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Issue: Vol. 5249 No. 5: 6 Mar. 2023
Type: Article
Published: 2023-03-06
DOI: 10.11646/zootaxa.5249.5.1
Page range: 501-539
Abstract views: 1380
PDF downloaded: 581

Taxonomy of the Speckled Dace Species Complex (Cypriniformes: Leuciscidae, Rhinichthys) in California, USA

Center for Watershed Sciences & Department of Wildlife, Fish and Conservation Biology, University of California, 425 LaRue Road, Davis CA 95626 USA .
California Department of Fish & Wildlife,787 Main Street, Bishop, CA 93514 USA.
Department of Animal Science, University of California, Davis, 1 Shields Avenue, Davis CA 95616 USA.
cryptic species genomics Great Basin fishes endangered species desert fishes Pisces

Abstract

The Speckled Dace, Rhinichthys osculus (Girard), is a small species of fish (Cypriniformes, Leuciscidae) that has the widest geographic range of any freshwater dispersing fish in western North America. The dynamic geologic history of the region has produced many isolated watersheds with endemic fish species. However, Speckled Dace from these watersheds cannot be differentiated readily by morphometrics and meristics. This has led to the widely accepted hypothesis that the dace’s adaptability and ability to cross geologic barriers has resulted in interbreeding among neighboring populations, maintaining the dace as a single species. We investigate this hypothesis by looking at Speckled Dace populations in California which are the result of at least three separate colonization events of isolated watersheds. We synthesize results from taxonomic, genetic, and zoogeographic studies in combination with the findings of a recent genomics study, to show that there are distinctive evolutionary lineages within the Speckled Dace complex. These lineages are used to designate multiple species and subspecies. We back up these designations by examining how well these lineages fit with the geologic history of the isolated basins they inhabit and with the presence of other endemic fishes. We conclude the following nine taxa can be recognized within the Speckled Dace complex in California.

References

  1. Ardren, W.R., Baumsteiger, J. & Allen, C.S. (2010) Genetic analysis and uncertain taxonomic status of threatened Foskett Spring Speckled Dace. Conservation Genetics, 11 (4), 1299–1315. https://doi.org/10.1007/s10592-009-9959-0

  2. Armbruster, J.W. (2012) Standardized measurements, landmarks, and meristic counts for cypriniform fishes. Zootaxa, 3586 (1), 8–16. https://doi.org/10.11646/zootaxa.3586.1.3

  3. Bangs, M.R., Douglas, M.R. Brunner, P.C. & Douglas, M.E. (2020) Reticulate evolution as a management challenge: patterns of admixture with phylogenetic distance in endemic fishes of western North America. Evolutionary Applications, 13 (6), 1400–1419. https://doi.org/10.1111/eva.13042

  4. Baumsteiger, J. & Moyle, P.B. (2019) A reappraisal of the California Roach/Hitch (Cypriniformes, Cyprinidae, Hesperoleucus/Lavinia) species complex. Zootaxa, 4543 (2), 2221–2240. https://doi.org/10.11646/zootaxa.4543.2.3

  5. Baumsteiger, J., Moyle, P.B., Aguilar, A., O’Rourke, S.M. & Miller, M.R. (2017) Genomics clarifies taxonomic boundaries in a difficult species complex. PLoS ONE, 12 (12), e0189417. https://doi.org/10.1371/journal.pone.0189417

  6. Bickford, D., Lohman, D.J., Sodhi, N.S., Ng, P.K., Meier, R., Winker, K., Ingram, K.K. & Das, I. (2007) Cryptic species as a window on diversity and conservation. Trends in Ecology & Evolution, 22 (3), 148–155. https://doi.org/10.1016/j.tree.2006.11.004

  7. Billman, E.J., Lee, J.B., Young, D.O., McKell, M.D., Evans, R.P. & Shiozawa, D.K. (2010) Phylogenetic divergence in a desert fish: differentiation of Speckled Dace within the Bonneville, Lahontan, and upper Snake River basins. Western North American Naturalist, 70 (1), 39–48. https://doi.org/10.3398/064.070.0105

  8. Center for Biological Diversity (2020) Petition to List Three Populations of Speckled Dace (Rhinichthys osculus) in the Death Valley Region under the Endangered Species Act: Amargosa Canyon Speckled Dace, Long Valley Speckled Dace, Owens Speckled Dace. Submitted to US Fish and Wildlife Service, 8 June 2020, 1–48.

  9. Chen, Y., Parmenter, S. & May, B. (2007) Introgression between Lahontan and endangered Owens Tui Chubs, and apparent discovery of a new Tui Chub in the Owens Valley, California. Conservation Genetics, 8 (1), 221–238. https://doi.org/10.1007/s10592-006-9164-3

  10. Colman, S.M., Platt Bradbury, J. & Rosenbaum, J.G. (2004) Paleolimnology and paleoclimate studies in Upper Klamath Lake, Oregon. Journal of Paleolimnology, 31 (2), 129–138. https://doi.org/10.1023/B:JOPL.0000019235.72107.92

  11. Cope, E.D. (1872) Report on the recent reptiles and fishes of the survey, collected by Campbell Carrington and C. M. Dawes. In: Part IV: Zoology and Botany of Preliminary Report of the U. S. Geological Survey of Montana and Adjacent Territories. Annual Report of Progress, 5, pp. 467–476.

  12. Cope, E.D. (1883) On the fishes of the recent and Pliocene lakes of the western part of the Great Basin and of Idaho Pliocene Lake. Academy of Natural Sciences, Philadelphia, Proceedings, 35, 134–166.

  13. Cope, E.D. & Yarrow, H.C. (1875) Report upon the collection of fishes made in portions of Nevada, Utah, California, New Mexico, and Arizona during the years 1871, 1872, 1873, and 1874. Report on Geographical and Geological Explorations and Surveys, W100th Meridian (Wheeler Survey), Chapter VI, 637–703.

  14. Cornelius, R.H. (1969) The Systematics and Zoogeography of Rhinichthys osculus (Girard) in Southern California. M.A. Thesis, California State University, Fullerton, California, 195 pp.

  15. Culver, G.B. & Hubbs, C.L. (1917) The fishes of the Santa Ana system of streams in southern California, Lorquinia, 1 (11), 82–83.

  16. Daniels, R.A. & Moyle, P.B. (1984) Geographic variation and a taxonomic reappraisal of the Marbled Sculpin, Cottus klamathensis. Copeia, 1984, 949–959. https://doi.org/10.2307/1445339

  17. Deacon, J.E. & Williams, J.E. (1984) Annotated list of the fishes of Nevada. Proceedings of the Biological Society of Washington, 97 (1), 113–118.

  18. Dixon, R.B. (1908) Notes on the Achomawi and Atsugewii Indians of northern California. American Anthropologist, 10 (2), 208–220. https://doi.org/10.1525/aa.1908.10.2.02a00030

  19. Dorsey, R.J. & Roering, J.J. (2006) Quaternary landscape evolution in the San Jacinto fault zone, Peninsular ranges of Southern California: transient response to strike-slip fault initiation. Geomorphology, 73 (1–2), 16–32. https://doi.org/10.1016/j.geomorph.2005.06.013

  20. Egger, A.E. & Miller, E.L. (2011) Evolution of the northwestern margin of the Basin and Range: The geology and extensional history of the Warner Range and environs, northeastern California. Geosphere, 7 (3), 756–773. https://doi.org/10.1130/GES00620.1

  21. Evermann, B.W. & Meek, S.B. (1898) A report upon salmon investigations in the Columbia River and elsewhere on the Pacific Coast in 1896. Bulletin of the U.S. Fish Commission, 17, 15–84. https://doi.org/10.5962/bhl.title.35601

  22. Evermann, B.W. & Clark, H.W. (1931) A distributional list of freshwater fishes known to occur in California. Division of Fish and Game, Fish Bulletin, 35, 1–67.

  23. Fattaruso, L.A., Cooke, M.L., Dorsey, R.J. & Housen, B.A. (2016) Response of deformation patterns to reorganization of the southern San Andreas fault system since ca. 1.5 Ma. Tectonophysics, 693, 474–488. https://doi.org/10.1016/j.tecto.2016.05.035

  24. Freudenstein, J.C., Broe, M.B., Folk, R.A. & Sinn, B.T. (2017) Biodiversity and the species concept -lineages are not enough. Systematic Zoology, 66, 644–656. https://doi.org/10.1093/sysbio/syw098

  25. Garcia & Associates (2000) The Distribution and Abundance of the Benthic Macroinvertebrate Fauna and Fish Populations in Tributaries Leading into and including the North Fork Mokelumne and Mainstem Mokelumne Rivers. Final Report Volume 3. Prepared for Pacific Gas and Electric Company by Garcia and Associates, San Anselmo, California. [unknown pagination]

  26. Gesch, D., Oimoen, M., Greenlee, S., Nelson, C. Steuck, M. & Tyler, D. (2002) The National Elevation Dataset. Photogrammetric Engineering and Remote Sensing, 68 (1), 5–11.

  27. Gilbert, C.H. (1893) Report on the fishes of the Death Valley expedition collected in southern California and Nevada in 1891 with descriptions of new species. U.S. Department of Agriculture, North American Fauna, 7, 229–234. https://doi.org/10.3996/nafa.7.0003

  28. Gilbert, C.H. & Evermann, B.W. (1894) A report upon investigations in the Columbia River basin, with descriptions of four new species of fishes. Bulletin of the U. S. Fish Commission, 14, 169–207.

  29. Girard, C.F. (1856) Researches upon the cyprinoid fishes inhabiting the fresh waters of the United States, west of the Mississippi Valley, from specimens in the museum of the Smithsonian Institution. Proceedings of the Academy of Natural Sciences of Philadelphia, 8, 165–213.

  30. Girard, C.F. (1858) Report upon fishes collected on the survey. In: U.S. War Department, Reports of Explorations and Surveys, to Ascertain the most Practicable and Economical Route for a Railroad from the Mississippi River to the Pacific Ocean 10. U.S. Government Printing Office, Washington, D.C., pp. 83–91.

  31. Grantham, T.E., Fesenmyer, K.A., Peek, R., Holmes, E., Quiñones, R.M., Bell, A., Santos, N., Howard, J.K., Viers, J.H. & Moyle, P.B. (2017) Missing the boat on freshwater fish conservation in California. Conservation Letters, 10, 1077–1085. https://doi.org/10.1111/conl.12249

  32. Grayson, D.K. (1993) The Desert’s Past: A Natural Prehistory of the Great Basin. Smithsonian Institution Press, Washington, xx + 356 pp.

  33. Gudde, E.G. & Bright, W. (1998) California Place Names. University of California Press, Berkeley, California, xxviii + 467 pp. https://doi.org/10.1525/9780520920545

  34. Harrison, S.P. (2013) Plant and Animal Endemism in California. University of California Press, Berkeley, California, ix + 189 pp. https://doi.org/10.1525/california/9780520275546.001.0001

  35. Hildreth, W. & Fierstein, J. (2016) Long Valley Caldera Lake and Reincision of Owens River Gorge. U.S. Geological Survey Scientific Investigations Report, 5120, 1–63. https://doi.org/10.3133/sir20165120

  36. Hoekzema, K. & Sidlauskas, B.L. (2014) Molecular phylogenetics and microsatellite analysis reveal cryptic species of Speckled Dace (Cyprinidae: Rhinichthys osculus) in Oregon’s Great Basin. Molecular Phylogenetics and Evolution, 77, 238–250. https://doi.org/10.1016/j.ympev.2014.04.027

  37. Hubbs, C.L., Follett, W.I. & Dempster, L.J. (1979) List of the fishes of California. Occasional Papers of the California Academy of Sciences, 133, 1–54.

  38. Hubbs, C.L. & Kuhne, E.R. (1937) A new fish of the genus Apocope from a Wyoming warm spring. Occasional Papers of the Museum of Zoology, University of Michigan, 343, 1–21.

  39. Hubbs, C.L. & Miller, R.R. (1948a) The Great Basin, with emphasis on glacial and postglacial time. The zoological evidence: correlation between fish distribution and hydrographic history in the desert basins of western United States. University of Utah Bulletin, 38 (20), 17–166.

  40. Hubbs, C.L. & Miller, R.R. (1948b) Two new relict genera of fishes from Nevada. Occasional Papers of the Museum of Zoology, University of Michigan, 507, 1–31.

  41. Hubbs, C.L. & Miller, R.R. (1972) Diagnoses of new cyprinid fishes of isolated waters in the Great Basin of western North America. Transactions, San Diego Natural History Society 17, 101–106.

  42. Hubbs, C.L., Miller, R.R. & Hubbs, L.C. (1974) Hydrographic history and relict fishes of the North-central Great Basin. San Francisco: California Academy of Sciences Memoir, 7, 1–259.

  43. Jelks, H.L., Walsh, S.J., Burkhead, N.M., Contreras-Balderas, S., Diaz-Pardo, E., Hendrickson, D.A., Lyons, J., Mandrak, N.E., McCormick, F., Nelson, J.S. & Platania, S.P. (2008) Conservation status of imperiled North American freshwater and diadromous fishes. Fisheries, 33 (8), 372–407. https://doi.org/10.1577/1548-8446-33.8.372

  44. Jordan, D.S. (1885) Identification of the species of Cyprinidae and Catostomidae, described by Dr. Charles Girard in Proceedings Academy of Natural Sciences of Philadelphia for 1856. Proceedings, US National Museum, 8, 118–127. https://doi.org/10.5479/si.00963801.8-500.118

  45. Jordan, D.S. & Evermann, B.W. (1896) The Fishes of North and Middle America. Bulletin, US National Museum, 47, Part 1, 1–1240.

  46. Kimsey, J.B. & Fisk, L.O. (1960) Keys to the freshwater and anadromous fishes of California. California Fish and Game, 46, 453–479.

  47. Kinziger, A.P., Nakamoto, R.J., Anderson, E.C. & Harvey, B.C. (2011) Small founding number and low genetic diversity in an introduced species exhibiting limited invasion success (Speckled Dace, Rhinichthys osculus). Ecology and Evolution, 1 (1), 73–84. https://doi.org/10.1002/ece3.8

  48. La Rivers, I. & Trelease, T.J. (1952) An annotated checklist of the fishes of Nevada. California Fish and Game, 38, 113–123.

  49. Leidy, R.A. (2007) Ecology, Assemblage Structure, Distribution, and Status of Fishes in Streams Tributary to the San Francisco Estuary. San Francisco Estuary Institute Contribution, 530 pp.

  50. Leidy, R.A. & Moyle, P.B. (2021) Keeping up with the status of freshwater fishes: a California (USA) perspective. Conservation Science and Practice, 3 (8), e474. https://doi.org/10.1111/csp2.474

  51. Markle, D.F. (2016) A Guide to the Freshwater Fishes of Oregon. Oregon State University Press, Corvallis, Oregon, 140 pp.

  52. McClay, K. & Bonora, M. (2001) Analog models of restraining stepovers in strike-slip fault systems. American Association of Petroleum Geologists Bulletin, 85, 233–260. https://doi.org/10.1306/8626C7AD-173B-11D7-8645000102C1865D

  53. Miller, R.R. (1973) Two new fishes, Gila bicolor snyderi and Catostomus fumeiventris from the Owens River Basin California. Occasional Papers of the Museum of Zoology, University of Michigan, 667, 1–19.

  54. Miller, R.R. (1984) Rhinichthys deaconi, a new species of dace (Pisces: Cyprinidae) from southern Nevada. Occasional Papers of the Museum of Zoology, University of Michigan, 707, 1–21.

  55. Minckley, W.L., Hendrickson, D.A. & Bond, C.E. (1986) Geography of western North America freshwater fishes: Description and relationship to intercontinental tectonism. In: Hocutt, C.H. & Wiley, E.O. (Eds.), The Zoogeography of North American Freshwater Fishes. New York, John Wiley and Sons, pp. 519–614.

  56. Minckley, W.L. & Marsh, P.C. (2009) Inland Fishes of the Greater Southwest: Chronicle of a Vanishing Biota. University of Arizona Press, Tucson, Arizona, 576 pp.

  57. Moyle, P.B. (1976) Inland Fishes of California. University of California Press, Berkeley, California, 405 pp.

  58. Moyle, P.B. (2002) Inland Fishes of California, Revised and Expanded. University of California Press, Berkeley, California, 504 pp.

  59. Moyle, P.B., Katz, J.V.E. & Quiñones, R.M. (2011) Rapid decline of California’s native inland fishes: a status assessment. Biological Conservation, 144, 2414–2423. https://doi.org/10.1016/j.biocon.2011.06.002

  60. Moyle, P.B. & Campbell, M.A. (2022) Cryptic species of freshwater sculpin (Cottidae: Cottus) in California. Zootaxa, 5154 (5), 501–507. https://doi.org/10.11646/zootaxa.5154.5.1

  61. Moyle, P.B. & Cech, J.J. (2004) Fishes: An Introduction to Ichthyology. 5th Edition. Prentice-Hall, Upper Saddle River, New Jersey, xvi + 726 pp.

  62. Moyle, P.B. & Davis, L.H. (2000) A list of freshwater, anadromous, and euryhaline fishes of California. California Fish and Game, 86, 244–258.

  63. Moyle, P.B., Quiñones, R.M., Katz, J.V.E, & Weaver, J. (2015) Fish Species of Special Concern in California. 3rd Edition. Sacramento, California Department of Fish and Wildlife. Available from: https://wildlife.ca.gov/Conservation/SSC/Fishes (accessed 8 February 2023)

  64. Murphy, G.L. (1941) A key to the fishes of the Sacramento-San Joaquin Basin. California Fish and Game, 27, 165–171.

  65. Mussmann, S.M. (2018) Diversification Across a Dynamic Landscape: Phylogeography and Riverscape Genetics of Speckled Dace (Rhinichthys osculus) in Western North America Ph.D. Dissertation, University of Arkansas, Fayetteville, Arkansas, 166 pp.

  66. Mussmann, S.M., Douglas, M.R., Oakey, D.D. & Douglas, M.E. (2020) Defining relictual biodiversity: conservation units in Speckled Dace (Leuciscidae: Rhinichthys osculus) of the Greater Death Valley Ecosystem. Ecology and Evolution, 10, 10798–10817. https://doi.org/10.1002/ece3.6736

  67. Oakey, D.D., Douglas, M.E. & Douglas, M.R. (2004) Small fish in a large landscape: diversification of Rhinichthys osculus (Cyprinidae) in western North America, Copeia, 2004 (2), 207–221. https://doi.org/10.1643/CG-02-264R1

  68. O’Brien, J.W., Hansen, H.K. & Stephens, M.E. (2011) Status of fishes in the Upper San Gabriel River Basin, Los Angeles County, California. California Fish and Game, 97, 149–163.

  69. Orme, A.R. (2008) Pleistocene pluvial lakes of the American West: a short history of research. Geological Society, London, Special Publications, 301 (1), 51–78. https://doi.org/10.1144/SP301.4

  70. Page, L.M. & Burr, B.M. (2011) Peterson Guide to Freshwater Fishes of North American North of Mexico. 2nd Edition. Houghten Mifflin Harcourt, Boston, Massachusetts, 663 pp.

  71. Page, L.M., Espinoza-Perez, H., Findley, L.T., Gilbert, C.R., Lea, R.N, Mandrake, N.E., Mayden, R.L. & Nelson, J.S. (2013) Common and Scientific Names of Fishes from the United States, Canada, and Mexico. 7th Edition. American Fisheries Society Special Publication 34. American Fisheries Society, Bethesda, Maryland, 243 pp.

  72. Patterson, T. &. Kelso, N.V. (2021) Natural Earth Data. https://www.naturalearthdata.com/ (accessed 30 December 2022)

  73. Peden, A.E. & Hughes, G.W. (1988) Sympatry in four species of Rhinichthys (Pisces), including the first documented occurrences of R. umatilla in the Canadian drainages of the Columbia River. Canadian Journal of Zoology, 66 (8), 1846–1856. https://doi.org/10.1139/z88-267

  74. Pfrender, M.E., Hicks, J. & Lynch, M. (2004) Biogeographic patterns and current distribution of molecular-genetic variation among populations of Speckled Dace, Rhinichthys osculus (Girard). Molecular Phylogenetics and Evolution, 30 (3), 490–502. https://doi.org/10.1016/S1055-7903(03)00242-2

  75. Reheis, M.C., Stine, S. & Sarna-Wojcicki, A.M. (2002) Drainage reversals in Mono Basin during the late Pliocene and Pleistocene. Geological Society of America Bulletin, 114 (8), 991–1006. https://doi.org/10.1130/0016-7606(2002)114%3C0991:DRIMBD%3E2.0.CO;2

  76. Rutter, C. (1903) Notes on fishes from streams and lakes of northeastern California not tributary to the Sacramento basin. United States Fish Commission Bulletin, 22, 145–148. [1902]

  77. Rutter, C. (1908) The fishes of the Sacramento-San Joaquin basin, with a study of their distribution and variation. United States Bureau of Fisheries Bulletin, 27, 105–152. [1907]

  78. Sada, D.W. (1989) Status, distribution, and morphological variation of Speckled Dace in the Owens River system. Final Report. California Department of Fish and Game, Sacramento, California, 37 pp.

  79. Sada, D.W., Britten, H.B. & Brussard, P.F. (1993) Morphometric and genetic differentiation among Death Valley System Rhinichthys osculus. CDFG Inland Fisheries Report, Contract, FG0524, 1–52.

  80. Sada, D.W., Britten, H.B. & Brussard, P.F. (1995) Desert aquatic ecosystems and the genetic and morphological diversity of Death Valley system Speckled Dace. In: Nielsen, J.L. (Ed.), Evolution and the Aquatic System. American Fisheries Society Symposium, 17, pp. 350–359.

  81. Sada, D.W. & Vinyard, G.L. (2002) Anthropogenic changes in biogeography of Great Basin biota. In: Hershler, R., Madsen, D.B. & Curry, D.R. (Eds.), Great Basin Aquatic Systems History. Smithsonian Contributions to the Earth Sciences, 33, pp. 277–293.

  82. Santos, N.R., Katz, J.V.E., Moyle, P.B. & Viers, J.H. (2013) A programmable information system for management and analysis of aquatic species range data in California. Environmental Modeling & Software, 53, 13–26. https://doi.org/10.1016/j.envsoft.2013.10.024

  83. Schönhuth, S., Shiozawa, D.K., Dowling, T.E. & Mayden, R.L. (2012) Molecular systematics of western North American cyprinids (Cypriniformes: Cyprinidae), Zootaxa, 3586 (1), 281–303. https://doi.org/10.11646/zootaxa.3586.1.27

  84. Schönhuth, S., Vukić, J., Šanda, R., Yang, L. & Mayden, R.L. (2018) Phylogenetic relationships and classification of the Holarctic family Leuciscidae (Cypriniformes: Cyprinoidei). Molecular Phylogenetics and Evolution, 127, 781–799. https://doi.org/10.11646/zootaxa.3586.1.27

  85. Scoppettone, G.G., Hereford, M.E., Rissler, P.H., Johnson, D.M. & Salgado, J.A. (2011) Relative Abundance and Distribution of Fishes within an Established Area of Critical Environmental Concern, Inyo and San Bernardino Counties, California. Open-File Report 2011-1161. U.S. Department of the Interior, U.S. Geological Survey, Reston,Virginia. [unknown pagination] https://doi.org/10.3133/ofr20111161

  86. Scoppettone, G.G. & Smith, J.J. (1978) Additional records on the distribution and status of native fishes in Alameda and Coyote Creeks, California. California Fish and Game, 64, 62-65.

  87. Shapovalov, L. & Dill, W.A. (1950) A check list of the fresh-water and anadromous fishes of California. California Fish and Game, 36, 382–391

  88. Shapovalov, L., Dill, W.A. & Cordone, A.J (1959) A revised check list of the freshwater and anadromous fishes of California. California Fish and Game, 45, 159–180.

  89. Shapovalov, L., Cordone, A.J. & Dill, W.A. (1981) A list of the freshwater and anadromous fishes of California. California Fish and Game, 67, 4–38.

  90. Smith, G.R., Chow, J., Unmack, P.J., Markle, D.F. & Dowling, T.E. (2017) Evolution of the Rhinichthys osculus complex (Teleostei: Cyprinidae) in Western North America. Miscellaneous Publications Museum of Zoology University of Michigan, 204 (2), 1–83

  91. Smith, G.R., Dowling, T.E., Gobalet, K.T., Lugaski, T., Shiozawa, D. & Evans, P. (2002) Biogeography and timing rates of evolutionary events among Great Basin fishes. In: Hershler, R., Madsen, D.B. & Curry, D.R. (Eds.), Great Basin Aquatic Systems History, Smithsonian Contributions to the Earth Sciences, 33, pp. 175–233.

  92. Smith, G.R. & Dowling, T.E. (2008) Correlating hydrographic events and divergence times of Speckled Dace (Rhinichthys: Teleostei, Cyprinidae), in the Colorado River drainage. In: Reheis, M., Hershler, R. & Miller, D. (Eds.), Late Cenozoic drainage history of the Southwestern Great Basin and Lower Colorado River region: geologic and biotic perspectives. Geological Society of America Special Publication, 439, pp. 301–317. https://doi.org/10.1130/2008.2439(13)

  93. Snyder, J.O. (1908) Relationships of the fish fauna of the lakes of Southeastern Oregon. U.S. Bureau of Fisheries Bulletin, 27, 69–102.

  94. Snyder, J.O. (1913) The fishes of the streams tributary to San Francisco Bay. Bulletin of the United States Bureau of Fisheries, 32, 49–72.

  95. Snyder, J.O. (1917) An account of some fishes from Owens River, California. Proceedings, United States National Museum, 54, 201–205. https://doi.org/10.5479/si.00963801.2233.201

  96. Snyder, J.O. (1918) The fishes of the Lahontan system of Nevada and northeastern California. United States Bureau of Fisheries Bulletin, 35, 33–86.

  97. Spencer, J.E., Smith, G.R. & Dowling, T.E. (2008) Middle to late Cenozoic geology, hydrography, and fish evolution in the American Southwest. In: Reheis, M.C., Hershler, R. & Miller, D. (Eds.), Late Cenozoic Drainage History of the Southwestern Great Basin and Lower Colorado River Region: Geologic and Biotic Perspectives. Geological Society of America Special Paper, 439, pp. 279–299. https://doi.org/10.1130/2008.2439(12)

  98. Stewart, O.C. (1939) The Northern Paiute Bands. Anthropological Records, 2 (3), 127–149.

  99. Stout, C.C., Tan, M., Lemmon, A.R., Lemmon, E.M. & Armbruster, J.W. (2016) Resolving Cypriniformes relationships using an anchored enrichment approach. BMC Evolutionary Biology, 16 (1), 1–3. https://doi.org/10.1186/s12862-016-0819-5

  100. Su, Y., Moyle, P.B., Campbell, M.A., Finger, A.J., O’Rourke, S., Baumsteiger, J. & Miller, M.R. (2022) Population genomic analysis of the Speckled Dace species complex identifies three distinct lineages in California. Transactions of American Fisheries Society, 151, 695–710. https://doi.org/10.1002/tafs.10388

  101. Swift, C.C., Haglund, T.R., Ruiz, M. & Fisher, R.N. (1993) The status and distribution of the freshwater fishes of southern California. Bulletin, Southern California Academy Sciences, 92, 101–167.

  102. Tan, M. & Armbuster, J.W. (2018) Phylogenetic classification of extant genera of fishes of the order Cypriniformes (Teleostei: Ostariophysi). Zootaxa, 4476 (1), 6–39. https://doi.org/10.11646/zootaxa.4476.1.4

  103. Taylor, B.L., Perrin, W.F., Reeves, R.R., Rosel, P.E., Wang, J.Y., Cipriano, F., Scott Baker, C. & Brownell Jr., R.L. (2017) Why we should develop guidelines and quantitative standards for using genetic data to delimit subspecies for data‐poor organisms like cetaceans. Marine Mammal Science, 33 (S1), 12–26. https://doi.org/10.1111/mms.12413

  104. Unmack, P.J., Dowling, T.E., Laitinen, N.J., Secor, C.L., Mayden, R.L., Shiozawa, D.K. & Smith, G.R. (2014) Influence of introgression and geological processes on phylogenetic relationships of western North American mountain suckers (Pantosteus). PLoS One, 9 (3), e90061. [11 March] https://doi.org/10.1371/journal.pone.0090061

  105. U.S. Census Bureau, Department of Commerce (2001) Cartographic Boundary File (U.S. States). Available from: https://www.census.gov/geographies/mapping-files/time-series/geo/carto-boundary-file.html. (accessed 30 December 2007)

  106. U.S. Fish and Wildlife Service (2019) Removing the Foskett Speckled Dace from the list of Endangered and Threatened Wildlife, 84 FR 48290, 48290–48308. Docket No. FWS-R1-ES-2017–0051. Available from: http://www.regulations.gov (accessed 8 February 2023)

  107. U.S. Geological Survey (2004) National Hydrography Dataset. U.S. Department of the Interior, Reston, Virginia. Available from: https://www.usgs.gov/national-hydrography (accessed 30 December 2022)

  108. U.S. Geological Survey and U.S. Department of Agriculture, Natural Resources Conservation Service (2013) National Watershed Boundary Dataset (WBD). 4th Edition. Available from: https://www.usgs.gov/national-hydrography/watershed-boundary-dataset (accessed 30 December 2022)

  109. Wiesenfeld, J.C., Goodman, D.H. & Kinziger, A.P. (2018) Riverscape genetics identifies Speckled Dace (Rhinichthys osculus) cryptic diversity in the Klamath–Trinity Basin. Conservation Genetics, 19 (1), 111–127. https://doi.org/10.1007/s10592-017-1027-6

  110. Wilke, P.J. (1980) Prehistoric weir fishing on recessional shorelines of Lake Cahuilla. Proceedings of the Desert Fishes Council, XI, 101–103.

  111. Williams, J.E., Stern, M.A., Munhall, A.V. &. Anderson, G.A. (1990) Conservation status of threatened fishes in the Warner Basin, Oregon. Great Basin Naturalist, 50 (3), 243–248. [http://www.jstor.org/stable/41712597]

  112. Woodman, D.A. (1992) Systematic relationships within the cyprinid genus Rhinichthys. In: Mayden, R.L. (Ed.), Systematics, Historical Ecology and North American Freshwater Fishes. Stanford University Press, Stanford, California, pp. 374–391.

  113. Wydoski, R.S. & Whitney, R.R. (2003) Inland Fishes of Washington. 2nd Edition. University of Washington Press, Seattle, Washington, 547 pp.

  114. Yang, L., Sado, T., Hirt, M.V., Pasco-Viel, E., Arunachalam, M., Li, J., Wang, X., Freyhof, J., Saitoh, K., Simons, A.M. & Miya, M. (2015) Phylogeny and polyploidy: resolving the classification of cyprinine fishes (Teleostei: Cypriniformes). Molecular Phylogenetics and Evolution, 85, 97–116. https://doi.org/10.1016/j.ympev.2015.01.014