THE EXOTIC ICHTHYOFAUNA OF THE CONTIGUOUS UNITED STATES WITH PRELIMINARY OBSERVATIONS ON INTRANATIONAL TRANSPLANTS

W.R. Courtenay, Jr. and J.N. Taylor
Department of Biological Sciences, Florida Atlantic University
Boca Raton, Florida, U.S.A.

ABSTRACT

We record 39 species of exotic (foreign) fishes as established in open waters of the 48 contiguous United States. Included are 13 cichlids, 7 cyprinids, 7 poeciliids, 3 loricariids, 2 gobiids, 2 sciaenids and one species each for the families Anabantidae, Clariidae, Cobitidae, Osmeridae and Salmonidae. All but nine of these fishes were introduced since the second world war. Only the brown trout (Salmo trutta), bairdiella (Bairdiella icistia) and orangemouth corvina (Cynoscion xanthulus) are generally accepted as having been beneficial introductions.

In addition, we record another 57 non-established exotic fishes known from U.S. waters. Fourteen of these were formerly established and another nine were intentionally released but failed to establish.

A preliminary review indicates that at least 168 native fishes have been transplanted beyond their natural ranges, with the majority appearing to have been moved as game and released bait.

RESUME

Trente-neuf espèces de poissons exotiques sont établies dans les eaux libres des 48 Etats contigus des Etats-Unis: treize cichlidés, sept cyprinidés et sept poécillidés, trois loricariidés, deux gobies et deux sciénidés et une espèce de chacune des familles suivantes: Anabantidae, Clariidae, Cobitidae, Osmeridae et Salmonidae. Tous ces poissons, à l'exception de neuf, ont été introduits après la deuxième guerre mondiale. Seule la truite (Salmo trutta) est en général considérée comme une introduction bénéfique par les aménagistes et les biologistes des pêches.

Quinze espèces de poissons exotiques ont des populations en expansion aux Etats-Unis, quatorze ont une répartition localisée, trois ont une aire de répartition assez vaste et plus ou moins stabilisée, quatre semblent avoir des populations en diminution; enfin, on dispose de trop peu d'informations pour pouvoir juger de la situation de trois autres espèces.

1. INTRODUCTION

The continental United States now hosts 39 species of exotic fishes, established as reproducing populations. Most are aquarium fishes.

Courtenay and Robins (1973, 1975) and Courtenay et al. (1974) discussed several reasons for and sources of introductions of foreign (exotic) fishes in the U.S. These included releases for establishing sport or food fishes; biological control of one or more pest aquatic plants (many of which are also exotic); and releases from culture facilities and by hobbyists. Other introductions have been made for purposes of aquaculture, insect control and forage. Two species have become established as a result of accidental transoceanic movement.

In this paper, we summarize the status of established exotic fishes in the continental U.S., in phylogenetic order after Robins et al. (1980), in the categories listed above. A list of non-established exotic fishes known from U.S. waters is included (Table 1) and preliminary observations on intranational transplants of native fishes are presented.

2. SPORT AND FOOD FISHES

Only three exotic fishes are regarded as beneficial introductions in this category: the brown trout (Salmo trutta, section 2.1), bairdiella (Bairdiella icistia, section 2.6) and orangemouth corvina (Cynoscion xanthulus, section 2.7).

2.1 Salmo trutta Linnaeus (brown trout)

First introduced in the lower peninsula of Michigan in 1883 (Mather, 1889; Goode, 1903; Laycock, 1966), this salmonid is now established in suitable waters in Arizona, Arkansas, California, Colorado, Connecticut, Delaware, Georgia, Idaho, Illinois, Indiana, Iowa, Maine, Maryland, Massachusetts, Michigan, Minnesota, Missouri, Montana, Nebraska, Nevada, New Hampshire, New Jersey, New Mexico, New York, North Carolina, North Dakota, Oregon, Pennsylvania, Rhode Island, South Carolina, South Dakota, Tennessee, Utah, Vermont, Virginia, Wisconsin and Wyoming (MacCrimmon and Marshall, 1968). Kentucky recently (1980) reactivated stocking toward establishing self-sustaining populations. As of 1980, only Arkansas, Illinois and Montana had discontinued annual stocking of this fish. Illinois indicated that stocking would be reactivated upon completion of a new hatchery.

The brown trout is one of three exotic fishes in the U.S. that is generally regarded as beneficial. Although reported as more difficult to catch than native trouts, it remains popular with anglers. Brown trout often occupy waters not inhabited by native trouts that prefer cooler waters. Nevertheless, there have been problems. Brown trout have adversely impacted, by predation, the golden trout (Salmo aguabonita), the “state fish” of California, in the Little Kern River, and a brown trout eradication programme is underway there (E.P. Pister, pers. comm.). The National Park Service is also eradicating brown trout from the Great Smoky Mountain National Park because of its negative impact, again by predation, on native brook trout (Salvelinus fontinalis; Reiger, 1981).

2.2 Cyprinus carpio Linnaeus (common carp)

DeKay (1842) records the first release of this cyprinid fish in the Hudson River, New York, in 1831. Self-sustaining populations now exist in the 48 contiguous states, mostly from federal stockings conducted in the late 1800s (Baird, 1879; Laycock, 1966).

Population densities are greatest in the midwestern states but only scattered populations exist in portions of the extreme southeastern U.S. In Florida, for example, this species occurs in only two rivers in the panhandle.

Introduced as a popular food fish from Europe for aquaculture and releases in natural waters, this species has never become a popular food or sport fish in the U.S. It is considered a pest fish in nearly every state where it is established. It is claimed to have disturbed waters by its feeding habits, thereby displacing preferred native fishes and damaging waterfowl habitat (Laycock, 1966). As of 1980, 15 states conducted annual common carp control or eradication programmes at a total cost of over U.S. $ 300 000.

2.3 Leuciscus idus (Linnaeus) (ide)

Now restricted to one pond in Connecticut (Whitworth et al., 1968), the ide was formerly established in New York (Bean, 1901), Pennsylvania (E.C. Cooper, pers. comm.) and the Potomac River adjoining the District of Columbia, Maryland, and Virginia (Smith and Bean, 1899; Bean and Fowler, 1929; Schwartz, 1963; Musick, 1972). Reasons for its decline and apparent absence in New York and the upper Potomac drainage are unknown; its disappearance from two sites in Pennsylvania was due to eradication and the filling of a pond for highway construction. The ide was imported for distribution (apparently with common carp) in 1877, probably as a food and ornamental (aquarium) species.

2.4 Scardinius erythrophthalmus (Linnaeus) (rudd)

Due to its popularity as a sport and food species in Europe, the rudd was also imported in the late 1870s. It was introduced and became established in New Jersey (Myers, 1925), New York (Myers, 1925; Greeley, 1937) and Wisconsin (Cahn, 1927; Greene, 1935). C.R. Robins (pers. comm.) reported this fish as present in Cascadilla Creek near the Cornell University hatchery ponds in New York in the fifties.

To our knowledge, the rudd is no longer extant in New Jersey or Wisconsin; reasons for its decline and disappearance there are unknown. In New York, it now appears restricted to two tributaries of the Hudson River.

2.5 Tinca tinca (Linnaeus) (tench)

Imported in 1877 with common carp, the tench was distributed to many states. It became established in California (Moyle, 1976), Colorado (Beckman, 1974), Connecticut (Whitworth et al., 1968), Delaware (Schwartz, 1963), Idaho (Simpson and Wallace, 1978), Maryland (Schwartz, 1963), New Mexico (Koster, 1957) and Washington (Wydoski and Whitman, 1979). It was introduced to Arizona (Minckley, 1973) and Missouri (Baughman, 1947) but failed to establish.

The tench apparently is now absent from New York (although it was collected from Cascadilla Creek, Ithaca, in the fifties; C.R. Robins, pers. comm.), New Mexico and Oregon (E.C. Raney, J.E. Johnson and C.L. Bond, pers. comm.) and probably also from Delaware and Maryland. There are no recent records from those states.

2.6 Bairdiella icistia (Jordan and Gilbert) (bairdiella)

The Salton Sea in southern California was created in 1905–7 by a closure of the lower Colorado River through sediment deposition, blocking its natural flow path to the Gulf of California. The river took the path of least resistance and invaded an almost-completed irrigation canal into the Salton Sink, about 82 m below sea level (Sykes, 1937). Freshwater fishes of the lower Colorado River were washed into the sink with the creation of this inland sea. In subsequent years, the salinity rose in the Salton Sea, extirpating the freshwater species. By 1956, the salinity had risen to 33‰ (Carpelan, 1961). Many introductions were made as the salinity rose and most failed (Walker et al., 1961).

In the early fifties it was decided to introduce marine species from the Pacific Ocean, collected from the Mexican coast. Among these were two sciaenids, the bairdiella, orangemouth corvina (Cynoscion xanthulus) and a haemulid, the sargo (Anisotremus davidsoni). The sargo is native in marine waters of extreme southern California (Walker et al., 1961). These fishes adapted and became established, along with some marine plants and invertebrates that became established before these fishes were introduced.

The bairdiella has been a popular angling species since its introduction. It has also provided forage for the orangemouth corvina. There is some evidence, however, that bairdiella populations have been adversely impacted by introductions of tilapia in the late seventies (G.L. Black and F.G. Hoover, pers. comm., see section 3.4).

2.7 Cynoscion xanthulus Jordan and Gilbert (orangemouth corvina)

The orangemouth corvina is the most popular angling species in the Salton Sea. It is common for large adults to weigh 11 kg or more (Walker et al., 1961). See account 2.6 for its history in the Salton Sea.

3. BIOLOGICAL CONTROL

Fishes in this category were released for control of pest aquatic plants or insects. Evaluation studies prior to releases were conducted primarily by agronomists, entomologists or plant pathologists and plant physiologists; these evaluations, however, were target-oriented (= pest control). Only within the latter part of the past decade have studies of impact on native fishes or their habitats been included in several studies. Many introductions were conducted without prior testing; we consider that fisheries “mis-management”.

In the U.S. the use of exotic fishes as biological control agents remains controversial. Fishes are not monophagous as are several insects (Courtenay, 1979a). Preferential herbivores in test situations have performed differently after release. Thus, non-target aquatic plants that shelter the young of native fishes or provide substrate for their food organisms often become food of the exotic used for biological control. If a fish introduced as a biological control reproduces in great numbers, it becomes a biological control “out of control”.

3.1 Ctenopharyngodon idella (Valenciennes) (grass carp)

Imported in 1963 from stocks in Malaysia and Taiwan (Guillory and Gasaway, 1978), grass carp were subsequently distributed to research agencies and companies with research capabilities in 11 states (Provine, 1975). The first release of this cyprinid was in Arkansas, probably in the late sixties. The first specimens caught in the wild were from the White River, Arkansas, in 1970 (Bailey, 1972); others were subsequently caught in the Mississippi River in Illinois in 1971 (Greenfield, 1973). Those specimens were of the 1966 age-class, suggesting the source of release to have been either the U.S. Fish and Wildlife Service Fish Farming Experimental Station at Stuttgart, Arkansas, or the Arkansas Game and Fish Commission hatchery at Lonoke (Guillory and Gasaway, 1978) or both. This species has been reported as established in the Mississippi River drainage since 1975 (Connor et al., 1980).

3.2 Tilapia aurea (Steindachner) (blue tilapia)

We include this cichlid in the biological control category because two, possibly three, states (Arizona, Florida and Georgia) introduced it, primarily for algal or other aquatic plant control. There is some question as to the identity of the tilapia in Georgia. The blue tilapia was imported from Israel via the late Dr. Earl Herald of the Steinhart Aquarium by Auburn University in 1954 for studies on its potential in aquaculture and as a sport fish (Swingle, 1960). In 1961, Florida introduced stocks from Auburn in an experimental area to examine the ability of blue tilapia to control aquatic vegetation; this introduction apparently was made under the guise of testing its sport potential, already disproven at Auburn (Swingle, 1960). Fishermen subsequently moved specimens out of the experimental area, and this species is now established in 18 counties of peninsular Florida. There is some evidence that since 1972, introductions of this fish have been made by commercial fishermen for the purpose of creating new fisheries (Harris, 1978). Commercial fishing for blue tilapia was legalized in 1972 in Florida.

Arizona has introduced this fish since 1975 for algal control (W.L. Minckley, pers. comm.). It is now dominant in the lower Colorado River near Yuma, where it appears to have replaced most of an earlier-introduced, largely Mississippian fish fauna. We assume its reported introduction into golf course ponds at Sea Island, Georgia, was for aquatic vegetation control.

The blue tilapia is also established in thermal springs in Colorado (for aquaculture), in one heated lake in North Carolina (introduced as a sport fish), a heated effluent in an Oklahoma river (for aquaculture; Pigg, 1978) and in heated reservoirs and the Rio Grande in Texas (for aquaculture, “inadvertently released”, and bait releases; Noble et al., 1976; C. Hubbs, pers. comm.).

3.3 Tilapia hornorum Trewavas (Wami tilapia)

The Wami tilapia was introduced into three counties in southern California for aquatic plant, mosquito and chironomid midge control in the mid seventies (Hauser et al., 1976; Legner and Pelsue, 1977; Legner, 1979; Legner et al., 1980). It was reported as introduced with Mozambique tilapia (Tilapia mossambica) and hybrids of the Mozambique and Wami tilapias.

3.4 Tilapia mossambica (Peters) (Mozambique tilapia)

This fish is established in Arizona, California, Florida and Texas. It was purposely introduced in Arizona for aquatic plant control (Mickley, 1973) and in southern California for the same reason (see section 3.3). It is also reported as established in heated effluents in raceways on a commercial catfish farm in Colorado where it is used for algal control (R.J. Behnke, pers. comm.). It became established at two localities in Florida after escaping from aquarium fish farms and by releases by hobbyists, and in a river in Texas after escaping from a public aquarium (Brown, 1961).

This species, or a hybrid between the Mozambique tilapia and an unknown congener, invaded the Salton Sea in southern California after 1976 and has become the dominant fish in this saline lake (G.L. Black, F.G. Hoover and J.A. St. Amant, pers. comm.). It, or a hybrid with the Wami tilapia, is now dominant in the San Gabriel River near Los Angeles, California (Knaggs, 1977).

3.5 Tilapia zilli (Gervais) (redbelly tilapia)

The redbelly tilapia is established near Phoenix, Arizona, and perhaps elsewhere in that state (Minckley, 1973), in irrigation canals in three southern California valleys (Pelzman, 1973; Moyle, 1976) and in a golf course pond in a valley west of Las Vegas, Nevada (Courtenay and Deacon, 1982). C. Hubbs (pers. comm.) has reported its possible establishment in southern Texas. To our knowledge, all of these populations were introduced for aquatic vegetation control. Those released into some California waters were also stocked for mosquito and chironomid midge control (Legner and Pelsue, 1977). This species is reported as being cultured in warm springs in Idaho (for aquaculture) and in heated effluents in North and South Carolina (for vegetation control and aquaculture).

4. AQUACULTURE

The utilization of exotic fishes for aquaculture in the U.S. was, in part, a reason for importing common carp in the 1870s (see section 2.2). Since the second world war, several tilapias have become aquaculture subjects in warm climates or where thermal springs or warm effluents are available (see sections 3.2 and 3.5). Except in sub-tropical or extreme warm temperate zones, survival of tilapias escaping from culture facilities is unlikely. In several localities where climate has permitted establishment, population explosions of tilapia have been accompanied with moderate to dramatic declines of native fishes, in some cases including transplanted native species. When blue tilapia populations reached 1 842 kg/ha in one Texas reservoir, most native fishes ceased reproducing (Noble et al., 1976; Noble, 1977).

5. FORAGE

To our knowledge, only one exotic fish has been introduced in the U.S. as a forage species.

5.1 Hypomesus nipponensis McAllister (wakasagi)

This osmerid was released into six reservoirs in California as forage for trout (Wales, 1962; Moyle, 1976). Moyle (1976) mentioned subsequent releases into other California reservoirs for the same purpose. The current status of this fish in California is unknown.

6. AQUARIUM FISHES

Aquarium fishes are very popular in the U.S. Most of our established exotic aquarium fishes were unintentionally (rarely intentionally) released from culture facilities (Courtenay and Robins, 1973, 1975; Courtenay et al., 1974; Courtenay and Hensley, 1980). Most such introductions have occurred in states within a sub-tropical climatic zone (e.g., southern Florida where 80 percent of the U.S. aquarium fishes are cultured and southern California). Areas where most of these releases resulted in establishment contain disturbed habitats and ecosystems, a combination that has facilitated establishment.

Hobbyists have been responsible for releasing aquarium fishes. Several of these releases have become established in canal systems in and around metropolitan areas. Other hobbyist releases have occurred in thermal springs in northwestern states and in remote desert ecosystems in the American southwest where they have shown adverse impacts on endemic, often endangered native fishes (Miller and Alcorn, 1946; Miller, 1961; La Rivers, 1962; Hubbs and Brodrick, 1963; Deacon et al., 1964; Hubbs and Deacon, 1965; Minckley, 1973; Courtenay et al., 1974; Deacon, 1979; Hardy, 1980; Courtenay and Deacon, 1982, 1983). No introduced aquarium fish has been demonstrated to have beneficial value in open waters of the U.S. We regard their introduction as needless but to be expected. These stock additions cannot be considered stock enhancement.

6.1 Carassius auratus (Linnaeus) (goldfish)

Introduced in the late 1600s (DeKay, 1842), this cyprinid was the first exotic fish to be released and become established in North America (Courtenay and Hensley, 1980). Self-sustaining populations exist at scattered locations in a majority of states.

6.2 Leuciscus idus (Linnaeus) (ide)

See section 2.3.

6.3 Rhodeus sericeus (Pallas) (bitterling)

First noted in the Sawmill River north of New York City in the twenties (Dence, 1925; Myers, 1925; Bade, 1926), the bitterling also became established in the adjacent Bronx River (Greeley, 1937). It is now apparently confined to one small area of the Bronx River where it is expected to be extirpated by industrial pollution through demise of its molluscan host (Schmidt et al., 1981).

6.4 Misgurnus anguillicaudatus (Cantor) (oriental weatherfish)

This cobitid is established in one river in the southeastern portion of the southern peninsula of Michigan (Schultz, 1960; M.L. Smith, pers. comm.) and some flood control channels in southern California (St. Amant and Hoover, 1969; M.H. Horn, pers. comm.).

6.5 Clarias batrachus (Linnaeus) (walking catfish)

After escaping from a fish farm in southeastern Florida in the mid sixties, followed by subsequent intentional releases in the Tampa Bay area in 1968 (Courtenay and Miley, 1975; Courtenay, 1978, 1979), this fish has spread over 20 counties in peninsular Florida (Courtenay, 1979). There have been large build-ups of local populations periodically. This fish has become a problem to aquarium fish farmers in recent years through predation on their culture fishes (Courtenay and Miley, 1975).

6.6 Hypostomus spp. (suckermouth catfishes)

At least three morphologically distinct but as yet unidentified species of this loricariid genus are established in the U.S.: one in western peninsular Florida and perhaps a second near Miami (Courtenay et al., 1974); another in a thermal spring in southern Nevada (Minckley, 1973; Courtenay and Deacon, 1982); and the third in southern Texas (Barron, 1964; Hubbs et al., 1978).

6.7 Belonesox belizanus Kner (pike killifish)

This poeciliid has been established south of Miami, Florida, since November 1957 (Belshe, 1961; Rivas, 1965; Lachner et al., 1970; Miley, 1978).

6.8 Poecilia mexicana Steindachner (shortfin molly)

The shortfin molly is established in southern California (St. Amant, 1966; St. Amant and Sharp, 1971; Mearns, 1975; Hubbs et al., 1978), in thermal springs in Montana (Brown, 1971) and Nevada (Deacon et al., 1964; Hubbs and Deacon, 1965; Courtenay and Deacon, 1982, 1983).

6.9 Poecilia reticulata Peters (guppy)

Populations of the guppy are established in thermal springs in Arizona (Minckley, 1973), Florida (F.W. King, pers. comm.), Idaho (Simpson and Wallace, 1978), Nevada (Deacon et al., 1964; Williams et al., 1980; Courtenay and Deacon, 1982, 1983; Texas (Hubbs et al., 1977; Hubbs, 1982) and Wyoming (Baxter and Simon, 1970). It may be established locally in sewage treatment ponds in California (Moyle, 1976).

6.10 Poeciliopsis gracilis (Heckel) (porthole livebearer)

This fish has been established since at least 1965 in a canal north of the Salton Sea, southern California (Mearns, 1975; Moyle, 1976; Hubbs et al., 1979; Shapovalov et al., 1981).

6.11 Xiphophorus helleri Heckel (green swordtail)

The green swordtail is established near Tampa Bay (Courtenay et al., 1974) and at Satellite Beach, south of Cape Canaveral, Florida (Dial and Wainright, 1983) and in a thermal spring in Montane (Brown, 1971). A hybrid of this species and the southern platyfish (Xiphophorus maculatus) is established in a thermal spring in southern Nevada (Courtenay and Deacon, 1982).

6.12 Xiphophorus maculatus (Günther) (southern platyfish)

The southern platyfish is established in two localities (Gulf and Atlantic drainages) in peninsular Florida (Courtenay et al., 1974; Dial and Wainright, 1983). A hybrid with the green swordtail is established in a southern Nevada spring.

6.13 Xiphophorus variatus (Meek) (variable platyfish)

The variable platyfish is established in Gainesville (Burgess et al., 1977) and perhaps elsewhere in Florida.

6.14 Astronotus ocellatus (Agassiz) (oscar)

Introduced in the late fifties, this cichlid is established in four counties of southern Florida (Rivas, 1965; Lachner et al., 1970; Courtenay and Hensley, 1980). This species has been accepted locally in Florida by some anglers, although it is never abundant.

6.15 Cichlasoma bimaculatum (Linnaeus) (black acara)

Established since the late fifties or early sixties, this fish is now dominant in many south Florida canals (Courtenay et al., 1974) and recently has entered Lake Okeechobee.

6.16 Cichlasoma citrinellum (Günther) (Midas cichlid)

The presence of the Midas cichlid in southeastern Florida was discovered by personnel of the Florida Game and Freshwater Fish Commission in 1981 (Shafland, in press). It is restricted to some canals south of Miami and apparently is expanding its range.

6.17 Cichlasoma meeki (Brind) (firemouth)

The firemouth is established in the Tamiami Canal system, Miami, Florida (Courtenay and Hensley, 1980; Robins et al., 1980).

6.18 Cichlasoma nigrofasciatum (Günther) (convict cichlid)

This exotic has been established in two thermal spring systems in southern Nevada since the early sixties (Deacon et al., 1964; Hubbs and Deacon, 1965; Courtenay and Deacon, 1982, 1983).

6.19 Cichlasoma octofasciatum (Regan) (Jack Dempsey)

The Jack Dempsey is established in three localities in east-central and southern peninsular Florida (Courtenay et al., 1974; Hogg, 1976, 1976a; Dial and Wainright, 1983).

6.20 Hemichromis bimaculatus Gill (jewelfish)

The jewelfish has been established since the early sixties in canals around Miami International Airport, Florida (Rivas, 1965; Courtenay et al., 1974).

6.21 Tilapia mariae (Boulenger) (spotted tilapia)

The spotted tilapia is established in three counties in extreme southern Florida (Courtenay and Hensley, 1979) and in a thermal spring in southern Nevada (Courtenay and Deacon, 1982, 1983).

6.22 Tilapia melanotheron (Ruppell) (blackchin tilapia)

The blackchin tilapia has been established since the late fifties or early sixties along the eastern shore of Tampa Bay, Florida (Springer and Finucane, 1963; Finucane and Rinckey, 1965). It has become established recently south of Cape Canaveral on Florida's east coast (Dial and Wainright, 1983). This species appears in the marine commercial catch where it has become established.

6.23 Tilapia mossambica (Peters) (Mozambique tilapia)

See section 3.4.

6.24 Trichopsis vittata (Kuhl and Van Hasselt) (croaking gourami)

This anabantid is established in a weedy area on one side of a Palm Beach County drainage canal, southeastern Florida (Hensley and Courtenay, 1980).

7. EXOTIC FISHES FROM OTHER SOURCES

7.1 Acanthogobius flavimanus (Temminck and Schlegel) (yellowfin goby)

Ballast pumped from transoceanic ships entering the San Francisco Bay area, California, appears to have been the source of this introduction (Brittan et al., 1963; Brittan et al., 1970). The U.S. range of the yellowfin goby has expanded greatly since its introduction (Kukowski, 1972; Miller and Lea, 1972; Haacker, 1979; Usui, 1981); it has been reported as far south as San Diego in 1980 (C. Usui, pers. comm.).

7.2 Tridentiger trigonocephalus (Gill) (chameleon goby)

The occurrence of this fish in California waters was first noted by Hubbs and Miller (1965). They theorized that its initial introduction into San Francisco Bay may have been as fertilized eggs on the introduced Japanese oyster (Crassostrea gigas). It is now established in San Francisco Bay and a fresh to brackish water lake in Oakland (Shapovalov et al., 1980) and in Los Angeles harbour (Moyle, 1976; Haacker, 1979).

8. PRELIMINARY OBSERVATIONS CONCERNING INTRANATIONAL TRANSPLANTS

8.1 Primary source of information

Lee et al. (1980) included distributional maps and information for 743 native fishes known to inhabit the fresh waters of Canada and the U.S. Our review of the data contained therein shows that at least 151 species have been transplanted via human activities beyond their natural ranges of distribution in the continental U.S. New information supplied by W.L. Minckley and J.R. Stauffer, Jr. indicates yet another 17 transplanted fishes.

The exact natural ranges for several of these fishes will never be known. Early surveys were incomplete and, in some instances, drainage basins were altered or disturbed prior to thorough surveys. Moreover, what appear to be transplants may be relict populations or transfers via stream capture for a few species.

The species accounts in Lee et al. (1980) typically and unfortunately did not indicate the reasons for these transplants. It is known that many fishes of the families Salmonidae, Esocidae, Catostomidae, Ictaluridae, Percichthyidae and Centrarchidae were transplanted for game (sport and food) purposes. Small fishes, particularly cyprinids and percids, were released from bait buckets.

8.2 Summary regarding transplants

Table 2 is a list of U.S. fishes transplanted beyond their natural ranges through introductions, a few via manmade canals or river diversions or as results of other human activities. For many species, we used a “best guess” as to the reason for the transplant and those are indicated by a question mark (?) in Table 2.

A major complicating factor in attempting to ascertain reasons and distributions for certain transplants has been the movement of mixed fishes. This has occurred when portions of lakes, rivers or streams were seined for the purpose of transplanting one or more target game fishes or salvaging fishes from drying habitats. Non-target fishes (we call these incidentals) were often transplanted in this manner in unknown and unreported numbers. Some (perhaps many) species we listed as bait fish releases may be incidentals.

Locally-seined bait fishes often are of several species; these, like commercially-sold bait, are often transported to other drainages for use. Even commercially-sold bait fishes, shipped from many hundreds of kilometres away to bait sellers, may contain mixed species. Some of these lots occasionally contain fishes belonging to several families, sometimes including juvenile centrarchids. Therefore, some of the smaller centrarchids listed in Table 2 as introduced as game fishes (the expected reason), in some instances, may represent bait releases. Many perhaps were also introduced as forage species.

Our preliminary analysis indicates the overwhelming importance of sport fishing as a major factor in transplants in the U.S. with 53 species probably moved as game fishes and another 58 probably released from bait buckets.

We identify 11 that were moved beyond their native ranges for species survival purposes (i.e., to prevent their immediate or expected future extirpation). For some, their native habitat subsequently was destroyed or so altered that extirpation without transplantation would have been certain; for others, survival continues within the native habitat or range and these fishes remain listed as threatened or endangered.

Native fishes have never matched the popularity of exotic fishes as aquarium or ornamental species, but as many as three transplants may have been from releases of native ornamental fishes. Two species were introduced to expand their ranges. Other transplants were made for the probable reasons listed in Table 2.

We again emphasize that these data are preliminary and will require extensive analysis of published state fish guides, as well as federal and state introduction records, and consultation with specialists for further refinement. It would be impossible to compile a truly accurate documentation because many agency records are incomplete, sometimes inaccurate, or perhaps no longer extant in some cases. Nevertheless, our summary shows that transplants of native fishes, planned or otherwise, far outnumber introductions of exotic fishes in the U.S. Perhaps this reflects an opinion of past and present fishery managers that moving native fishes is somehow safer than introducing exotics.

8.3 Environmental concerns

For intentional transplants, we know of no studies prior to release on potential impacts on non-game fishes or habitat. With the exception of a few studies beginning in the mid-to-late seventies on potential impacts of grass carp, we find few earlier studies on potential impacts of exotic fishes. Nearly all previous intentional introductions seem goal-oriented only. Those persons emptying bait buckets have no environmental concerns for that activity beyond perhaps a perception that the released bait may provide forage for game species.

An introduction is an introduction, whether the source is intranational, intracontinental or foreign. Impacts, ranging from negligible to major, are to be expected.

Fishery managers and biologists are under continual public or political pressure to improve or enhance sport fisheries and to correct problems in aquatic ecosystems. Since introductions long have served as a management tool, they have all too often been accepted, mostly without question, as potential “cure-alls”. For example, fishes are often introduced to fill a so-called “vacant niche”; in fishery management terms, this means a perceived vacant trophic level. Thus, fishes may be released to control phytoplankton blooms; their high reproductive success may then lead to the introduction of one or more predatory fishes, usually game species, to control the phytoplankton consumers, now regarded as forage. The predators may control this forage or eliminate it. There has been at least one instance where two predatory game fishes were introduced to reduce such a forage base; the forage was destroyed, one predator eliminated the other (Allen and Roden, 1978), and then its population collapsed. Manipulating open waters in such a manner is unwarranted, needless and costly. Open waters, including reservoirs, are not farm or culture ponds and deserve better management policies.

Proper fishery management has to (and must) be based on the best available knowledge of aquatic ecology, fishery biology and ichthyology, and not on the “introduce anything” psychology that has developed over the past century. Introductions remain an extremely viable tool in fishery management of open waters when conducted properly.

9. ACKNOWLEDGEMENTS

We thank J.A. McCann and J.R. Stauffer, Jr. for their helpful comments on the manuscript and C.R. Gilbert, W.L. Minckley, C.R. Robins and J.R. Stauffer, Jr. for their input on our preliminary review of transplants.

This effort was largely supported by Contract 14-16-0009-78-021 and Cooperative Agreement 14-16-0009-80-952 from the Department of the Interior, U.S. Fish and Wildlife Service, through the National Fishery Research Laboratory, Gainesville, Florida.

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Table 1 A list of exotic fishes known from waters of the United States

* Unconfirmed report

Table 2 A preliminary list of transplanted native fishes within the contiguous United States, including the likely reason for release

FAMILY	SPECIES	COMMON NAME	REASON
Petromyzontidae	Petromyzon marinus Linnaeus	sea lamprey	accidental via canal
Acipenseridae	Acipenser transmontanus Richardson	white sturgeon	game
Amiidae	Amia calva Linnaeus	bowfin	incidental?
Anguillidae	Anguilla rostrata (Lesueur)	American eel	commercial?
Clupeidae	Alosa aestivalis (Mitchill)	blueback herring	forage?
	A. pseudoharengus (Wilson)	alewife	forage
	A. sapidissima (Wilson)	American shad	commercial
	Dorosoma cepedianum (Lesueur)	gizzard shad	forage
	D. petenense (Günther)	threadfin shad	forage
Salmonidae	Coregonus artedii Lesueur	cisco/lake herring	game?
	C. clupeaformis (Mitchill)	lake white fish	commercial
	Oncorhynchus gorbuscha (Walbaum)	pink salmon	game
	O. keta (Walbaum)	chum salmon	game
	O. kisutch (Walbaum)	coho salmon	game
	O. nerka (Walbaum)	sockeye salmon	game; forage
	O. tshawytscha (Walbaum)	chinook salmon	game
	Salmo aguabonita Jordan	golden trout	game
	S. apache Miller	Apache trout	species survival?
	S. clarki Richardson	cutthroat trout	game
	S. gairdneri Richardson	rainbow trout	game
	S. gilae Miller	Gila trout	species survival
	S. salar Linnaeus	Atlantic salmon	game
	Salvelinus alpinus (Linnaeus)	Arctic char	game
	S. fontinalis (Mitchill)	brook trout	game
	S. namaycush (Walbaum)	lake trout	game
	Thymallus arcticus (Pallas)	Arctic grayling	game
Osmeridae	Osmerus mordax (Mitchill)	rainbow smelt	commercial; forage?
Umbridae	Dallia pectoralis Bean	Alaska blackfish	bait release?
Esocidae	Esox americanus americanus Gmelin	redfin pickerel	game? accidental?
	E. americanus vermiculatus Lesueur	grass pickerel	game? accidental?
	E. lucius Linnaeus	northern pike	game; forage control
	E. masquinongy Mitchill	muskellunge	game
	E. niger Lesueur	chain pickerel	game
Characidae	Astyanax mexicanus (Filippi)	Mexican tetra	bait; ornamental fish release
Cyprinidae	Agosia chrysogaster Girard	longfin dace	bait release; accidental
	Campostoma anomalum (Rafinesque)	central stoneroller	bait release?
	Clinostomus funduloides Girard	rosyside dace	bait release?
	Gila atraria (Girard)	Utah chub	bait release
	G. bicolor (Girard)	tui chub	unknown
	G. coerulea (Girard)	blue chub	unknown
	G. copei (Jordan & Gilbert)	leatherside chub	bait release
	G. orcutti (Eigenmann & Eigenmann)	arroyo chub	bait release?
	G. pandora (Cope)	Rio Grande chub	bait release?
	G. purpurea (Girard)	Yaqui chub	species survival
	Lepidomeda mollispinis Miller	Virgin spinedace	accidental
	Nocomis micropogon (Cope)	river chub	bait release?
	Notemigonus crysoleucas (Mitchill)	golden shiner	bait; ornamental fish release
	Notropis albeolus Jordan*	white shiner	bait release
	N.baileyi Suttkus & Raney	rough shiner	bait release?
	N. buccula Cross	smalleye shiner	bait release?
	N. cerasinus (Cope)*	crescent shiner	bait release?
	N. chiliticus (Cope)*	redlip shiner	bait release?
	N. chrosomus (Jordan)	rainbow shiner	bait release?
	N. coccogenis (Cope)	warpaint shiner	bait release?
	N. cornutus (Mitchill)*	common shiner	bait release?
	N. galacturus (Cope)*	whitetail shiner	bait release?
	N. gibbsi Howell & Williams	Tallapoosa shiner	bait release?
	N. girardi (Hubbs & Ortenburger)	Arkansas River shiner	bait release?
	N. heterodon (Cope)	blackchin shiner	bait release?
	N. heterolepis (Eigenmann & Eigenmann)	blacknose shiner	bait release?
	N. leuciodus (Cope)*	Tennessee shiner	bait release?
	N. lutrensis (Baird & Girard)	red shiner	bait release
	N. oxyrhynchus Hubbs & Bonham	sharpnose shiner	bait release?
	N. procne (Cope)	swallowtail shiner	bait release
	N. rubricroceus (Cope)	saffron shiner	bait release?
	N. spilopterus (Cope)	spotfin shiner	bait release?
	N. stramineus (Cope)	sand shiner	bait release?
	N. telescopus (Cope)*	telescope shiner	bait release?
	N. volucellus (Cope)	mimic shiner	bait release?
	N. xaenacephalus (Jordan)	Coosa shiner	bait release?
	N. zonistius (Jordan)	bandfin shiner	bait release?
	Orthodon microlepidotus (Ayres)	Sacramento blackfish	forage; food fish
	Phoxinus oreas (Cope)	mountain redbelly dace	bait release?
	Pimephales notatus (Rafinesque)	bluntnose minnow	bait release
	P. promelas Rafinesque	fathead minnow	bait release
	P. vigilax (Baird and Girard)	bullhead minnow	bait release
	Relictus solitarius Hubbs & Miller	relict dace	unknown
	Rhinichthys cataractae (Valenciennes)	longnose dace	bait release?
	Richardsonius balteatus (Richardson)	redside shiner	bait release
Catostomidae	Carpiodes carpio (Rafinesque)	highfin carpsucker	accidental via canal
	Catostomus catostomus (Forster)	longnose sucker	bait release?
	C. fumeiventris Miller	Owens sucker	accidental via aqueduct
	C. plebeius Baird & Girard	Rio Grande sucker	bait release
	C. santaanae (Snyder)	Santa Ana sucker	incidental?
	Ictiobus bubalus (Rafinesque)	smallmouth buffalo	food fish
	I. cyprinellus (Valenciennes)	bigmouth buffalo	food fish; accidental
	I. niger (Rafinesque)	black buffalo	food fish
	Moxostoma cervinum (Cope)	black jumprock	incidental?
	M. erythrurum (Rafinesque)	golden redhorse	bait release?
	M. rhothoecum (Thoburn)	torrent sucker	incidental?
	M. rupiscartes Jordan & Jenkins	striped jumprock	incidental?
Ictaluridae	Ictalurus catus (Linnaeus)	white catfish	game
	I. furcatus (Lesueur)	blue catfish	game
	I. melas (Refinesque)	black bullhead	game
	I. natalis (Lesueur)	yellow bullhead	game
	I. nebulosus (Lesueur)	brown bullhead	game
	I. pricei (Rutter)	Yaqui catfish	game
	I. punctatus (Rafinesque)	channel catfish	game
	Noturus gilberti Jordan & Evermann	orangefin madtom	incidental?
	N. gyrinus (Mitchill)	tadpole madtom	incidental?
	N. insignis (Richardson)	margined madtom	incidental?
	N. phaeus Taylor	brown madtom	accidental via river diversion
	Pylodictus olivaris (Rafinesque)	flathead catfish	game
Amblyopsidae	Chologaster agassizi Putnam	spring cavefish	experimental?
Cyprinodontidae	Crenichthys nevadae Hubbs	Railroad Valley springfish	species survival
	Cyprinodon diabolis Wales	Devils Hole pupfish	species survival
	C. macularius Baird & Girard	desert pupfish	range expansion; species survival
	C. rediosus Miller	Owens pupfish	species survival
	C. rubrofluviatilis Fowler	Red River pupfish	bait release?
	C. salinus Miller	Salt Creek pupfish	range expansion?
	Empetrichthys latos Miller	Pahrump killifish	species survival
	Fundulus heteroclitus (Linnaeus)	mummichog	bait release?
	F. parvipinnis Girard	California killifish	bait release?
	F. stellifer (Jordan)	southern studfish	bait release?
	F. zebrinus Jordan & Gilbert	plains killifish	bait release?
	Jordanella floridae Goode & Bean	flagfish	ornamental/bait release?
	Lucania goodei Jordan	bluefin killifish	bait release?
	L. parva (Baird)	rainwater killifish	from ship ballast; incidental
Poeciliidae	Gambusia affinis(Baird & Girard)	mosquitofish	mosquito control
	G. amistadensis Peden	Amistad gambusia	species survival
	G. gaigei Hubbs	Big Bend gambusia	species survival
	G. geiseri Hubbs & Hubbs	largespring gambusia	unknown
	Poecilia formosa (Girard)	Amazon molly	unknown
	P. latipinna (Lesueur)	sailfin molly	ornamental fish release
	Poeciliopsis occidentalis (Baird & Girard)	Gila topminnow	mosquito control; species survival
Atherinidae	Labidesthes sicculus (Cope)	brook silverside	forage
	Menidia beryllina (Cope)	inland silverside	bait release; pest control; incidental
Gasterosteidae	Apeltes quadracus (Mitchill)	fourspine stickleback	bait release
	Culaea inconstans(Kirtland)	brook stickleback	incidental?
Percichthyidae	Morone chrysops (Rafinesque)	white bass	game
	M. mississippiensis Jordan & Eigenmann	yellow bass	game
	M. saxatilis (Walbaum)	striped bass	game
Centrarchidae	Ambloplites cavifrons Cope	Roanoke bass	game
	A. constellatus Cashner & Suttkus	Ozark bass	game
	A. rupestris (Rafinesque)	rock bass	game; forage
	Archoplites interruptus (Girard)	Sacramento perch	game
	Centrarchus macropterus (Lacépède)	flier	game; forage
	Enneacanthus gloriosus (Holbrook)	bluespotted sunfish	game; forage
	Lepomis auritus (Linnaeus)	redbreast sunfish	game; forage
	L. cyanellus Rafinesque	green sunfish	game; incidental
	L. gibbosus (Linnaeus)	pumpkinseed	game; forage
	L. gulosús (Cuvier)	warmouth	game; forage
	L. humilis (Girard)	orangespotted sunfish	bait release; incidental?
	L. macrochirus Rafinesque	bluegill	game; forage
	L. megalotis (Rafinesque)	longear sunfish	game; forage
	L. microlophus (Günther)	redear sunfish	game; forage
	L. punctatus (Valenciennes)	spotted sunfish	game; forage
	Micropterus coosae Hubbs & Bailey	redeye bass	game
	M. dolomieui Lacépède	smallmouth bass	game
	M. punctulatus (Rafinesque)	spotted bass	game
	M. salmoides (Lacépède)	largemouth bass	game
	M. treculi (Vaillant & Bocourt)	Guadalupe bass	game
	Pomoxis annularis Rafinesque	white crappie	game
	P.nigromaculatus (Lesueur)	black crappie	game
Percidae	Ammocrypta bifascia Williams	Florida sand darter	bait release?
	Etheostoma chlorosomum (Hay)	bluntnose darter	accidental via canal
	E. edwini (Hubbs & Cannon)	brown darter	bait release?
	E. exile (Girard)	Iowa darter	bait release?
	E. fusiforme (Girard)	swamp darter	bait release?
	E. zonale (Cope)	backwater darter	bait release
	Perca flavescens (Mitchill)	yellow perch	game; perhaps forage
	Percina macrolepida Stevenson	bigscale logperch	bait release? incidental
	P. roanoka (Jordan & Jenkins)	Roanoke darter	bait release?
	P. tanasi Etnier	snail darter	species survival
	Stizostedion canadense (Smith)	sauger	game
	S.vitreum (Mitchill)	walleye	game
Haemulidae	Anisotremus davidsoni (Steindachner)	sargo	game
Cichlidae	Cichlasoma cyanoguttatum (Baird & Girard)	Rio Grande cichlid	ornamental fish release
Cottidae	Cottus rhotheus (Smith)	torrent sculpin	incidental

* Upper New River drainage, North Carolina, Virginia and West Virginia; transplanted status questionable

Questionable transplant