The Classification and Scientific Names of Rainbow and Cutthroat Trouts Gerald R
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The Classification and Scientific Names of Rainbow and Cutthroat Trouts Gerald R. Smith and Ralph F. Stearley ABSTRACT Two unambiguous discoveries involving rainbow trout require scientific name changes. First, the rainbow trout has been demonstrated to be the same species as the Kamchatka trout. Second, studies of osteology and bio- chemistry of trout and salmon show that rainbow and cutthroat trout, and their close relatives, the golden, Mexican golden, Gila, and Apache trouts, are more closely related to Pacific salmons (Oncorhynchus) than to brown trout and Atlantic salmon (Salmo). The different names required by these two discoveries will cause some confusion in communications in which the formal classification is used, so we present evidence to acquaint biologists and managers with the rationale for the changes. The species name of the rainbow trout becomes mykiss, an older Latinized indigenous name of the Kamchatka trout. The generic designation of rainbow and cutthroat trout poses a more subjective problem, involving four possibilities: Salmo, Oncorhynchus, Rhabdofario, and Parasalmo. The balance of evidence indicates to us that the generic name for Pacific trouts and salmons should be Oncorhynchus. We suggest recognition of two divergent sister lineages, (1) Atlantic trout and salmon, and (2) Pacific trouts and salmons, as the genera Salmo and Oncorhynchus, respectively. Alternative generic classifications considered include the following: (a) Enlarge Salmo to include all Atlantic and Pacific trouts and salmons. This would be well supported by morphological and biochemical characters, but would fail to emphasize the dis- tinctions between the Pacific and Atlantic groups. (b) Use a separate generic name, Rhabdofario, for rainbow and cutthroat trout, and their inland relatives. This would be valid, but would fail to recognize the gradation between Pacific trouts and Pacific salmons. (c) Continue to assign Pacific trout to the genus Salmo, separate from On- corhynchus. This would be stable, but at the expense of evolutionary information in the classification-rainbow and cutthroat trout are on the same branch of evolution as the Pacific salmon. To reflect these biological re- lationships in the classification of trouts and salmons will contribute to better understanding of their life histories and better predictions for their management. The AFS Names of Fishes Committee has the salmon. If the classification is not neric and specific names of trout and adopted several changes in North American clear it is impossible to trace even salmon are partly the result of his- trout names as outlined in the following the distribution of this group. Still toric sequence of publication of de- article. The American FisheriesSociety has less is it possible to elucidate the scriptions by early pioneers in fish implemented use of the new names in all question of the origin (freshwater or biology, and partly the result of AFS publications produced after January marine) of the salmon; of the bio- modifications necessitated by new 1, 1989. See also, Fisheries 13(6):24.- logical reason for the inevitable death information on species relationships. The Editor. after spawning of all specimens in At any point in history, people some of the species of this tribe; of working on classification try to sat- IlX arge quantities of Salmo the origin of the migratory habits of isfy two criteria: (1) stability of no- Pass through the market the salmon; of the return of these menclature and (2) correspondence and hundreds of thousands of tons fish for spawning purposes into their between classification and knowl- of Oncorhynchus are caught annually. native streams, and many other edge about evolutionary relatedness. Many laboratories investigate these problems which interest everyone These criteria come into conflict when fishes, yet there is no agreement on who studies the life history of the new information compels re-align- simple and fundamental questions, salmon. ment of species groupings. In the such as: (1) Does the genus Salmo "Therefore, the first thing that must present case, information on the re- extend to the Pacific? (2) Does there be done is to attempt to find a nat- lationships of Pacific trouts and exist a distinct genus Oncorhynchus ural arrangement of this group." salmons has been available for 70 distinct from Salmo? Such an uncer- V. Tchernavin (1937 p. 236) years (since the work of C. Tate Re- tainty inevitably renders it impossi- gan in 1914), but people have been ble to clarify the many obscure ques- The Problem reluctant to destabilize names that so tions in regard to the life history of The internationally accepted ge- many biologists and managers use in their reports and communications. A Gerald R. Smith is the curator of fishes at the Museum of Zoology and Museum of number of workers have recently de- Paleontology, University of Michigan, Ann Arbor, MI 48109. His primary research is cided to correct the classification, but on the evolution of catostomids, salmonids, sculpins, and other freshwater fishes, with have unfortunately chosen an invalid special emphasis on speciation in whitefishes and sculpins. Ralph F. Stearly is a doctoral name, Parasalmo, for classifying Pa- student in fish paleontology and evolution in the Museum of Paleontology at the Univer- cific trouts. We review the history of sity of Michigan. trout and salmon names and suggest 4 Fisheries, Vol. 14, No. 1 choices which we think will be sta- Jordan and Evermann (1896), influ- ble in the future. enced by characteristics given by At the June 1988 meeting of The genus Salmo was proposed by Richardson (1836) and Suckley (1860), the American Society of Ichthy- Linnaeus in 1758. It originally in- applied the name mykiss (Walbaum's ologists and Herpetologists, the cluded the Atlantic salmon and trout 1792 Kamchatka trout) to the cut- APS-ASIH Committee on Names (Salmo salar and Salmo trutta, respec- throat trout, which they mistakenly of Fishes was able to review thor- tively) as well as huchen, chars, thought to extend to Kamchatka. In oughly the study of Pacific trouts grayling, and other soft-rayed fishes. 1898, Jordan and Evermann correctly outlined by Smith and Stearley Several decades later, with the ex- assigned the name clarki (Richardson in the accompanying article. After ploration of the lands bordering the 1836) to the cutthroat trout when they examining the evidence, the North Pacific, many new trouts and discovered that it ranged only to Committee voted unanimously salmons were discovered. Most of Alaska, not to the Bering Sea, and to accept Oncorhynchus as the these species (e.g., mykiss, tshawyts- that mykiss from Kamchatka lack or- proper generic name for these cha, gorbuscha, nerka, keta, kisutch) were ange on the throat ordan and Ev- fish, leaving Salmo as the genus described in Latin by Johann Wal- ermann 1898). But they mistakenly of trouts (including the Atlantic baum (1792) on the basis of speci- concluded that mykiss was a relative salmon) native to Europe, west- mens collected by Thomas Pennant's of Salmo salar because their specimen emn Asia, and the Atlantic basin. expedition in Kamchatka. In 1836, had larger scales (125 lateral series) A change in the specific name of John Richardson described most of than the steelhead, gairdneri. rainbow trout from gairdner to them again (e.g., gairdneri, quinnat, Robert Behnke (1966) corrected the mykiss has been advocated on scouleri, paucidens, consuetus, tsup- misunderstanding of scale size and good evidence by ichthyologists pitch) on the basis of specimens from provided convincing morphological for several years, and the Com- northwestern North America. (Wal- evidence that the Kamchatka trout mittee had already accepted it. baum included the new trouts, salm- and the rainbow trout were similar. The Committee has not en- ons, and chars in Linnaeus' genus Okazaki (1984) added data from 37 dorsed these changes lightly, for Salmo. Richardson did the same, ex- electrophoretic loci and demon- it has an overriding interest in cept that he separated chars out as strated beyond doubt that coastal the stability of fish names. We the genus Salvelinus.) steelheads from western North consider the evidence supporting Most of the species currently rec- America are genetically more similar the changes too strong to dis- ognized were captured repeatedly by to Kamchatka trout (genetic distance pute. Only the scientific names explorers on both sides of the Pacific - .0026) than to inland gairdneri (ge- are affected; the common names and were named again by naturalists netic distance = .0073). Okazaki long recommended by the Com- attempting to describe the spectac- summarized all of the available evi- mittee still remain unchanged. ular diveristy (e.g., Suckley 1860, dence, including the identity of ver- The new nomenclature will be 1862). Why were the new discov- tebral counts (Savvaitova 1975) and reflected in two of the Commit- eries so often thought to represent chromosome data (Vasil'yev 1975; tee's forthcoming books: "World new species? Partly because they Thorgard 1977) and concluded that List of Fishes ... of Interest to sometimes were, partly because the the rainbow trout and the Kam- North Americans," scheduled for species concept was less inclusive chatka trout are the same biological publication in 1989, and "List of then, but mostly because the nu- species, for which the oldest avail- Common and Scientific Names merous species of trout and salmon able name is mykiss (which is Wal- of Fishes from the United States show remarkable variation in sexual baum's spelling for an indigenous and Canada," 5th edition, 1990. dimorphism, breeding shapes, colors, Kamchatkan name for the rainbow C. Richard Robins, Chair and life history. trout). Committee on Names of Fishes Walbaum's, Richardson's, and The facts are solid and the choice Suckley's names for the same spe- seems unambiguous. Although North has established a stable set of ver- American biologists could appeal to cies, above, were mostly sorted out nacular names to assist unambigu- the International Commission on Zo- by Jordan and Gilbert (1883).