1. From a Frogs Perspective, is a Dragonfly a Dragonfly? Evolutionary Avenues for and Constraints on the Transmission of Frog Lung Flukes (Haematoloechus spp.). Matthew G. Bolek*, Scott D. Snyder, and John Janovy Jr. School of Biological Sciences, University of Nebraska- Lincoln

2. Archetypical and Paradigmatic Life of a Frog Lung Fluke

3. Real World Lives of Frog Lung Flukes 6 spp. 21 spp. 511 spp. 12 spp.

4. The Problem of Parasite Flow Through an Ecosystem Actually Looks Something Like This:

5. (1) Two species are specialists only infecting dragonflies. The Situation: 2 nd Intermediate Host Specificity

6. (1) Two species are specialists only infecting dragonflies. (2) One species is intermediate in its host specificity and can infect dragonflies and damselflies. The Situation: 2 nd Intermediate Host Specificity

7. (1) Two species are specialists only infecting dragonflies. (2) One species is intermediate in its host specificity and can infect dragonflies and damselflies. (3) Two other species are generalists infecting dragonflies, damselflies, and other aquatic arthropods. The Situation: 2 nd Intermediate Host Specificity

8. N. Leopard Frog P. Leopard Frog Bullfrog

9. Two Generalists X N. Leopard Frog P. Leopard Frog Bullfrog

10. Two Generalists One Specialist N. Leopard Frog P. Leopard Frog Bullfrog X X

11. Two Generalists One Specialist One Intermediate One Specialist N. Leopard Frog P. Leopard Frog Bullfrog X X

12. Two Generalists One Specialist One Intermediate Ecological Host Specificity One Specialist Ecological Host Specificity N. Leopard Frog P. Leopard Frog Bullfrog X X

13. What is the role of second intermediate dragonfly hosts and their parasite interactions in the transmission of frog lung flukes? ?

14. Nickol Pond, Cass Co., NE

15. Eastern Pondhawk teneral

16. Eastern Pondhawk post-teneral

17. P < 0.05

18. The Eastern Forktail

19. N = 80 N = 28 N = 46 P < 0.05

22. Eastern Forktail larva

23. P > 0.05

24. Eastern Pondhawk larva

25. Eastern Pondhawk castskin

27. P < 0.001 P > 0.05 Mean Abundance of Lung Flukes Recovered from the Body Cavity and Branchial Basket of 100 Larvae and Lab Metamorphosed Tenerals of the Eastern Pondhawk

28. Experimental Infections I experimentally infected dragonflies with parasites from lab reared snails infected with four species of lung fluke and examined these dragonflies for the location and survival of these parasites through metamorphosis.

29. Table I. Prevalence and location of four lung fluke parasites in experimentally exposed eastern pondhawk dragonflies PrevalenceHeadThoraxBranchial Basket Generalist 1 10/10Yes Generalist 2 6/10Yes H. varioplexus 5/9 __ Yes H. longiplexus 6/9 __ Yes

30. Mean Abundance of Four species of Frog Lung Flukes in Experimentally Infected Eastern Pondhawk Larvae and Tenerals Mean Abundance + 1SD Life Stage P < 0.05

31. Natural History of Frog Hosts Aquatic Amphibians Semi-terrestrial Amphibians

32. Haematoloechus varioplexus N = 4 N = 4 N = 3 N = 3 N. Leopard Bullfrog P. Leopard Bullfrog Frog Frog N. Leopard Bullfrog P. Leopard Bullfrog Frog Frog

33. X Haematoloechus varioplexus X N = 4 N = 4 N = 3 N = 3 Yes N. Leopard Bullfrog P. Leopard Bullfrog Frog Frog N. Leopard Bullfrog P. Leopard Bullfrog Frog Frog

34. Morphological Studies of the specialist Haematoloechus varioplexus, or, is it really H. varioplexus? Haematoloechus varioplexus, a parasite of northern leopard, was synonymized with five other species of Haematoloechus including H. parviplexus which infects bullfrogs (Kennedy, 1981). The question arises as to whether Kennedy was justified, or correct, in making this synonymy.

35. Molecular Studies

37. Implications In terms of transmission, depending on the frog species, its habitat, and the trematode species involved, a dragonfly is not a dragonfly.

38. H. coloradensis (NA) H. complexus (NA) H. varioplexus (NA) H. abbreviatus (E) H. variegatus (E) H. floedae (NA) H. breviplexus (NA) H. parviplexus (NA) H. medioplexus (NA) H. longiplexus (NA) H. asper (E) H. similis (E) Plagiorchiidae OutGroup North America Europe 3 2 1 Bolek, Snyder, and Janovy In Prep

39. Life cycle unknown H. coloradensis (NA) H. complexus (NA) H. varioplexus (NA) H. abbreviatus (E) H. variegatus (E) H. floedae (NA) H. breviplexus (NA) H. parviplexus (NA) H. medioplexus (NA) H. longiplexus (NA) H. asper (E) H. similis (E) Plagiorchiidae Various Arthropods, Gastropods, Amphibians Damselflies North America Europe OutGroup

40. Damselflies Life cycle unknown H. coloradensis (NA) H. complexus (NA) H. varioplexus (NA) H. abbreviatus (E) H. variegatus (E) H. floedae (NA) H. breviplexus (NA) H. parviplexus (NA) H. medioplexus (NA) H. longiplexus (NA) H. asper (E) H. similis (E) Plagiorchiidae Various Arthropods, Gastropods, Amphibians Damselflies and Dragonflies North America Europe OutGroup

41. Damselflies Life cycle unknown H. coloradensis (NA) H. complexus (NA) H. varioplexus (NA) H. abbreviatus (E) H. variegatus (E) H. floedae (NA) H. breviplexus (NA) H. parviplexus (NA) H. medioplexus (NA) H. longiplexus (NA) H. asper (E) H. similis (E) Plagiorchiidae Various Arthropods, Gastropods, Amphibians Specialist North America Europe OutGroup Damselflies and Dragonflies

42. Damselflies Life cycle unknown H. coloradensis (NA) H. complexus (NA) H. varioplexus (NA) H. abbreviatus (E) H. variegatus (E) H. floedae (NA) H. breviplexus (NA) H. parviplexus (NA) H. medioplexus (NA) H. longiplexus (NA) H. asper (E) H. similis (E) Plagiorchiidae Various Arthropods, Gastropods, Amphibians Specialist Generalists North America Europe OutGroup Damselflies and Dragonflies

43. What About the Amphibian Host?

44. Damselflies Life cycle unknown H. coloradensis (NA) H. complexus (NA) H. varioplexus (NA) H. abbreviatus (E) H. variegatus (E) H. floedae (NA) H. breviplexus (NA) H. parviplexus (NA) H. medioplexus (NA) H. longiplexus (NA) H. asper (E) H. similis (E) Plagiorchiidae Various Arthropods, Gastropods, Amphibians Dragonfly Generalists Damselflies and Dragonflies

45. Damselflies Small to Medium Frogs Life cycle unknown H. coloradensis (NA) H. complexus (NA) H. varioplexus (NA) H. abbreviatus (E) H. variegatus (E) H. floedae (NA) H. breviplexus (NA) H. parviplexus (NA) H. medioplexus (NA) H. longiplexus (NA) H. asper (E) H. similis (E) Plagiorchiidae Various Arthropods, Gastropods, Amphibians Dragonfly Generalists Damselflies and Dragonflies

46. Small to Medium Frogs Life cycle unknown H. coloradensis (NA) H. complexus (NA) H. varioplexus (NA) H. abbreviatus (E) H. variegatus (E) H. floedae (NA) H. breviplexus (NA) H. parviplexus (NA) H. medioplexus (NA) H. longiplexus (NA) H. asper (E) H. similis (E) Plagiorchiidae Various Arthropods, Gastropods, Amphibians Dragonfly Damselflies Generalists Large Frogs Damselflies and Dragonflies

47. Damselflies Small to Medium Frogs Life cycle unknown H. coloradensis (NA) H. complexus (NA) H. varioplexus (NA) H. abbreviatus (E) H. variegatus (E) H. floedae (NA) H. breviplexus (NA) H. parviplexus (NA) H. medioplexus (NA) H. longiplexus (NA) H. asper (E) H. similis (E) Plagiorchiidae Various Arthropods, Gastropods, Amphibians Dragonfly Generalists Medium Frogs Large Frogs Damselflies and Dragonflies

48. Damselflies Small to Medium Frogs Life cycle unknown H. coloradensis (NA) H. complexus (NA) H. varioplexus (NA) H. abbreviatus (E) H. variegatus (E) H. floedae (NA) H. breviplexus (NA) H. parviplexus (NA) H. medioplexus (NA) H. longiplexus (NA) H. asper (E) H. similis (E) Plagiorchiidae Various Arthropods, Gastropods, Amphibians Dragonfly Generalists Small to Medium Frogs Medium Frogs Large Frogs Damselflies and Dragonflies

49. Adult North American Bullfrog and Northern Leopard Frog M. Bolek

51. Bolek and Janovy 2007

53. Mean Prey Size and Gape Size of 5 North American Frog Species Bullfrog Bolek unpublished.

54. M. Bolek

55. Diet of Bullfrogs Amphibians and Reptiles 13.9% Fish 3.1% Crustaceans 31.6% Misc. Inverts. 3.5% Mammals 15.3% Other 6.2% Insects 26.4% Data from various sources.

56. Conclusion Our data on closely related trematode species clearly suggest that their complex life cycles are dynamic and there are numerous avenues for and constraints on transmission to their definitive hosts.

57. Acknowledgments Dr. John Janovy Jr. Dr. Scott Snyder (UNO) Randy Peterson Dr. Brent Nickol Dr. Virginia Leon, and Rosario Mata (UNAM) Dr. Agustin Jimenez Cedar Point Biological Station UNL School of Biological Sciences Special Funds, Initiative for Ecological and Evolutionary Analysis UNL, Center for Great Plains Studies Jaclyn Helt, Jill Detwiler, Samana Schwank

58. Questions?