Amphibian Predation and Defense

Home , Clinotarsus, Clinotarsus alticola, Lithobates clamitans, Mount Lyell salamander, Pseudotriton

2/27/2017

By: Katie Harris

“Each stage of development from egg to adult faces it own particular dangers, and at no time of day or night are frogs and their offspring safe from attack.” –Tyler 1976

“almost anything will eat an amphibian.” – Porter 1972

Why Eat Amphibians?

Amphibian Predators

Evolutionary Response

Future Directions

Summary

1 2/27/2017

Why Eat Amphibians?

Amphibian Predators

Evolutionary 1. Egg Response 2. Larvae 3. Adult

Future Directions

Summary

Why eat an amphibian?

• Small, slow-moving, defenseless?

• High population densities

• Good source of protein

• Lack indigestible material

Why eat an amphibian?

• Small, slow-moving, defenseless?

• High population densities

• Good source of protein

• Lack indigestible material

2 2/27/2017

Why eat an amphibian?

• Small, slow-moving, defenseless?

• High population densities

• Good source of protein

• Lack indigestible material

Why eat an amphibian?

• Small, slow-moving, defenseless?

• High population densities

• Good source of protein

• Lack indigestible material

Common Predators?

3 2/27/2017

Common Predators?

• Invertebrates • eggs and larvae • temporary ponds • caddisfly larvae • parasites • trophic loop

Lethocerus americanus Pseudacris regilla

Common Predators?

• Invertebrates • eggs and larvae • temporary ponds • caddisfly larvae • parasites • trophic loop

Ambystoma maculatum Trichoptera

Common Predators?

• Invertebrates • eggs and larvae • temporary ponds • caddisfly larvae • parasites • trophic loop

Lithobates sylvaticus Macrobdella

4 2/27/2017

Common Predators?

• Invertebrates • eggs and larvae • temporary ponds • caddisfly larvae • parasites • trophic loop

Common Predators?

• Invertebrates • Fish • Eggs, larvae, adults

Common Predators?

• Invertebrates • Fish • Reptiles

5 2/27/2017

Common Predators?

• Invertebrates • Fish • Reptiles • Birds

Common Predators?

• Invertebrates • Fish • Reptiles • Birds • Amphibians

Common Predators?

• Invertebrates • Fish • Reptiles • Birds • Amphibians • Mammals

6 2/27/2017

Option 1: Option 2: Reduce the Reduce the chances likelihood of being of being consumed detected once detected

Chemical Behavioral Morphological Physiological Phenological

Ambystoma maculatum

Option 1: Option 2: Reduce the Reduce the chances likelihood of being of being consumed detected once detected

Chemical Behavioral Morphological Physiological Phenological

Ambystoma maculatum

Option 1: Option 2: Reduce the Reduce the chances likelihood of being of being consumed detected once detected

Chemical Behavioral Morphological Physiological Phenological

Ambystoma maculatum

7 2/27/2017

Option 1: Option 2: Reduce the Reduce the changes likelihood of being of being consumed detected once detected

Chemical Behavioral Morphological Physiological Phenological

Ambystoma maculatum

Eggs: Mechanical and Chemical

• Egg capsule and jelly

• Toxic and distasteful compounds

• Effectiveness depends on predator • Eggs of R. calmitans & R. catesbianus

• distasteful to newts and larval ambystomatids (Walters 1975)

• readily consumed by leeches (Howard 1978)

Eggs: Oviposition Site

• Areas without fish • temporary ponds • on land

• Areas void of predators • Anaxyrus americanus avoid oviposition in areas that contained Lithobates sylvaticus eggs (Pentranka et al. 1994) • L. sylvaticus didn’t lay eggs in ponds with predatory

sunfish (Hopey and Petranka 1994)

Lithobates sylvaticus

8 2/27/2017

Eggs: Oviposition Site

• Areas without fish • temporary ponds • on land

Lithobates sylvaticus

Eggs: Timing of Reproduction

• Early spring • before predatory species reach peak densities

• Synchronized reproduction • earlier hatchlings consume eggs deposited later • swamp predators increasing individual survival

Lithobates sylvaticus

Eggs: Timing of Reproduction

• Early spring • before predatory species reach peak densities

• Synchronized reproduction • earlier hatchlings consume eggs deposited later • swamp predators increasing individual survival

Lithobates sylvaticus

9 2/27/2017

Eggs: Adaptive Plasticity

• Red-eyed tree frog (Agalychnis callidryas) • egg mass hatches immediately if clutch is attacked (Warkentin 1995, Warkentin 2000, Warkentin et al. 2001) • survive by dropping in water • trade off!  small, not very mobile • wasp predation – single hatching (Warkentin 2000) • fungus – eggs closet to fungus hatch first

Agalychinis callidryas

Eggs: Adaptive Plasticity

Leptodeira

Eggs: Adaptive Plasticity

10 2/27/2017

Eggs: Adaptive Plasticity

• Hatching is delayed in the presence of predators (Sih and Moore 1993, Moore et al. 1996)

• Allows larvae to hatch at a more advanced stage

Ambystoma opacum

Larval Defense:

Lithobates clamitans

11 2/27/2017

Larval Defense: Color Patterns

• First line of defense

• Drab colors

• Countershading

• Disruptive coloration

• Reflective or transparent

• Deflection marks

Lithobates clamitans

Larval Defense: Color Patterns

• First line of defense

• Drab colors

• Countershading

• Disruptive coloration

• Reflective or transparent

• Deflection marks

LithobatesStorfer etclamitans al. 1999

Larval Defense: Color Patterns

• First line of defense

• Drab colors

• Countershading

• Disruptive coloration

• Reflective or transparent

• Deflection marks

Lithobates catesbianus

12 2/27/2017

Larval Defense: Color Patterns

• First line of defense

• Drab colors

• Countershading

• Disruptive coloration

• Reflective or transparent

• Deflection marks

Larval Defense: Color Patterns

• First line of defense

• Drab colors

• Countershading

• Disruptive coloration

• Reflective or transparent

• Deflection marks

Xenopus longipes

Larval Defense: Color Patterns

• First line of defense

• Drab colors

• Countershading

• Disruptive coloration

• Reflective or transparent

• Deflection marks

Acris sp.

13 2/27/2017

Larval Defense: Color Patterns

• First line of defense

• Drab colors

• Countershading

• Disruptive coloration

• Reflective or transparent

• Deflection marks

Clinotarsus alticola

Larval Defense: Phenotypic Plasticity • Acris

• tail tip coloration (Caldwell 1982) • Hyla chrysoscelis dragonfly naiads present • dragonfly naiads present • dark orange, deep tail fins,

dark (Van Buskirk & McCollum 1999) • predators absent fish present • Shallow, unmarked tail fins (Van Buskirk & McCollum 1999) • early age exposure = gradual development (LaFiandra & Babbitt 2004) • direct contact not Acris sp. necessary

Larval Defense: Phenotypic Plasticity • Acris

• tail tip coloration (Caldwell 1982) Present • Hyla chrysoscelis • dragonfly naiads present • dark orange, deep tail fins,

dark (Van Buskirk & McCollum 1999) • predators absent Absent • Shallow, unmarked tail fins (Van Buskirk & McCollum 1999) • early age exposure = gradual development (LaFiandra & Babbitt 2004) • direct contact not necessary

14 2/27/2017

Larval Defense: Phenotypic Plasticity • Acris

• tail tip coloration (Caldwell 1982) Present • Hyla chrysoscelis • dragonfly naiads present • dark orange, deep tail fins,

Larval Defense: Phenotypic Plasticity • Acris

• tail tip coloration (Caldwell 1982) Present • Hyla chrysoscelis • dragonfly naiads present • dark orange, deep tail fins,

Larval Defense: Behavioral

• Schooling (Rodel and Linsenmair 1997) • decreases chance of individual capture • Time of day

• shift active when predators are not active (Sih et al. 1992) • Ambystoma barbouri – tend to be nocturnal in

presence of fish (Taylor 1983) • Refuge habitats: less food

• water column – Ambystoma talpoideum (Holomuzki 1986) • trade-offs  vulnerability versus foraging efficiency

• Limit activity (Altwegg 2003)

Lithobates sylvaticus

15 2/27/2017

Larval Defense: Behavioral

• Schooling (Rodel and Linsenmair 1997) • decreases chance of individual capture • Time of day

• shift active when predators are not active (Sih et al. 1992) • Ambystoma barbouri – tend to be nocturnal in

presence of fish (Taylor 1983) • Refuge habitats: less food

• water column – Ambystoma talpoideum (Holomuzki 1986) • trade-offs  vulnerability versus foraging efficiency

• Limit activity (Altwegg 2003)

Lithobates sylvaticus

Larval Defense: Behavioral

• Schooling (Rodel and Linsenmair 1997) • decreases chance of individual capture • Time of day

• shift active when predators are not active (Sih et al. 1992) • Ambystoma barbouri – tend to be nocturnal in

presence of fish (Taylor 1983) • Refuge habitats: less food

• water column – Ambystoma talpoideum (Holomuzki 1986) • trade-offs  vulnerability versus foraging efficiency

• Limit activity (Altwegg 2003)

Lithobates sylvaticus

Larval Defense: Behavioral

• Schooling (Rodel and Linsenmair 1997) • decreases chance of individual capture • Time of day

• shift active when predators are not active (Sih et al. 1992) • Ambystoma barbouri – tend to be nocturnal in

presence of fish (Taylor 1983) • Refuge habitats: less food

• water column – Ambystoma talpoideum (Holomuzki 1986) • trade-offs  vulnerability versus foraging efficiency

• Limit activity (Altwegg 2003)

Lithobates sylvaticus

16 2/27/2017

(Strangel and Semlitsch 1987)

Lithobates sylvaticus

Larval Defense: Sensory Cues

• Tactile • Visual • Chemical • discrimination between dangerous and less dangerous genetic response-previous exposure is not required (Lefcourt 1996,1998, Gallie et al. 2001, Mathis et al. 2003) • discrimination between predators of conspecific larvae (Chivers and Mirza 2001)

• not always present, can develop rapidly (Kiesecker and Blaustein 1997)

Lithobates sylvaticus

(Chivers, Kiesecker, Blaustein 1996)

Predatory

17 2/27/2017

Larval Defense: Sensory Cues

• not always present, can develop rapidly (Kiesecker and Blaustein 1997)

Lithobates sylvaticus

Larval Defense: Alarm Responses

• Cues from injured conspecifics (Hews and Blaustein 1985)

• use these cues to avoid areas where predators are likely (Hews 1988)

• increase activity (different than most)

• beneficial if predator has already been detected (Hews 1998))

Lithobates sphenocephalus

Larval Defense: Growth & Development • Rapid growth • Predators have limited prey size

• Sprint speed increases with size (Richards and Bull 1990)

• Higher growth rate= higher survivorships (Travis 1983) • Growth and predator exposure

• Bufo boreas – increased growth rate (Chivers et al. 1999) • Ambystoma macrodactylum – decreased growth rate

• foraging decreased (Wildy et al 1999)

18 2/27/2017

Larval Defense: Growth & Development • Rapid growth • Predators have limited prey size

• Sprint speed increases with size (Richards and Bull 1990)

• Higher growth rate= higher survivorships (Travis 1983) • Growth and predator exposure

• Bufo boreas – increased growth rate (Chivers et al. 1999) • Ambystoma macrodactylum – decreased growth rate

• foraging decreased (Wildy et al 1999)

Larvae Defense: Chemical Defenses • Distasteful and toxic; particularly anuran tadpoles • Bufo- palatability varies among predators

• Why not in all species?

Bufo bufo

Larvae Defense: Chemical Defenses • Distasteful and toxic; particularly anuran tadpoles • Bufo- palatability varies among predators

• Why not in all species? • expensive to produce • temporary ponds: too costly, rapid growth • permanent ponds: worth the energy costs, slow growth

Bufo bufo

19 2/27/2017

Adult Defense: Cryptic Coloration

• Matching dorsal coloration to environment (Edmunds 1974)

• Reflect light in visible and infrared spectrum (Schalm et al. 1977, Emerson et al. 1990)

• Color change (Iga and Bagnara 1975)

• Cryptic patterning (Cott 1940)

Hyla chrysoscelis

Adult Defense: Cryptic Coloration

• Matching dorsal coloration to environment (Edmunds 1974)

• Reflect light in visible and infrared spectrum (Schalm et al. 1977, Emerson et al. 1990)

• Color change (Iga and Bagnara 1975)

• Cryptic patterning (Cott 1940)

Hyalinobatrachium taylori

Adult Defense: Cryptic Coloration

• Matching dorsal coloration to environment (Edmunds 1974)

• Reflect light in visible and infrared spectrum (Schalm et al. 1977, Emerson et al. 1990)

• Color change (Iga and Bagnara 1975)

• Cryptic patterning (Cott 1940)

Hyla cinerea

20 2/27/2017

Adult Defense: Cryptic Coloration

• Matching dorsal coloration to environment (Edmunds 1974)

• Reflect light in visible and infrared spectrum (Schalm et al. 1977, Emerson et al. 1990)

• Color change (Iga and Bagnara 1975)

• Cryptic patterning (Cott 1940)

Ceratobatrachus guentheri - Soloman Island Frog

Adult Defense: Color Polymorphism

• Polymorphism = different forms

• Genetically controlled (Merrell 1965, Fogelman et al. 1980)

• can be environmentally influenced (Travis and Trexler 1984, Harkey and Semlitch 1988)

• Pacific tree frog (Pseudacris regilla)

• color-changing morph (brown/green) (Wente and Phillips 2003) • slow  not effective to immediate predation • adaptive  seasonal shifts • light intensity and temperature  thermoregulation (Stegen et al. 2004)

Acris crepitans

Adult Defense: Color Polymorphism

• Polymorphism = different forms

• Genetically controlled (Merrell 1965, Fogelman et al. 1980)

• can be environmentally influenced (Travis and Trexler 1984, Harkey and Semlitch 1988)

• Pacific tree frog (Pseudacris regilla)

Acris crepitans

21 2/27/2017

Adult Defense: Color Polymorphism

• Polymorphism = different forms

• Genetically controlled (Merrell 1965, Fogelman et al. 1980)

• can be environmentally influenced (Travis and Trexler 1984, Harkey and Semlitch 1988)

• Pacific tree frog (Pseudacris regilla)

Acris crepitans

Adult Defense: Predator Avoidance

• Avoid locations where predators present

• Chemical cues • predators • injured conspecifics

• Newts (Woody and Mathis 1998)

• did not avoid chemical stimuli from fish (Marvin and Hutchinson 1995, Woody and Mathis 1997) • avoided chemical stimuli from fish if associated with injured conspecific (Woody and Mathis 1998)

Plethodon vandykei

Adult Defense: Predator Avoidance

• Avoid locations where predators present

• Chemical cues • predators • injured conspecifics

• Newts (Woody and Mathis 1998)

• did not avoid chemical stimuli from fish (Marvin and Hutchinson 1995, Woody and Mathis 1997) • avoided chemical stimuli from fish if associated with injured conspecific (Woody and Mathis 1998)

Plethodon vandykei

22 2/27/2017

Adult Defense: Predator Avoidance

• Avoid locations where predators present

• Chemical cues • predators • injured conspecifics

• Newts (Woody and Mathis 1998)

• did not avoid chemical stimuli from fish (Marvin and Hutchinson 1995, Woody and Mathis 1997) • avoided chemical stimuli from fish if associated with injured conspecific (Woody and Mathis 1998)

Plethodon vandykei

Adult Defense: Behavioral

• Jumping – slender bodies (Heinen and Hammond 1997)

• Flipping – Plethodontids (Whiteman and Wissinger 1991, Azizi and Landberg 2000)

• Coiling – Mount Lyell salamander Hydromantes playcephalus (Garcia- Paris and Deban 1995)

• Death feigning – leaf litter frog Ischnocnema aff. henselii (Vaz-Silva et. al 2004)

• Vocalization (Weber 1978)

• Biting (Taylor and Ludlam 2013)

• Tail display – Plethodontidae, Salamandridae (Wake and Dresner 1967)

• Unken reflex (Kuchta 2005)

Rana so.

Adult Defense: Behavioral

• Jumping – slender bodies (Heinen and Hammond 1997)

• Flipping – Plethodontids (Whiteman and Wissinger 1991, Azizi and Landberg 2000)

• Coiling – Mount Lyell salamander Hydromantes playcephalus (Garcia- Paris and Deban 1995)

• Death feigning – leaf litter frog Ischnocnema aff. henselii (Vaz-Silva et. al 2004)

• Vocalization (Weber 1978)

• Biting (Taylor and Ludlam 2013)

• Tail display – Plethodontidae, Salamandridae (Wake and Dresner 1967)

• Unken reflex (Kuchta 2005)

Plethodon cinereus

23 2/27/2017

Adult Defense: Behavioral

• Jumping – slender bodies (Heinen and Hammond 1997)

• Flipping – Plethodontids (Whiteman and Wissinger 1991, Azizi and Landberg 2000)

• Coiling – Mount Lyell salamander Hydromantes playcephalus (Garcia-Paris and Deban 1995)

• Death feigning – leaf litter frog Ischnocnema aff. henselii (Vaz- Silva et. al 2004)

• Vocalization (Weber 1978)

• Biting (Taylor and Ludlam 2013)

• Tail display – Plethodontidae, Salamandridae (Wake and Dresner 1967)

• Unken reflex (Kuchta 2005)

Ischnocnema aff. henselii

Adult Defense: Behavioral

• Jumping – slender bodies (Heinen and Hammond 1997)

• Flipping – Plethodontids (Whiteman and Wissinger 1991, Azizi and Landberg 2000)

• Coiling – Mount Lyell salamander Hydromantes playcephalus (Garcia-Paris and Deban 1995)

• Death feigning – leaf litter frog Ischnocnema aff. henselii (Vaz- Silva et. al 2004)

• Vocalization (Weber 1978)

• Biting (Taylor and Ludlam 2013)

• Tail display – Plethodontidae, Salamandridae (Wake and Dresner 1967)

• Unken reflex (Kuchta 2005)

Lithobates catesbianus

24 2/27/2017

Adult Defense: Behavioral

• Jumping – slender bodies (Heinen and Hammond 1997)

• Flipping – Plethodontids (Whiteman and Wissinger 1991, Azizi and Landberg 2000)

• Coiling – Mount Lyell salamander Hydromantes playcephalus (Garcia-Paris and Deban 1995)

• Death feigning – leaf litter frog Ischnocnema aff. henselii (Vaz- Silva et. al 2004)

• Vocalization (Weber 1978)

• Biting (Taylor and Ludlam 2013)

• Tail display – Plethodontidae, Salamandridae (Wake and Dresner 1967)

• Unken reflex (Kuchta 2005)

Amphiuma tridactylum

Adult Defense: Behavioral

• Jumping – slender bodies (Heinen and Hammond 1997)

• Flipping – Plethodontids (Whiteman and Wissinger 1991, Azizi and Landberg 2000)

• Coiling – Mount Lyell salamander Hydromantes playcephalus (Garcia-Paris and Deban 1995) • Death feigning – leaf litter frog Ischnocnema aff. henselii (Vaz-Silva et. al 2004)

• Vocalization (Weber 1978)

• Biting (Taylor and Ludlam 2013)

• Tail display – Plethodontidae, Salamandridae (Wake and Dresner 1967)

• Unken reflex (Kuchta 2005)

Batrachoseps relictus

Adult Defense: Behavioral

25 2/27/2017

Adult Defense: Behavioral

• Jumping – slender bodies (Heinen and Hammond 1997)

• Flipping – Plethodontids (Whiteman and Wissinger 1991, Azizi and Landberg 2000)

• Coiling – Mount Lyell salamander Hydromantes playcephalus (Garcia-Paris and Deban 1995)

• Death feigning – leaf litter frog Ischnocnema aff. henselii (Vaz- Silva et. al 2004)

• Vocalization (Weber 1978)

• Biting (Taylor and Ludlam 2013)

• Tail display – Plethodontidae, Salamandridae (Wake and Dresner 1967)

• Unken reflex (Kuchta 2005)

Taricha granulosa

Adult Defense: Chemical

• Granular glands • Found in all amphibians, relatively similar structure • Distributed throughout the skin or concentrated • Defensive secretions 1. Widespread, naturally occurring compounds. • Can serve other functions  microbes and fungi • Defensive when secreted in large quantities 2. Evolved for defense  toxic • Origin

• Synthesized within the animal (Smith et al. 2002)

• Derived from chemicals of other organisms (Daly et al. 1994)

Anaxyrus sp.

Adult Defense: Chemical

• Synthesized within the animal (Smith et al. 2002)

• Derived from chemicals of other organisms (Daly et al. 1994)

Anaxyrus sp.

26 2/27/2017

Adult Defense: Chemical

• Synthesized within the animal (Smith et al. 2002)

• Derived from chemicals of other organisms (Daly et al. 1994)

Anaxyrus sp.

Adult Defense: Aposematic Coloring

• Coloration to warn or deter predators

• Only beneficial when predator can see colors

• Dendrobatidae • Color brightness correlates to toxicity (Summers and Clough 2001)

HylaOophagachrysoscelispumilio

Adult Defense: Mimicry

• Predators learn to avoid the aposematic patterns • Batesian mimicry: edible species benefit by evolving pseudoaposematic colors • Pseudotriton ruber & Notophthalmus viridescens

• readily eaten before exposure to red efts (Brodie & Howard 1972)

• less palatable than Desmognathus species (Brandon et al. 1979) • produce toxins  Mullerian • Mullerian mimicry: toxic species share a common warning coloration

Notophalmus viridescens Pseudotriton ruber

27 2/27/2017

Future Directions

• Impact on populations • demographic impact of predation • impact of specific predators • invasive predators

• Genetics

• Relationships between toxicity and predators

• Caecilians and salamanders

Lithobates sylvaticus

Summary

• Subject to vast array of predators • Predation is greatest on egg and larval stages • Most predators are generalists • Protection against predators is varied • Predation lessened by: • reduce the change of detection • reduce the changes of being consumed once detected • chemical, phenological, behavioral, physiological, morphological

Ensatina eschscholtzii

References

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Ensatina eschscholtzii

28 2/27/2017

References

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Photo Credits

• Anna Spyrka • Kristiina Hurme • Amber Hart • Larry Kimball • Audobaun Society • Mike Benard • Barbara Magnuson • Montgomery Co. Dept. of Environmental Protection • Brandon Ballengee • Nancy Tray • Brian Gratwicke • Natalle McNear • Christina Rollo • Nathan Shepard • David Huth • Oliver Kratz • Douglas Mills • Peter May • Dwight Kuhn • Ricky Relya • Gary Nafis • Robert Clontz Photography • Greg Pecold • Rod Williams • Harikrishnan S. • Ryan Photographic • Jacob Loyacano • The Amphibain Foundation • Jason Gibson • Warren Photographic • Johnathan Mays • Werther Pereira Ramalho • John Whit • John White • Jones Fish Hatcher

Ensatina eschscholtzii

Questions?

Katie Harris 240 EPS Building [email protected]