The Systematics of the Prickly Sculpin, Coitus asper Richardson,a Polytypic Species

Part I. Synonymy, Nomenclatural His tory, and Distribu tion'

RICHARD J. KRE] SA2

ABSTRACT : Th e prickly sculpin, Cottus asper, is a geographically widespread,polytypic species characteristically represented by very pr ickly, nonmigratory, fresh­water spawning " inland" forms, and less prickly, catadromous, brackish-waterspawni ng "coastal" for ms. Part I , the first contri bution in a series on the systematicsof this species, presents a synonymy complete for the period 1836-1936, with aresume of the most important citations from 1936 to 1965. A nomenclatural historyof the species is given. The distributional range is listed and also presented inillustration.

THE PRICKLY SCULPIN ranges over about 3,000miles of Pacific N orth Temperate coastline andinland as far as 300 miles. The species existsin two primary modes of morph ological vari­ability: one, a nonmi gratory, fresh-w aterspawner , has extensive squamation on certainregions of the body; the other, a catadromous,brackish-water spawner, has little or none.Pr ickly sculpin eggs are spawned naturally inenvironments which are known to vary in atleast one major factor, i.e., salinity. The mar­phological, behavioral, and ecological varia­tions existing within this species make it anexcellent subject for systematic analysis.

SYNONYMY

It has been 130 years since Cattus asper wasfirst described by Sir John Richardson . Priorto the present study, regional systematic treat­ments of this widespread species resulted in aproliferation of generic and specific taxa, allreferable to C. asper. Early revisionary work byGirard in 1851 and 1852 was incompletebecause of lack of specimens. Recent reg ional

1 The data for this paper are taken from a thesissubmitted in partial fulfillm ent of the requir ementsfor the degree of D octor of Philosophy at the Uni­versi ty of Britis h Columbia. Manuscript receivedMarch 23, 1966.

2 Institute of Fisheries, University of BritishColumbia, Vancouver, Canada. Present address: D e­partment of Anatomy, College of Physicians and Sur­geons, Columbia University, New York.

works (Robins and Miller, 1957 ; McAll ister,1957; McAllister and Lindsey, 1959 ; and Bond ,1963) have included the species as part of ageographical area or river drainage system, butthere has never been a comprehensive treatme ntof the species throughout its entire range. Inthe period 1836-1 936 , 41 reports of C. asperwere cited in the literature. In the same period,32 addi tiona l citations occurred which wereeither misidentifications or synonyms properlyreferable to C. asper. In none of the systematictreatments published since the original descrip­tion in 1836 has there been a synonymy con­taining more than 7 citations. McAllister(1 957) listed 15 citations in h is unpublishedM.A. thesis.

Th e present synonymy consists of 73 citationspubli shed during the period 1836-1936, and isthought to be comp lete for that period. In thepast 30 years, the species has been cited in­cidentally in so many fishery journals andpubli cations that only the major systematic, orotherwise noteworth y, citations have been re­por ted in the remaining synonymy.

Couus asper Richardson, 1836

Cottus asper. Richardson, 1836 :295, pI. 95,fig. 1 (o riginal description and figure ; Colum­bia R. ; collected by Dr. Gairdner, proba blynear Fort Van couver, Washington Territory) .Storer, 1846a:260 , and 1846b :8 (northwesterncoast of N . America) . Girard, 1850 :409, and1851a :189 ( discusses propriety of present

241

242

nomenclature) . Eigenmann, 1895: 118 (ab un­dant in Fraser system from tidewater to1,900 ft ; Mission, Sicamous, Kamloops, andGriffin L., British Columbia ; and Umatilla,Oregon). Gilb ert and Evermann, 1895:201(description; comparison with Sacramento Rform; Walla W alla R. at W allula, and LakeWashing ton, Washington). Seale, 1896 :854(Lake Washington) . Gilbert, 1896 :418 (de­scription; stream entering Departure Bay, Van­couver Island ). Jordan and Evermann, 1896:439 (synonymy; streams of the Cascade Range,from Vancouver Island to Oregon ) . Gilbert,1898 :1 (Co lumbia R. ; notes absence in Kla­math R.). Jordan and Evermann, 1898 :1944(descript ion; synonymy; W alla W alla ; De­par ture Bay ; about Port Townsend ; streams ofthe Cascade Range, from Vancouver Island toOregon ) . Evermann and Meek, 1898:83 ( LakeWashington) . Meek, 1899 :231 ( Lake Souther­land , Olympic Peninsula, W ashington ) . Jordan,1905:445 (strea ms of the Pacific coast ) .Snyder, 1905:337 (description; affinities;habitat preference; San Franciscito, Madera,San Antonio, Guadalupe, Coyote, and Alamedacreeks, all flowing into San Francisco Bay) .Evermann and Goldsborough, 1907a:306(characters; prickling descriptions ; Deep Bay,Naha R , and Steelhead Cr., Loring, Alaska;H unter Bay, Yes Bay, and McDonald L.,Alaska) . Evermann and Goldsborough , 1907b :110 ( Fraser R. at Mission, Shuswap L. at Sica­mous, Th ompson R. at Kamloops, and GriffinL. ) . Rutter, 1908 :145 (Cottopsis parvlIs firstplaced in synonymy with asper,o synonymy, inpart, except Uraniden semiscaber (sic) centro­pleura Eigenmann and Eigenmann; summary of9 localities in Sacramento R. system) . Snyder,1908a :269 (Russian R., California) . Snyder,1908b :184 (characters; prickling description ;summary of 41 localities: from Lake Was hing­ton, Columbia and Sacramento R., and riverbasins in between) . Ni chols, 1909: 172 (h eadof Chilkoot L., Alaska). Evermann and Lati­mer, 1910:138 (4 localities in Marin Co., and2 localities in San Francisco Bay, Califo rnia ; 12localities in Olympic Peninsula, W ashington) .Snyder, 1913:72 (characters; Pajaro R., Cali­fornia). Snyder, 1916 :381 (Papermill andWa lker creeks, California). Kermode, 1917 :20( Hancevi lle, British Columbia [Chik otin R.

PACIFIC SCIENCE, Vol. XXI, April 1967

drainage] ) . Jordan, 1919 :249 (designatesCottus aspel' Richardson as the orthotype ofCottopsis Girard ) . Bean and We ed, 1920: 76(mouth of Fraser R.) . Hubbs, 1921:7 ( re­identification of San Luis Cr., Califo rnia, speci­mens misidentified by Jordan as C. gtllosllS[ 1895:141] ; range extension to Ventura R.,Californ ia) . Fowler, 1923: 282 (Hancevi lle,and Shawnigan L., British Columbia; Shawni­gan L. specimen misidentified by Kermode[1909 :87] as Uranidea glllosa). Crawford,1927 :177 (strea ms flowing into Puget Sound) .Schultz , 1929:48 (l isting only) . Schultz , 1930 :14 (most streams and lakes of western Was h­ington ) . Jordan, Evermann, and Clark, 1930 :383 (synonymy; streams of the Cascade Range,southeastern Alaska to Oregon ; south to Sacra­mento R.). Kermode, 1931:19 (Cowichan L.,Vancouver Island ) . Evermann and Clark, 1931 :56 (summary of 32 recorded localities in Cali­fornia). Taft , 1934 :251 (spawning migra­tion). Schultz and DeLacy, 1936a:128 (syn­onymy ; coastal streams from Alaska to VenturaCo., California ; fresh water and brackish water;review of most records from Puget Sound toOregon, and add ition of 26 more localit ies).Schultz and DeLacy, 1936b :213 ( additionalsynonymy ; 3 new localities) . Schultz, 1936: 179(keys to species of Cottns; coastal streams fromAlaska to Ventura Co., Califo rnia; fresh-waterand brackish) . Dymond, 1936 :71 (descr iption;throughout southwestern British Columbia, in­cluding southern Vancouver Island ; 16 locali­ties listed ) . Sumner, 1942: 1- 25 ( common intidewater areas along the Oregon coast). Hubbsand Wa llis, 1948 :141 (identification of "Cot­tns sp." recorded by D ill, 1946:5 4). Baileyand Dimick, 1949 :14 (comparison with Cottnshllbbsi). Shapovalov and D ill, 1950 :387 (li st­ing only) . Wi limovsky, 1954: 285 (sou theastAlaska to Califo rnia) . Robeck, et al., 1954:B-65 (Co lumbia R , above T rinid ad, Was hing­ton ; cited as "p rickly sculpins Cottus sp.," thesemay include Cottus rbotbeus in part) . Lindsey,1956 :777 (Pacific Slope of N. America fromAlaska to California; British Columbia main­land from Columbia, Fraser, and Skeena sys­tems, Stikine R. headwaters, and Peace R.[Summit L., Heart L., Angusmac Cr., andMcLeod L.l) . Robins and Miller, 1957 :229(Cottopsis pan'lls again removed from syn-

Systematics of Prickly Sculpin, I-KREJ sA

onymy of C. gnlo sus ) , Lindsey, 195 7:657(Britis h Columbia : Columbia R. ; Fraser R.;Skeena R.; coast drainages south of Skeena ;N ass R.; Stikine R. ; Peace R.) . Wilimovsky,1958:62 (key to species in Alaska ; southeastAlaska to California) . Shapovalov, Dill, andCordone, 19 59:1 73 (listing only) . Carl, Clem­ens, and Lindsey, 1959 :158 (d escription;Pacific drainages from Chilkoot 1. , Alaska, toVentura R., California. In British Columbia :lakes and rivers of the Columbia, Fraser, Dean,Skeena, Nass, and Stikine systems; coastalrivers of the mainland and Vancouver Island,and Queen Charlotte Islands; headwaters ofPeace R. system from Summit 1. to McLeod1. ) . McAllister and Lindsey, 1959:70 ( de­scription; synonymy ; intraspecific variation;localities as in Carl, Clemens, and Lindsey,1959) . McAllister, 1960 :42 (collection in saltwater, Pt. Atkinson, British Columbia) . Bond,1961 :36 (key to species in Oregon ; pricklingvariation; Columbia R. drainage) . Bond, 1963:79 (synonymy; life history observations; oxy­gen, temper ature , and salinity tolerance ofadults ; fish associates; habitat preference; lists35 new collection localities in coastal Oregon,30 localities from Columbia R. drain age inOregon, and also some lakes in the southwestcorner of Rainier National Park, Washington) .Bailey and Bond, 1963: 19 (recognition ofseveral species groups within Cottus in westernN. Amer ica; characters and list of species inthe "ns per species group"). Krejsa , 1965:1- 109(synonymy ; nomenclatural history; distribution ;life history ; morphological variation; salinitytolerance of eggs ; phylogeny of C. asper andclosely related species) . Bohn and Hoar, 1965 :977 (salinity effects on iodine metabolism;physiological divergence of inland and coastalC. asper) .

Centridermicbtb ys asper. Richardson,1844 :76 (River Oregon [= Columbia R.]) .Gunther, 1860: 170 ( descript ion ; synonymy;fresh waters of the Oregon and WashingtonTerritories). Lord 1866a:130 (life history ;spawning behavior , in part ; Puget Sound ;"streams flowing throug h the Sumass andChilukweyuk prairies" [Sumas and ChilliwackR. ?], British Columbia; in part, all streamseast and west of the Cascades) .

243

Cottopsis asper. Girard, 1851b:303 (intro­duction of Cottopsis gen. nov. ; synonymy;limited to River Oregon [= Columbia R.] ) .Girard, 1851c:185 (not seen) . Gira rd, 1852 :61(definition of Cottopsis gen. nov., based onRichardson's description of Cottus asper; syn­onymy; Columbia R. ). Girard, 1859 :51 (de­scription; synonymy; based on 8 specimensfrom Astoria and Fort Dalles, Oregon, andFort Steilacoom, Puget Sound, Was hingtonTerritory) . Suckley (in Cooper and Suckley,1859 ) 18 59:351, and Suckley, 1860 :35 1 (de­scription; synonymy; small fresh-water streamsemptying into Puget Sound; Ft. Steilacoom ;and Columbia R., 200 miles above mouth ) .Jord an and Jouy, 1882 :5 (Puget Sound; Co­lumbia R. ; Mare Island and Sacramento R.,California) .

Uranidea aspera. Jordan and Gilbert,1883 :694 (d escription; synonymy; streamswest of the Sierra N evada and Cascade Moun­tains) . Jordan, 1885: 110 (subgenus Cottopsisand a list of species therein) .

T racbidermus ricbardsonii. Heckel, 1840 :162(synonymy; Columbia R.) , (not Cottus richard­soni of Agassiz, 1850). Note: Girard, 185 2:62,erred in repor ting the date of H eckel's paperas 183 7, and in the spelling of Tracbidermns.

Cottopsis parvm . Gira rd, 1856b:144 (orig­inal description ; Presidio on San FranciscoBay, California) . Girard, 1857: 11 (descrip­tion ; Presidio, and Monterery, California) .Girard, 1859:54 (descri ption ; synonymy ;Monterey, Presidio, Fort Reading, and Peta­luma, California). Cooper, 1868 :492 (listi ngonly) . Jordan, 1877: 5 (as the young of Cottop­sis asper) .

Centriderniicbtb ys parvtls. Gunther, 1860:170 (description; synonymy; fresh waters ofCalifornia) . Lord, 1866b:352 (lis ting; "fre­quenting the same localities as . . ." Centri­dermicbtbys asper) .

Urauidea aspera var. parvm. Jordan andGilbert , 1883:694 (Sacramento R. forms) .

Cottus gttloStls parvtfs. Jordan and Ever­mann, 1898 :1945 , and Jordan, Evermann , andClark, 1930 :383 (Cottopsis parvm: Monterey,Presid io, Fort Reading, and Petaluma, Cali­fornia) .

244

Uranidea gulosa, in part. Jordan and Gilbert,1883 :695 (misidentifications: all specimensfrom "Vancouver' s Island" and probably thosefrom "about Port Townsend," d . Jord an andEvermann, 1898:1 944) . Kermode, 1909 :87(li sting only; misidentification: Shawnigan L.,Vancouver Island , d. Fowler, 1923 :282) .

Cottus gulosus, in part. Jordan, 1895:141(mis identification : San Luis Cr., near Avila,California, d. Hubbs, 1921 :7) . Jordan andEvermann, 1898: 1945 (m isidentification : allspecimens from San Franciscito Cr., Santa ClaraCo., Californ ia). Jordan, Evermann, and Clark,1930: 383 (probable misidentifications: speci­mens from Loring and Boca de Quadra,Alaska) . Evermann and Clark, 1931 :57 (mis­identifications : Presidio, Monterey, Fort Read­ing ' Petaluma, and San Luis Cr., California) .Evermann and Clark, 1931:12, 13 (misidenti­fications : Monterey, Presidio, Fort Reading, andPetaluma) . Bean and W eed, 1920 :76 ( ques­tionable identification: 4 specimens fromVictoria, Vancouver Island, Brit ish Columbia) .W ilimovsky, 1954: 285 (doubts validity ofsoutheast Alaska record) .

Centrid ermicbtb ys gllloslls. Lord , 1866 b:352 (l isting; " frequenting the same localitiesas . . ." Centridermicbtbys asper) .

Coitus sp. Dill, 1946 :54 (San Joaquin R.,near Fr iant, Californi a; identification as asperby Hubbs and W allis, 1948 :141) .

N OMENCLAT URAL H ISTORY

The specific name asper is currently wellfounded in the genus Cottus, to which it wasoriginally designated by Richardson in 1836.But, as shown in the preceding synonymy, thebinomen was extremely unstable for the first100 years after its int roduction. After an initialper iod of uncertainty regardin g its affinity tomarine or to fresh-water Cottoids, three mainnomenclatural difficulties are encount ered : theoften-rep eated misidenti fication as Cottus glllo­SlIS ( Girard); the failure to recognize thatCottopsis parvlIs and Cottns asper are con­specific; and the failure to recognize the specificrelationship of asper to other species in thegenus Coitus.

PACIFIC SCIENCE, Vol. XXI, April 1967

The almost immediate placement of asperinto T racbidermus by Heckel (1 840) , and theninto the synonymous Centridennicbtbys byRichardson ( 1844) , reflects the early opinionthat asper was more closely allied to the marineCottoids. Girard (1851b, 1852) recognized itsaffinities with the fresh-water genus Cottus, butdistinguished it from that genus by erecting thegenus Cott opsis, based on the presence ofpalatine teeth and the "skin beset with prickles,instead of being smooth and scaleless." Lackingany specimens, Girard defined Cott opsis on thebasis of Richardson's original description but,on p. 63, where he quoted Richardson' s entirediscussion of prickles (p . 295), he misquotedRichardson by attributing to him the statement,"There are no scales." Girard 's lack of speci­mens proved unfortun ate since soon thereafter(1856b) he named and described CottopsisparvlIS from the Presidio (in San Francisco) ,California, comparing it not with C. asper butwith Cottopsis gllloslls Girard , also newlydescribed ( 1856a ) from the San Joaquin R.,California. In his later report (185 9), Girardhad 8 specimens of C. asper in his possession,from the Columbia R. and Puget Sound.Obviously he again failed to recogn ize the con­specificity of asper and parvlls, and he followedRichardson's original description rather thancomparing them with specimens of parvllS,which he seems to have reserved for com­parison with guloms.

Jordan ( 1877) referred, in passing, to Cot­topsis parvlls as the young of C. asper. Jordanand Jouy ( 1882) , however, l'sted specimens ofCottopsis asper from Mare Island and Sacra­mento R., California, and from Puget Soundand the Columbia R. Less than a year later ,Jordan and Gilbert ( 1883) placed asper inthe genus Uranidea DeKay, subgenus Cottopsis,based on the presence of palatine teeth and thegill membranes being broadly united to th eisthmus . In the same report, they referred tothe Sacramento R. form of U. aspera as "var.parous, smaller in size, paler in color and withthe interorbital space concave, narrower thaneye."

Eigenmann (1895) used the valid name todescribe specimens from the Fraser and Colum­bia rivers, as also did Gilbert and Evermann(1895), who suggested that the nominal spe-

Systematics of Prickly Sculpin, I-KRE]SA

cies was separable "at least subspecifically fromthe Sacramento River form." Seale (1896) andGilbert ( 1896) used the valid name for north­ern specimens. But, obviously, Jordan (18 95)and Jordan and Evermann (1896) still thoughtin terms of a distinct Californian species (gll/o­SIlS ) and a distinct northern species (asper).Jordan misidentified a specimen of asper f romSan Luis Cr., near Avila, Calforn ia, as gll/OSIIS.Jordan and Evermann listed the range of thenominal species from Vancouver Island toOregon, and of gu/osus, from California CoastRange streams and inland in the San Joaquin R.Gilbert (1 896) referred to Cottus asper of theColumbia and Coitus gu/osus of the Sacramentoas "two species so extremely similar that it isdifficult to distinguish them." Jordan and Ever­mann (1 898) repeated the suggestion of Gil ­bert and Evermann that the nominal species isseparable, at least subspecifically, from theSacramento R. form, "Coitus gttfosllS."

That Jordan and Evermann perceived neitherthe conspecific relationship of parvus to aspernor the limits of the valid species Cot/us gll/o­SIIS becomes more fully evident on the nextpage (p . 1945) of their 1898 report. Theirdescript ion of Coitus gulosus (Girard) is takenfrom misidentified specimens of C. asper col­lected in San Franciscito Cr., Santa Clara Co.,California. Th ese were large specimens " 3 to 7inches in length" and, most significantly, thecount for anal rays is given as "A. 16 to 18."Both of these characters separate asper fromgu/OStls . Furthermore, they include Cottopsisparvtls Girard , from Monter ey, the Presidio,Fort Reading, and Petaluma, California, in thesynonymy of gttfosus.

Snyder (1905) collected and correctly identi­fied Coitus asper from the same locality, SanFranciscito Cr. He was probably the only oneof his time to understand and explain the truerelationships of nsper, parvus, and gu/osus. Onp. 337, he stated :Recent authors have identified the common Sacra­mento form which represents the Coitus asper of theColumbia River with the Cottopsis gulosus of Girard.Th ey have sometimes considered the Sacramento formas identical with C. asper and have placed the namegil/oms in the synonymy of the latter. At other timesthey have considered the species as a slightl y differ­entiated form wort hy of recognit ion in nomencla ture,and have used the name gil/oms to designa te it. Th e

245

former view concerning the species is probably cor­rect. Th e association of the name gulasus with it,however , is without warrant. The latter belongs to aspecies easily distinguished from C. asper, differingnotably in having a much shor ter anal fin. Th ereare usually fewer dorsal spines and rays, a morelimited dis tribu tion of prickles, and an almost uni­form absence of palatine teeth. In C. asper the dor ­sal has 8 to 10 spines and 19 to 22 articulated rays,the anal 16 to 18 rays, whi le in C. gulosus the dorsalhas 7 to 9 spines, 17 to 18 rays, the anal 12 to 14rays.

Snyder then continues with a note on habitatpreference:

In its distr ibution C. asper appears to be largelyconfined to the lower courses of the streams, beingespecially abundant near tide water, while C. gulosusis found further up, where the water is clear and thecurrent rap id.

Rutter (1908) correctly placed Cottopsisparvus into the synonymy of Coitus asper, pre­sumably recognizing that the two were con­specific. However, he incorrectly synonymizedUranidea semiscabra centropleura Eigenmannand Eigenmann, which is properly referable toCoitus gu/osus.

Snyder was the first to consider a series ofspecimens of the nominal species throughout itsentire range, as then known, and, in the sameissue of the Bulletin of the Bureau of Fisheriesin which Rutter had correctly synonymizedpal'vlIS, he noted the extreme variation of pr ick­ling investment. While recognizing the varia­tion between streams, he also noted that theprickling variation is common among individ­uals from the same stream.

Although explicitly aware of Snyder's com­ments on asper and gttfosus, Evermann andGoldsborough (1907a) identified 16 specimensof gs losus from Loring and Boca de Quadra,Alaska. Th e reliability of these identificationsis questionabl e. Kerrnode's listing (1909) ofUraniden gu/osa from Shawnigan 1., VancouverIsland is probably a misidentification, sinceFowler (19 23:282) listed the same specimenas Cottns asper.

Snyder (1913, 1916) again recorded theoccurrence of C. asper and gu/oSt/s in differinghabitats of the same stream. Hubbs (19 21)recognized Jordan's earlier misident ification ofgulosns from San Luis Cr., California. He alsocommented on the variability of prickl ing inC. asper from several streams.

246

It would seem that with the accession ofSnyder 's insight into the problem, the validname was destined for stability. H owever,Jordan, Evermann, and Clark (1930) repeatedthe earlier error of Jordan and Evermann(1898) by including Cottopsis parVlIS as asynonym of Coitus gnlosus. In a similar man­ner , they also incorporated the error of Ever­mann and Gold sborough ( 1907a), previouslycited, by including the misident ified specimensof asper from Loring and Boca de Quadra,Alaska, in the list of records for gltlosttS. Inthe same work, Jordan and Evermann extendedthe range of asper (c ited in 1898 as: "streamsof the Cascade Range, from Vancouver Islandto Or egon" ) by appending the phrase "southto Sacramento River." Evermann and Clark( 1931) also retained Cottopsis parotrs in thesynonymy of C. gltlosttS and perpe tuated Jor­dan's misidentification of the San Luis Cr.gltlosttS, which Hubbs had correctly re-identi­fied as asper ten years before (192 1) .

Schultz and D el.acy's catalogue (193 6) in­cluded a comprehensive listing of Was hingto nand Oregon localities for C. asper. H owever,some remain doub tfu l since Schultz and DeLacyfrequently misidentified C. asper as gl/losttSand/or perplexes. Th ey also incorrectly rmin­tained the pr esence of C. gltloslts in Alaska.

Robins and Mill er (19 57) presumably over­looked the earlier citation of Rutt er ( 1908)and removed Cottopsis parvttS fro m the syn­onymy of gltlosttS, placing it in the synonymyof asper, supposedly for the first time.

McAllister and Lindsey (1959) first sug­gested the pro bable existence of "coastal" and"non-coastal" populations of COtt/IS asper onthe basis of morph ological and, perhaps, be­havioral differences.

Bond (1961) h inted at the possibility ofpolytypy in Couus asper when he stated in hiskey that the body is "well covered with pr ickles,especially in inland waters and in young in­dividuals from coastal waters." Bond ( 1963)gave the 'most comprehensive treatm ent yetrecorded for Cottus asper and 12 other speciesin the genus. Hi s study, however, was moreconcerned with interspecific rather than intra­specific relationships within the genus. Baileyand Bond (1963) ind icated their concern forthe supraspecific relationships within the genus

PACIFIC SCIENCE, Vol. XX I, April 1967

Coitus by their recognition of several speciesgroups, one of which is the "asper speciesgroup."

Krejsa ( 1965) offered morphological, be­havioral, and distributional evidence for genet:edivergence between "coastal" and "inland"populations of C. asper. Bohn and Ho ar(1 965) offered physiological evidence in sup­port of Krejsa's hypothesis. Unfor tunately,their brief introductory remarks regarding thelife h istories and prick ling patterns are some­what inaccurate, and therefore misleading, in­terpretat ions of Kr ejsa's unpublished thesis.These minor points will be clarified in a futurepublication.

DISTRIBUTION

Range. Pacific Slope drainage of N orth Am ­erica : coastal streams from Seward , Alaska, toVentura R., Californ ia ; lakes and streams ofthe Qu een Charl otte Islands and VancouverIsland; and all major Pacific drainages fromthe headwaters of the Stikine R. in BritishColumbia, to the Kern R., San Joaquin R.drainage, California. The following are ex­ceptions: Fraser R. in the area of the RockyMountain Trench, east of Prince George,British Columbia (a rea not yet collected);Kootenay Lake drainage of the Columbia R. inBritish Columbia; Upper Snake R. of the Co­lumbia R. drainage in W ashington and Ore­gon ; Middle Fork of the Wi llamette R. inOregon, above Oakridge ; Klamath R. Basinin Oregon ; and Sacramento R. drainages aboveLake Shasta, California. Arctic Slope drainageof N orth America: headwaters of the Peace R.in British Columbia: from Summit L. toMcLeod L., Crooked R. drainage; fromTacheeda L., Parsnip R. drainage; fromTchentlo L., Na tion R. drainage.

The presen t distributional range of Cottnsasper is illustrated in Figur e 1.

REFERENCES

BAILEY, R. M., and C. E. BOND. 1963. Fournew species of freshwater sculpins, genusCottns, from western N orth America. Occas.Papers Mus. Zool. Un iv. Mich. 634: 1-27.

- - - and M. F. DIMICK. 1949. Cottus

COTTUS ASPERPRESENT

DISTRIBUTION

Systematics of Prickly Sculpin, I-KRE]sA

iII

iIiiI

i!II;;;i;;iI;iIiiI

._ ......i.._ .._ .._ .._ .._ ._

\\\

\\ -,

\\

"\'.

\\ '1- --1

FIG. 1. Distributional range of Conus asper.

24 7

248

bubbsi, a new cottid fish from the ColumbiaRiver system in W ashington and Idaho.Occas. Papers Mus. Zool. Uni v. Mich.513:1-18.

BEAN, B. A., and A. C. W EED. 1920. N otes ona collection of fishes from Vancouver Island ,B. C. Trans. Roy. Soc. Canada ( 3) 13(1919) :69-83.

BOHN, A., and W. S. HOAR. 1965. The effectof salinity on the iodine metabolism ofcoastal and inland pr ickly sculpins, Cottusasper Richardson. Canad ian J. Zool. 43( 1965) :977-985.

BOND, C. E. 1961. Keys to Oregon freshwaterfishes. Agr . Expt. Sta., Oregon State Univ.,Tech. Bull. 58 :1-42.

- - - 1963. Distribution and ecology of fresh ­water sculpins, genus Coitus, in Oregon.Unpublished Ph.D. thesis, Un iv. Mich.186 pp.

CARL, C. G., W. A. CLEMENS, and C. C.LINDSEY. 1959. The fresh-water fishes ofBrit ish Columbi a. Brit ish Columbia ProvoMus. H andb. 5(3rd ed. rev.) :1- 192.

COOPER, J. G. 1868. Fishes. In : J. G. Cooperand T. F. Cronise, The N atural W ealth ofCaliforn ia. Chap. 7, pp . 487-498. H. H .Bancroft and Co., San Francisco.

CRAWFORD, D. R. 1927. Extension of the rangeof C OttJlS ale/dims. Copeia 1927(160) :177­178.

DILL, W . A. 1946. A preliminary report on thefishery of Millerton Lake, Californi a. Calif.Fish and Game 32( 2) :49- 70.

DYMOND, J. R. 1936. Some fresh-water fishesof British Columbia. Rept. British ColumbiaComm. Fish. (for 1935) :60-73.

EIGENMANN, C. H. 1895. Results of explora­tions in western Canada and northwesternUnited States. Bull. U. S. Fish Comm. 14(for1894) :101-132.

EVERMANN, B. W ., and H. W . CLARK. 1931. Adistributional list of the species of freshwaterfishes known to occur in California. Calif.Div. Fish and Game, Fish Bull. 35: 1- 67.

.- - - and E. L. GOLDSBOROUGH. 1907a. Thefishes of Alaska. Bull. U. S. Bur. Fish. 26(for 1906) :219-360.

___ - - - 1907b. A check list of thefreshwater fishes of Canada. Proc. BioI. Soc.Washington 20:89-1 20.

PACIFIC SCIENCE, Vol. XXI, April 1967

--- and H. B. LATIMER. 1910. On a col­lection of fishes from the Olympic Penin sula,together with notes on other Wes t Coastspecies. Proc. BioI. Soc. W ashington 23:131-140.

--- and S. E. MEEK. 1898. A report uponsalmon investigations in the Columbia RiverBasin and elsewhere on the Pacific Coastin 1896. Bull. U. S. Fish Comm. 17(for1897) :15- 84.

FOWLER, H. W. 1923. Records of W est Coastfishes. Proc. Acad. Nat. Sci. Ph iladelphi a75 :279-301.

GILBERT, C. H. , 1896. The ichthyological col­lections of the U. S. Fish CommissionSteamer Albatross during the years 1890 and1891. Rept. U. S. Comm. Fish and Fish.(for1893) , Appendix 6 :393-476.

--- 1898. Fishes of the Klamath Basin.Bull. U. S. Fish Comm. 17(for 1897) :1- 13.

--- and B. W . EVERMANN. 1895. A reportupon investigations in the Columbia RiverBasin, with descript ions of four new speciesof fishes. Bull. U. S. Fish Comm. 14 (for1894) :169-207.

GIRARD, C. 1850. A monograph of the fresh­water Cottus of N orth America. Proc. Am.Assoc. Adv. Sci. 2:409-411.

--- 1851a. On the genus C Ot/lIS Auct. Proc.Boston Soc. N at. Hist. 3 (1 848-1851) :183­190.

--- 1851b. Some additional observations onthe nomenclatur e and classification of thefishes of the genus Coitus. Proc. Boston Soc.N at. Hist. 3( 1848- 1851) :302-305.

--- 185 1c. Revision du genre Cottus desAuteurs. N ouv. Mem. Soc. Hel vet. Sci. N at.12(1851) :185. [Not seen.]

--- 1852. Cont ribut ions to the natural his­tory of the freshwater fishes of N orth Am­erica. I. A monograph of the cottoids. Smith­sonian Contr. Knowledge 3 (a rt. 3): 1-80.

--- 1856a. Descriptions of new fishes, col­lected by Dr. A. L. Heermann, naturalistattached to the Survey of the Pacific RailroadRoute, under Lieut. R. S. W illiamson, U.S.A.Proc. Acad. N at. Sci. Philadelphi a 7( 1854­1855) :129- 141.

--- 1856b. Observations upon a collectionof fishes made on the Pacific coast of theUnited States, by Lieut. W. P. Trowbridge,

Systematics of Prickly Sculpin, I-KRE]SA

U.S.A., for the museum of the SmithsonianInstitution. Proc. Acad. N at. Sci. Philadelphia7(1854-1855): 142-1 56.

--- 1857. Report upon fishes collected onthe Survey. 6(Part IV, No.1 ) :9-34. In :House of Representatives, 33rd Congr ., 2ndSess., Exec. Doc. 91, Reports of explorationsand surveys for a railroad route from theMississippi River to the Pacific Ocean. Vol.6 : Report of Lieut. H enry 1. Abbot uponexplorations for a railroad route from theSacramento Valley to the Columbia River,made by Lieut. R. S. Williamson, assisted byLieut. Henry 1. Abbot. 1854-1855. W ash­ington, D .C.

--- 1859. Fishes. In : General report uponthe zoology of the several Pacific railroadroutes. U. S. Pac. R. R. Expl. and Surv. 10(p art 4) :1-400. Also found as Part IV, 33rdCong., 2nd Sess., Senate Exec. Doc. 78.

GUNTHER, A. C. 1. G. 1860. Catalogue of theAcanthopterygian Fishes in the Collectionof the British Museum, Vol. 2, pp. 1-548.

HECKEL, J. J. 1840. Ichthyologische Beitragezu den Familien der Cottoiden , Scorpae­noiden, Gobioiden und Cyprinoiden. Ann.Wien. Mus. N aturgeschichte 2(1840) :143­164.

HUBBS, C. 1. 1921. N otes on Coitus asper andCotms aleuticus. Copeia 1921 (9) :7- 8.

--- and O. 1. WALLIS. 1948. Th e nativefish fauna of Yosemite N ational Park and itspreservation. Yosemite N ature N otes 27(1 2) :131- 144.

JORDAN, D. S. 1877. Contributions to NorthAmerican ichthyology, No.2. Bull. U. S.Natl. Mus. 10 :1-1 20.

- - - 1885. A catalogue of the fishes knownto inhabit the waters of North America,north of the Tropic of Cancer, with notes onthe species discovered in 1883 and 1884. In :Rept. U. S. Comm. Fish and Fish. ( for1884) :1- 185. Also appeared as AppendixE, Rept. U. S. Comm. Fish. 1885 (18 87):789-973.

- - - 1895. N otes on the fresh-water speciesof San Luis Obispo County, California. Bull.U. S. Fish Comm. 14(for 1894) :141- 142.

--- 1905. A Guide to the Study of Fishes.Vol. II , pp. 1- 599. Henry Holt and Co.,New York.

249

- - - 1919. The genera of fishes. Part II .From Agassis to Bleeker, 1833-1858, twenty­six years, with the accepted type of each. Le­land Stanford , Jr . Univ. Publ., Univ. Ser,1919:1-284.

--- and B. W . EVERMANN. 1896. A checklist of the fishes and fish-like vertebrates ofN orth and Middle America. Rept. Comm.,U. S. Comm. Fish and Fish. (for 1895),App endix 5:207- 584.

- -- - -- 1898. Th e fishes of North andMiddle America. Part II. Bull. U. S. Natl.Mus. 47 :1241-2183.

--- B. W. EVERMANN, and H. W . CLARK.1930. Check list of the fishes and fish-likevertebrates of North and Middle Americanorth of the northern boundary of Venezuelaand Colombia. Rept. U. S. Comm. Fish. ( for1928) , Part 2:1- 670.

- - - and C. H . GILBERT. 1883. Synopsis ofthe fishes of N orth America. Bull. U. S. Natl.Mus. 16(1882) :1-1018.

- - - and P. 1. JOUY. 1882. Check list ofduplicate fishes from the Pacific Coast ofNo rth America, distributed by the Smith­sonian Institut ion in behalf of the UnitedStates N ational Museum. Proc. U. S. Natl.Mus. 4 ( 1881) :1-18.

KERMODE, F. 1909. Visitor's guide to the nat­ural history and ethnological collections in theProvincial Museum. Provo Mus. N at. Hi st.and Ethnol., Victoria, British Columbia 1909 :1- 92.

--- 1917. Fishes. In : Rept. Provo Mus. N at.Hi st., ProvoBritish Columbia (1 916 ) :18-22.

- - - 1931. Accessions : Ichthyology. In:Rept. Provo Mus. N at. Hi st., Prov. BritishColumbia (1930) :17-23.

KRE]SA, R. J. 1965. Th e systematics of theprickly sculpin, Cottns asper : an investigationof genetic and non-genetic variation within apolytypic species. Unpublished Ph.D. thesis,Univ. British Columbia, June, 1965. 109 pp .

LINDSEY, C. C. 1956. Dist ribution and taxon­omy of fishes in the Mackenzie drainage ofBrit ish Columbia. J. Fish. Res. Bd. Canada13(6 ) :759- 789.

--- 1957. Possible effects of water diver­sions on fish distribut ion in British Columbia.J. Fish. Res. Bd. Canada 14(4) :651-668.

250

LORD, J. K. 1866a. Th e Naturalist in VancouverIsland and British Columbia. Vol. 1. RichardBentley, London. 358 pp.

--- 1866b. The N aturalist in Vancouver Is­land and British Columbia. Vol. 2. RichardBentley, London. 375 pp.

McALLISTER, D. E. 1957. The systematics ofthe freshwater sculpins of British Columbia.Unpublished M.A. thesis, Univ. British Co­lumbia, April, 1957. 109 pp.

--- 1960. List of the marine fishes of Can­ada. N atl. Mus. Canada Bull. 168, BioI. Ser.62:1- 76.

- - - and C. C. LINDSEY. 1959. Systematicsof the freshwater sculpins ( Cot/tts) of BritishColumbia. N atl. Mus. Can. Contr. Zool., Bull.172:66-89.

MEEK, S. E. 1899. Notes on a collection of cold­blooded vertebrates from the Olympic Moun­tains. Field Mus. N at. Hist. Publ. 31, Zool.Ser. ( 1897) 1( 12) :225-236.

NICHOLS, J. T. 1909. A small collection ofAlaskan fishes. Proc. BioI. Soc. W ashington21:171-1 74.

RICHARDSON, SIR J. 1836. Fauna Boreali-Amer­icana. Part 3. The Fish. Richard Bentley,London. 327 pp.

- - - 1844. Ichthyology. In: R. B. Hinds, ed.,Voyage of the H.M.S. "Sulphur," under thecommand of Captain Sir Edward Belcher,during the years 1836-42. [Not seen.]

ROBECK, C. G., C. H ENDERSON, and R. C.PALANGE. 1954. Water quality studies on theColumbia River. U. S. Dept. HEW , Publ.Health Serv., R. A. Taft Sanitary Engr. Cen­ter, Cincinnati. 99 pp. + 4 appendices.

ROBINS, C. R., and R. R. MILLER. 1957. Clas­sification, variation and distribution of thesculpins, genus Coitus, inhabiting the PacificSlope waters in Californ ia and southern Ore­gon, with a key to the species. Calif. Fish andGame 43(3) :213-233.

RUTTER, C. 1908. The fishes of the Sacramento­San Joaquin Basin, with a study of their dis­tribution and variation. Bull. U. S. Bur. Fish.27(for 1907) :103- 152.

SCHULTZ, 1. P. 1929. Check-list of the fresh­water fishes of Oregon and W ashington.Publ. Fish., Un iv. W ashington Call. Fish.2(4) :43-50.

PACIFIC SCIENCE, Vol. XXI, Apr il 1967

--- 1930. Notes on the species of Cottus inwestern W ashington. Copeia 1930 (1 ) :14-1 5.

- -- 1936. Keys to the fishes of W ashington,Oregon and closely adjoining regions. Univ.Washington PubI. BioI. (Dec., 1936 ) , 2(4) :103-228.

--- and A. C. DELACY. 1936a. Fishes ofthe American Northwest. A catalogue of thefishes of W ashington and Oregon, with dis­tributional records and a bibliography. In­stallment 3. Mid-Pacific Mag. (April- June)1936:127- 142.

--- - - - 1936b. Fishes of the AmericanN orthwest. Installment 4. Mid-Pacific Mag.(july-Sept.) 1936:211-226.

SEALE, A. 1896. List of the fresh-water fishescollected in the vicinity of Seattle, Washing­ton, by Edwin C. Starks. Proc. Calif. Acad.Sci. (2) 5(for 1895) , part 2 :852-854.

SHAPOVALOV, 1. , and W . A. DILL. 1950. Acheck list of the fresh-water and anadromousfishes of California. Calif. Fish and Game36 (4) :382-391.

- - - - - - and A. J. CORDONE. 1959. Arevised check list of the fresh water and anad­romous fishes of Californ ia. Calif. Fish andGame 45(3) :159-180.

SNYDER, J. O. 1905. Notes on the fishes of thestreams flowing into San Francisco Bay. Rept.Bur. Fish. (for 1904) , App. 5:327-338.

- - - 1908a. The fauna of the Russian River,California, and its relation to that of theSacramento. Science, N . S. 27:269-271.

--- 1908b. The fishes of the coastal streamsof Oregon and northern California. Bull.U. S. Bur. Fish. 27(for 1907) :153-189.

- - - 1913. The fishes of the streams tr ibutaryto Monterey Bay, Californ ia. Bull. U. S. Bur.Fish. 32(for 1912) :47- 72.

- - - 1916. Fishes of the streams tributary toTomales Bay. Bull. U. S. Bur. Fish. 34(for1914) :375- 381.

STORER, D. H. 1846a. A synopsis of the fishesof N orth America. Mem. Am. Acad. Artsand Sci., N . S. 2:253-550.

- - - 1846b. A Synopsis of the Fishes ofN orth America. Metcalf and Co., Cambridge.298 pp.

SUCKLEY, G. 1859. Report upon the fishes col­lected on the Survey. Chap. II: Report uponthe fishes exclusive of the Salmonidae, pp.

Systematics of Prickly Sculpin, I-KRE] SA

350-368. In: J. G. Cooper and G. Suckley,The N atural H istory of Washington Terr i­tory. Part III. BailU:re Brothers, Ne w York.399 pp .

--- 1860. Report upon the fishes collectedon the Survey. Chap. 2: Report upon thefishes exclusive of the Salmonidae. U. S.Pacif. R.R. Expl. and Surv. (for 1853-55) ,12(2) part 3 ( 5) :350-368. Also appears as:House of Representatives Exec. Doc. 56,Washington, 1860, pp . 350-368.

SUMNER, F. H . 1942. A reconnaissance surveyof Oregon coastal streams. Rept. Oregon

251

State Game Comm., pp. 1-25 (mimeo). In:C. E. Bond, 1963, p. 81.

- - - 1953. Migrations of salmonids in SandCreek, Oregon. Tr ans. Am. Fish. Soc. 82( 1952) :139-1 50.

TAFT, A. C. 1934. Californ ia steelhead experi­ments. Trans. Am. Fish. Soc. 64 (1 933) :248­251.

WILIMOVSKY, N. J. 1954. List of the fishes ofAlaska. Stanford Ichthyol. Bull. 4 :279-294.

- - - 1958. Provisional Keys to the Fishes ofAlaska. U. S. Fish and Wildlife Serv., Fish.Res. Lab., Juneau. 113 pp.