotaphic versus Zygokrotaphic

Video

Caecilian families

1) Caeciliidae (Common Caecilians)

•95 species (57%)

•Primary Annuli•Most Fossorial•No true tail

•Stegokrotaphic skull

Boulengerula boulengeri

Boulengerula fischeri

Dermophis mexicanus Gegeneophis mhadeiensis

Gegeneophis danieli •Can reach 5 ft in length

Caecilian families

2) Ichthyophiidae (Fish Caecilians)

•38 species (23%)

•Primary annuli with secondary & tertiary annuli•True tail•Females attend eggs

•Stegokrotaphic skull

Ichthyophis kohtaoensis Ichthyophis kohtaoensis

•Primitive family

55

Caecilian families

3) Typhlonectidae (Aquatic Caecilians)

•14 species (8%)

•Primary annuli•No true tail•Strongly aquatic

•Zygokrotaphic skull

Chthonerpeton indistinctum

•Viviparous

Chthonerpeton indistinctum

Typhlonectes compressicauda

Caecilian families

4) Rhinatrematidae (Beaked Caecilians)

•9 species (5%)

•Primary annuli with secondary & tertiary annuli•True tail•Oviparous

•Zygokrotaphic skull•Primitive family

Epicrionops bicolorRhinatrema bivittatum

Epicrionops bicolor

Caecilian families5) Scolecomorphidae (Tropical Caecilians)

•6 species (4%)•Primary annuli•No true tail•Some are viviparous

•Zygokrotaphic skull

6) Uraeotyphlidae (Indian Caecilians)•5 species (3%)

•Primary annuli with secondary annuli•True tail

•Stegokrotaphic skull

•Calcified spines on phallodea

Scolecomorphus vittatus

Uraeotyphlus

66

Caecilian phylogeny

Rhinatrematidae

Ichthyophidae

Uraeotyphlidae

Scolecomorphidae

Caecilidae

Typhlonectidae

Primitive

Derived

1

Loss of tailLoss of secondary annuli

Viviparous/Direct DevelopmentStegokrotaphic skull

2 4

4

3

Caecilian morphology and ecologySome other topics to explore on your own

1. Modes of locomotion

2. Burrowing

3. Sensory systems

Caudata (Salamanders)Characteristics:

•Tails and superficially segmented bodies•Well-developed limbs (except aquatic)•Internal fertilization (most)

•Pheromones (mucous glands)

•Larval development external (most)

•Lack tympanum & middle ear

Mostly Temperate Distribution

10 Families

•Regenerate lost limbs

77

Salamander families

1) Plethodontidae (Lungless salamanders)

•378 species (68%)

•Found in the US and New World Tropics•Cutaneous respiration•Nasolabial groove (chemoreception)•Reduced skull

•Eggs usually guarded

Bolitoglossa adspersa Hydromantes supramontisOedipina gracilis

Thorius papaloae

Pseudoeurycea bellii

MovieMovie

Salamander families

2) Salamandridae (True salamanders)

•74 species (13%)

•Found in the US and SE Asia•Lungs•Toxic skin with bright coloration•Free-swimming larvae in most species

Triturus pygmaeus

Tylototriton shanjing

Neurergus crocatus

Neurergus kaiseri

Triturus dobrogicus

Salamander families

3) Hynobiidae (Asian salamanders)

•51 species (9%)

•Found in Asia•External fertilization•Reduced lungs in most species•Teeth are in patches

Salamandrella keyserlingii Ranodon sibiricus

Pachyhynobius shangchengensis

Onychodactylus japonicus

Hynobius tsuensis

Batrachuperus pinchonii

88

Salamander families

4) Ambystomatidae (Mole salamanders)

•32 species (6%)

•Found in North America•Highly terrestrial adults•Primarily winter and spring breeders•Aquatic courtship

•Hybridization common

(Jefferson’s and Blue-Spotted)

Ambystoma opacum

Ambystoma annulatum Ambystoma jeffersonianum

Ambystoma mavortium

Ambystoma cingulatumSpecimen

Salamander families5) Proteidae (Mudpuppies and Waterdogs)

•6 species (1%)•Found in eastern US and Europe•Obligate paedomorphs•Depressed body and external gills

•Cave dwelling species

6) Sirenidae (Sirens)•4 species (0.7%)•Found in southern US (still water)•Obligate paedomorphs

•External fertilization (lack spermatheca)•Lack pelvic girdles, eye lids, and pedicellate teeth

•Pre-maxillary beak

•External gills

Siren intermedia intermedia

Pseudobranchus striatus

Necturus maculosus

Proteus anguinus

Salamander families7) Rhyacotritonidae (Torrent Salamanders)

•4 species (0.7%)

•Found in costal NW US•Semi-aquatic•No operculum or opercular muscle•Reduced lungs

•Bright yellow abdomen

Rhyacotriton variegatus

Rhyacotriton kezeri

8) Dicamptodontidae (Giant Salamanders)•4 species (0.7%)

•Found in costal NW US and Canada•Large terrestrial salamander (20 cm SVL)•Vomerine teeth (shape of M)•Larvae develop for 2-5 years

Dicamptodon ensatus

Dicamptodon aterrimus

99

Salamander families9) Amphiumidae (Amphiumas)

•3 species (0.5%)

•Found in SE US•Obligate paedomorphs•No external gills or gill slits•Spermatophore directly deposited into spermatheca

•Toe number used for species ID

10) Cryptobranchidae (Hellbenders)•3 species (0.5%)•Found in eastern US, China & Japan•Obligate paedomorphs•No external gills, excessive skin folds

•External fertilization•No eye lids or tongue pad

Andrias japonicus

Amphiuma tridactylum

Amphiuma means

Salamander phylogenyPlethodontidae

AmphiumidaeRhyacotritonidae

Ambystomatidae

Dicamptodontidae

Salamandridae

Proteidae

Sirenidae

Hynobiidae

Cryptobranchidae Primitive

Derived

Salamander morphology and ecology

Some other topics to explore on your own

1. Adaptations

2. Paedomorphosis

3. Evolution of lunglessness

1010

Anura (Frogs and Toad)Characteristics:

Global Distribution

29 Families

•Shortened presacral vertebrate (usually 8)

•Ribs are reduced or absent (2nd or 4th)

•Presacral vertebrae firmly articulated

•External fertilization (usually)

•Large hind limbs, no tail (except 1 family)

•Flat heads and large mouths (usually)

•Vocal sacs in males (usually)

Saltatorial

2-10X BLVideo

Anuran families

1) Leptodactylidae (Southern Frogs)

•1283 species (24%)

•Found in the New World Tropics•Males brood eggs•Foam nests•Varied life history (tadpoles, direct dev., viviparous)

•Eggs usually guarded

Lithodytes lineatus

Eleutherodactylus coqui Eleutherodactylus coqui

Leptodactylus mystacinus

Pleurodema thaulPleurodema thaul

Ceratophrys ornata

Telmatobius culeus

Anuran families

2) Hylidae (Tree Frogs)

•835 species (15%)

•Global distribution•Toe discs•Good climbers and jumpers•Free swimming tadpoles (most)

Anotheca spinosa

Cruziohyla calcarifer

Dendropsophus berthalutzae

Hylomantis lemur

Pachymedusa dacnicolor

Triprion petasatus

Scinax garbei

1111

Anuran families

3) Ranidae (True Frogs)

•799 species (14.7%)

•Global distribution (Africa and Asia most)•Well-developed legs and webbed feet•Free swimming tadpoles (most)

Ceratobatrachus guentheri

Micrixalus saxicola

Tomopterna tuberculosaStaurois natator

Rana warszewitschii

Pyxicephalus adspersus

Platymantis vitiensis

Anuran families

4) Bufonidae (True Toads)

•493 species (9%)

•Global distribution•Cutaneous glands•Teeth nearly absent•Bidder’s organ

Nectophrynoides asperginis

Atelopus crucigerPedostibes hosii

Melanophryniscus stelzneri

Crepidophryne chompipe

Bufo superciliaris

Anuran families

5) Microhylidae (Narrow-mouthed Frogs)

•449 species (8%)

•Mostly tropical and subtropical•2-3 palatal folds•Stout hind legs, short snouts, and globose bodies

•Breviceps - males produce secretions to stick to females

Scaphiophryne madagascariensis Rhombophryne testudo

Relictivomer pearsei

Phrynomantis bifasciatus

Myersiella microps

1212

Anuran families

6) Rhacophoridae (Asian Tree Frogs)

•288 species (5%)

•Africa, India, SE Asia•Hanging foam nests•“Flying Frogs”•Flash coloration on inner thigh

Polypedates cruciger

Polypedates otilophus Polypedates otilophus

Rhacophorus reinwardtii

Theloderma corticale

Nyctixalus pictus

Specimen

Anuran families

7) Hyperoliidae (African Tree Frogs)

•261 species (4.8%)

•Africa, Madagascar, Seychelles•Pupil vertically elliptical•Toe discs•Brightly colored

Afrixalus fornasini

Heterixalus tricolor

Hyperolius marmoratus

Kassina kuvangensis

Leptopelis vermiculatus

Anuran families8) Dendrobatidae (Poison Arrow Frogs)

•252 species (4.7%)

•Found in Central and South America•Brightly colored and toxic skin•Most very small (<1 inch)•Males wrestle for dominance

•Tadpole ride on males back

Dendrobates tinctorius

Video

Dendrobates imitator

9) Mantellidae (Mantellas)•164 species (3%)•Madagascar only•Brightly colored and toxic skin•Most very small (<1 inch)

•Some convergent with Dendrobatidae

Mantella madagascariensis

Tsingymantis antitra

Laliostoma labrosum

1313

Anuran families10) Centrolenidae (Glass Frogs)

•143 species (2.3%)

•Central and South America•Transparent skin, no ribs•Most very small (<1 inch)•Males guard eggs

11) Megophryidae (Cryptic Frogs)•138 species (2.5%)•Asia & Indonesia (streams)•Leaf-like appearence•Nocturnal and poor jumpers

•Tadpoles feed at surface

Brachytarsophrys intermedia Megophrys nasuta

Brachytarsophrys carinensis

Centrolene buckleyi

Centrolene ilex

Cochranella spinosa

Anuran families12) Myobratrachidae (Water Frogs)

•126 species (2.3%)•New Guinea, Australia, & Tasmania•Foam nests in water•Unique egg brooding (pouch, mouth)•Lack toe discs

Limnodynastes dumerilii

•Rheobatrachus (extinct)

Lechriodus fletcheri

13) Arthroleptidae (Squeakers)•51 species (0.9%)•Sub-Saharan Africa (forests)•Direct development (some tadpoles)

Cardioglossa aureoli

Arthroleptis reichei

Cardioglossa gracilis

Anuran families14) Pipidae (Tongueless Frogs)

•31 species (0.6%)•Africa & South America•Fully aquatic (webbed feet, lateral line system)•Pipa (eggs embedded on female)

•Eyes dorsal and no tongue

15) Astylosternidae (Astylosternids)•29 species (0.5%)•Sub-Saharan Africa (forests)•Often grouped with Arthrolepidae•Toe discs•Hairy projections

Pipa parvaPipa parva

•Xenopus (tadpoles with barbs)

Xenopus laevisXenopus muelleriXenopus muelleri

•“Claws”Trichobatrachus robustusTrichobatrachus robustus

Leptopelis kivuensisLeptopelis kivuensis

1414

Anuran families16) Discoglossidae (Disc-tongued Frogs)

•12 species (0.2%)

•SW Europe; Northern Africa

•Females vocalize some

•Toad-like

•Terrestrial and life in burrows

Alytes cisternasii

Discoglossus sardusDiscoglossus sardus

17) Pelobatidae (Spadefoots)

•11 species (0.2%)

•US, Mexico, Europe, Eastern Asia•Xeric environments

•Fast developing larvae(cannibalistic larvae)

•Explosive breeders

Pelobates cultripes

18) Brachycephalidae (Pumpkin Toads)•11 species (0.2%)•SE Brazil

•Reduced digits•All direct development

•Bright orange (tetrodotoxin)•B. didactylus: smallest tetrapod insouthern hemisphere (3/8”)

Brachycephalus nodoterga

Brachycephalus ephippium

Anuran families20) Hemisotidae (Shovel-nosed Frogs)

•9 species (0.17%)

Hemisus marmoratus

•Sub-Saharan Africa•Burrows head first

•Females dig ditch ortransport tadpoles

•Lay eggs in burrow

21) Heleophrynidae (Ghost Frogs)•6 species (0.1%)

Heleophryne regis

•Southern Africa

•Fast-flowing streams

•Well-developed toe discs, spines, sucker-like oral disc (tadpoles)

22) Sooglossidae (Seychelles Frogs)•29 species (0.5%)

Sooglossus pipilodryas

•Madagascar•Inguinal amplexus (only Neobatrachid )

•Secretive: litter and rocks•Direct development & tadpoles on back

19) Bombinatoridae(Fire-bellied Toads & Barbourulas)•10 species (0.2%)

•Europe; East Asia

•Toxic Skin (unken reflex)

•Barbourulas: Rockystreams Bombina orientalis

Anuran families

25) Ascaphidae (Tailed frogs)

•2 species (<0.1%)

Ascaphus montanus

•NW US, British Columbia

•Most primitive extant family(don’t call, 7 yrs to maturation)

•Tail: Cloacal Extension•Internal fertilization

•Fast moving streams

•4 species (0.1%)

Leiopelma archeyi

•New Zealand•Primitive group

•Do not call (no T, ME, VS)•Inscriptional ribs

23) Leiopelmatidae (Leiopelmatids)

•3 species (0.1%)

Pelodytes punctatus

•Black & Caspian Seas, S. Europe•Bulging Eyes; Parsley Color•Females reported vocalizingwhen amplexed

24) Pelodytidae (Parsley Frogs)

26) Rhinodermatidae (Mouth-brooding frogs)

•2 species (<0.1%)•Southern South America (Chile)•Tadpoles Develop in Vocal Sac (male)

•“Rhinoceros nosed”

Rhinoderma darwinii

1515

Anuran families27) Allophrynidae

(Ruthven’s Frog)

•1 species (<0.1%)

Allophryne ruthveni

•NE South America

•Little known about its ecology

•Centrolenidae (related?; foot muscle morphology)

28) Nasikabatrachidae(Purple Frog)•1 species (<0.1%)

Nasikabatrachussahyadrensis

•India (discovered 2003)•Fossorial•Explosive breeders•Little known

29) Rhinophrynidae(Mexican Burrowing Toad)

•1 species (<0.1%)

•Costa Rica to Rio Grande•Fossorial

•Explosive breeders•Termite and ant specialists

•No teeth Rhinophrynus dorsalis

Anuran phylogenyPrimitive Derived

Anuran morphology and ecology

Some other topics to explore on your own

1. Habitat associations of adult anurans

2. Body size and ecology of adult anurans

3. Adaptations for different life styles

1616

Evolutionary history of Amphibia

Events in Geologic History

Fish to Tetrapods

Tetrapods to Amphibians

Geologic history

Alfred WegenerContinental drift

Events in geologic history

(Mississippian)

350 MYA

First Amphibians

ModernAmphibians

(late Permian)

250 MYA

Carboniferous

1717

Ecological historyFirst tetrapods appeared in the Devonian (400 MYA)What were the conditions at this time?

• Tropical/subtropical latitudes - relatively warm and stable• Primitive plants and arthropods

Pangaea

Fish to tetrapod transition

Sarcopterygian (lobe-finned fishes)

Panderichthys-Long snout-Dorsal eyes-Reduced median fins-Flattened bodies

Tiktaalik-More developed limbs with wristand finger bones (body postures)-Robust rib cage-Lungs and gills-Neck separated from body

Eusthenopteron-Pelagic-Internal nostrils-Distinct humerus, ulna, and radiusand femur, tibia, and fibula

Fish to tetrapod transitionIchthyostega

1 m

Acanthostega

0.6 m

Greenland

•Piscivorous

•Limbs likely used fornavigating

•Lungs

•Tail for balance

•Skeletal structure

forelimbs

•Piscivorous

•Fish-like

•Limbs likely used forpaddling

Elbow could not bend

•Skeletal structure

•Gills and Lungs

•8 digits

1818

Fish to tetrapod transition

Evolutionary gap

30 million year gap in thefossil record for tetrapods

Relationships betweenDevonian and Carboniferous

tetrapods obscured

Age of amphibiansCarboniferous (360-300 MYA)

Many species were large and protected by armor

Probably lacked cutaneous respiration but well developed lungs

Retained aquatic reproduction

Pangaea starting to split apart

Amphibian lineage split from reptile lineage 360 MYA

(diversity in form and habitat)

1919

Temnospondylous amphibians

Eryops megacephalus

•1.5-2 m long

•Bicuspid pedicellate teeth

•Lungs & Cutaneous Respiration

•Likely piscivorous

•Engulfed prey

•Not a strong swimmer or fast tetrapod

(likely hunted by stealth or opportunity)

•Shoulder disconnected from skull

Lepospondylous amphibians

Microsauria•“Small Lizard”

•Terrestrial and fossorial

Nectridia•Mostly aquatic

•Resembled newts (flat tails)

•Some with triangular heads

0.5-1 m

•Small fish and aquaticinvertebrates

Lysorophia•Elongate body

•Diminutive limbs

•Fenestrated skulls

No larval forms known

Fossil record of LissamphibiaTriassic, Jurassic and Cretaceous Periods

1) Anurans

Triadobatrachus massinoti

2) Salamanders 3) Caecilians

Karuaus sharovi

•Late Cretaceous•Origin: Gondwanaland (SA)

•Origin: Kazakstan•Late Jurassic

•Origin: Madagascar•Early Triassic (100 mya)

(170 mya)(230 mya)

Apodops pricei

Probably Triassicfor All Orders Video

Paleobatrachus

2020

Lecture summary

General characteristics of amphibians

Characteristics of extant amphibian orders

Diversity of extant amphibian orders

Evolutionary history of amphibians