Vertebrate Evolution Chapter #1 – The Diversity, Classification, and Evolution of Vertebrates, pg. 2 – 15. Chapter #2 – Vertebrates and Basic Structure, pg. 19 – 45. Chapter #3 – Early Vertebrates, Jawless and Jawed Vertebrates, pg. 47 – 73. Amphibians What is a (Liss)Amphibian? Liss = smooth (greek – referring to skin) Amphibia = double life (greek – referring to metamorphosis) Jurassic Long hind limbs (before break-up of Pangea) HOP Short, stiff bodies Equal-sized limbs Long, slender bodies WADDLE No limbs Long, slender bodies 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 2) Pelvic Patch: Highly vascularized skin; absorbs majority of water Warm, dry nights 3) Permeable Bladder: Store dilute urine; “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) • Poison glands: • Concentrated on dorsal surface Tarichatoxin (newts) Water-soluble alkaloid (blocks Na+ channels) • Produce diverse array of chemicals Epibatidine (dart-poison frogs) Lipid-soluble Alkaloid (Blocks Ach receptors) Non-addictive painkiller Batrachotoxin (dart-poison frogs) 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 exchange) • Gills – Larval forms and neotenics (neoteny = adults that retain juvenile characteristics) 3) Pedicellate Teeth: • Crown and base of tooth separated by uncalcified dentine / fibrous tissue • 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 window) • 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 unit (ground) • Vibration via leg bones / scapula (activated via opercularis muscle) • 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 species 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 cutaneous) • Cryptobrachidae = large skin folds • Terrestrial = lungs (50%) / skin (50%) Adults retain: laterally compressed tail / flattened head functional lateral line system external gills (not always…) • Anguilliform locomotion (walking-trot / swim) Plethodontidae = lungless skin (10%) & buccopharynx (90%) Hypothesis: Evolved in fast, cold water ( O2) Lungs disadvantageous (buoyancy) Amphibians Salamanders (10 families; ~ 515 species): Diet / Prey Capture: • Generalist carnivores (primarily invertebrates) • Aquatic = mouth gape • Suction pulls prey into oral cavity Cryptobrachus alleganiensis movie • Works will with gills / gill slits (one-way flow) • Tongue = broad, flat, immobile • Terrestrial = Thick, sticky tongue grabs prey • Plethodontidae: Projective tongue Why 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 nares • Aids in collection / delivery of chemical cues (capillary action) • Sexually dimorphic (esp. during reproductive season) • Hearing – limited high frequency (no ear drum); primarily ground vibration • 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) Reproduction: • External fertilization; female nest guarding Cryptobranchidae (Hellbenders): Aquatic: SE United States / NE Mexico • largest individuals • 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 (Amphiums): • terrestrial egg-laying • 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) Habitat: • Prefers cool forests with litter / dead trees • Tolerates logging better than most plethodons Terrestrial: North / Central / South America & Europe 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 • Elaborately patterned courtship “dance” • “Tail straddle-walking” (unique to plethodons) • ♂ repeats dance (~ 1 – 5 hours) • Pathway / movement unique to each species Pheromone: Chemical cue that affects the behavior and / or physiology of a conspecific 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 & Europe 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 Growth / Maturation: • Sexual Maturation ~ 4 years • Life Span ~ 10 years (best guess) Anti-predator Display Terrestrial: NW United States Amphibians Larval Anatomical Specializations: Pond Larvae Ambystomatidae Salamandridae • High body profile • broad tail fin extending up to head • long, plume-like gills • Only front legs at hatching Stream Larvae Dicamptodontidae Rhyacotritonidae Plethodontidae • Depressed body profile • Narrow tail fin (not onto trunk) • Short, thread-like / curly gills • All four 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) • Neurotoxin – blocks NA+ channels (paralysis) • Small dose lethal to birds / mammals • Large dose lethal to humans Red-spotted Garter Snake “The Arms Race” North America / Europe / NW Africa / Asia Assignment: http://evolution.berkeley.edu/evolibrary/article/biowarfare_01 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) • ♀ 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 Terrestrial: North America / Europe / NW Africa / Asia Amphibians Dicamptodontidae: (example = Dicamptodon tenebronsus) Anatomy: • Full metamorphosis / facultative paedomorphosis • Large (up to 30 cm SVL) • Well-developed lungs / blade-like teeth Habitat: • Damp coniferous forests (near streams / seeps) • 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… Terrestrial: NW United States / SW Canada 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) Terrestrial: Western North America • Enter torpor during winter (below frost line) 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 Life History: • 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) Order: Gymnophiona (“Naked snake”) Tropics (S. America / Africa / S.E. Asia) Fossorial / Aquatic 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 species 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 Short limbs / Robust body Semi-aquatic Frogs Webbed feet / stream-lined Arboreal Frog 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 Direct Development (egg protection) (tadpole protection) Guard Eggs Vegetation over water Foam Nest (egg protection) (egg protection) Bromeliad Nest (egg protection) (tadpole protection) • Parental Care (common): Carry Eggs (Mid-wife Toad ♂) Carry Eggs / tadpoles (Sirinam Toad ♀) Internal Brooding (♀ Gastric Brooders) Viviparous (Morogoro Tree Toad) Amphibians Still Water: Ovoid bodies Tails with large fins Frogs (29 families; ~ 4800 species): Tadpoles: • Aquatic larval stage of frogs • Advantages: 1) Exploit non-adult resources • Tadpoles = aquatic / herbivores • Adults = terrestrial / carnivores 2) Efficient eating machines (rapid growth) Fast Water: Stream-lined bodies Small tail fins Metamorphosis: (3 stages) • Triggered by thyroid hormone 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 formation Amphibians Rhinophrynidae: (Burrowing Toad) • Single species • Unique tongue Morphology: • Short, powerful limbs; tubercle present • Robust body; pointed head (cornified tip) Habitat: • Sub-humid areas; surface after heavy rains Fossorial Reproduction: • Explosive breeder; aquatic tadpole North America / Central America Leptodactylidae (Neotropical frogs): • Among largest Morphology: • Highly variable (10 mm • Many nocturnal 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: (Microhylid frogs) Morphology: • Highly variable (10 mm • Commensal assoc. with spiders 100 mm) Habitat: • Variable (Arid deserts Wet rain forests) Reproduction: • Explosive breeders / prolonged breeders • Aquatic tadpoles / direct development Terrestrial / Fossorial / Arboreal Ubiquitous Dendrobatidae (Poison-dart frogs): Morphology: • Relatively small; highly aposematic Habitat: • Tropical rainforests Reproduction: • Most lack amplexus • Small clutches; parental care • Poisonous • Introduced to Hawaii 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 Aquatic South America / Africa Cane Toad (Bufo marinus) “… one of the 100 worst invasive Invasive Species species worldwide.” Specialist Group 1) Prolific breeders (~ 33,000 eggs) 2) Tolerance for temperature / salinity 3) All stages of life poisonous (bufotoxin) 4) Generalist diet (including cat food) • Trap / hand catch frogs • Biocontrol (virus) No native toad species 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 Habitat: • 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 Aquatic: Northwest United States Amphibians Pelobatidae (example = Scaphiopus intermontana): Anatomy: • Keratinous, spade-like metatarsal tubercle • Rotund body; short legs; large eyes • Glandular, tuberculate skin Habitat: • 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 Terrestrial: North America / Europe / Asia / Africa Amphibians Bufonidae (example = Bufo boreas): Anatomy: • Bidder’s 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 Habitat: • 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 ~ ??? Primarily Arboreal: North / Central / South America Amphibians Ranidae (example = Rana Aurora ): Red located on lower abdomen / hind limbs Anatomy: • Mid-sized body: 2 - 3 inches in length • Long rear legs; exceptional leaping ability • Presence of vocal sacs dependent on sub-species Habitat: • Require riparian vegetation near ponds / streams • Move into upland forest outside of breeding season Terrestrial: Cosmopolitan 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 • Life Span ~ 10 years Believed to be inspiration for Twain’s "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 Global Distribution of Amphibians • Eastern Australia • Central / Southern China • West Africa Why are populations declining? (7 major hypotheses…) Global Distribution of Threatened Amphibians Amphibians Amphibian Decline - Reasons: 1) Habitat Changes: California redlegged 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: 1) Habitat Changes: California redlegged 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 Illicit Pet Trade (~ 80,000 harvested / year) (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 Nitrogen Pollution: Reduced feeding activity, swim less vigorously, Display disequilibrium, development malformations Synergisms most likely at work… 7) Disease: Common Frog Virus Yellow-legged Frog Bacteria Western Toad Water Mold Pacific Tree Frog Trematodes