extinctions of north american fishes during the past century

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Extinctions of North American Fishes During the pastCenturyRobert R. Miller a , James D. Williams b & Jack E. Williams ca Museum of Zoology, University of Michigan, Ann Arbor, MI, 48109, USAb Fish and Wildlife Service, National Fisheries Research Center, Gainesville, FL, USAc Bureau of Land Management

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To cite this article: Robert R. Miller, James D. Williams & Jack E. Williams (1989): Extinctions of North American Fishes Duringthe past Century, Fisheries, 14:6, 22-38

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Extinctions of North American FishesDuring the Past Century

Robert R. Miller, James D. Williams, and Jack E. Williams

ABSTRACT

Extinctions of 3 genera, 27 species, and 13 subspecies of fishes from North America are documented during the past100 years. Extinctions are recorded from all areas except northern Canada and Alaska. Regions suffering the greatestloss are the Great Lakes, Great Basin, Rio Grande, Valley of Mexico, and Parras Valley in Mexico. More than one factorcontributed to the decline and extinction of 82% of the fishes. Physical habitat alteration was the most frequently citedcausal factor (73%). Detrimental effects of introduced species also were cited in 68% of the extinctions. Chemical habitatalteration (including pollution) and hybridization each were cited in 38% of the extinctions, and overharvesting adverselyaffected 15% of the fishes. This unfortunate and unprecedented rate of loss of the fishery resource is expected toincrease as more of the native fauna of North America becomes endangered or threatened.

RESUMEN

Durante los 100 afios pasados se registraron extinciones de tres generos, 27 especies, y 13 subespecies de peces enNorte America. Extinciones fueron registrados de todas partes fuera de la parte norte de Canada y Alaska. Las regionesque sufrieron las perdidas mas grandes fueron los Lagos Grandes (E.U. y Canada), el Bolson Grande, Rio Bravo, Vallede Mexico, y el Valle de Parras de Mexico. Mas de un factor contribuy6 a 82% de las extinciones. Alteraci6n fisica delhabitat fue el factor causal citada con mayor frecuencia (73%). Efectos dafiosos de introducciones de especies ex6ticastambien fueron citados como factores contribuyentes en una mayoria de las extinciones (68%). Ambas alteraci6n quimicadel habitat (incluyendo la contaminaci6n) e hibridaci6n fueron citadas en 38% de las extinciones, y sobre explotaci6nafect6 adversamente a 15% de las especies extinctas. Se espera que esta desafortunada, y anteriormente desconocida,tasa de extinci6n se aumentara ain mas mientras mayor parte de la fauna nativa de Norte America Ilega a encontrarseen peligro, o en amenaza, de extinci6n.

'M \any rivers, lakes, and springsMr.in North America have beenfragmented, polluted, or destroyed.These losses have been documentedthrough growing lists of endangeredfishes (Deacon et al. 1979; Campbell1985,1988; USD11989) and endangeredecosystems (Williams et al. 1985). TheAmerican Fisheries Society now lists103 fish taxa as endangered, 114 asthreatened, and 147 others as of specialconcern in North America (Williams etal. 1989, this issue).

Despite the increasing threat to theaquatic resources, extinctions havebeen poorly documented. Part of theproblem is the difficulty in determining

when a species is extinct (Diamond1987). Should a species be consideredextinct when it is not found in a singlesurvey, or are many more negativesearches required before extinctionshould be considered confirmed? Ex-tinctions are particularly hard to doc-ument for small fishes or species thateasily evade casual observation or oc-cupy habitats that are difficult to sam-ple.

Some taxa may persist at perilouslylow numbers for many years, only tobe rediscovered in remote comers oftheir habitats or after populations haverecovered sufficiently to again be no-ticed. There are several examples of

Robert R. Miller is curator emeritus of fishes at the Museum of Zoology, Universityof Michigan, Ann Arbor, MI 48109. His career in the study of fishes from westernUnited States, Mexico, and Guatemala spans more than 5 decades. James D. Williamsconducts research on endangered and threatened fishes, and impacts of exotic species onnative fishes at the Fish and Wildlife Service, National Fisheries Research Center inGainesville, FL. Jack E. Williams conducted research on endangered fishes for the Fishand Wildlife Service and the University of California at Davis. He is currently fisheriesprogram manager for the Bureau of Land Management. All three authors have servedas chairman of the AFS Endangered Species Committee. The present paper is part of amonograph on extinct fishes in preparation by the authors.

fishes that were considered to be extinctonly to be rediscovered by surprisedscientists. The Owens pupfish (Cypri-nodon radiosus) was rediscovered in aremote comer of Fish Slough, Califor-nia, in 1964 (Miller and Pister 1971)after being presumed extinct by Miller(1961) and others. Similarly, the Sho-shone pupfish (C. nevadensis shoshone)was rediscovered in 1986 after numer-ous surveys failed to find a populationthat had dwindled to such a small sizethat the remaining individuals expe-rienced a genetic bottleneck (Taylor etal. 1988). For such endangered specieswhere remnant populations survive,much of the original genetic variationhas been lost (Wilson 1988). Unfortu-nately, chances of rediscovery are pastfor an increasing number of fishes.

Our purposes here are to provide acomprehensive list of recently extinctfishes of the North American continentand detailed accounts of each extinc-tion. We include fishes where extinc-tions are presumed on the basis ofnumerous negative surveys and thosetaxa that are poorly known but whereextinction is presumed and rediscovery

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Figure 1. Miller Lake lamprey, Lampetra minima. An adult male collected near the northern shore of Miller Lake,Klamath County, Oregon, 17 July 1952. Drawing by Sara V. Fink.

appears impossible. All fish extinctionssuspected to have occurred during thepast 100 years are included.

The Extinct FishesMiller Lake Lamprey

Lampetra minima Bond and KanFigure 1

This dwarf species of parasitic lam-prey was endemic to Miller Lake, Kla-math County, Oregon (Bond and Kan1973). It is thought to have evolvedfrom a stock of the Pacific lamprey (L.tridentata) isolated in Miller Lake afterthe explosion of Mt. Mazama (nowCrater Lake), which probably de-stroyed virtually all aquatic life in thearea 6,600 years ago. Kan and Bond(1981) attributed evolution of thisunique species to a reduced food sup-ply, short generation time, and thesmall numbers of lampreys survivingin the area after the mountain's erup-tion.

Forty-five adult specimens examinedby Bond and Kan (1973) ranged from72 to 129 mm TL, making this thesmallest species of parasitic lampreyknown. In addition to its small size,the species was notable in having rel-atively high fecundity, indicating agreater use of energy for gamete pro-duction than for spawning migrations(Kan and Bond 1981).

Because it preyed on introducedtrout, the Miller Lake lamprey waspurposely poisoned with ichthyocidesduring the 1950s by the state of Oregon.The last collection of the species wasmade from Miller Lake on 17 July 1952.Subsequent investigations have con-firmed the extinction of this species.

Longjaw CiscoCoregonus alpenae (Koelz)

The longjaw cisco, described fromLake Michigan off Charlevoix, Michi-

gan, was most abundant in Lakes Mich-igan and Huron. The species was rarein Lake Erie, where the last record ofcapture was in November 1957 (Scottand Smith 1962). It was one of thelargest ciscoes in the Great Lakes;adults often attained a length of 38 cmand a weight of 1 kg (Koelz 1929).

Stocks of this large and commerciallyvaluable species were severely de-pleted by overfishing in Lakes Michi-gan and Huron by the turn of thecentury (Smith 1968). As the catchesbegan to decrease, the fishery becamemore intense and selective. The severedecline caused by commercial overex-ploitation probably was aided in itslater stages by predation from the exoticsea lamprey (Petromyzon marinus), whichbecame established in the 1940s. Asthe longjaw cisco became increasinglyrare, it is believed to have becomeextinct through hybridization withmore common ciscoes (G. R. Smith,University of Michigan, personal com-munication). The last known record ofcapture in Lake Michigan was fromGrand Traverse Bay in 1967 and in LakeHuron from Georgian Bay, Ontario, on12 June 1975 (T. N. Todd, Fish andWildlife Service, Ann Arbor, MI, per-sonal communication).

Deepwater CiscoCoregonus johannae (Wagner)The deepwater cisco was one of the

two most important species in the earlycisco fisheries of the Great Lakes. Thedeepwater cisco occurred in the deeperparts of Lakes Michigan and Huron,at depths of 55 to 183 m (Koelz 1929).The last known record of capture forthe species in Lake Michigan was fromGrand Traverse Bay in June 1951, andin Lake Huron from Wolfsell, Ontario,on 4 August 1952 (T. N. Todd, Fishand Wildlife Service, Ann Arbor, MI,

personal communication).Heavy exploitation from the com-

mercial fishery produced a marked de-cline in deepwater cisco populationsduring the early 1900s (Smith 1970).Continued intensive fishing for alllarger ciscoes, coupled with predationby exotic sea lampreys, led to dwin-dling stocks of this species (Smith 1964).As populations of the deepwater ciscowere diminished, introgressive hybrid-ization with more common ciscoes mayhave contributed to final extinction.

Lake Ontario KiyiCoregonus kiyi orientalis KoelzIn his monograph on coregonid fishes

of the Great Lakes, Koelz (1929) rec-ognized the kiyi of Lake Ontario as adistinct subspecies (Coregonus kiyi or-ientalis). It was one of the smallestciscoes in the Great Lakes and occurredthroughout Lake Ontario at depths of37 to 137 m (Koelz 1929).

Along with other ciscoes, the LakeOntario kiyi supported an importantcommercial fishery from the 1800sthrough the 1930s. Although fisherystatistics did not distinguish among thespecies, the Lake Ontario kiyi wasprobably the most important speciesin the Lake Ontario fishery (Pritchard1931).

The last known collection of this fishwas made on 19 September 1964 nearOswego, New York (T. N. Todd, Fishand Wildlife Service, Ann Arbor, MI,personal communication). Like the otherextinct coregonid fishes of the GreatLakes, the decline and ultimate dis-appearance of the Lake Ontario kiyiwas caused by a combination of events.Overfishing was a major factor. Estab-lishment of exotic and non-native fishesand deterioration of water quality fromeutrophication and release of toxicchemicals also were contributing fac-tors.

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Blackfin CiscoCoregonus nigripinnis (Gill)

The blackfin cisco was one of thelargest and most commercially valuableof the coregonid fishes in the GreatLakes. It occurred in the deeper waters(110 to 146 m) of Lakes Michigan andHuron (Koelz 1929).

The decline and eventual extinctionof the blackfin cisco were caused bythe same factors (overfishing, sea lam-prey predation, and introgressive hy-bridization) as those described for thelongjaw cisco. The last blackfin ciscoknown from Lake Huron was taken atWiarton, Ontario, on 26 June 1923(Koelz 1929). The last record of thespecies from Lake Michigan was on 26November 1969 at Marinette, Wiscon-sin (T. N. Todd, Fish and WildlifeService, Ann Arbor, MI, personal com-munication).

Yellowfin Cutthroat TroutOncorhynchus clarki macdonaldi

(Jordan and Evermann)

The yellowfin cutthroat trout wasknown only from Twin Lakes on LakeCreek, a tributary to the Arkansas Riverin Lake County, Colorado. The pres-ence of the greenback cutthroat trout(0. clarki stomias) in Twin Lakes and itsearly extinction clouded the taxonomichistory of the yellowfin cutthroat trout.Behnke (1979) examined yellowfin andgreenback cutthroat trouts of the samesize from Twin Lakes and concludedthat macdonaldi was a valid taxon, prob-ably most closely related to the Colo-rado cutthroat trout (0. c. pleuriticus):

Apparently a favorite of local anglersduring the late 1800s, the yellowfin

cutthroat attained a weight of at least5 kg (Jordan 1922). Yellowfin cutthroattrout were propagated at Leadville Na-tional Fish Hatchery and may havebeen introduced into Colorado's GrandMesa (Behnke 1979) and even intoFrance (Jordan 1922). Attempts at es-tablishment and long-term propaga-tion failed.

Non-native trouts were extensivelyintroduced into Twin Lakes and land-locked salmon (Salmo salar sebago), rain-bow trout (Oncorhynchus mykiss), laketrout (Salvelinus namaycush), and brooktrout (S. fontinalis) all were establishedin the lakes by the turn of the century(Juday 1907). Introgressive hybridiza-tion with rainbow trout and competi-tion from deep-water lake trout ap-parently eliminated yellowfin cutthroattrout as well as the local population ofgreenback cutthroat trout from TwinLakes around 1910. The Forest Serviceapparently propagated a silvery "yel-lowfin" trout from Island Lake, Colo-rado, during the early 1930s, but theirgenetic purity was doubtful.

Alvord Cutthroat TroutOncorhynchus clarki ssp.

Figure 2The Alvord cutthroat trout was en-

demic to streams in the Alvord Basinof southeastern Oregon and north-western Nevada. It was known onlyfrom Trout Creek in Oregon and VirginCreek in Nevada, although it may havelived in several of the larger AlvordBasin streams during recent times(Hubbs and Miller 1948). The Alvordcutthroat's nearest relative appears tobe another undescribed subspecies

of cutthroat trout from Willow andWhitehorse creeks in Oregon, just eastof the Alvord Basin.

Hybridization and introgression withintroduced rainbow trout apparentlycaused extinction of the Alvord cut-throat trout. Rainbow trout charac-teristics were already noticeable inthe first collection of trout from VirginCreek made by Carl L. Hubbs andfamily in 1934. Cutthroat trout pig-mentation was present in Trout Creekfish collected by Carl E. Bond in 1953and 1957, but no basibranchial teeth(typically present in 0. clarki) werefound (C. E. Bond, Oregon State Uni-versity, personal communication; Wil-liams and Bond 1983). Although nocollections of trout from Virgin Creekare known from 1957 to 1970, speci-mens collected in 1971, 1978, and 1979showed typical rainbow trout charac-teristics or appeared to be rainbow xcutthroat hybrids (Williams and Bond1983). Electrophoretic analyses of troutcollected from isolated portions of Vir-gin Creek in 1984 to 1986 showed largeportions of rainbow trout alleles in allsamples (Tol and French 1988). Genet-ically pure Alvord cutthroat trout ap-pear to be extinct, and may have beenextirpated soon after stockings of rain-bow trout during the early 1930s.

Silver TroutSalvelinus agassizi (Garman)

Figure 3The silver trout from Dublin Pond,

New Hampshire, was recognized as aunique, distinctive trout as early as1849 (Bigelow 1850), although it wasnot formally described as a new species

Figure 2. Alvord cutthroat trout, Oncorhynchus clarki ssp. An adult female, 106.5 mm SL, collected from Virgin Creek,Humboldt County, Nevada. Drawing by Sara V. Fink.


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Figure 3. Silver trout, Salvelinus agassizi. A specimen 21.6 cm TL collected fromMonadnock Lake, Cheshire County, New Hampshire, 2 December 1887. Drawing bySherman F. Denton.

until 1885. In addition to Dublin Pond(also called Center Pond or MonadnockLake), silver trout occurred in ChristineLake (Warfel et al. 1939). Both were inthe Connecticut River drainage. Mor-phologically, silver trout are similar tobrook trout, but the two are distinctlydifferent in coloration. Both speciesoccurred sympatrically, and the silvertrout was relegated by many earlyworkers to subspecific status, Salvelinusfontinalis agassizi (Jordan 1885).

Adult silver trout typically weighedbetween 1 and 2 kg, but individualsup to 3 kg were known (Kendall 1914).They were taken by fishermen using avariety of baits, including worms, min-nows, grasshoppers, and artificial flies.

The species began to decline in abun-dance during the late 1800s. This de-cline appears to be attributable tooverharvesting during spring and au-tumn, when silver trout were in shal-low water. Although populations werereduced by excessive fishing, it wasthe introduction of non-native fishesthat caused extinction of the species.Warfel et al. (1939) reported the follow-ing introduced fishes in Dublin Pond:Atlantic salmon, rainbow trout, laketrout, chinook salmon, rainbow smelt(Osmerus mordax), brown bullhead (Ic-talurus nebulosus), yellow perch (Percaflavescens), and white perch (Moroneamericana). Both Dublin Pond andChristine Lake received hatchery stocksof brown trout in addition to the nativeform. Hybridization with hatchery-reared brook trout may have contrib-uted to extinction of the silver trout.The last record of silver trout fromChristine Lake was in 1926 (Warfel etal. 1939) and from Dublin Pond in 1930(Behnke 1980).

Maravillas Red ShinerCyprinella lutrensis blairi (Hubbs)

Hubbs (1940) described the Mara-villas red shiner from two localities inthe Big Bend region of Texas. The typeseries consists of two collections of 41specimens taken from Garden Springson 16 April 1937 and 19 July 1938, andone collection of 10 specimens fromPefia Colorado Creek taken on 16 April1937. Both localities are in the Mara-villas Creek drainage, tributary to theRio Grande. The last known collectionof this subspecies was apparently madeby Clark Hubbs in 1954, when redshiners were collected from upperMaravillas Creek at its confluence withPefia Colorado Creek.

By 1954, Garden Springs was dom-inated by an introduced population ofplains killifish (Fundulus zebrinus) andno red shiners could be found (Mat-thews 1987). Matthews (1987) sug-gested that the subspecies was extinctbecause of loss of the Garden Springspopulation and unsuccessful efforts tocollect the shiner from Pefia ColoradoCreek in 1978.

Mexican DaceEvarra spp.

The Mexican cyprinid genus Evarrawas described by Woolman (1895) fromcanals in the endorheic Valley of Mex-ico, where Mexico City and its suburbsnow stand. The genus Evarra, an Indianname meaning "maker of gods in landsbeyond the sea" (Jordan and Evermann1896-1900), includes three species: E.eigenmanni Woolman, E. tlahuacensisMeek, and E. bustamantei Navarro.

Unfortunately, relatively few mu-seum specimens of these small (max-

imum SL = 80 mm) minnows still exist.Alvarez and Navarro (1957) had largeseries of two of the species, but thesehave largely disappeared. Evarra is pre-sumed to be derived from the Mexicangenus Algansea (Barbour and Miller1978).

Alvarez and Navarro (1957) believedall three species to be nearing extinctionthree decades ago, and all were be-lieved to be extinct by 1983 (E. Diaz-Pardo, Instituto Politecnico Nacional,Mexico City, personal communica-tion). The extinctions coincided withdrying of the lakes, spring-fed ponds,and canals of the valley floor, due toagricultural practices, persistentgroundwater removal, and the devel-opment of Mexico City and suburbs.

Independence Valley Tui ChubGila bicolor isolataHubbs and Miller

Gila bicolor isolata was endemic toWarm Springs of Independence Valleyin Elko County, Nevada. It was quitepossibly extinct by the time it wasoriginally diagnosed as a new subspe-cies by Hubbs and Miller (1972). In thefirst collection of the subspecies on 25August 1965, Hubbs et al. (1974) de-scribed the fish as abundant in thespring complex. They greatly outnum-bered another endemic, the Indepen-dence Valley speckled dace (Rhinichthysosculus lethoporus). A second collectionof the tui chub was made on 3 April1966 by S. H. Berwick and others. Since1966, no collections of the Indepen-dence Valley tui chub are known. Re-cent surveys of the Warm Springssystem conducted by Vinyard (1984)yielded a few speckled dace but no tuichub.

Between 1960 and 1966, largemouthbass (Micropterus salmoides) and blue-gills (Lepomis macrochirus) were intro-duced into the Warm Springs systemand the largemouth bass is now abun-dant (Vinyard 1984). Hubbs et al. (1974)reported the introduced bullfrog (Ranacatesbeiana), common carp (Cyprinus car-pio), and largemouth bass in the springsin 1965 and expressed concern for sur-vival of the native fishes. Vinyard (1984)speculated that the introduced cen-trarchids were responsible for elimi-nation of the tui chub. This appearsto be a logical hypothesis, as G. b. iso-lata preferred midwater habitats thatwere easily invaded by largemouth


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bass, whereas the speckled dace mayhave largely escaped extirpation byfinding refuge in shallow water withdense aquatic vegetation.

Extinction of the Independence Val-ley tui chub probably occurred duringthe late 1960s or early 1970s. Surveysof other springs in the valley for fisheshave been unsuccessful (Vinyard 1984).

Thicktail ChubGila crassicauda

Baird and Girard

At one time the thicktail chub, whichattained lengths of 250 mm, was oneof the most common fishes in Califor-nia's Central Valley and at the turn ofthe century was sold in markets in SanFrancisco. Examination of fish remainsfrom historic Indian middens revealedthe abundance of the chub. In onemidden approximately 1 km west ofthe Sacramento River in Colusa County,thicktail chub was the most abundantspecies, contributing 41% of the 1,497fish recovered (Schulz 1979). Museumrecords for the species during the late1800s and early 1900s indicated a broaddistribution throughout lower-eleva-tion streams and rivers of the CentralValley, tributary streams of San Fran-cisco Bay, central California coastal riv-ers, and Clear Lake (Rutter 1908; Miller1963; Mills and Mamika 1980). In pe-riods of heavy runoff, the species alsoinhabited fresh surface waters of SanFrancisco Bay (Ayres 1862). Preferredhabitats consisted of rivers, marshes,sloughs, lakes, and slow-flowingreaches of rivers.

Primary causes of the decline ofthicktail chub can be traced to conver-sion of much of the Central Valley tointensive agricultural production.Many of the sloughs and marshes weredrained during the late 1800s, and theremaining streams and rivers werelargely dredged or channelized for nav-igation or flood control. Of the 34 mu-seum lots of thicktail chub with collec-tion dates, only 11 (32%) were collectedafter 1900. Habitat loss, combined withcompetition and predation pressuresfrom a myriad of introduced non-native fishes, probably combinedto cause extinction of the thicktailchub. It was last collected from ClearLake in 1938 and from the SacramentoRiver in 1950. The last known collectionof the species was made from Steam-boat Slough along the San Joaquin River,

Solano County, on 16 April 1957. Re-peated efforts to find the species sincethat time have failed.

Pahranagat SpinedaceLepidomeda altivelisMiller and Hubbs

The Pahranagat spinedace wasknown from only two localities, theoutflow of Ash Springs and UpperPahranagat Lake, in Pahranagat Valley,Nevada. Miller and Hubbs (1960) re-ported collecting the spinedace inabundance at the two sites during 1938.It was not present in Crystal or Hikosprings or Ash Springs proper, indi-cating an avoidance of constant, warmwater.

Extinction of the Pahranagat spine-dace is most likely attributable to com-petition or predation by introducedspecies. In 1938, only a few commoncarp were collected with the spinedacefrom the outflow of Ash Springs,whereas this exotic was plentiful inUpper Pahranagat Lake. Common carpand another introduced species, themosquitofish (Gambusia affinis), wereabundant during a 1959 survey thatyielded no Pahranagat spinedace(Miller and Hubbs 1960). The intro-duction of bullfrogs also may havecontributed to the decline of the nativefish by predation. The 1938 collectionof Pahranagat spinedace is the lastknown, as numerous attempts to col-lect the species by Miller and Hubbs,and J. E. Deacon from the Universityof Nevada at Las Vegas, have failed.

Ameca ShinerNotropis amecae

Chernoff and MillerThe Ameca shiner was described

from upper parts of the Rio Amecadrainage in Jalisco, Mexico (Chernoffand Miller 1986). The Rio Teuchitlan,from which the type specimens werecollected, lies about 75 km west ofGuadalajara. Its waters are heavily uti-lized for drinking, irrigation, and otherdomestic uses and have been heavilypolluted by humans and livestock.

The species was not common duringthe early 1900s, as only 46 of 2,875fishes collected from the upper RioAmeca basin between 1939 and 1969were Notropis amecae. The last knowncapture of the Ameca shiner was on 21April 1969. Intensive collecting in the

drainage during the past 20 years hasuncovered no shiners. Many habitatsformerly occupied by native fishes arenow populated by introduced fishessuch as shortfin molly (Poecilia mexi-cana), platyfish (Xiphophorus), and ti-lapia (Tilapia). Thus, we fear that thisRio Ameca endemic is extinct.

Durango ShinerNotropis aulidion

Chernoff and Miller

The Durango shiner was native tothe Rio Tunal, which forms the head-waters of the Rio Mezquital, a Pacific-slope river rising near Durango City,Durango, Mexico (Chernoff and Miller1986). It was taken there only in 1951and 1961. All specimens of this specieswere collected in a long, narrow res-ervoir on the Rio Tunal at an elevationof 1,865 m. Like its closest relatives, allof which occur south of Durango, thisspecies evidently preferred quiet water.Collections farther upstream in the RioTunal (where the current is notable)failed to yield any specimens.

Since the mid-1960s, Durango Cityhas grown considerably, with the resultthat streamflows have become reducedand domestic and industrial pollutionhave adversely affected surface waters.In addition, introduced species haveseriously impacted native fishes bycompetition and predation. We believethat only 6 of the original 11 speciesknown from the drainage still occurthere. None of these six is a cyprinid.Introduced largemouth bass, in partic-ular, have seriously affected the nativebiota. Attempts to locate the Durangoshiner as recently as 1985 have failed,and we therefore doubt that the speciessurvives.

Phantom ShinerNotropis orca Woolman

The phantom shiner, formerly con-fused with the bluntnose shiner, No-tropis simus (Cope), has recently beenrecognized as a valid species. It wasendemic to the Rio Grande from themouth of the river at Boca Chica Beach,Texas, to central New Mexico, includ-ing the mouth of the Pecos River (Cher-noff et al. 1982). Little is known aboutthe biology of this species, as it hasbeen collected only once in the past 38years-a single adult taken in 1975 fromthe Mexican side of the lower Rio


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Grande.Habitat alteration of the Rio Grande

has been extensive as a result of waterdiversions, dams and their impound-ments, chemical pollution and an in-crease in salinity that is correlated withirrigation projects. These factors, alongwith an abundance of introducedfishes, brought about a significant de-cline in abundance of both the phantomand bluntnose shiners, especially after1940. Historically, the Rio Grande wascharacterized by floods and droughts,thus providing an unstable flow re-gime. The limited fish fauna may wellhave been adapted to these long-termunstable conditions. Introduced fishes,which often are greatly diminished dur-ing floods and droughts, flourished inthe more stable flows.

Notropis simus and N. orca are knownto have hybridized in the upper RioGrande at Juarez, Mexico, as long agoas 1891 (Chernoff et al. 1982). By 1939,some individuals of N. simus taken inthe river north of the Valencia (NewMexico) County line exhibited intro-gressive hybridization. The precisecause of hybridization is unknown, butlikely was brought on by the reductionin number of spawning sites and/orrarity of one or both species. What rolehybridization played in the decline ofthese species is unknown because weknow so little about their biology.

Rio Grande Bluntnose ShinerNotropis simus simus Cope

Figure 4The bluntnose shiner was recently

recognized as comprising two subspe-

cies, N. s. simus from the upper RioGrande between the Chama River andEl Paso, Texas; and N. s. pecosensis fromthe Pecos River in New Mexico (Cher-noff et al. 1982). Adult Rio Grandebluntnose shiners inhabited the swift,relatively deep, main river channel, butwere occasionally taken from irrigationditches or shallow riffles.

The last known Rio Grande blunt-nose shiner was caught in the RioGrande north of Pefia Blanca, SandovalCounty, New Mexico, on 28 July 1964by R. D. Suttkus and class. As dis-cussed above for the phantom shiner,a number of factors may have contrib-uted to the extinction of the Rio Grandebluntnose shiner, including modifica-tions of the river by dams and theirimpoundments, irrigation practices,and channelization. Loss of spawningsites by desiccation, perhaps in con-junction with competition from intro-duced species, may have been espe-cially critical.

Clear Lake SplittailPogonichthys ciscoides HopkirkPogonichthys ciscoides was restricted

to Clear Lake and its tributary streamsin Lake County, California. This la-custrine cyprinid once formed largeschools in Clear Lake, especially in thelittoral zones. During April and May,the splittail ascended streams tributaryto the lake to spawn (Moyle 1976).Hopkirk (1973) reported the speciesfrom several localities in Clear Lakeduring the early and mid-1960s, and asingle specimen that apparently waswashed downstream into CacheCreek, 0.6 km upstream from State

Highway 16 (1965).The decline of the once abundant

splittail parallels the drastic change inClear Lake itself. Prior to the 1880s,the lake was a cool, clear habitat witha fishery dominated by rainbow troutand Sacramento perch (Archoplites in-terruptus) (Murphy 1951); by the early1940s, it had been transformed into awarm, turbid lake dominated by cen-trarchids, common carp, and other in-troduced fishes. Goldman and Wetzel(1963) documented the "extreme eu-trophication" of Clear Lake by the late1950s and attributed the change to rapidand poorly planned development ofthe watershed for agriculture, whichwashed sediment, fertilizers, and sew-age into the lake. The Clear Lake split-tail was abundant until 1942 or 1943,but during three summers of intensiveseining from 1946 to 1950, Murphy(1951) collected only a few juveniles.Murphy (1951) concluded that the lowrainfall years of the early 1940s, coupledwith diversion of tributary streams foragriculture, eliminated most of the hab-itat for the remaining splittails. Thedemise of the species may have beenhastened by the application of largequantities of chlorinated hydrocarbonpesticides used to control swarms ofClear Lake gnats (Chaoborus astictopus).During the 1940s and 1950s, increas-ing use of DDD (dichloro-diphenyl-dichloroethane) resulted in the ap-pearance of this toxin in fishes andpiscivorous birds (Hunt and Bischoff1960) and massive die-offs of westerngrebes (Aechmophorus occidentalis). Also,the abundance of introduced littoralfishes, such as bluegills and inland

Figure 4. Rio Grande bluntnose shiner, Notropis s. simus. A specimen 65.2 mm SL, collected from Rio Grande 7 kmnorth of Pena Blanca, Sandoval County, New Mexico. Drawing by Sara V. Fink.


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silverside (Menidia beryllina) probablywas detrimental. The last specimen ofClear Lake splittail was taken fromClear Lake in 1970. Recent collectionefforts have failed and the species ispresumed to be extinct (Moyle 1976).

Banff Longnose DaceRhinichthys cataractae smithi

Nichols

The Banff longnose dace was en-demic to the marsh created by Caveand Basin Hotsprings 1.7km southwestof Banff, Alberta, Canada. Based oncollection records during the late 1800sand early 1900s, the Banff longnosedace was quite abundant in its re-stricted habitat (Lanteigne 1988). Afterthe early 1900s, surveys revealed fewspecimens. In May 1971, 16 individualswere collected at the inflows, and 7more from the same site in May 1981(Lanteigne 1988). A September 1981trip yielded only two specimens froma pool at the outlet of the marsh intothe Bow River, but others were sightedin the same locality. Decreasing pop-ulation size by 1981 indicated that theBanff longnose dace was endangeredat that time, and more recent unsuc-cessful collection efforts indicate thesubspecies to be extinct (Lanteigne1988).

Initial decline of the Banff longnosedace can be attributed to introductionsof numerous exotic and non-nativefishes, including mosquitofish, whichwere stocked as early as 1924 (Mc-Allister et al. 1985), and various tropicalfishes such as sailfin mollies, guppies(Poecilia reticulata), convict cichlids(Cichlasoma nigrofasciatum), and greenswordtails (Xiphophorus helleri), (Lan-teigne 1988). Other factors contributingto the decline of the longnose dace areperiodic cessation of flows related touse of the hotsprings as public baths,and the periodic spillage of sewagefrom the public facilities. Once theBanff longnose dace declined in num-bers, it was susceptible to introgressivehybridization with the eastern subspe-cies of longnose dace (R. c. cataractae),which apparently caused its final ex-tinction (Renaud and McAllister 1988).

Las Vegas DaceRhinichthys deaconi Miller

Between 1891 and 1940, several col-lections of a small minnow were made

Figure 5. Stumptooth minnow, Stypodon signifer. The syntype, 40.0 mm SL, collectedfrom a spring near Parras, Coahuila. Drawing by Sara V. Fink.

from Las Vegas Creek, near Las Vegas,Nevada. This dace was described byMiller (1984) and distinguished fromits closest relative, the speckled dace(R. osculus).

In May 1844, John Fremont describedLas Vegas Creek as "a delightfulbathing place." In 1891, Vernon Baileymentioned the numerous little fish inthe springs and stream. Cottonwoodsand willows grew along the banks. Inhigh water, flow from Las Vegas Creekreached the Colorado River, some 40km distant, but by 1947, flow downthe creek had already been reduced bydevelopment of the Las Vegas com-munity. Nothing is left of the springand creek complex today, except a holein the ground with stagnant water atthe bottom. At one time, this creek andmarsh also supported the endemic Ve-gas Valley leopard frog (Rana fisheri),which is also extinct. Both species wereextirpated as the system was convertedfor agricultural and urban develop-ment.

The last known collection of the LasVegas dace was made on 30 July 1940,at Lorenzi Ranch in Las Vegas. Thespecies probably survived in one of thesprings and its outflow until about1955, when there was a marked in-crease in the amount of water with-drawn from the artesian basin beneathLas Vegas. No fish were found 10 yearslater when the area was searched byJ. E. Deacon and colleagues from theUniversity of Nevada, Las Vegas.

Grass Valley Speckled DaceRhinichthys osculus reliquus

Hubbs and Miller

The Grass Valley speckled dace isknown only from a single collection

made by C. L. Hubbs and R. R. Milleron 9 August 1938. The collection localitywas given by Hubbs and Miller (1972)as a "spring-fed creek in a grassymeadow in the partly enclosed south-western arm of Grass Valley, 13 kmeast of Mt. Callaghan, in the course ofCallaghan (Woodward) Creek, . . ineastern Lander County, Nevada."

The speckled dace was common whencollected in 1938. The single series con-sisted of 474 specimens. At that time,one trout was seen in the stream. Asecond collecting trip by Hubbs andMiller in 1969 produced only rainbowtrout and brook trout from the stream(Hubbs et al. 1974). Ira La Rivers (Uni-versity of Nevada, Reno, personal com-munication, to Hubbs) also reportedfinding no speckled dace, but only astream "full of trout" during his visitto Grass Valley in about 1959. Effortsto collect this speckled dace from GrassValley during the 1970s also failed andthe fish appears to be extinct. Theintroduction of trout probably is theprimary cause of extinction, but use ofthe water for agriculture may havecontributed to the loss.

Stumptooth MinnowStypodon signifer Garman

Figure 5

This species represents a monotypicgenus known only from six speci-mens-two collected in 1880 and fourin 1903. All specimens came from aspring or spring complex near Parras,in southern Coahuila, Mexico. The spe-cies was named for its large, molari-form pharyngeal teeth, which evi-dently were specialized for feeding onthe abundant mollusks.


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A 2-day search of Parras Valley in1953, by C. L. and L. C. Hubbs, re-vealed Gila as the only surviving nativefish. This was confirmed by Contreras-Balderas (1969). Habitat modificationand loss of springs, and/or water pol-lution and irrigation practices that car-ried water onto cultivated fields wereall important causal factors in the de-cline. The snails that were the food ofthe stumptooth minnow may havebeen particularly sensitive to changesin water level and deteriorating waterquality.

June SuckerChasmistes liorus liorus JordanThe June sucker (Chasmistes liorus),

Utah sucker (Catostomus ardens), andtheir hybrids historically occurred insuch great abundance in Utah Lakethat it was referred to as the "greatestsucker pond in the universe" (Jordanand Gilbert 1881). June suckers wererestricted to Utah Lake, except duringlate spring (typically in June-hencethe common name), when they as-cended tributary streams for spawning.Commercial fishermen harvested thesuckers during their spawning runs inthe Provo and Spanish Forks rivers aswell as in Utah Lake during winter.

Diversion of inflowing streams forirrigation decreased the lake's com-mercial fisheries for the sucker andnative cutthroat trout as early as 1860(Winter, personal communication, inHeckmann et al. 1981). Diversion damson tributary rivers also blocked spawn-ing runs and caused hybridization be-tween the two sucker species as theywere concentrated in the dam's tail-waters. Introductions of 25 exotic andnon-native fishes, also contributed to

decline of the native fishes. Eleven ofthese were established in Utah Lake orits tributaries prior to 1930 (Heckmannet al. 1981).

The final collapse of the June suckerappeared to result from the severedrought of the mid-1930s. Tanner (1936)succinctly described the final demiseof genetically pure June suckers asfollows: "At this writing Jan. 1936,practically all the suckers as well asother fish in Utah Lake have been killedby the severe drought of the past fouryears. . . During the winter of 1934-35 the water was so shallow thathundreds of tons of suckers and carpwere killed due to freezing and crowd-ing in the few deep holes. . . In thespring of 1935 there were no suckersto run up Provo River, something thathas never happened before in the his-tory of Utah Lake."

A Chasmistes now exists in Utah Lake,but it differs significantly from typespecimens of C. liorus. The true-breed-ing population of Chasmistes from UtahLake was described as a new subspe-cies, C. liorus mictus, that resulted fromhybridization between the June andUtah suckers (Miller and Smith 1981).Unlike previous hybrids in the lake,this subspecies of June sucker com-prised a large breeding population withconsistent, distinguishable characters.The original form of June sucker, C. .liorus, is extinct (Miller and Smith 1981).

Snake River SuckerChasmistes murieiMiller and Smith

Figure 6The Snake River sucker is known

only from a single specimen collectedby Dr. Olaus J. Murie from the Snake

River below Jackson Lake Dam, Wyo-ming, on 13 October 1927 (Miller andSmith 1981). The species is presumedto have occupied Jackson Lake andperhaps other lakes on the upper SnakeRiver. Like other members of the genusChasmistes, the Snake River sucker wasa lacustrine species with numerousdendritic gill rakers for feeding onplankton.

Chasmistes muriei may have been ex-tinct in genetically pure form when theonly specimen was collected in 1927.By this time, introgression of the Utahsucker with the Snake River sucker wasevident. A rock-filled dam was con-structed on the outflow of Jackson Lakein 1905-1906. After this dam waswashed out in 1910, construction beganthe next year on a concrete dam thatwould eventually almost double thesurface area of Jackson Lake (J. Daugh-erty, National Park Service, personalcommunication). Hybridization withthe Utah sucker probably occurred be-cause such dams on the upper SnakeRiver blocked spawning migrations andforced remaining suckers to spawn inthe dams' tailwaters. Had it not beenfor the diligence of wilderness advocateDr. Murie, who collected the only spec-imen, the passage of this species intoextinction would have gone unnoticed.

Harelip SuckerLagochila lacera

Jordan and BraytonFigure 7

During the late 1800s, the harelipsucker was widespread in the easternUnited States, where it occurred in atleast eight states. Museum records existfor the Chickamanga River (GA), ElkRiver (TN), Scioto River and Maumee

Figure 6. Snake River sucker, Chasmistes muriei. An adult female, 371 mm SL, collected from the Snake River belowJackson Lake Dam, Teton County, Wyoming, 13 October 1927. Drawing by Sara V. Fink.


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Figure 7. Harelip sucker, Lagochila lacera. Specimen collected at Fairview, WilliamsonCounty, Tennessee, sometime before May 1880. Drawing by H. L. Todd.

River (OH), Tennessee River drainage(Al, TN, VA), Cumberland River drain-age (KY, TN), and Tippecanoe River(IN) (Jordan 1882, 1890; Evermann andJenkins 1889; Woolman 1892; Kirsch1896). Although the distribution of theharelip sucker was widespread, its hab-itat appears to have been restricted todeep pools between shoal areas withclear water over a bedrock or rockybottom in moderate to large streams.Most collectors reported the speciesas "abundant," "common," or "notrare."

Some life history data are availablefrom studies of catostomid brain andlips (Miller and Evans 1965) that notedthe large optic brain lobes of the harelipsucker and its highly modified lips withfew taste buds. These characteristicsindicate visual location of foods. Ex-amination of stomach contents of nineharelip suckers revealed that about 90%of the food items were mollusks (R. E.Jenkins, Roanoke College, personalcommunication).

The decline and ultimate extinctionof the species may be attributable toincreased turbidity and siltation of itsclearwater pool habitats as a result ofagricultural practices. Siltation and pol-lution could have greatly reduced mol-lusk populations and impaired the abil-ity of the sucker to visually locate foods.The species was last collected in 1893,but it probably did not become extinctuntil the early 1900s based on reportsof the harelip sucker as "common" inthe 1880s and 1890s.

Parras PupfishCyprinodon latifasciatus Garman

The Parras pupfish is known onlyfrom two collections made in 1880 and

1903, from a spring or spring complexnear Parras in southern Coahuila, Mex-ico. The species was collected at thesame time as the stumptooth minnow,now also extinct. Until the early 1930s,Parras Valley contained magnificentspring systems. Subsequent develop-ment of extensive wine factories, a flourmill, rubber mill, and textile factory(Imlay 1936) seriously reduced andgreatly modified available surfacewater. By 1953, when an extensiveattempt was made to collect or observethis species again, no trace of it wasfound. The extinction of the Parraspupfish, like that of the stumptoothminnow, was probably caused pri-marily by the failure of spring flows.Industrial and domestic pollution alsolikely contributed to the decline.

Tecopa PupfishCyprinodon nevadensis calidae

MillerThe Tecopa pupfish was known from

outflows of two hot springs near Te-copa, Inyo County, California (Miller1948). The species was fairly commonin water of 36.5°C and abundant inwater of 32 to 36°C.

Modification of North and SouthTecopa Hot Springs for use as bath-houses resulted in the decline andeventual extinction of the Tecopa pup-fish. Pools were enlarged and outflowcreeks were ditched and diverted.Channelization of the outflow allowedthe Amargosa pupfish (Cyprinodon n.amargosae) from the nearby AmargosaRiver to gain access to habitat of theTecopa pupfish. This resulted in hy-bridization and continued loss of thecalidae genome by introgression. Theintroduction of mosquitofish also con-

tributed to the decline.Extinction of the Tecopa pupfish oc-

curred in late 1970 or soon thereafter.Six pupfish collected by D. V. Hemphillon 28 March 1954 from "Tecopa HotSprings Well" represent this subspe-cies. A more recent collection from thesame place by R. Papp and others on2 February 1970 comprises nine pupfishthat are clearly referable to C. n. calidae,three that are probably referable to C.n. calidae, and three that are intro-gressed with C. n. amargosae. This isthe last known collection of the Tecopapupfish. Surveys in 1972 by R. R. Millerand in 1977 by D. A. Selby confirmedextinction of the subspecies. This pup-fish was the first creature to be declaredofficially extinct under provisions ofthe Endangered Species Act of 1973.

Monkey Spring PupfishCyprinodon sp.

This undescribed species of pupfishwas restricted to Monkey Spring, atributary to the Santa Cruz River systemin Santa Cruz County, Arizona (Minck-ley 1973). The species occurred in abun-dance in the spring and an extensivecienega formed by its outflow.

Attempts to impound and deepenthe cienega apparently "broke the seal,"which resulted in draining of the marsh(Minckley 1973). The pupfish remainedabundant in the spring until large-mouth bass were introduced in 1968and 1969. Minckley (1973) reported thata few pupfish escaped predation bylargemouth bass until late 1969, whenthe species was extirpated in its naturalhabitat. Introduction of the bass alsoextirpated the local stock of Gila chub(Gila intermedia), which "was markedlydifferent from other stocks" of the spe-cies (Minckley 1973). Captive stocks ofMonkey Spring pupfish failed in 1971,thus eliminating any hope for survivalof Arizona's only single-spring en-demic fish species.

Raycraft Ranch PoolfishEmpetrichthys latos concavus

MillerFigure 8

Miller (1948) described Empetrichthyslatos from Pahrump Valley, Nevada,and recognized three subspecies fromsprings located along the east side ofthe valley. The Raycraft Ranch poolfishoccurred only in one spring on Raycraft


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Figure 8. Raycraft Ranch poolfish, Empetrichthys latos concavus. The holotype, a 39.0 mm SL adult female, collectedfrom Raycraft Ranch, Nye County, Nevada. Drawing by Sara V. Fink.

Ranch, about 0.8 km north of PahrumpRanch.

When collecting the type series in1942, Miller (1948) noted that the sub-species "was not common" and sug-gested that reduced numbers werecaused by introduced carp. Sokol (1954)reported the last known collection ofthe Raycraft Ranch poolfish in late Sep-tember 1953. Twelve specimens werecollected at that time and Sokol (1954)commented on the presence of intro-duced carp and bullfrogs. Miller (1961)reported correspondence from Ira LaRivers who surveyed Pahrump Valleyin about 1958 and found that only onespring formerly inhabited by poolfishin the valley was relatively undis-turbed. It appears that the RaycraftRanch poolfish became extinct in about1960. The spring was subsequentlyfilled to control mosquitoes.

Pahrump Ranch PoolfishEmpetrichthys latos pahrump

Miller

The Pahrump Ranch poolfish wasknown only from two springs locatedabout 180 m east of the ranch houseon Pahrump Ranch, Nye County, Ne-vada (Miller 1948). When Robert R. andFrances H. Miller collected the typeseries in October 1942, they noted thatthe northernmost of the two springshad been greatly altered by dredgingand no poolfish were observed there.The other spring still supported a pool-fish population in a marsh formed bythe spring outflow.

Sokol (1954) visited the springs inSeptember 1953, but was unable to findany poolfish. Ira La Rivers reported toMiller (1961) that all subspecies were

extant during his surveys of about 1958,but that carp and bullfrogs were abun-dant and that the spring sources wereperiodically pumped dry. Extinction ofthe Pahrump Ranch poolfish probablyoccurred in 1958, when one of thesprings failed because of excessivepumping.

Ash Meadows PoolfishEmpetrichthys merriami Gilbert

The genus Empetrichthys and the AshMeadows poolfish (E. merriami) weredescribed by Gilbert (1893) on the basisof specimens collected from Ash Mead-ows, Nevada, on 3-5 March 1891. Itappears that the poolfish may havebeen rare even at the time it was dis-covered; only seven specimens werecollected, whereas numerous speci-mens of speckled dace were taken. TheAsh Meadows poolfish was next col-lected in 1930, when George S. Myersand Joseph H. Wales obtained threespecimens. Despite concerted effortsfrom 1936 to 1942, Ralph G. and RobertR. Miller collected only 22 specimensfrom Big, Jack Rabbit, Point of Rocks,Forest, and Rogers springs (Miller1948). The last known individual wascollected from Big Spring (=DeepSpring of Miller 1948) on 7 September1948 by W. Hildemann and John A.Kopec. Sokol (1954) attempted unsuc-cessfully to collect Ash Meadows pool-fish from Big Spring in 1953. Numerousefforts since then have failed.

Introductions of bullfrogs and cray-fish (Procambarus clarkii) were first no-ticed in habitats of the Ash Meadowspoolfish in 1937 (Miller 1948). Bothintroduced species undoubtedlypreyed on the poolfish and may have

been the primary causal factor in itsextinction, which probably occurred inthe early 1950s. Habitat alteration alsomay have been a contributing factor insome springs.

Whiteline TopminnowFundulus albolineatus Gilbert

The whiteline topminnnow occurredonly in Spring Creek at Huntsville,Alabama. The only known specimensare the original 20 type specimens col-lected on 27 May 1889 by P. H. Kirsch,plus 5 others taken on the same date.The species appeared to be restrictedto Big Spring and its 0.3 km long out-flow, Spring Creek.

Armstrong and Williams (1971) de-scribed unsuccessful efforts to collectthe whiteline topminnow in 1966. In-stead they found numerous commoncarp and goldfish (Carassius auratus) ina spring system modified by smalldams, channelization, and surroundedby concrete walls. According to localresidents, Big Spring historically wasused as a water supply for the cityof Huntsville and was pumped dryon several occasions. More than 25springs and their outflows in the vi-cinity of Huntsville have been sampledduring the past 7 years but the specieshas not been found (D. A. Etnier,University of Tennessee, personal com-munication). It appears that the white-line topminnow is extinct.

Opal AllotocaAllotoca maculataSmith and Miller

The opal allotoca was described fromcollections made on 22 February 1970


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at Laguna de Santa Magdalena, about82 km west of Guadalajara in Jalisco,Mexico (Smith and Miller 1980). Thespecies also is known from a reservoirnear Etzatlan. Both waters are withinthe isolated Magdalena Basin. Thissmall goodeid inhabited shallow lakemargins and adjacent marshes.

Attempts by ichthyologists to collectthis species in recent years have failed.Alfred Radda (Instituit fiir Virologie,University of Vienna, personal com-munication) tried three times and BruceJ. Turner once. The exact cause of itsdisappearance is not known but maybe related to shrinkage of the restrictedhabitat from increasing use of ground-water in the basin by the expandinghuman population in Magdalena. Noexotic species have been reported fromthe Laguna de Santa Magdalena area.

Austin. On 30 November 1974, about35 fish from this stock were transferredto Dexter National Fish Hatchery inNew Mexico for propagation. Unfor-tunately, examination of both stocksrevealed that they were contaminatedwith mosquitofish, which resulted inextinction of the Amistad gambusia(Hubbs and Jensen 1984). The univer-sity stock may have been contaminatedbefore the fish were transferred to Dex-ter.

mental plant, elephant ears (Colocasiaasculenta), restricted suitable habitat inthe river to the areas sprayed withherbicides. All of these factors contrib-uted to the decline of the species. Asnumbers of San Marcos gambusia werereduced, hybrids with Gambusia affinisbecame more predominant. Geneti-cally pure G. georgei have not beencollected since 1983 and the species ispresumed to be extinct (Johnson andHubbs 1989).

Parras CharacodonCharacodon garmani

Jordan and EvermannThis goodeid, only recently recog-

nized as a valid species (Smith andMiller 1986), is known only from asingle specimen collected from ParrasValley in southern Coahuila, Mexico.The adult female was obtained by Ed-ward Palmer, probably in the 1880s.Parras Valley supported a highly en-demic fish fauna, which largely dis-appeared near the turn of the century(Williams et al. 1985). The extinctionof the Parras characodon, like that ofthe stumptooth minnow and Parraspupfish, was probably caused by thephysical alteration of spring habitats.

Amistad GambusiaGambusia amistadensis PedenThe Amistad gambusia was endemic

to Goodenough Spring and its outflowin Val Verde County, Texas (Peden1973). When the first specimens ofAmistad gambusia were collected on11 April 1968, their habitat was alreadythreatened by waters rising behind thenewly constructed Amistad Dam onthe Rio Grande. By 3 August, risingwaters covered the spring, but Alex E.Peden collected 30 to 40 live Amistadgambusia from flooded prickly pearand mesquite. By April 1969, the springwas covered to a depth of 23.2 m.

The stock was successfully propa-gated at The University of Texas at

Figure 9. San Marcos gambusia, Gambusia georgei. A male (holotype), 29.3 mm SL,collected from the San Marcos River at the 1-35 bridge, Hays County, Texas, on 7 March1968. Drawing by Sara V. Fink.

San Marcos GambusiaGambusia georgei

Hubbs and PedenFigure 9

Type specimens of the San Marcosgambusia were collected by ClarkHubbs and Michael M. Stevenson on7 March 1968 from the San MarcosRiver in Hays County, Texas (Hubbsand Peden 1969). They occurred onlyin thermally constant, shallow areas ofthe river characterized by mud bottomsand a lack of vegetation. Hubbs andPeden (1969) remarked on the fragilenature of its existence because of itsnarrow habitat requirements and seg-regation from two other native speciesof Gambusia in the San Marcos River.

Springs on the San Marcos Riverhave been altered by increased pump-ing of groundwater for urbanization.Pollution, including the use of herbi-cides along the river, appears to haveadversely affected the habitat of theSan Marcos gambusia. In addition, thefollowing introduced fishes have beenrecorded in habitats typically occupiedby G. georgei: Mexican tetra (Astyanaxmexicanus), sailfin molly, Amazon molly(Poecilia formosa), smallmouth bass (Mi-cropterus dolomieui), redear sunfish (Le-pomis microlophus), and others (Ed-wards et al. 1984). Finally, and perhapsmost critically, the spread of an orna-

Blue PikeStizostedion vitreum glaucum

HubbsThe blue pike was primarily known

from Lake Erie, but also occurred inLake Ontario, the lower Niagara River,and rarely in Lake Huron (Deason 1936;S. H. Smith, Fish and Wildlife Service,Ann Arbor, MI, personal communi-cation). It was an important species inthe commercial fishery of Lakes Erieand Ontario, especially in Lake Eriewhere the annual catch until the mid-1950s was between 5 and 10 million kg(Deason 1936). By 1950 the fishery wasin a substantial decline and by 1959the total catch was only 35,834 kg. Thelast confirmed specimen was takenfrom Lake Erie in 1965, and no reli-able reports of this fish exist past1970.

The physical, chemical, and biolog-ical environments of Lakes Erie andOntario deteriorated measurably fromthe 1940s to the 1960s. Enrichment ofthe lakes and depletion of dissolvedoxygen created conditions unfavorablefor pike eggs and young. Impacts fromintroduced fishes and overharvestingby selective commercial and sport fish-eries also contributed to the declineduring this period. Hybridization andintrogression between the blue pikeand walleye (S. v. vitreum) also were


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Figure 10. Utah Lake sculpin, Cottus echinatus. An adult female, 64.5 mm SL, collected from Utah Lake at the mouthof the Provo River, Utah County, Utah, during April 1928. Drawing by Suzanne Runyan.

involved in the demise of the blue pike(Regier et al. 1969). All of the abovefactors contributed to the extinction ofthe blue pike.

Utah Lake SculpinCottus echinatus Bailey and Bond

Figure 10The original description of the Utah

Lake sculpin was based on seven spec-imens collected from Utah Lake, Utah,between 1880 and 1928 (Bailey andBond 1963). The species apparently wasstrictly lacustrine as it has never beentaken outside the lake.

As described earlier for the Junesucker, increased pollution and salinityby developing agriculture greatly di-minished water quality in Utah Lakeduring the late 1800s and early 1900s.In addition, at least 11 non-native andexotic fishes became established inUtah Lake or its tributaries before 1930(Heckmann et al. 1981). Extremedrought and low water conditions ofthe mid-1930s apparently extinguishedthe remaining Utah Lake sculpins. Thelast known specimen was collected byVasco M. Tanner in April 1928.

SummaryDuring the past 100 years, activities

of mankind have caused the extinctionof 40 taxa of North American fishes: 27species, and 13 subspecies (as well as3 genera) have been lost (Table 1). Theminnow family (Cyprinidae) has lostmore members (16 taxa) than any otherfamily of fishes. A large number ofsalmonids and cyprinodontids, seventaxa each, are also extinct. Regions thathave lost a substantial proportion oftheir native fish fauna are the Great

Lakes (three species and two subspe-cies), Great Basin (one species and sixsubspecies), Parras Valley (three spe-cies including one monotypic genus),Valley of Mexico (one genus and threespecies), and Rio Grande (two speciesand two subspecies).

For most extinct fishes, more than asingle factor was responsible for theirdecline and subsequent extirpation (Ta-ble 2). The most common cause ofextinction was habitat loss, a contrib-uting factor for at least 73% of the 40taxa. The second most common causalfactor was the effects of introducedspecies (cited for 68% of the 40 taxa).These factors were followed by chem-ical alteration or pollution (38%), hy-bridization (38%), and overharvesting(15%).

Unfortunately, the rate of extinctionof North American fishes is likely toincrease. Of the 40 taxa treated here,19 have disappeared since 1964. Pres-ent laws and recovery activities appearinadequate to stem the increasing tideof endangered fishes (Williams et al.1989). We are especially concerned forthe fate of the Mexican ichthyofauna,where the fishes are poorly known,environmental protection laws are in-adequate, and the human populationis rising at a staggering rate.

AcknowledgmentsWe greatly appreciate the assistance

of E. Diaz-Pardo, K. E. Hartel, S. L.Jewett, G. R. Smith, and T. N. Todd.Dean A. Hendrickson assisted in prep-aration of the resumen. Funding forillustrations by Sara V. Fink was kindlyprovided by Mrs. Virginia Ullman. Ad-ditional support was provided by aFaculty Assistance Fund grant to Rob-

ert Miller and by a grant from theCalifornia-Nevada Chapter of AFS. Theillustrations of the harelip sucker andsilver trout are courtesy of the U.S.National Museum of Natural Historyand are deposited in the files of theDivision of Fishes, Department of Ver-tebrate Zoology, Smithsonian Institu-tion. The U.S. Fish and Wildlife Serviceprovided publishing funds.

ReferencesAlvarez, J., and L. Navarro G. 1957. Los

peces del Valle de Mexico. Sec. de Marina.Dir. Pesca Indus. Conex., Mx. 1957:1-62.

Andreasen, J. K. 1975. Systematics andstatus of the family Catostomidae insouthern Oregon. Doctoral dissertation.Oregon State Univ., Corvallis. 76 pp.

Armstrong, J. G., and J. D. Williams. 1971.Cave and spring fishes of the southernbend of the Tennesee River. J. Tenn.Acad. Sci. 46:107-115.

Ayres, W. 0. 1862. Occurrence of freshwa-ter fishes in San Francisco Bay. Proc. Calif.Acad. Nat. Sci. 2:163.

Bailey, R. M., and C. E. Bond. 1963. Fournew species of freshwater sculpins,genus Cottus, from western North Amer-ica. Occas. Pap. Mus. Zool. Univ. Mich.634:1-27.

Bailey, R. M., and W. J. Richards. 1963.Status of Poecilichthys hopkinsi Fowler andEtheostoma trisella, new species, percidfishes from Alabama, Georgia, and SouthCarolina. Occas. Pap. Mus. Zool. Univ.Mich. 630:1-21.

Barbour, C. D., and R. R. Miller. 1978. Arevision of the Mexican cyprinid fish ge-nus Algansea. Misc. Publ. Mus. Zool. Univ.Mich. 155:1-72.

Behnke, R. J. 1979. Monograph of the na-tive trouts of the genus Salmo of westernNorth America. USDA Forest Service,Lakewood, CO. 163 pp.


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Table 1. Recently extinct fishes of North America. Location codes follow those given by Williams et al. (1989). Dates of extinction areapproximate for most taxa (see text).

Common name, scientific name Location Year of extinction

Whiteline topminnow, Fundulus albolineatusParras characodon, Characodon garmaniYellowfin cutthroat trout, Oncorhynchus clarki macdonaldiHarelip sucker, Lagochila lacera

Snake River sucker, Chasmistes murieiUtah Lake sculpin, Cottus echinatusSilver trout, Salvelinus agassiziStumptooth minnow, Stypodon signiferParras pupfish, Cyprinodon latifasciatusJune sucker, Chasmistes 1. liorusAlvord cutthroat trout, Oncorhynchus clarki ssp.Pahranagat spinedace, Lepidomeda altivelisGrass Valley speckled dace, Rhinichthys osculus reliquusMiller Lake lamprey, Lampetra minimaAsh Meadows poolfish, Empetrichthys merriamiDeepwater cisco, Coregonus johannaeLas Vegas dace, Rhinichthys deaconiThicktail chub, Gila crassicaudaPahrump Ranch poolfish, Empetrichthys latos pahrumpMaravillas red shiner, Cyprinella lutrensis blairiRaycraft Ranch poolfish, Empetrichthys . concavusRio Grande bluntnose shiner, Notropis simus simusDurango shiner, N. aulidionLake Ontario kiyi, Coregonus kiyi orientalisBlackfin cisco, C. nigripinnisMexican dace, Evarra bustamanteiMexican dace, E. eigenmanniMexican dace, E. tlahuacensisIndependence Valley tui chub, Gila bicolor isolataAmeca shiner, Notropis amecaeClear Lake splittail, Pogonichthys ciscoidesBlue pike, Stizostedion vitreum glaucumTecopa pupfish, Cyprinodon nevadensis calidaeMonkey Spring pupfish, Cyprinodon sp.Amistad gambusia, Gambusia amistadensisPhantom shiner, Notropis orcaLongjaw cisco, Coregonus alpenae

Banff longnose dace, Rhinichthys cataractae smithiSan Marcos gambusia, Gambusia georgeiOpal allotoca, Allotoca maculata

ALCOACO

AL, AR, IN, KY,GA, OH, TN, VA

WYUTNH

COACOAUT

NV, ORNVNVORNV

IL, IN, MI, WI; ONNVCANVTXNV

NM, TX; CHIDGO

NY; ONIL, IN, MI, WI; ON

DFEDFEDFENVJALCA

MI, NY, OH, PA; ONCAAZTX

NM, TX; CHI, TAMIL, IN, MI, NY,OH, PA, WI; ON

ABTXJAL

. 1980. A systematic review of thegenus Salvelinus. Pages 441-480 in E. K.Balon, ed. Charrs salmonid fishes of thegenus Salvelinus. W. Junk Publ., The Neth-erlands.

Bigelow, S. L. 1850. Observations on someof the habits of Salmo fontinalis. Boston J.Nat. Hist. 6:49.

Bond, C. E., and T. T. Kan. 1973. Lampetra(Entosphenus) minima n. sp., a dwarfedparasitic lamprey from Oregon. Copeia1973:568-574.

Campbell, R. R. 1985. Rare and endangeredfishes and marine mammals of Canada:COSEWIC fish and marine mammals

subcomittee status reports: II. Can. Field-Nat. 99:404-408.

. 1988. Rare and endangered fishesand marine mammals of Canada: CO-SEWIC fish and marine mammal subcom-mittee status reports: IV. Can. Field-Nat.102:81-86.

Chernoff, B., and R. R. Miller. 1986. Fishesof the Notropis calientis complex with akey to the southern shiners of Mexico.Copeia 1986:170-183.

Chernoff, B., R. R. Miller, and C. R. Gil-bert. 1982. Notropis orca and Notropis simus,cyprinid fishes from the American South-west, with description of a new subspe-

cies. Occas. Pap. Mus. Zool. Univ. Mich.698:1-49.

Contreras-Balderas, S. 1969. Perspectivesde la ictiofauna en las zonas aridas delnorte de M6xico. Pages 293-304 in Mem.Simp. Intern. sobre el aumento de lapr6duccion de alimentos en zonas aridas.ICASALS Pub. 3.---. 1978. Speciation aspects and man-

made community composition changes inChihuahuan desert fishes. Pages 405-431in R. H. Wauer and D. H. Riskind, eds.Transactions of the symposium on thebiological resources of the ChihuahuanDesert region. U.S. Natl. Park Serv.

November - December 1989

1900190019101910

192819281930193019301935194019401950195319531955195519571958196019601964196519671969197019701970197019701970197019711971197319751978

198219831984

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Table 2. Summary of factors responsible for decline and eventual loss of extinct fishes. A plus sign (+) indicates that the factor at leastcontributed to the extinction; a minus sign (-) indicates no known effect.

Physical habitat Introduced Chemical alterationScientific name alteration species or pollution Hybridization Overharvesting

Lampetra minimaCoregonus alpenaeC. johannaeC. kiyi orientalisC. nigripinnisOncorhynchus clarki macdonaldi0. clarki ssp.Salvelinus agassiziCyprinella lutrensis blairiEvarra bustamanteiE. eigenmanniE. tlahuacensisGila bicolor isolataG. crassicaudaLepidomeda altivelisNotropis amecaeN. aulidionN. orcaN. simus simusPogonichthys ciscoidesRhinichthys cataractae smithiR. deaconiR. osculus reliquusStypodon signiferChasmistes . liorusC. murieiLagochila laceraCyprinodon latifasciatusC. nevadensis calidaeC. sp.Empetrichthys latos concavusE. 1. pahrumpE. merriamiFundulus albolineatusAllotoca maculataCharacodon garmaniGambusia amistadensisG. georgeiStizostedion vitreum glaucumCottus echinatus

_ _ + - _

++++++++

+++

+++++++++

+

++++++++

++

+++++++++++++++

++

++++

+

++++ +

+++

++

++

++ +

++

+ ++

++

++

+++

+++

+++

+++ +

Trans. Proc. Ser. No. 3.Cook, K. D. 1979. Fish population study of

West Cache Creek with emphasis onsearch for the Wichita spotted bass, Mi-cropterus punctulatus wichitae Proc. Okla.Acad. Sci. 59:1-3.

Day, D. 1981. The doomsday book of ani-mals, a natural history of vanished spe-cies. Viking Press, New York. 288 pp.

Deacon, J. E., G. Kobetich, J. D. Williams,S. Contreras-Balderas, and others. 1979.Fishes of North America endangered,threatened or of special concern: 1979.Fisheries (Bethesda) 4(2):29-44.

Deason, H. J. 1936. Morphometric and lifehistory studies of the pike-perches (Sti-

zostedion) of Lake Erie. Doctoral disser-tation. Univ. Michigan, Ann Arbor. 368pp.

Diamond, J. M. 1987. Extant unless provenextinct? Or, extinct unless proven extant?J. Conserv. Biol. 1:77-79.

Edwards, R. J., H. E. Beaty, G. Longley,D. H. Riskind, D. D. Tupa, and B. G.Whiteside. 1984. The San Marcos recov-ery plan. U.S. Fish Wildl. Serv., Albu-querque, NM. 109 pp.

Evermann, B. W., and 0. P. Jenkins. 1889.Notes on Indiana fishes. Proc. U.S. Natl.Mus. 11:43-57.

Gilbert, C. H. 1893. Report on the fishesof the Death Valley expedition collected

in southern California and Nevada in1891, with descriptions of new species.N. Am. Fauna 7:229-234.

Goldman, C. R., and R. G. Wetzel. 1963.A study of the primary productivity ofClear Lake, Lake County, California.Ecology 44:283-294.

Greene, W. S. 1937. Colorado trout. DenverMus. Nat. Hist. Popular Ser. 2. 48 pp.

Heckmann, R. A., C. W. Thompson, andD. A. White. 1981. Fishes of Utah Lake.Great Basin Nat. Mem. 5:107-127.

Hopkirk, J. D. 1973. Endemism in fishes ofthe Clear Lake region. Univ. Calif. Publ.Zool. 96:1-160.

Hubbs, C. 1957. Distributional patterns of


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Appendix. Erroneously reported extinctions of fishes from North American waters. References indicate the first author or authors thatlisted the taxon as extinct. The Status shows the correct classification of the taxon as follows: E = endangered, T = threatened, SC =of special concern as determined by Williams et al. 1989; C = common if the taxon was valid but not listed by Williams et al. 1989; I =invalid if it is not recognized as a valid taxonomic entity.

Taxon References Status

Greenback cutthroat trout, Oncorhynchus clarki stomias Greene 1937 TSan Gorgonio trout, 0. evermanni USDI 1966 IRoyal silver trout, 0. mykiss regalis La Rivers 1962 IEmerald trout, 0. m. smaragdus La Rivers 1962 ISaltillo roundnose minnow, Dionda episcopa punctifer Day 1981 IClear Lake minnow, Endemichthys grandipinnis Opler 1977 ISummer Basin tui chub, Gila bicolor ssp. ONHDB 1985 EBig Spring spinedace, Lepidomeda mollispinis pratensis Miller and Hubbs 1960 EBlackmouth shiner, Notropis melanostomus Layne 1976 TRio Conchos sucker, Catostomus conchos Contreras-Balderas 1978 TShortnose sucker, Chasmistes brevirostris Day 1981 EBigmouth lakesucker, C. stomias Andreasen 1975 IDusky mountain sucker, Pantosteus sp. Miller 1961 ISmoky madtom, Noturus baileyi Opler 1977 EYellowfin madtom, N. flavipinnis Taylor 1969 TLeon Springs pupfish, Cyprinodon bovinus Hubbs 1957 EShoshone pupfish, C. nevadensis shoshone Pister 1974 EOwens pupfish, C. radiosus Miller 1961 EBig Bend gambusia, Gambusia gaigei Matthiessen 1959 ESpring pygmy sunfish, Elassoma sp. Ramsey 1976 TWichita spotted bass, Micropterus punctulatus wichitae Cook 1979 ISharphead darter, Etheostoma acuticeps Bailey and Richards 1963 SCTrispot darter, E. trisella Bailey and Richards 1963 TSnail darter, Percina tanasi Robins et al. 1980 TTilefish, Lopholatilus chamaeleonticeps Lucas 1891 C

Texas fresh-water fishes. Southwest. Nat.2:89-104.

Hubbs, C., and B. L. Jensen. 1984. Extinc-tion of Gambusia amistadensis, an endan-gered fish. Copeia 1984: 529-530.

Hubbs, C., and A. E. Peden. 1969. Gambusiageorgei sp. nov. from San Marcos, Texas.Copeia 1969:357-364.

Hubbs, C. L. 1940. Fishes from the Big Bendregion of Texas. Trans. Tex. Acad. Sci.23:3-12.

Hubbs, C. L., and R. R. Miller. 1948. Thezoological evidence: correlation betweenfish distribution and hydrographic historyin the desert basins of western UnitedStates. Pages 17-166 in The Great Basinwith emphasis on glacial and postglacialtimes. Bull. Univ. Utah. Vol. 38.

- . 1972. Diagnoses of new cyprinidfishes of isolated waters in the Great Basinof western North America. Trans. SanDiego Soc. Nat. Hist. 17:101-106.

Hubbs, C. L., R. R. Miller, and L. C. Hubbs.1974. Hydrographic history and relictfishes of the north-central Great Basin.Mem. Calif. Acad. Sci. 7:1-259.

Hunt, E. G., and A. I. Bischoff. 1960.Inimical effects on wildlife of periodicDDD application to Clear Lake. Calif. FishGame 46:91-106.

Imlay, R. W. 1936. Evolution of the CoahuilaPeninsula, Mexico. Part IV. Geology of

the western part of the Sierra de Parras.Bull. Geol. Soc. Am. 47:1091-1152.

Johnson, J. E., and C. Hubbs. 1989. Statusand conservation of poeciliid fishes. Pages301-317 In G. Meffe and E Snelson, eds.Ecology and evolution of livebearingfishes (Poeciliidae). Prentice Hall, NewYork.

Jordan, D. S. 1882. Report on the fishes ofOhio. Geol. Surv. Ohio 4:735-1002.-___. 1885. Note on Mr. Garman's paper

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. 1890. Report of explorations madeduring the summer and autumn of 1888,in the Allegheny Region of Virginia,North Carolina, Tennessee and in westernIndiana, with an account of the fishesfound in each of the river basins of thoseregions. Bull. U.S. Fish. Comm. 8:97-173.

-- . 1922. The days of a man, Vol. 1.World Books Co., New York. 710 pp.

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Jordan, D. S., and C. H. Gilbert. 1881.Notes on a collection of fishes from UtahLake. Proc. U.S. Natl. Mus. 3 (for 1880):459-465.

Juday, C. 1907. A study of Twin Lakes,Colorado, with special consideration ofthe food of the trouts. Bull. U.S. Bur.

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the biology of the Miller Lake lamprey,Lampetra (Entosphenus) minima. NorthwestSci. 55:70-74.

Kendall, W. C. 1914. The fishes of NewEngland. The salmon family. Part I: Thetrout or chars. Mem. Boston Soc. Nat.Hist. 8:103 pp.

Kirsch, P. H. 1896. A report upon investi-gations in the Maumee River basin duringthe summer of 1893. Bull. U.S. Fish.Comm. 14:315-334.

Koelz, W. 1929. Coregonid fishes of theGreat Lakes. Bull. U.S. Bur. Fish. 43 (for1927):297-643.

La Rivers, I. 1962. Fishes and fisheries ofNevada. Nevada State Fish and GameComm. 782 pp.

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Lucas, F. A. 1891. Animals recently extinctor threatened with extermination, as rep-resented in the collections of the U.S.National Museum. Rept. U.S. Natl. Mus.1888-1889. 41 pp.

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---. 1961. Man and the changing fishfauna of the American Southwest. Pap.Mich. Acad. Sci. Arts. Lett. 46:365-404.

---. 1963. Synonymy, characters, andvariation of Gila crassicauda, a rare Cali-fornian minnow, with an account of itshybridization with Lavinia exilicauda. Calif.Fish Game 49:20-29.

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extinctions of north american fishes during the past century

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