Phone: 974-0831 (JH)
Office: 230 Ellington PSB (JH)
Meeting Time and Place: 5:057:05 pm T,R 113 PBB (2 field trips required: 18 March and 17 April)
Course Goal: To expose students to the life history, diversity, ecology, conservation, and management of amphibians through a combination of lectures, readings, class discussions, labs, and field experiences.
Expected Outcomes: Students that successfully complete WFS 433/533 will have a basic understanding of amphibian identification, anatomy, life history, and ecology. They will be aware of potential mechanisms of amphibian declines, and understand how to identify, collect and ship diseased amphibians to a diagnostic lab. Students will understand how to sample amphibian populations and be aware of conservation strategies.
Required Text: The Ecology and Behavior of Amphibians, 2007 (ISBN 9780226893341)
Author: Kentwood D. Wells
Journal Papers: Occasionally journal papers will be assigned instead of or to supplement the required text. Papers will be provided in class or on the course website.
Weights of Academic Assessments:
● Test #1
● Test #1
● Test #2
● Test #2
● Amphibian ID Exam
● Amphibian ID Exam
1Participation includes attendance on the 2 field trips and participation in the 3 laboratory activities. You will earn 4% and 2% for attending each field trip and participating in each laboratory activity for WFS 433 and 533, respectively.
2If you miss a field trip or laboratory activity, you can either: (1) write a 10-page scientific paper on an amphibian topic of your choice or (2) accept the percent deduction (4% or 2%) in your final grade corresponding with the appropriate course number.
3One hour lecture on an amphibian topic please see Dr. Hoverman before 2 March 2010 to discuss potential topics.
Your course grade will be determined using the following scale:
Final Weighted Percent
Final Weighted Percent
You can positively influence your final grade as much as 4.5% by volunteering for extra credit. Volunteer work must be related to herpetofauna, and can include work on university projects or with government agencies or NGOs. For every 8 hours of volunteer work, your final grade will be increased by 1.5% up to 4.5% (or 24 hours total). All volunteering must be completed by 29 April 2010. A volunteer form (see below) must be filled out by the supervising individual. Scott Dykes and Pete Wyatt (TWRA non-game biologists) often are looking for volunteer assistance (Region 4 Office: 1-800-332-0900; email@example.com, firstname.lastname@example.org). Also, Wayne Schacher (consulting biologist for Seven Islands Wildlife Refuge) frequently needs help with herptofaunal surveys (457-4355 home, email@example.com). Nathan Haislip (WFS 433/533 TA), Kevin Hamed (WFS 433/533 TA), and Matthew Niemiller (PhD Candidate in EEB, firstname.lastname@example.org) may need assistance collecting amphibians or conducting experiments during the semester. Lastly, students can volunteer with the Tennessee Amphibian Monitoring Program for extra credit; interested students should contact Dr. Gray (email@example.com, 974-2740).
Tadpole Morphology (Haislip Lecture)
TEST #1 Material
1) Amphibian Diversity and Evolution
Wells: pp. 1-15, 41-58, 65-74, 77-80
NOTE: Questions given in lecture slides.
Wells: pp. 16-41, 59-65, 75-77
2) Amphibian Phylogenetics and Phylogeography
3) Amphibian Declines
a. What makes amphibians especially vulnerable to declines?
Wells: pp. 787-792
b. What are some species in North America with relic populations?
Wells: pp: 793-794
c. What is the region of the United States with the greatest number of species declining? Also, be able to list a few species with distributions east of the Mississippi that are declining.
Wells: pp: 800-803
d. Know which island in the South Pacific likely has the highest diversity of amphibian species per unit area in the world.
Wells: p. 795
e. Be able to provide an argument for why we should care that amphibians are declining.
Wells: pp. 850-853
Wells: pp. 816-850 (hypotheses for declines)
4) Reproductive Strategies
a. Be able to describe different strategies for anuran oviposition in standing water.
Wells: pp. 465-468
b. Be able to describe the differences between bubble and foam nests used in anuran oviposition and their adaptive significance.
Wells: pp: 472-478
c. Be able to describe the differences between two modes of salamander reproduction and their adaptive significance.
Wells: pp. 488-493
d. Be able to provide an explanation why some salamanders that deposit eggs in still water lay their eggs in clumps while others scatter single eggs.
Wells: pp. 788-489
e. Know which family of salamanders only exhibits direct development.
Wells: p. 491
f. Know the two salamander genera that exhibit ovoviviparity and viviparity.
Wells: p. 493
Wells: pp. 465-493 (modes of anuran reproduction)
Zug Handout (pp. 169-189, Zug 1993, Herpetology, Academic Press).
5) Salamander Courtship and Mating
a. Know the median home range for anurans and salamanders, and how they compare with birds and mammals.
Wells: pp: 230-231
b. Know the 4 ways that salamanders communicate, and be able to describe their primary function(s).
Wells: pp. 404-411
c. Know the 3 locations of chemical receptors in salamanders, and the 2 chemosensory organs in the nasal cavity. Also, know which sex the chemosensory organs are usually larger and why this may occur.
Wells: pp. 417-418
d. In the work performed by Robert Jaeger and Alicia Mathis on red-backed salamander territoriality, know the most important determinant of territory quality.
Wells: pp. 424-425
e. Be able to describe the adaptive significance of internal fertilization via a spermatophore, and the difference between the duration that sperm survive in the spermatheca in ambystomatids vs plethodontids.
Wells: pp. 459-461
f. Understand the relationships between female body size and clutch size, female body size and egg size, egg size and clutch size, egg size and hatchling size, egg size and development rate, and developmental rate and temperature.
Wells: pp. 497-500
g. Be able to describe selective advantages of species that produce small vs. large eggs, and the environmental constraints that may drive these relationships.
Wells: pp: 502-504
h. Know which mode of development has species that produce the largest eggs relative to body size.
Wells: p: 504
Wells: pp. 254-266 (orienteering)
Wells: pp. 452-461 (external vs. internal fertilization)
6) Anuran Movements, Courtship, and Mating
a. Understand the difference between home range, migration and dispersal.
Wells: pp. 230-231
b. Know which age class disperses most frequently in an amphibian population.
Wells: pp. 243-244
c. Be able to provide some hypotheses for the adaptive significance of homing (i.e., site fidelity).
Wells: p. 253
d. Know the principal site of extraocular photoreception and how amphibians use polarized light to orienteer.
Wells: pp. 261-264
e. Be able to describe the 2 auditory channels in amphibians, and know which is sensitive to low frequencies (<300 Hz).
Wells: pp. 322-324
f. Be able to provide a couple examples of anurans that do not have vocal sacs, and an explanation of why this may have occurred.
Wells: p. 277
g. Be able to describe unison bout calling and be able to provide some explanations why it may occur.
Wells: p. 297
h. Schwartz (1991) proposed 3 hypotheses for the duration of unison bouts. Be able to describe those and know which is most plausible.
Wells: p. 297
i. Understand the difference between explosive vs. prolonged breeders, and be able to provide some reasons why cold-weather breeders and species that inhabit xeric environments may breed explosively.
Wells: pp. 339-341
j. Understand factors that influence sexual selection in prolonged vs. explosive breeding systems. Also, understand how energy allocation differs between prolonged and explosive breeders, and a few strategies that prolonged breeders may use to reduce energy spent on calling.
Wells: pp. 342-343
k. Be able to provide at least 2 explanations why inguinal amplexus is considered more primitive than axillary amplexus.
Wells: pp. 454-456
Wells: pp. 269-304, 314-316 (anuran calls)
7) Tadpole Ecology and Metamorphosis
What do tadpoles eat? Altig et al. (2007)
a. Understand how limb development is different between larval salamanders and tadpoles.
Wells: p. 559
b. Be able to describe a few oral adaptations of larval salamanders that assist in suction feeding.
Wells: pp: 561-562
c. Know whether salamander larvae are herbivorous, omnivorous or carnivorous.
Wells: p. 562
d. Be able to explain some of the costs and benefits of cannibalism for larval salamanders. What are some explanations for cannibalizing different species?
Wells: pp. 563-564
e. Be able to provide some examples of tadpole species that delay lung development and why this may occur.
Wells: pp. 565
f. Be able to provide some explanations why the limbs of tadpoles emerge late in development compared to salamanders.
Wells: pp. 565-566
g. Understand the mechanism that induces carnivory in spadefoots found in the southwestern United States. Also, understand how frequency of cannibalistic and omnivorous phenotypes differs between spadefoot species (S. bombifrons and S. multiplicata) and why this may occur.
Wells: pp. 575-576
h. Although most amphibian larvae are solitary, be able to provide some benefits of aggregation behavior and what may be the adaptive significance of each benefit.
Wells: pp. 588-590
i. Be able to describe the selfish herd effect and provide an example.
Wells: p. 589
j. Be able to describe kin recognition and offer some explanations why this may be advantageous.
Wells: pp. 591, 593-596
k. Be able to explain the factors that drive a larval amphibian to decide when to initiate metamorphosis according to Werner (1986, 1988) and Rowe and Ludwig (1986, 1988). NOTE: Your explanation should discuss differential mortality and growth rates in the aquatic and terrestrial environments and the amount of time between metamorphosis and the age of first reproduction.
Wells: pp. 601-602
l. Know what hormone likely is responsible for initiating metamorphosis in response to an environmental stressor.
Wells: p. 608
Petranka and Kennedy 1999: Tadpoles: Macrophagous Predators
Wells: pp. 564-566, 604-608
8) Phenotypic Plasticity
Wells pp. 601-603, 609-610, 618-628, 632-642
Gotthard and Nylin 1995. Oikos 74:3-17
Relyea 2007. Oikos 152:389-400
Wells pp. 563-564, 573, 575, 596-597, 693-728
Basics of plasticity (Gotthard & Nylin 1995)
1) Be able to define phenotypic plasticity
2) Know the difference between adaptive and non-adaptive plasticity
3) Know what 4 conditions favor the evolution of plasticity
Metamorphosis and phenotypic plasticity
1) Know what factors (e.g., environmental conditions) can affect the decision to metamorphose (from lecture)
2) Understand the Wilbur and Collins model of metamorphosis and its predictions (Wells pp. 609-610)
3) Leips and Travis 1994 - know the experiments, results, and interpretation with respect to the Wilbur and Collins model (Wells pp. 618-619)
4) Be able to list the costs and benefits of plasticity in timing and size at metamorphosis (Wells pp. 625-628)
5) Know the specific environmental cues used to detect pond drying (Wells pp. 621-622)
Facultative paedogenesis (paedomorphosis)
1) Know the salamander families that display facultative paedogenesis and the % occurrence across salamanders (Wells pp. 632-637)
2) Be able to list the costs and benefits of paedogenesis (Wells pp. 639-642)
3) Understand the factors that affect the paedogenesis (Wells p. 638)
4) Understand the models developed to understand paedogenesis (Wells p. 638)
5) Semlitsch 1987 - be able to describe the experiment and the results (Wells p. 640)
1) Be able to describe the morphological changes that occur in cannibalistic tadpoles and salamanders (Wells pp. 563-564, 573, 575)
2) Know the types of environments that are frequently associated with cannibalistic phenotypes (Wells pp. 563-564)
3) Know the groups that display cannibalism (Wells p. 563)
4) Know the mechanisms that trigger cannibalism in tadpoles and salamander larvae (Wells pp. 563-564)
5) Be able to list the costs and benefits of cannibalism (Wells pp. 596-597)
6) Know how kin relationships impact the decision to cannibalize (Wells pp. 596-597)
Egg hatching plasticity (Wells pp. 696-697)
1) Know the work of Karen Warkentin (http://people.bu.edu/kwarken/) on this topic
2) Know the costs and benefits of egg hatching plasticity
3) Know the differences in predation strategy between snakes and wasps
Predator-induced plasticity Directly from the lecture unless otherwise noted
1) Know the types of cues tadpoles use to detect predators
2) Know which cue is most useful and reliable for detecting predators
3) Know the distinction between alarm cues and kairomones
4) Be able to describe how to design experiments to examine predator-induced plasticity
5) Be able to list and describe (e.g., the direction) the behavioral and morphological responses to predators
6) Be able to describe the costs and benefits of predator-induced plasticity
7) Be able to describe the theoretical predictions for how tadpoles will responded to combined predators (e.g., multiple predators encountered simultaneously)
8) Know how tadpoles respond to combined predators and what drives this decision
9) Know whether tadpoles rely on alarm cues or kairomones to detect predators
10) Know how predator diet impacts the formation of predator-induced defenses
11) Be able to explain the Werner m/g model of metamorphosis and its predictions with respect to predation risk (Wells p. 601)
12) Know whether there is support for the Werner m/g model and be able to explain your answer (Relyea 2007)
13) Know the adaptive reasoning behind the reversibility of predator-induced plasticity
Competitor-induced plasticity Directly from the lecture
1) Be able to list the responses of tadpoles to competitors
2) Be able to describe the interactive effects of competition and predation on the expression of tadpole phenotypes
Predator defenses in adults Directly from the lecture and Wells pp. 709-727
1) Be able to list and describe the behavioral responses of adults to predators
2) Be able to list and describe the function significance of cryptic coloration in adults
TEST #2 Material
1) Amphibian Sampling Techniques: Populations and Disease
· SE PARC Collection and Shipping Information Sheet #9 (Miller and Gray)
· SE PARC Disinfection Information Sheet #10 (Miller and Gray)
· Chapter 26 (Green et al. 2009: remaining sections): Disease Monitoring and Biosecurity (in Dodd 2009)
2) Community Ecology
· Wells pp. 694-696, 745-758, 768-778
· Wellborn et al. (1996): Annual Review of Ecology and Systematics 27:337-363
· Werner et al. (2007): Oikos 116:1697-1712
· Wells pp. 729-744, 779-783
Basics of Community Ecology Directly from the lecture
1) Be able to define community ecology
2) Know the goals of community ecology
3) Know the difference between direct and indirect effects
4) Be able to describe the hierarchical structure of ecological communities
Amphibian Community Ecology Directly from the lecture unless otherwise noted
1) Know the consequences of a biphasic life cycle for amphibian communities (Wells pp. 754-755)
2) Be able to define:
a. Priority effects
b. Asymmetric competition
c. Character displacement
d. Keystone predation
3) Know the factors that influence the oviposition decision of adults (Wells pp. 694-696)
4) Be able to describe how environmental conditions impact the competitive interactions between spadefoot toad species and the role of character displacement in understanding the interaction
5) Know the role of breeding phenology in understanding priority effects and be able to provide an example in amphibian communities (Wells pp. 768-769)
6) Be able to discuss the importance of predation, pond hydroperiod, and species traits in the driving the distribution of larval green frogs and bullfrogs (Wells pp. 777-778)
7) Be able to discuss the roles of predators and pathogens in altering competitive interactions between tadpole species (Wells p. 769)
8) Be able to discuss the role of keystone predators in structuring tadpole communities and an example of keystone predation (Wells pp. 772-773)
9) Be able to discuss how environmental conditions (e.g., hydroperiod) interact with keystone predators to structure tadpole communities
10) Be able to discuss the conceptual model of Wellborn et al. (1996) that explains the mechanisms structuring aquatic communities
11) Be able to discuss patterns in amphibian species richness along environmental gradients (Werner et al. 2007)
3) Graduate Student #1 Lecture
4) Graduate Student #2 Lecture
5) Invasive Species
6) Ranaviruses and Bd
a. What was the first country to report die-offs from Bd?
Wells: p. 831
b. Be able to describe some weaknesses of the Out of Africa hypothesis.
Wells: p: 834
c. Be able to describe how global climate change may be contributing to the emergence of Bd.
Wells: p. 834
d. Be able to describe how differences in antimicrobial peptides (AMPs) associated with the skin of amphibians may be responsible for differences in susceptibility to Bd that have been observed among amphibian species.
Wells: pp. 835-837
· Gray et al. (2009): Ecology and pathology of amphibian ranaviruses
· Kilpatrick et al. (2010): The ecology and impact of chytridiomycosis
Wells: pp. 831-838
7) Climate Change
· Wells: pp. 848-850
· No Required Readings
· Recommendations for Riparian Buffers: Salamanders (Crawford and Semlitsch 2007)
· Recommendations for Wetlands Buffers: Amphibians and Reptiles (Semlitsch and Bodie 2003)
Podcasts: (MP3 Format)
1. iTunes Instructions: (you must first download iTunes to Listen to Podcasts)
1) Go to http://itunesu.utk.edu
2) Click on "Download iTunes & Quick Time"
3) Click on Download iTunes Free
4) Save iTunes to your hard drive and install.
2. Link to iTunes to Listen to Podcasts: Launch Podcasts in iTunes U
Slides: (PDF Format)
Lectures: Test 1
Tennessee Salamander Identification: 2009 Lecture (Guest Lecturer: Matthew Niemiller, UTK)
Diversity of Amphibia (Kevin Hamed, UTK)
Amphibian Phylogenetics and Phylogeography (Guest Lecturer: Matthew Niemiller, UTK)
Tadpole Ecology and Metamorphosis (Dr. Matt Gray, UTK)
Phenotypic Plasticity (Dr. Jason Hoverman, UTK)
Lectures: Test 2
Amphibian Sampling Techniques (Kevin Hamed, UTK)
Community Ecology (Dr. Jason Hoverman, UTK)
Landscape Ecology (Andrew West, UTK, WFS 533 Presentation)
Zoos and Captive Breeding (Guest Lecturer: Tim Herman, Toledo Zoo)
Invasive Species (Nathan Haislip, UTK)
Climate Change (Kevin Hamed, UTK)
Role of infectious disease in amphibian population decline and extinction (Guest Lecturer: Dr. Jim Collins, ASU)
Amphibian Conservation (Dr. Matt Gray, UTK)
Graduate Student Lectures: Test 2
2) Andrew West
Diversity and Evolution Lecture
1. Parental care in caecilians. Female caecilian feeding her young sloughed skin that is high in fat content. From Life in Cold Blood. BBC production. Hosted by David Attenborough. Link: http://www.bbc.co.uk/sn/tvradio/programmes/lifeincoldblood/video.shtml?licbtt03
2. Plethodontid salamander feeding. Hydromantes platycephalus is able to project its tongue more than half its body length to capture prey. From Dr. Stephen Debans website. Dr. Deban is at the University of South Florida and studies evolutionary biology and functional morphology (http://autodax.net/index.html). Link: http://autodax.net/hydromovie.html
3. Hellbender feeding. Cryptobranchus alleganiensis uses suction feeding to consume prey within its aquatic environment. From Dr. Stephen Debans website. Link: http://autodax.net/Cryptomovie.html
4. Japanese Giant Salamander Andrias japonicas introduction and conservation. From National Geographics Wild Chronicles. Link: http://www.youtube.com/watch?v=VN60DCHHQ50
5. Bolitoglossa schizodactyla walking. Bolitoglossa schizodactyla has unique adaptations for climbing. One of your assignments is to read about this in your book. From Amphibia Web website. Link: http://amphibiaweb.org/sounds/Bolitoglossa_schizodactyla.mov
6. Male poison arrow frogs wrestling over territories. From Life in Cold Blood. BBC production. Hosted by David Attenborough. Link: http://www.bbc.co.uk/sn/tvradio/programmes/lifeincoldblood/video.shtml?licbtt02
7. Male poison arrow frogs and their tadpoles. Males carry their tadpoles on their backs to bromeliads to complete their larval development. From Life in Cold Blood. BBC production. Hosted by David Attenborough. Link: http://www.youtube.com/watch?v=64y15Ho6d84
NOTE: Video 7 did not play during lecture. Please watch it.
1. Worldwide Amphibian Declines: How big is the problem, what are the causes and what can be done? Interview with Vance Vredenburg, Robert Drewes, Tyrone Hayes, and Karen Swaim. Video at: http://amphibiaweb.org/declines/declines.html.
1. Spotted Salamander Migration: http://www.youtube.com/watch?v=UkGEXrL8udU
2. Red-legged Salamander head slapping (1st video on the page): http://plethodon.science.oregonstate.edu/behavior.html#
3. Red-legged Salamander spermatophore deposition (2nd video on the page): http://plethodon.science.oregonstate.edu/behavior.html#
4. Alpine Newt Courtship: http://www.youtube.com/watch?v=aRvCbSHLJwE
5. Marbled Salamander Courtship and Natural History: http://www.youtube.com/watch?v=rk4h7QbSzLM
Anuran Courtship Lecture
1. Dr. Carl Gerhardt interview about anuran calls. From Frog Calls. Midwest Frogs: http://www.midwestfrogs.com/media/soundcommunication.wmv
2. Spring Peeper Courtship and Aggressive call. From Frog Calls. Midwest Frogs: http://www.midwestfrogs.com/media/peeperagressioncall2.wmv
3. Coqui frog calling: http://www.youtube.com/watch?v=54-FzuE-w0U
4. Spring Peepers Calling. From You Tube: http://www.youtube.com/v/ZX9uODHi0zg
5. American Toads Calling. From You Tube: http://www.youtube.com/watch?v=A6-8pC8o5fw
6. Gray Treefrog female preference for calling rates. From Frog Calls. Midwest Frogs: http://www.midwestfrogs.com/media/treefrogchoice.wmv
7. Frog-eating Bat. From You Tube: http://www.youtube.com/watch?v=Pa9xn0w9JBk
8. Frog-eating Bat capturing a Tungara Frog: From Dr. Mike Ryans Website: http://www.sbs.utexas.edu/ryan/Multimedia/bat-frog%20close3.wmv
9. Frog-eating Bat responding to calls of a Tungara Frog: From Dr. Mike Ryans Website: http://www.sbs.utexas.edu/ryan/Multimedia/toad%20then%20frog%201.WMV
10. Blood sucking flies responding to Tungara Frog Calls: From Dr. Mike Ryans Website: http://www.sbs.utexas.edu/ryan/Multimedia/720%20x%20480_30_cynepack%202clip%20fly%20bout.avi
11. Golden Frog waving to rival males. From Life in Cold Blood. BBC production. Hosted by David Attenborough: http://www.youtube.com/watch?v=A1FWQvaBoRg
12. A male Dendrobates tinctorius fertilizing eggs. From You Tube: http://www.youtube.com/watch?v=wHo2m8Ci04k
13. Wood Frog ovipositing. From Frog Calls. Midwest Frogs: http://www.midwestfrogs.com/media/woodfrogeggs.wmv
14. Red-eyed Treefrogs ovipositing in the tree canopy. From You Tube: http://www.youtube.com/watch?v=6dtRsrYkCeE (watch only the initial 60 seconds)
TWRA Amphibian Identification: http://www.state.tn.us/twra/tamp/frogs.html
LEAPS Anuran Identification: http://www.leaps.ms/Tn.%20Frogs%20ID%20Page.htm
Video Clips of Frog Calls: http://www.midwestfrogs.com/
TWRA Salamander Identification: http://www.state.tn.us/twra/tamp/salamanders.htm
Amphibians of the
Southeast Tadpole Identification: http://fl.biology.usgs.gov/armi/Guide_to_Tadpoles/guide_to_tadpoles.html
ARMI 5-year Report (Amphibian Declines): http://www.fort.usgs.gov/products/publications/21733/21733.pdf
Global Amphibian Assessment: http://www.globalamphibians.org
USGS Field Guide to Malformations of Frogs and Toads: http://www.nwhc.usgs.gov/publications/fact_sheets/pdfs/frog.pdf
PARC Habitat Management Guidelines
for Amphibians and Reptiles of the
Previous WFS 433/533 Websites
Spring 2008: http://fwf.ag.utk.edu/mgray/wfs493/493home2008.htm
Spring 2007: http://fwf.ag.utk.edu/mgray/wfs493/493home2007.htm