Abstract
A new species of crayfish, Cambarus emegi sp. nov., is described from the upper James and Roanoke River basins in Virginia and the lower Delaware River basin in Pennsylvania, where it occurs in streams and small rivers. The species was formerly part of the Cambarus acuminatus species complex (Cambarus species C complex), a long-standing taxonomic enigma that is widely distributed across the Atlantic slope drainages of eastern North America. Cambarus emegi sp. nov. can be distinguished from Cambarus acuminatus due to the absence of a well-defined subpalmar tubercle, greater number of punctations across its areola, and differences in the mesial process of the form I male gonopod. Coloration and pattern in life differ between the two species. Additionally, the distributions of C. emegi sp. nov. and C. acuminatus are separated by hundreds of kilometers. The ranges of two other undescribed members of the Cambarus species C complex border that of C. emegi sp. nov. in Virginia. These species do not appear to overlap geographically; C. emegi sp. nov. is confined to the Ridge and Valley physiographic province, whereas the two undescribed species are found in the Blue Ridge and northern inner Piedmont regions. Cambarus emegi sp. nov. is morphologically similar but visually and genetically different from the undescribed species directly to the south in the upper Roanoke River system. Cambarus emegi sp. nov. is both visually and morphologically similar but genetically distinct from the undescribed species immediately downstream in the James River basin. Cambarus emegi sp. nov. is threatened by several potentially interacting factors including invasive crayfishes, narrow range, and urban and agricultural development. Alarmingly, the species is being rapidly replaced by invasive Faxonius species across a substantial part of its range. This paper represents an important step in providing the information needed to conserve and manage Cambarus emegi sp. nov.
References
- Buhay, J.E., Moni, G., Mann, N. & Crandall, K.A. (2007) Molecular taxonomy in the dark: evolutionary history, phylogeography, and diversity of cave crayfish in the subgenus Aviticambarus, genus Cambarus. Molecular Phylogenetics and Evolution, 42 (2), 435–448. https://doi.org/10.1016/j.ympev.2006.07.014
- Chernomor, O., Von Haeseler, A. & Minh, B.Q. (2016) Terrace aware data structure for phylogenomic inference from supermatrices. Systematic Biology, 65 (6), 997–1008. https://doi.org/10.1093/sysbio/syw037
- Cooper, J.E. (2001) Cambarus (Puncticambarus) hobbsorum, a new crayfish (Decapoda: Cambaridae) from North Carolina. Proceedings of the Biological Society of Washington, 114 (1), 152–161.
- Cooper, J.E. (2006) A new species of crayfish of the genus Cambarus Erichson, 1846 (Decapoda: Cambaridae) from the eastern Blue Ridge foothills and western Piedmont Plateau of North Carolina. Proceedings of the Biological Society of Washington, 119 (1), 67–80. https://doi.org/10.2988/0006-324X(2006)119[67:ANSOCO]2.0.CO;2
- Cooper, J.E. & Cooper, D.G. (2003) A new crayfish of the genus Cambarus Erichson, 1846 (Decapoda: Cambaridae) from the Cape Fear River basin in the Sandhills of North Carolina. Proceedings of the Biological Society of Washington, 116 (4), 920–932.
- Cooper, J.E. & Price, J.E. (2010) A new spinose crayfish of the genus Cambarus, subgenus Puncticambarus (Decapoda: Cambaridae), from South Carolina. Proceedings of the Biological Society of Washington, 123 (4), 335–344. https://doi.org/10.2988/10-20.1
- Erichson, W.F. (1846) Uebersicht der Arten der Gattung Astacus. Archiv fur Naturgeschichte, 12 (1), 86–103.
- Fabricius, J.C. (1798) Supplementum entomologiae systematicae. Proft et Storch, Hafniae, 572 pp. https://doi.org/10.5962/bhl.title.65803
- Faxon, W. (1884) Descriptions of new species of Cambarus; to which is added a synonymical list of the known species of Cambarus and Astacus. Proceedings of the American Academy of Arts and Sciences, 20, 107–158. https://doi.org/10.2307/25138768
- Fetzner, J.W., Jr. & Crandall, K.A. (2013) Assessing the genetic distinctiveness of Cambarus (Puncticambarus) acuminatus: a recently discovered crayfish in Valley Creek within Valley Forge National Historical Park. Natural Resource Technical Report NPS/VAFO/NRTR—2013/752. National Park Service, Fort Collins, Colorado. https://doi.org/10.13140/RG.2.2.16232.52483
- Folmer, O.F., Black, M.B., Hoeh, W.R. & Vrijenhoek, R.C. (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3 (5), 294–299.
- Girard, C. (1852) A revision of the North American Astaci, with observations on their habits and geographical distribution. Proceedings of the Academy of Natural Sciences of Philadelphia, 6, 87–91.
- Hagen, H.A. (1870) Monograph of the North American Astacidae. Illustrated Catalogue of the Museum of Comparative Zoology at Harvard College, Number 3, viii + 109 pp.
- Hoang, D.T., Chernomor, O., von Haeseler, A., Minh, B.Q. & Vinh, L.S. (2018) UFBoot2: Improving the ultrafast bootstrap approximation. Molecular Biology and Evolution, 35 (2), 518–522. https://doi.org/10.1093/molbev/msx281
- Hobbs, H.H., Jr. (1942) A generic revision of the crayfishes of the subfamily Cambarinae (Decapoda, Astacidae) with the description of a new genus and species. American Midland Naturalist, 28 (2), 334–357. https://doi.org/10.2307/2420820
- Hobbs, H.H., Jr. (1972) Crayfishes (Astacidae) of North and Middle America. Biota of Freshwater Ecosystems. Identification Manual No. 9 (U.S. Environmental Protection Agency), i–x + 173 pp.
- Hobbs, H.H., Jr. (1974) A checklist of the North and Middle American crayfishes (Decapoda: Astacidae and Cambaridae). Smithsonian Contributions to Zoology, 166, 1–161. https://doi.org/10.5479/si.00810282.166
- Hobbs, H.H., Jr. (1989) An illustrated checklist of the American crayfishes (Decapoda: Astacidae, Cambaridae, and Parastacidae). Smithsonian Contributions to Zoology, 480, 1–236. https://doi.org/10.5479/si.00810282.480
- Hobbs, H.H., Jr. & Hobbs, H.H., III. (1990) A new crayfish (Decapoda: Cambaridae) from southeastern Texas. Proceedings of the Biological Society of Washington, 103 (3), 608–613.
- Hobbs, H.H., Jr. & Peters, D.J. (1977) The entocytherid ostracods of North Carolina. Smithsonian Contributions to Zoology, 247, 1–77. https://doi.org/10.5479/si.00810282.247
- Kalyaanamoorthy, S., Minh, B.Q., Wong, T.K.F., von Haeseler, A. & Jermiin, L.S. (2017) ModelFinder: Fast model selection for accurate phylogenetic estimates. Nature Methods, 14, 587–589. https://doi.org/10.1038/nmeth.4285
- Katoh, K. & Standley, D.M. (2013) MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Molecular Biology and Evolution, 30 (4), 772–780. https://doi.org/10.1093/molbev/mst010
- Katoh, K., Misawa, K., Kuma, K. & Miyata, T. (2002) MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform. Nucleic Acids Research, 30 (14), 3059–3066. https://doi.org/10.1093/nar/gkf436
- Leray, M., Yang, J.Y., Meyer, C.P., Mills, S.C., Agudelo, N., Ranwez, V., Boehm, J.T. & Machida, R.J. (2013) A new versatile primer set targeting a short fragment of the mitochondrial COI region for metabarcoding metazoan diversity: application for characterizing coral reef fish gut contents. Frontiers in Zoology, 10, 34. https://doi.org/10.1186/1742-9994-10-34
- Lieb, D.A., Carline, R.F., Rosenberger, J.L. & Mengel, V.M. (2008) The discovery and ecology of a member of the Cambarus acuminatus complex (Decapoda: Cambaridae) in Valley Creek, southeastern Pennsylvania. Journal of Crustacean Biology, 28 (3), 439–450. https://doi.org/10.1651/07-2877R.1
- Lieb, D.A., Bouchard, R.W. & Carline, R.F. (2011a) The crayfish fauna of southeastern Pennsylvania: Distributions, ecology, and changes over the last century. Journal of Crustacean Biology, 31 (1), 166–178. https://doi.org/10.1651/10-3287.1
- Lieb, D.A., Bouchard, R.W., Carline, R.F., Nuttall, T.R., Wallace, J.R. & Wengert, C.B. (2011b) Conservation and management of crayfishes: Lessons from Pennsylvania. Fisheries, 36 (10), 489–507. https://doi.org/10.1080/03632415.2011.607080
- Lindqvist, O.V. & Huner, J.V. (1999) Life history characteristics of crayfish: What makes some of them good colonizers? In: Gherardi, F. & Holdich, D.M. (Eds.), Crustacean Issues. Vol. 11. Crayfish in Europe as Alien Species: How to Make the Best of a Bad Situation?. A.A. Balkema, Rotterdam, The Netherlands, pp. 23–30. https://doi.org/10.1201/9781315140469-3
- Loughman, Z.J., Welsh, S.A. & Thoma, R.F. (2017) Cambarus (C.) appalachiensis, a new species of crayfish (Decapoda: Cambaridae) from the New River Basin of Virginia and West Virginia, USA. Zootaxa, 4243 (3), 432–454. https://doi.org/10.11646/zootaxa.4243.3.2
- Minh, B.Q., Schmidt, H., Chernomor, O., Schrempf, D., Woodhams, M., von Haeseler, A. & Lanfear, R. (2020) IQ-TREE 2: new models and efficient methods for phylogenetic inference in the genomic era. Molecular Biology and Evolution, 37 (5), 1530–1534. https://doi.org/10.1093/molbev/msaa015
- Nguyen, L.-T., Schmidt, H.A., von Haeseler, A. & Minh, B.Q. (2015) IQ-TREE: A fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution, 32 (1), 268–274. https://doi.org/10.1093/molbev/msu300
- Perkins, M.A., Williams, B.W. & Russ, W.T. (2019) Cambarus franklini, a new crayfish (Decapoda: Cambaridae) from the Catawba River Basin in western North Carolina, USA. Zootaxa, 4568 (3), 520–532. https://doi.org/10.11646/zootaxa.4568.3.6
- Perkins, M.A., Williams, B.W. & Russ, W.T. (2023) Integrative taxonomy reveals two new narrowly-endemic crayfish species (Decapoda: Cambaridae) from the Yadkin River Basin in western North Carolina, USA. Zootaxa, 5270 (2), 207–230. https://doi.org/10.11646/zootaxa.5270.2.3
- Rafinesque, C.S. (1817) Synopsis of four new genera and ten new species of Crustacea, found in the United States. American Monthly Magazine and Critical Revue, 2, 40–43.
- Richman, N.I., Böhm, M., Adams, S.B., Alvarez, F., Bergey, E.A., Bunn, J.J., Burnham, Q., Cordeiro, J., Coughran, J., Crandall, K.A., Dawkins, K.L., DiStefano, R.J., Doran, N.E., Edsman, L., Eversole, A.G., Füreder, L., Furse, J.M., Gherardi, F., Hamr, P., Holdich, D.M., Horwitz, P., Johnston, K., Jones, C.M., Jones, J.P.G., Jones, R.L., Jones, T.J., Kawai, T., Lawler, S., López-Mejía, M., Miller, R.M., Pedraza-Lara, C., Reynolds, J.D., Richardson, A.M.M., Schultz, M.B., Schuster, G.A., Sibley, P.J., Souty-Grosset, C., Taylor, C.A., Thoma, R.T., Walls, J., Walsh, T.S. & Collen, B. (2015) Multiple drivers of decline in the global status of freshwater crayfish (Decapoda: Astacidea). Philosophical Transactions of the Royal Society B: Biological Sciences, 370 (1662), 20140060. https://doi.org/10.1098/rstb.2014.0060
- Taylor, C.A., Warren, M.L., Jr., Fitzpatrick, J.F., Jr., Hobbs, H.H., III, Jezerinac, R.F., Pflieger, W.L. & Robison, H.W. (1996) Conservation status of crayfishes of the United States and Canada. Fisheries, 21 (4), 25–38. https://doi.org/10.1577/1548-8446(1996)021<0025:csocot>2.0.co;2
- Taylor, C.A. (2000) Systematic studies of the Orconectes juvenilis complex (Decapoda: Cambaridae), with descriptions of two new species. Journal of Crustacean Biology, 20 (1), 132–152. https://doi.org/10.1163/20021975-99990023
- Taylor, C.A., Schuster, G.A., Cooper, J.E., DiStefano, R.J., Eversole, A.G., Hamr, P., Hobbs, H.H., III, Robison, H.W., Skelton, C.E. & Thoma, R.F. (2007) A reassessment of the conservation status of crayfishes of the United States and Canada after 10+ years of increased awareness. Fisheries, 32 (8), 372–389. https://doi.org/10.1577/1548-8446(2007)32[372:AROTCS]2.0.CO;2
- Taylor, C.A., DiStefano, R.J., Larson, E.R. & Stoeckel, J. (2019) Towards a cohesive strategy for the conservation of the United States’ diverse and highly endemic crayfish fauna. Hydrobiologia, 846 (1), 39–58. https://doi.org/10.1007/s10750-019-04066-3
- Wang, Y., Tan, W., Li, B., Wen, L. & Lei, G. (2021) Habitat alteration facilitates the dominance of invasive species through disrupting niche partitioning in floodplain wetlands. Diversity and Distributions, 27 (9), 1861–1871. https://doi.org/10.1111/ddi.13376
- Williams, B.W., Delekta, E.M. & Loughman, Z.J. (2019) Redescription and circumscription of the Acuminate Crayfish, Cambarus acuminatus Faxon, 1884 (Decapoda: Cambaridae). Zootaxa, 4560 (1), 40–50. https://doi.org/10.11646/zootaxa.4560.1.2
- Williams, A.B. (1952) Six new crayfishes of the genus Orconectes (Decapoda: Astacidae) from Arkansas, Missouri and Oklahoma. Transactions of the Kansas Academy of Science, 55 (2), 330–351. https://doi.org/10.2307/3626240
- Williams, B.W., Loughman, Z.J. & Lieb, D.A. (2020) Taxonomy and conservation status of Cambarus (Puncticambarus) sp. in Pennsylvania. Final Report for Project Number F16AF01305. Submitted to the Pennsylvania Fish & Boat Commission, Bellefonte, Pennsylvania, 50 pp.
