Abstract
We describe as new species Micropterus pucpuggy Freeman & Freeman (Bartram’s Bass), sp. nov., and Micropterus calliurus Freeman & Freeman (Altamaha Bass), sp. nov., which occur allopatrically in four river systems draining the Atlantic Slope of the southeastern United States. In recent decades, biologists and anglers have acknowledged the existence of these two distinctive taxa of black bass, both of which were previously considered synonymous with M. coosae Hubbs & Bailey (Redeye Bass). However, introgression with non-native congeners that have been widely introduced for sport-fishing (including M. henshalli Hubbs & Bailey and M. dolomieu Lacepède) has confounded formal description of M. pucpuggy and M.calliurus. We examined mitochondrial (mtDNA) and nuclear gene sequences of candidate type-specimens of M. pucpuggy and M. calliurus. We then used reduced-representation, short-read sequencing of candidate types along with specimens of six other Micropterus species to identify a series of non-introgressed individuals for each of the two new species. Micropterus pucpuggy and M. calliurus are each reciprocally monophyletic in both mitochondrial and RADseq phylogenies and are diagnosable from all other Micropterus species and from each other in chromatic fin coloration, body pigmentation, and other morphological attributes.
References
- Bagley, J.C., Mayden, R.L., Roe, K.J., Holznagel, W. & Harris, P.M. (2011) Congeneric phylogeographical sampling reveals polyphyly and novel biodiversity within black basses (Centrarchidae: Micropterus). Biological Journal of the Linnean Society, 104 (2), 346–363. https://doi.org/10.1111/j.1095-8312.2011.01720.x
- Bailey, R.M. & Hubbs, C.L. (1949) The black bases (Micropterus) of Florida, with description of a new species. Occasional Papers of the Museum of Zoology, University of Michigan, 516, 1–43.
- Baker, W.H., Blanton, R.E. & Johnston, C.E. (2013) Diversity within the Redeye Bass, Micropterus coosae (Perciformes: Centrarchidae) species group, with descriptions of four new species. Zootaxa, 3635 (4), 379–401. https://doi.org/10.11646/zootaxa.3635.4.3
- Bangs, M. (2011) Decline of the Savannah River Redeye Bass (Micropterus coosae) Due to Introgressive Hybridization with Invasive Alabama Spotted Bass (Micropterus punctulatus henshalli). MS Thesis, University of South Carolina, Columbia, South Carolina, 39 pp.
- Bangs, M.R., Oswald, K.J., Greig, T.W., Leitner, J.K., Rankin, D.M. & Quattro, J.M. (2018) Introgressive hybridization and species turnover in reservoirs: a case study involving endemic and invasive basses (Centrarchidae: Micropterus) in southeastern North America. Conservation Genetics, 19, 57–69. https://doi.org/10.1007/s10592-017-1018-7
- Barwick, D.H., Oswald, K.J., Quattro, J.M. & Barwick, R.D. (2006) Redeye bass (Micropterus coosae) and Alabama spotted bass (M. punctulatus henshalli) hybridization in Keowee Reservoir. Southeastern Naturalist, 5 (4), 661–668. https://doi.org/10.1656/1528-7092(2006)5[661:RBMCAA]2.0.CO;2
- Barwick, D.H. & Moore, P.R. (1983) Abundance and growth of redeye bass in two South Carolina reservoirs. Transactions of the American Fisheries Society, 112 (2A), 216–219. https://doi.org/10.1577/1548-8659(1983)112%3C216:AAGORB%3E2.0.CO;2
- Bayona-Vásquez, N.J., Glenn, T.C., Kieran, T.J., Pierson, T.W., Hoffberg, S.L., Scott, P.A., Bentley, K.E., Finger, J.W., Louha, S., Troendle, N., Diaz-Jaimes, P., Mauricio, R. & Faircloth, B.C. (2019) Adapterama III: Quadruple-indexed, double/triple-enzyme RADseq libraries (2RAD/3RAD). PeerJ, 7, e7724. https://doi.org/10.7717/peerj.7724
- Berra, T.M. (1989) The significance of William Bartram (1739-1823) to North American ichthyology. American Midland Naturalist, 122 (2), 214–227. https://doi.org/10.2307/2425906
- Borror, D.J. (1960) Dictionary of Word Roots and Combining Forms. Mayfield Publishing Comp, Palo Alto, California, 134 pp.
- Boschung, H.T. & Mayden, R.L. (2004) Fishes of Alabama. Smithsonian Institution Press, Washington D.C., 736 pp.
- Catchen, J., Hohenlohe, P.A., Bassham, S., Amores, A. & Cresko, W.A. (2013) Stacks: an analysis tool set for population genomics. Molecular Ecology, 22 (11), 3124–3140. https://doi.org/10.1111/mec.12354
- Cox, C. (2022) Patterns of Occurrence of the Black Basses in the Upper Savannah River Tributaries of Georgia and South Carolina with Emphasis on Bartram’s Bass Micropterus sp. cf. coosae. MS Thesis, Clemson University, Clemson, South Carolina, 73 pp.
- Dakin, E.E., Porter, B.A., Freeman, B.J. & Long, J.M. (2015) Hybridization threatens Shoal Bass populations in the upper Chattahoochee River basin. In: Tringali, M.D., Long, J.M., Birdsong, T.W. & Allen, M.S. (Eds.), Black Bass Diversity: Multidisciplinary Science for Conservation. American Fisheries Society, Bethesda, Maryland, pp. 491–501.
- DeRaad, D.A. (2022) SNPfiltR: an R package for interactive and reproducible SNP filtering. Molecular Ecology Resources, 22, 2443–2453. https://doi.org/10.1111/1755-0998.13618
- Eroh, G. (2020) The bass menagerie: A Micropterus odyssey. American Currents, 45 (3), 6–13.
- Etnier, D.A. & Starnes, W.C. (1993) The Fishes of Tennessee. The University of Tennessee Press, Knoxville, Tennessee, 689 pp.
- Freeman, B.J., Taylor, A., Oswald, K.J., Wares, J., Freeman, M.C., Quattro, J. & Leitner, J. (2015) Shoal basses, a clade of cryptic identity. In: Tringali, M.D., Long, J.M., Birdsong, T.W. & Allen, M.S. (Eds.), Black Bass Diversity: Multidisciplinary Science for Conservation. American Fisheries Society, Bethesda, Maryland, pp. 449–466.
- Frichot, E., Mathieu, F., Trouillon, T., Bouchard, G. & François, O. (2014) Fast and efficient estimation of individual ancestry coefficients. Genetics, 196 (4), 973–983. https://doi.org/10.1534/genetics.113.160572
- Gunn, J.C., Taylor, A.T., Buckingham, J.J., Kern, A.I. & Long, J.M. (2023) Limited hybridisation and introgression despite stocking among endemic Interior Highlands black basses (Centrarchidae: Micropterus). Diversity and Distributions, 29 (10), 1299–1314. https://doi.org/10.1111/ddi.13759
- Harper, F. (1958) The Travels of William Bartram, Naturalist’s Edition. Yale University Press, New Haven, Connecticut, 727 pp.
- Henshall, J.A. (1881) Book of the Black Bass. Robert Clarke Co., Cincinnati, Ohio, 463 pp. https://doi.org/10.5962/bhl.title.40249
- Hubbs, C.L. & Bailey, R.M. (1940) A revision of the black basses (Micropterus and Huro) with descriptions of four new forms. Miscellaneous Publications, Museum of Zoology, University of Michigan. No. 48. Museum of Zoology, University of Michigan, Ann Arbor, Michigan, 51 pp. https://doi.org/10.2307/1437996
- Hubbs, C.L. & Lagler, K.F. (1958) Fishes of the Great Lakes region. 2nd edition. Cranbrook Institute Science Bulletin, 26, 1–213. https://doi.org/10.3998/mpub.12946839
- Huelsenbeck, J.P. & Ronquist, F. (2001) MrBayes: Bayesian inference of phylogeny. Bioinformatics, 17, 754–755. https://doi.org/10.1093/bioinformatics/17.8.754
- Jackson, C.R., Martin, J.K., Leigh, D.S. & West, L.T. (2005) A southeastern piedmont watershed sediment budget: Evidence for a multi-millennial agricultural legacy. Journal of Soil and Water Conservation, 60 (6), 298–310. https://doi.org/10.1080/00224561.2005.12435820
- Jackson, D.A. (2002) Ecological effects of Micropterus introductions: the dark side of black bass. In: Philipp, D.P. & Ridgway, M.S. (Eds.), Black Bass: Ecology, Conservation, and Management. American Fisheries Society, Bethesda, Maryland, pp. 221–232.
- Jombart, T. & Collins, C. (2015) An introduction to adegenet 2.0. 0. Available from: http://adegenet.r-forge.r-project.org/files/tutorial-basics.pdf (accessed 28 July 2025)
- Jombart, T., Devillard, S. & Balloux, F. (2010) Discriminant analysis of principal components: a new method for the analysis of genetically structured populations. BMC Genetics, 11, 1–15. https://doi.org/10.1186/1471-2156-11-94
- Judson, E.E. (2018) Conservation of Endemic Bartram’s Bass: Nesting Microhabitat Use and Spatial Distribution with Congeners in the Savannah River Basin. MS Thesis, Clemson University, Clemson, South Carolina, 70 pp.
- Judson, E., Darden, T.L., Farrae, D., Leitner, J. & Peoples, B.K. (2021) Nesting microhabitat use and hybridisation of endemic Bartram’s Bass in Savannah River tributaries. Ecology of Freshwater Fish, 30 (3), 306–319. https://doi.org/10.1111/eff.12585
- Kamvar, Z.N., Tabima, J.F. & Grünwald, N.J. (2014) Poppr: An R package for genetic analysis of populations with clonal, partially clonal, and/or sexual reproduction. PeerJ, 2, e281. https://doi.org/10.7717/peerj.281
- Kim, D., Taylor, A.T. & Near, T.J. (2022) Phylogenomics and species delimitation of the economically important Black Basses (Micropterus). Scientific Reports, 12 (1), 9113. https://doi.org/10.1038/s41598-022-11743-2
- Knaus, B.J., Grünwald, N.J., Anderson, E.C., Winter, D.J., Kamvar, Z.N. & Tabima, J.F. (2023) Package ‘vcfR’. Available from: https://github.com/knausb/vcfR (accessed 28 July 2025)
- Koppelman, J.B. & Garrett, G.P. (2002) Distribution, biology, and conservation of the rare black bass species. In: Philipp, D.P. & Ridgway, M.S. (Eds.), Black Bass: Ecology, Conservation, and Management. American Fisheries Society, Bethesda, Maryland, pp. 333–341.
- Lee, D.S., Gilbert, C.R., Hocutt, C.H., Jenkins, R.E., McAllister, D.E. & Stauffer Jr, J.R. (1980) Atlas of North American Freshwater Fishes. North Carolina Museum of Natural History, Raleigh, North Carolina, 867 pp. https://doi.org/10.5962/bhl.title.141711
- Leitner, J.K., Oswald, K.J., Bangs, M., Rankin, D. & Quattro, J.M. (2015) Hybridization between native Bartram’s Bass and two introduced species in Savannah drainage streams. In: Tringali, M.D., Long, J.M., Birdsong, T.W. & Allen, M.S. (Eds.), Black Bass Diversity: Multidisciplinary Science for Conservation. American Fisheries Society, Bethesda, Maryland, pp. 481–490.
- Long, J.M., Allen, M.S., Porak, W.F. & Suski, C.D. (2015) A historical perspective of black bass management in the United States. In: Tringali, M.D., Long, J.M., Birdsong, T.W. & Allen, M.S. (Eds.), Black Bass Diversity: Multidisciplinary Science for Conservation. American Fisheries Society, Bethesda, Maryland, pp. 99–122.
- Maddison, W.P. & Whitton, J. (2023) The species as a reproductive community emerging from the past. Bulletin of the Society of Systematic Biologists, 2 (1), 1–35. https://doi.org/10.18061/bssb.v2i1.9358
- Menhinick, E.F. (1991) The Freshwater Fishes of North Carolina. North Carolina Wildlife Resources Commission, Raleigh, North Carolina, 227 pp.
- Near, T.J. & Kim, D. (2021) Phylogeny and time scale of diversification in the fossil-rich sunfishes and black basses (Teleostei: Percomorpha: Centrarchidae). Molecular Phylogenetics and Evolution, 161, 107156. https://doi.org/10.1016/j.ympev.2021.107156
- Oswald, K.J. (2007) Phylogeography and Contemporary History of Redeye Bass (Micropterus coosae). Ph.D Thesis, University of South Carolina, Columbia.
- Oswald, K.J., Leitner, J., Rankin, D., Barwick, D.H., Freeman, B.J., Greig, T., Bangs, M. & Quattro, J.M. (2015) Evolutionary genetic diversification, demography, and conservation of Bartram’s Bass. In: Tringali, M.D., Long, J.M., Birdsong, T.W. & Allen, M.S. (Eds.), Black Bass Diversity: Multidisciplinary Science for Conservation. American Fisheries Society, Bethesda, Maryland, pp. 601–614.
- Page, L.M. & Burr, B.M. (1991) A Field Guide to Freshwater Fishes of North America North of Mexico. Houghton Mifflin Company, Boston, Massachusetts, 432 pp.
- Paris, J.R., Stevens, J.R. & Catchen, J.M. (2017) Lost in parameter space: A road map for Stacks. Methods in Ecology and Evolution, 8, 1360–1373. https://doi.org/10.1111/2041-210X.12775
- Peoples, B.K., Judson, E., Darden, T.L., Farrae, D.J., Kubach, K., Leitner, J. & Scott, M.C. (2021) Modeling distribution of endemic Bartram’s bass Micropterus sp. cf. coosae: Disturbance and proximity to invasion source increase hybridization with invasive Alabama bass. North American Journal of Fisheries Management, 41, 1309–1321. https://doi.org/10.1002/nafm.10637
- Power, M.E., Matthews, W.J. & Stewart, A.J. (1985) Grazing minnows, piscivorous bass, and stream algae: dynamics of a strong interaction. Ecology, 66 (5), 1448–1456. https://doi.org/10.2307/1938007
- Pritchard, J.K., Stephens, M. & Donnelly, P. (2000) Inference of population structure using multilocus genotype data. Genetics, 155, 945–959. https://doi.org/10.1093/genetics/155.2.945
- Pyron, R.A., O’Connell, K.A., Duncan, S.C., Burbrink, F.T. & Beamer, D.A. (2023) Speciation hypotheses from phylogeographic delimitation yield an integrative taxonomy for Seal Salamanders (Desmognathus monticola). Systematic Biology, 72 (1), 179–197. https://doi.org/10.1093/sysbio/syac065
- R Core Team (2023) R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna. Available from: https://www.R-project.org/ (accessed 28 July 2025)
- Robins, R.H., Page, L.M., Williams, J.D., Randall, Z.S. & Sheehy, G.E. (2018) Fishes in the Fresh Waters of Florida: An Identification Guide and Atlas. University of Florida Press, Gainesville, Florida, 488 pp. https://doi.org/10.2307/j.ctvx1ht6s
- Rochette, N.C., Rivera-Colón, A.G. & Catchen, J.M. (2019) Stacks 2: Analytical methods for paired-end sequencing improve RADseq-based population genomics. Molecular Ecology, 28 (21), 4737–4754. https://doi.org/10.1111/mec.15253
- Rohde, F.C., Arndt, R.G., Foltz, J.W. & Quattro, J.M. (2009) Freshwater Fishes of South Carolina. Univ of South Carolina Press, Columbia, South Carolina, 430 pp.
- Rohde, F.C., Arndt, R.G., Lindquist, D.G. & Parnell, J.F. (1994) Freshwater Fishes of the Carolinas, Virginia, Maryland, and Delaware. University of North Carolina Press, Chapel Hill, North Carolina, 222 pp.
- Ronquist, F., Teslenko, M., Van Der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61 (3), 539–542. https://doi.org/10.1093/sysbio/sys029
- Taylor, A.T., Long, J.M., Tringali, M.D. & Barthel, B.L. (2019) Conservation of black bass diversity: an emerging management paradigm. Fisheries, 44, 20–36. https://doi.org/10.1002/fsh.10187
- Thia, J.A. (2022) Guidelines for standardizing the application of discriminant analysis of principal components to genotype data. Molecular Ecology Resources, 23, 523–538. https://doi.org/10.1111/1755-0998.13706
- Toczydlowski, R.H., Drummond, C.P., Brennan, R.S., Goldstein, S. & Kartzinel, R.Y. (2025) Stacks on SLURM pipeline. [in preparation]
- Tracy, B.H., Rohde, F.C. & Hogue, G.M. (2020) An annotated atlas of the freshwater fishes of North Carolina. Southeastern Fishes Council Proceedings, 60, 1–198. https://doi.org/10.7290/sfcp60gca7
- Tracy, B.H., Rohde, F.C., Smith, S.A., Bissette, J.L. & Hogue, G.M. (2024) A Guide to North Carolina’s Freshwater Fishes. University of North Carolina Press, Chapel Hill, North Carolina, 453 pp.
- Trimble, S.W. (1974) Man-induced Soil Erosion on the Southern Piedmont, 1700-1970. Soil Water Conservation Society, Ankeny, Iowa, 188 pp.
- Walter, R.C. & Merritts, D.J. (2008) Natural streams and the legacy of water-powered mills. Science, 319 (5861), 299–304. https://doi.org/10.1126/science.1151716
- Wares, J.P. (2024) The genomic ghosts of Geukensia granosissima. Estuaries and Coasts, 47, 494–503. https://doi.org/10.1007/s12237-023-01296-6
- Williams, J.D. & Burgess, G.H. (1999) A new species of bass, Micropterus cataractae (Teleostei: Centrarchidae), from the Apalachicola River basin in Alabama, Florida, and Georgia. Bulletin of the Florida Museum of Natural History, 42, 80–114. https://doi.org/10.58782/flmnh.afti4206