1. Comparative phylogeography and molecular dating of various Western North American fish Nina J. Laitinen, Dennis K. Shiozawa, Peter J. Unmack, Derek D. Houston BRIGHAM YOUNG UNIVERSITY

2. Upper Missouri River Basin Columbia River Basin Lower Colorado River Basin Upper Colorado River Basin Lahontan Basin Eastern Snake River Plain Bonneville Basin

3. C. plebeius Catostomus (Pantosteus)

4. Pantosteus Gerald Smith (taxonomy and distribution) Currently 6 or 7 species Is Pantosteus a full genus or subgenus? Interesting biogeographically A lot of work done within several species but little done describing family relationships – Several undescribed forms are hypothesized to exist Introgression a potential issue

5. Distinguishing Characteristics of Pantosteus: Development of cartilaginous scraping edges on the jaws Lateral notches at the junctions of the upper and lower lips A shallow, posterior median incision of the lower lip A long intestine A black peritoneum

7. Cutthroat trout (Oncorhynchus clarkii) Combined, 3649 bp, MP 1000 reps

8. Cutthroat trout range

9. C. rhotheus C. bairdii (Columbia, Coastal WA, C. extensus (Bear), Bonneville, Colorado R. C. bairdii (Montana) C. bairdii kumlieni (WI) C. cognatus (WI) C. cognatus (Columbia) C. confusus C. leiopomus (Columbia), C. beldingii (Lahonton, Colorado, Bonneville) Outgroup species: C. asper, C. klamathensis Cottus (ATPase)

10. Cottus beldingii Cottus bairdii

11. Objective What does calibrating molecular clocks of several phylogenetic trees with fossils reveal about divergence dates of major geologic events?

12. Methods Sequenced various mitchondrial genes (ATPase, ND4L, ND4, ND5, ND6, CytB) for the species involved using standard PCR methods. Constructed maximum likelihood trees on each data set. Calibrated molecular clocks in BEAST using fossil data recorded in the literature.

13. Geologic background The field of phylogeography is concerned with the principles and processes governing the geographic distributions of genealogical lineages. In order to account for different lineages and interpret the dispersal history of these fish, we must first understand the geological history of Western North America

14. Geologic Time Scale

15. In the 1930’s Carl L. Hubbs from the University of Michigan along with student Robert R. Miller collected fish from western North America Their 1948 publication on the Great Basin established the foundation for interpreting biogeography of western fishes.

16. Fossil Studies of Western Fishes Most modern interior western fishes are of relatively recent origin—however, many are older than the Pleistocene. The oldest relevant fossils date to the early to middle Miocene (10-20 Ma)

17. Lake Clarkia, ca 12-15 mya Gila L. Idaho Glenns Ferry 3.7-2.4 mya Cottus, Gila, Ptychocheilus L. Idaho Chalk Hills 9-6 mya Gila, Ptychocheilus Rhabdofario Lake Bidahochi Formation 9-6 mya Ptychocheilus, Gila sp, Gila robusta Owens Lake core 700,000 ybp Trout, Prosopium, Gila Mopung Hills 2-3 mya O. clarkii, Gila Honey Lake 2-3 mya Rhabdofario Gila Deer Butte Formation 11 mya? Ptychocheilus Rhabdofario Blackjack Butte 3 mya Ptychocheilus Prosopium Grand Rhonde, ca 3.7-3.8 mya Ptychocheilus, Prosopium Ringold Formation - 15 mya Ptychocheilus; 5 mya Oncorhynchus; 4.5-3.7 mya Catostomus, Chasmistes, Ptychocheilus Duck Point, ca 9-0.2 mya Ptychocheilus, Gila Salt Lake group 9-6 mya Ptychocheilus

18. Major transfer events influencing dispersal of fishes 1) The connection between the upper Snake River, the Humboldt River, and the Pit-Sacramento River from 10 Ma to 2.5 Ma (Humboldt River formed 9 Ma) 2) The capture of the Upper Colorado River by the Lower Colorado River about 4.6-4.4 Ma—prior path of Upper Colorado River unknown 3) The Paleo-Wood River was captured into the snake River Basin 4 Ma from the Missouri River Basin 4) The transfer of the upper and middle Snake River to the Columbia River Basin 2.5 Ma 5) The Bear river capture from the Snake River to the Bonneville basin about 0.05 Ma and Bonneville Flood (about 0.015 Ma)

19. Sierra batholith rises— initiates dessication of the Great Basin 1) The connection between the upper Snake River, the Humboldt River, and the Pit-Sacramento River from 10 Ma to 2.5 Ma (Humboldt River formed 9 Ma) 1 10 mya (late Miocene)

20. 4 MYA 1 2) The capture of the Upper Colorado River by the Lower Colorado River about 4.6-4.4 Ma—prior path of Upper Colorado River unknown 3) The Paleo-Wood River was captured into the snake River Basin 4 Ma from the Missouri River Basin 3 2

21. 2-3 MYA 1 3 2 44) The transfer of the upper and middle Snake River to the Columbia River Basin 2.5 Ma

22. 2 MYA 1 3 2 4

23. 0.1 MYA 1 3 2 4 5 5) The Bear river capture from the Snake River to the Bonneville basin about 0.05 Ma and Bonneville Flood (about 0.015 Ma)

24. Catostomus

27. 4 MYA

28. 2-3 MYA 44) The transfer of the upper and middle Snake River to the Columbia River Basin 2.5 Ma

29. Catostomus

30. 0.1 MYA 1 3 2 4 5 5) The Bear river capture from the Snake River to the Bonneville basin about 0.05 Ma and Bonneville Flood (about 0.015 Ma)

31. Many species of fish follow a North-South division in the Bonneville Basin.These fish include the following: Leatherside chub Utah Chub Cutthroat Trout Speckled Dace Paiute sculpin Utah sucker

32. Cutthroat trout (ND4L, ND4, CB, ATPase, CR)

33. Catostomus

34. C. rhotheus C. bairdii (Columbia, Coastal WA, C. extensus (Bear), Bonneville, Colorado R. C. bairdii (Montana) C. bairdii kumlieni (WI) C. cognatus (WI) C. cognatus (Columbia) C. confusus C. leiopomus (Columbia), C. beldingii (Lahonton, Colorado, Bonneville) Outgroup species: C. asper, C. klamathensis Cottus (ATPase)

35. Additional data: Fossils of C. bairdii are known from Pleistocene fluvial sediments in the Glenns Ferry Formation, Idaho The absence of C. bairdii in the Lahontan Basin and Wood/Lost River systems indicate that it entered the Snake River Basin after the Late Pliocene capture of Lake Idaho. C. beldingii was already present in the Snake River Basin, but also invaded the Bonneville Basin with the C. bairdii transfers

36. 4 MYA 2) The capture of the Upper Colorado River by the Lower Colorado River about 4.6-4.4 Ma—prior path of Upper Colorado River unknown 2

37. Catostomus

38. Of five major, dated, transfer events that may have influenced modern fish community structure: 1 – 10 Ma to 2.5 Ma upper Snake River, the Humboldt River, and the Pit- Sacramento River pikeminnow, hardhead chub, tui chub, Paiute sculpin 2 –4.6-4.4 Ma Upper Colorado River capture by the Lower Colorado pikeminnow, Utah chub-roundtail chub, speckled dace, bluehead sucker, desert sucker 3 – 4 Ma Wood River capture into Snake River Basin mountain whitefish, mountain sucker, Paiute sculpin 4 – 2.5 Ma capture upper and middle Snake River to the Columbia River Basin redside shiner, least chub, mountain whitefish, mountain sucker, Paiute sculpin 5 – The Bear River capture from the Snake River to the Bonneville Basin mountain sucker, bluehead sucker, redside shiner, speckled dace, mottled sculpin, Paiute sculpin, mountain whitefish, cutthroat trout

39. Conclusions Pantosteus is more diverse than previously suggested. Our current taxonomy is limiting our understanding and conservation of the group Further comparison may reveal more concordance among various fish species and geologic events.

40. Acknowledgements: Collections: Aaron Webber Rob Gipson Pete Cavelli Matt Maples Heather Stutz Keith Tanner Kelli Boulter Mark Belk John Aedo Nicole LeDuc Rachel Barney DNA work: R. Paul Evans Michael Evans Katherine Yokoyama Nicholes Candelaria Andrew Johnson Sun-yeong Oh Kenny Eyring Derek Houston

41. Questions?