Range O. Bayer

P.O.Box 1457 Newport,Oregon 97365 (503)265 2965

ShinerPerch and PacificStaghorn Sculpins in YaquinaEstuary, Oregon

Abstract

Thc shincr pcfch (d:tzrdtogaster dggregata) ^nd Pacific slxghofn sculp\n (.Leplacollus armalus) were intertidxlly scincd xi an cnbaymcnt "r Yequinr Estuefy, Oregon ffom August 1975 through.lul) 1976. The shiner perch was rnost common fioflJunc thr()ugh Octobcr, but fic crtch ot' this declined after Augus!. Young shiners were born fromJune dlrou!{h micl August. X{ost shincrs cxught v"crc lcss ihrn 9 cm TL. Thc \\-as prcscnt during €vefy month. Young of:the year flrst appeared in Decenlber, and stxFihornsless than 2 cm TL wcrc collcctcd through Apfil Most stxghorns s'efe less dran 12 cm T1. The preopcrcuhr spincs of st:rghorns d() not appcxr to bc an cffective cleftnse against Great Blue llerons (Ardea heroclids). which swallowed the stxghorns \\'jthout first brcaking off thc sphcs.

Introduction The shiner perch (Clmatogaster agllregata) and the P)cific staghorn sculpin (1,?p- tocottus ermatus) are among the most numerous fish speciescollected in bays and estuariesin Oregon (Beardsley1969, Hostick 1975, Stout 1976, Dlrkln et .al. 1977, Bottom and Forsberg 1978, Hutchinson 1979, Mullen 1979, Myers 1980, Bayer 1981), washington (Anderson and Chew 7972, Crosset dl. 1979, Fresh 1979), and Califbrnia (Anderson and Bryan 1970, Fierstineet al. 1973, Allen and Horn 1975). These specieshave also been frequently studied in California (e.g.,Jones 1962, Anderson ancl Bryan 1970, Bane and Robinson 1970, Shaw et al. 1971, Odenweiler 1975, Tasto 1975, Eckmal.er 7979, D2lllng et a/. 1980),\f"shington (suomela 1931, weiss 1969), and British columbia (Gordon 1965, \treibe 1968). However, information concerninll the natural history of these species in Oregon, except \trilson and Nlilleman (1969), is either unpublished or in govefnmental publications of limited distribution. The objective of this paper is to outline aspects of the life histor)' of the shiner perch (henceforth termed shiner) ancl the Pacific staghorn sculpin (henceforth termed staghorn) in Yaquina Estuary,Oregon and to comparc these aspcctstbr shiners and staghorns at other Oregon csruarics.

Study Area and Methods The shiner perch and staghom sculpin were collected at studv sites shown in Bayer (1981)at the 15.8 km'?YaquinaEstuary (44'18'N, 12.1'01'!(/),a drowned river valley located .)n the mid Oregon coast.A beach seineJ.0 n long and 1.1 n high with a l0 mm stretchealmesh was used to capture fish in the upper intertidal ( + 0.9 to + 1.3 m abovc MLLIX/)on mudflats, and a speciallydesigled seine of the same dimensions (B?yer 1979) *'as useclto capture fish in the lower intertidal ( 0.2 to + 0.4 r.nabove MLL\tr) in a persistent eelgrassbed (Bayer 1980).

23O Northwest Science,Vol. 59, No. l, 1985 In August 1975, and from December 1975 through Julv 1976, five samplesn ere taken each month in thc lower it.rtertidalduring daylight hours. Five monthl\. daylight collections wcre also made in the upper intertidal in August 1975 ancl ftom October 1975 rluough Jul_y1976. Collecring during the nighr was carriecl our in August 1975 (fivc times in borh the upper ancllower interridal),September and December 1975(four rimeseach month in the lower interridal),October 1975 (two samplesin the lower intcrtidel), and Noyember 1975 (rnc sample in drc lower intertidal).Each samplew2s obtained from a seriesof 10 hauls, each abour 30 m long. Seinin€lvelocity was about 0.5 m/s in water 0.4 0.7 m deep. At upper inrer- tidal elevations,the seinewas pulled to shore at the end of a haul, but the anrour.rt of mecrophyton caught in the net in eelgrasssometimes made it necessaryto ter- ninate seining in water. \(/hen a haul s'as terminated in weter, the foot line of the net was raisedas rapidly as possible until it was levcl with the hcad line, and thc seiner.as carricd to shore. Thc area covered in a sample was about 600 m,. Gene|ally, the total lenllrh (T L) ancl I'et weight of each fish were measured. Exceptionswerc made for shincrsless than 9 cm in August through October 1975 andJulv 1976 and fbr staghornslcss than'1 cm from January through Ma_vl!76; thcse fish we.e all counted ancl thc total lengths and wet weillhts of a sample of at leest 100 fish were nreasurcdeach month. I also collcctecl shineis and stagirorns droppeci by Grcat Blue He )Ls (Ardea berodias) ^t the herolr's ncsting colonv near Yaquina Estuary during the summers of 1973 1980. The toral lengths of most of these fish were measured. Coefticients of Variafion (CV) were calcuhted as a percentage b_vdividing tlte standard dcvi2tion (SD) by the mean and muir4rlying bv 1OO.To converr rhe averagenunber of shiners/sampiecaugltt during darknessin the lower intertidal from Scptember throu€lh November 1975 to dal.light values, I mulriplied the Septcmber-Novembcr neans bv a conversion factor, which w?s the ayerage mrmber of shiners/sanlple in tlte lo$,'cr inteftidal in August alurinll daylight clivicled bv the average nurnbel of shiners/srmple in thc lo$'er intertidal in August in darkness.Al1 length measLrrementsare total length (TL) nnlessspecifically notecl as standard length (SL).

Resultsand Discussion Shincr Perch Although approxintatelv equal seining efforts q,'crc llrade in thc upper irrtertid2l as were nlade in cclgfass,95 percent of the 22,171 shincrs collected were in eelgrass.Fufil]er, significantly more shiners n'ere caught cluring darkness than dur ing dal,light irr eithcr the upper or lower intefridal (Bayer 1981) with a densiry at ni€lht in August 1975 as grearas 5.8 fish/m, 6 = 2.3, N : 5 sampies) and a nax imum st?nding stock of 11..1g/s'er weight m, (-='1.6 g/m,). Shiners werc absent in Januar),and February and were most corlnton from June through October (Figurc 1), but calcularionsfrom Mycrs (19130:Appendix f'able 1) indicatcclthar Yaquina shinersin 1978 were abundant fion] tr4avthrough Octobcr with a peak abundancefor all her siresconbined in 1977 in August and in 1978 in July. After August, catchesdcciined markedly in 1975 for da1'lightcor rected meens in eelEirass(Figure 1) and also for night catchesin eelgrassfiom August through Octobcri Mvers ( 1980)likewise seinedfewer shirrersafter August in 1977

Shiner Perch ard Pacific Sraghorn Sculpins 231 _o_o-o-o-o too o 300 I I <9 cM r 75 o I LrJ 200 I \, ;e- (L 50

(o a too

T

Lr- lr)rq9O o -F@N $oJooArF F l-.- o|r, ,^i F@i.) -o z drt

UJ STAGHORNS %< t2 cM 200 O-O-O-O-o-O-6-O roo

E. o'" 75 lrJ is too An

25 f\) a c) o n- o I to

A o M A M t975 t976

Figure 1 The relarive size offish caught/monrh, the averag€ number of shiners/sample collected dur ing daylight (or converted to daylight values for September November, see Merhods) in the lower int€rtidal, and rhe :rverage number of sraghorns/sample seinecl during daylight in the Lrpper intertidal. The rotal nunbers of shine$ or sraghorns caughr each month irregardless of tidal height or lighr condirions are along the baseline. a1r.d1978. Elsewhere in Oregon, shiners were abundant from May through Oc- tober with a peak in July at Youngs Bay (Higley and Holton 1975), from May through November at Tillamook Bay (Bottom and Forsberg 1978), and fromJune through October at the Salmon River (Mullen 1979). In \fashingron, shiners were common from June through November (Anderson and Chew 1972).

Bayer Largeshiners (i.e., >9 cfl and >8 g) were collecteclbct\\'een April and C)c- tober Llut were only conrmon between June and Octobcr (Figufe 2). In June. l0 cm of longer shinersfirst appearecl.There may be some yeady variation at the Yaquinx,however, asMyers (1980)found 12 cm SL or longcr shinersfron Merch through October 1978,and I found an 1 1 cm shiner caughtby a inlateApril 1980.Ar Youngs Ray,a l4 cm shinerwas seinedon S May 197,1(Higley and l{olton 1975),?nd ar Till?nook Bay, Borrom and Forsberg(1979) reporred that most adults of unspccilled size arrived in Ma). ancl left by September. The flrst young-ofthc y,earshiners werc seined on 17 Jul]e 1976, and thcy rangeclfronr 4.2-1.9 cn. Slliner young at Yaquina Estuarv rangc fron.r3.9,4.9 crl at parturirion ('WilsonaDCi X4illelnen 1969), and some were seincd rhat haclgrown to 5. 1 cm b)' 2 Jul-v1 976 and to 5.9 cm b1' 15 Julj, 1976.Thus, thc birth of shincrs at the Yaquina sccms to begiD in Junc each year, as cvidenced from rhe _voung slriners(5.0 cn on 1 Julv 1973,LO cm on 26 Junc 1975, 5.8 cm on 6 luly I 977, ancl5.3 cm on 9July 1978) I ibund capturcd by Great Blue Herons. In addition, N1yers(1980) founcl _vounginJune 1978 ar thc Yaquina. Elscwhere in Oregon, McConDaughey(1971) at Coos Estuer,vend Flutchinson (1979)ar SiuslawEsruafy also four.rclvoung first 2ppearinllin June, but t)urkin el al. (1977)ar rhc Columbia River:rDclBottom ?nd Forsberg(1978) at Tiliamook tse,vdid nor find young unril Jul,v.Thus, the commencell]ent of the ltirth of Oregon shincrs is more similar to the usualJuDeor Jul) onset ar Brirish Columbia (Wicbc 1968) aDd puget Sound (Andersonand Che$' 1972) than fbr the Mafch MaJ'onser il.l southcrn California (tsanexnd Robinson 1970, Ficrsrine et al. 1973, Odenweiler 1975). As slriners lcss dran 5 cm $'erc seined from Junc through August, young shiners at thc Yaqlrina appear to bc born ffom June through August. In 1975, howevcr, most shiners appearcd to have been born by August 15, as by that date onl\' 6 percent of the voung (N : 2,12)wcrc lessthan 5 cm. Similarl_v,Nlyers' (1980) clara indicate that she coliectealyollng shiners lessthan 5 cm TL onlv through August 1977 and throuElh-luly 1978, if hcr SL dara afe convcrted to TI- rtcasurenents using'iflilson and Milleman's (1969),Anderson and Bryan's (1970),Odcnweiler's (1975),or Eckma,ver's(tc)79) equations. Thus, the|e appcarsro be lirrleycrdy varia tion in the duration of p?rturition at rhe Y?quiDa. Nlost (95 percclf) shiners were lcss than 5 g, and most (>95 perccnt) n'ere lessthan 9 cm (and about 8 g) except in lhe April June pcriod (Figure 1). Because shincrsare about 8.7 cln 2t the enciofonc year (calculatedfrom C)clens,'eilcr1975 and Eckm?yer 1979), most Yaquina shinef llerch are a ycar old or less during perioclsr4rcn thel.are abundant (Figufe l). Shincrsr?nged from 4.0 15.0 cm and ftom lessth:rn 0.5 g to .19.1g. analGrcat Blue Herons cau€lhtshincrs ran€ling fron 4.0 15.2 cm (N : 79,1).Shincr weights increascdexponentiailv wirh lengrlt (FiEiure 3). Ilowcver, shine| s'eillhts lilr each I cllr classvaried markcdlr'(i.e., CV's for each cm class were about 15 pcrccnt for shiners ovcr 7 cm) witlt ovcrlep in I'eighrs bets'een cl:rsses(Figufe J). The prcdicted length wci€ahtequation (calculatedfor llsh l5 cm or less)bcst preclicted aycrageweigltts for cach lengtlt classfbr fish undef 14 cm (Figure3). Separatelength-\\'eight rclarionships tbr ntale and fenalc sl.rinersat the Yarluina \vcre not derernlincd, although separaterehrionships fbr males and fcmales were computed fbr California shiners (Anderson end Ilryan 1970, Eckmaver 1919).

Shiner Perch and P?cific Staghorn Sclrlpins 2aa SHINERPERCH 500 S SAMPLE M EASURED o = l-5 FtSH N = 400 TOTAL FISH

300

?oo

too

o l\ l\l\k----s e .--{-f--}------.- r () t50 R: tq^-l z r00 l\ lN=24sI l! 50

l NN o t ,-{.. \ l. . . . . a ?o - F lNovDE.l trJ N=24 t0F t_1_ | | E o t l--. I I

Ll- ?oF FAi- MAyr toF o t t50 l-\ lJ-un-JUL-T too N lru=sreI 50 r\N | ' --l---r-- t\J\-\ .l I L__l

5 t9 t5 TOTAL LENGTH(CM) "Sample Figure 2. Lengrh frequency for shiner perch. measured,' signifies that only a portion of fish less than I or 9 cm were measured_

231 SHINERPERCH

f uraru,naree O PR E DICTED WEIGHT ?o r+t++* to tT ol) 5 t- I (9 ;2 It' = I I 0.5 1 o.3

COEFFICIENT OF VAR IAT IO N a

t5% a a- -a d's a

ooooco _|r)\''o (o- oooo|r) l.)|r)',ToN

5 to TOTAL LENGTH(C M)

Figure 3. Length-weight relarionships for shiner perch. Coefficenrs ofVariarion (CV), means, and rang€s are for I cm intervals wirh sample sizes along the baseline. The lengrh-weight equarion was calculated from the measurements of 45 shiners with 4 fish for each cm class. exceDt there s/as only one shiner for the 15.0 15.9 cm ciass. Preclicted values were determined from mid, interval lengths from the lengrh weight equation (W:4.07x10 3TL3.5, r = 0.99).

Shiner Perch and Pacific Staghorn Sculpins 235 Pacific Staghorn SculpjDs Most (69 pelcer.rt)of the '{,'{73 staghorns seined were in the upper inrerridal (Bayer 1981).The nrulber of staghornscollecteal in darknessdid not diffcr significantly from thc number collected during deylighr in either cclgrassor rhe upper infer tidal (Baycr l98l ). Staghornswere prescnt each month bLtt\\.ere m()st common from Februaly throughJul_y1976 with thc peak abunclanceinJunc (Figure1). Similarly, staghorns v!ere preseDtnonthl,v il.rother stucliesat Yaquina Estuary(Bcardsley 1969, Bur reson 1973, Mycrs 1980). Calculati(Dsfrom a combinariur of all Myers' (1980, Appendix Table l) sitcsindicare rhat stallhornsat the Yaquina in 1978 $.ere most common from February through Aullust s'ith a peak inJune. At Tillamook Bay, staghorns were m()st common from March through October (Bottom and Forsberg 1978). Small( i .8 2.0 cm) steghornss,.ere first captured in December and were seincd throullh April. Staghornsless than 3 cm were found from December throupihJul,v, (Figure4). Likewise, Myers (1980) only found sraghornsless than 2 cm SL at the Yaquina fron December tl]rough Febluar,v1978 and in April 1978, and Bottom and Forsberg(1978) first found juvenile stallhornsat Tillamook Bay in December. Length-ffequencydata indic?te that sta€jhornsmay grow ft) abour 11-14 cm by the end of onc vear'(Figure .1). This agreeswith Jones'( 1962),Burfeson's (1973), and Tasto's(197i) length-frequcncvdara, but nor wirh the length,frequencydata of txreiss(1969). Since staghornsspawn during abour a 5 6 month pcriocl (|ones 1962)\\'ith larveeprescnt at the Yaquina from Augllst through Marct (pcarc1.and Ml.ers 1980).the multiple length frequency modcs for \ii/eiss'staghornsless than 12 cm may indicate multiple pcaks in spawninll or hatching rather than ditTerent year classesas suggestecl by \(/eiss(1959). weiss (1969)also useclotolirhs ro deter- mine sevcral year classesfor sraghorns less than 12 cm, but Jones' (1962) mcasurements of otoliths indic?ted that staghorns were about 1 1 13 cm when thcy were one year old. Most (63 pefcent) of the 4,473 sr:rgl]ornsweighecl less than 5 g; however, staghorns weighing over 5 €lrepresented 86 percent of the total sta€jhorn biomass seined(31.9 kg). Staghornsranged from 1.8 27.0 cm anciless rhan 0.1 gtoa25.3 cm staghorr.rweiglting 273.4 9,.As in California esruariesand bays (for.res1962, Tasto 1975), most (>75 percent, Figure 1) staghofl.rswere small (i.e., < 12 cm) and probablv lcss than one vear old. Staghorn weights increasedexponentially with length (Fi€ilr1'e5), but therc $'as much variation in weighrs for each cm class with CV's oi 20 38 percent for cach classof st:rgbornsless than 9 cm and about 1G22 percent lcrr staghorns over 9 cm (Figure 5). Predicted wcights for mid intcrval lengthsclosel,v matchcd average$'eights for eachcm classof staghornsfrom j 26 cnr (Figure 5). Staghornshave large uppcr preopercular spincs on both sides of the headj when a sta€lhornis caught the fish raisesthese spincs. T?sro (1c)75)and Forbes (1983) indicated that thesespincs m2y be useful for dcftnse; ho$'ever, the spinesdo not appcar to cleterat leastooe avian precl:rtor,the Gfeat tslueI.leron. I examined 472 stalllrorns,ranging from 3.J 2,1.3cm, that wcrc c?ptured, swallowed, and latcr regurgitatedbv thcse herons. In nonc of thesest:rgltorns was either preopcrcular spine brokcn, and for some staghorns 12 cm or lcss both spilteswcrc still raiscd

Baver STAGHORNS too .= l-5 FISH r N= TOTAL FISH 50 rF AUG-SEP N=495 o t_ oaaoaaaa 25 N=lOB t-tl OCT o t -lll-. aaa t5 NoV N=33 t-h| ,o o t a a o, o aa a 40 t f--L DEC N=90 o L I l-. a ao a o JAN-FEB N=513 400 z

tL, 200

l o L....aaa 600 MAR-APR N=879 LU 400

200 tL o a ?OQ L.. MAY N=773 I-h loo o -{l In-r-.... aaao 4 00 JUN-JUL N=1582

200

o EaooEaFoaao t,|t,,,L,,,|,,,,|,,r,|,, t.5 lo 15 2a )^ TOTAL LENGTH(CM)

Figru€ .1. l.ength-fr€quency for sraghorns.

Shiner Perch and P?cific Staghorn Sculpins STAGHO R N SCULPINS

+ MEAN,RANGE ri+f{+{ 50 O PREOICTEDWEIGHT

30 {{{" 20

3 to +l+T F

I ('

2 LIJ

= i+1'

o-l

40 COEFFICIENTOF VARIAT ION

,--t-2 a ------a --a-- a-a be c'

o @so\rt.)@o-@()(0F @ o o to t5 TOTAL LENGTH(CM)

Figure 5. Lengrb weight relationships for sraghorns. Coefficienrs ofvariarion (CV), means, and ranges are tor 1 cm inreFels with sample sizes along rhe baseline. The lengrh weight equation was calculated from the measuremenrs of 137 staghorns with 6 nsh for each cm inrerval, except there were 5 fish for rhe 2J.0-23.9 cm inreNal. predicted values were determined for mid- - interval lengths from the iength *'eighr equarion (W: 5.14x19 3113.31, 1= 6 99;

234 Bayer after herons had regurgitatecl them to feed their young. Additionally, I observed these herons capture several hundred sta€lhornswithout a sinEllestaghofl.r escap ing once the heron pickecl it up and held it long enough (i.e., about 5 s) ior me to identify it as a staghorn. Severalstudies (Kendall 1966, Tasto 1975) have indicated that sra€jhornsfollow the tide. Tllis resulted in staghorns sometimes being stranded on intertidal mudflats at low tide (personal observation). Staghorns have becn observed to bur)' themselvesin aquafia(Tasto 1975), but I also found a 17.5 cm staghorn buried in mud under a clam shell in an interticlal muclJlat at the Yaquina. After such a stranding, a staghorn is vulnereble to avian prcdation. I observecl Great BIue Herons capture and eat seyeralapproximatel]. 12 15 cm staghornsthat were shallowly buried in intertidal mudflats of the Yaquina.

Acknowledgments I am grateful to many colleagues at the Ore[ion State University Marine Science Center and Ore-Aqua Foods, hc. Their assistanceenableal me to complete this study.I thank F. Button, G. Brced Willeke,M. Evtchison,R. !(/isner,Jr.,and others who hclpecl seine. The constructive comments of C. Bond, \tr. G. Pearcy, and an anonvmous referee cnhancecl this manuscript.

LileralureCited

A]]cn. L G , end M . t L I lorn I 97 5 . Abundence, diversiry, and scasonaliry o I flshes in Coloraclo Lagoon, Alaniios Bxy, Calitbrnia. Est and Coasr.Mer. Sci. l:371 380 An.lerson, R. D., and C. F. Bfyen. 1970. Age nnd lirowth of rhree surfperchcs (rr?t /otocidde) f Jtn Humboldt Bay. C2liforde. Trxns. Amer. I.ish. Soc 99:'175482. , an.t K. \(. Che\1'. 1972 Prclininafy studv on the rransrcnt species of flsh in Bili Beef Harbor. -:( -2) f '. {rl e- l-..11.5 ). lnl Bane. G., and M. Rd)inson. I970. Srudics on dre shiner perch , Cymatagaster aggregdld clbbons, in uppcf l{ewpori Bay, California.\Yasmenn J. Bio. 2U:259-268. Bayer. R. D. 1979. An xnti-roll bexch seille. Cal. Fish came 65:189 190 . 1980. lnterridrl zorlation of Zaster.r marina in rhe Yaquinx Esluery, Orcgon. Syesis l2:t1l $1. . 19i11.Sh"llow \1iler intertidal ichfiyolaune of rhe Y:rclulnaEsruarl', Oregon. Norrhw. Sci. 55:18219J. Bcarclslcy, A J 1969 M()r.cmcnr aird angler use offour foodfishes in yaquina Bay, Oregon Oregon ":.. s,Jt. Ini\er.r.\.( ..r\ Di\\(fr.rtrorl B()t(nn, D., and B. Forsberg. 1978. The of Tillamook Ba_v.Oregon Dept. Fish \vjldlifi, poftllnLl. Projcct F 100 R. BurresoD, E. f,L 1973. Host p:rrxsite rchtionships of rhe stxghornSculpi]n Leptacottus armatus Gtt^rd in ()relion. Oregon Srare tlnl\.crsirv, Corvallis. Dissertaiion Cfoss,J.N.. K L Fresh,B. S.Ililler, C. A. Sinenstxd, S. N. Sreinfofl,anclJ C. Fcgle\. t978. Neafthorc flsh anci nracroinvcrtcbratc essemblagesalong thc Stmir ofjuen de Fuca, inclul:ling food habirs ()1thc common nearshorefish. NOAA Tech. Meflo. ERL NfES,q.32. Darling.J D S.. M. L Nd)ll:, and E Sha$..19U0. Rcproducrive straregics in th€ surfperches.L Multi ple insemin:rtron in naRrfal popuiations ()f rhc shiner perch, Cltnatugdster dggregata. E\toL 31:271 2l ,-. Dnrkin, I T., S.J Ljpovskl, c. R. Sn',der, and \'L E. Tutde. 1917 Enrironncntal Smdjes of Thrcc Columbia River lstuxrin. Bcxches. Enyiroo. (lonsc|l . Div., N\v and Al$ka Fish C€nter. Oregon Ficld Facilir_!,ilemnlond Ecknra,ver, l(.J 1979. Age and gron'th ()f fbuf surfperches (tmb ktocldae) fraa\ the outcr herbor of Anaheifl ll:ry, California Cal Irish c2lnc 65:265 272. Ficrsiinc, H. L., K Ir Klil]e, anll c R. Garmen. ltl3. Fish€scollected in tr{()rroBay, Caljfornia be- t*.een J,rnuar_y1968 2ncl Dcccmbcr 1970. Cel. Fish G"mc 59:72 89. Forbes. 1..S. 19E2.Prcy nanipularioD in thc cfcar BIue tteron. Murrcler 63r89 9,. Fresh-K. L. 1979. Distfiburion ,ncl abund?nccof lishesoccurrin!! ln rhe dearshoresur.f:rcc Eaters of northern Puget Souod, \{-ashington. LniYersirr of Wxshingron, Seaftle. Thcsis.

Shiner Perch and Pacific Staghorn Sculpins 239 Gordon, C. D. 1965. Aspects of the age and growrh of Cltzatogaster aggregata Gibbons. Universiry of British Columbia. Vancouver. Thesis. Higley, D. L., and R. L. Holroo. 1975. Biological baseline dare, Youngs Bay, Oregon, 1974. Oregon State University, School Oceanography, Ret 75 6. Hostick, G. A. 1975. Numb€rs of fish captured in beach seine hauls in fte Coos River Esruary, Oregon, Jun€ ihrough September 197O. Oregon Depr. Fjsh Wildlife, Coastal Rivers lnvesr., Info. Rep. 74-ll. Hutchinson, J. M. 1979. Seasonal distribution of fishes in Siuslaw Bay. Oregon Dept. Fish Wildlife, NW Regional Office, Corvallis. Jones, A- C. 1952- The biology of the euryhaline fish Leptoco us armatus Gn^rd (). \tniv. Cal. Publ. Zool. 67 :321-36a. KendaI, A. w., Jr. 1966. Sampling jrvenile fishes on some sandy beaches of Puget Sound, washington. University of Washingron, Searrle. Thesis. Mcconnaughey, E. A. 1971. Coos Bay study: an interdisciplinary study of man and rhe estuary. NSF Student Originated-Studies Program. University of Oregon, Eugene. Mullen, R. E. 1979. Fishes of the Salmon River Esruary, Oregon. Oregon Dept. Fish \ ildlife, tnfo. Rep. Ser. Fish., no. 79 5. Myers, K. W. 1980. An investigation of rhe urilizatiofl of four study arexs in Yaquina Bay, Oregon by hatchery and wild iuvenile salmonids. Oregon State Universiry, Corvallis. Thesis. Od€nweiler, D. B. 1975. The life hisrory of rhe shjner perch (Ctmatogaster aggregata) Gibbons jn Anaheim Bay, California. 1'?E. D. Lane and C. W. Hill (eds.),The marine resources ofAnaheim Bay. Cal. Fish Game, Fish Bull. 165. Pp. 107 116. Pearcy, W. G., and S. S. Myers. 1974. L?llal fishes ofYaquina Bay, Orcgonr a nursery ground for madne fishes? Fish. BUI|. 72:201-213. Shaw, 8., J. Allen, and R. Stone- 1974. Nores on collection of shiner perch, Cymatagaster atgregata in Bodega Harbor, California. Cal. Fish came 60:15-22. Stout, H. 1976. The natural resources and human urilizerion of Netarts Bay, Oregon. NSF Student Originated-Studies Program. Oregon State Universiry, Corvallis. Suomela, A. J. l93l. The age and growah of Clmatogaster aggregata Gibbons collected in Puger Soufld, Washington. Univ€rsity of Washington, Seattle. Thesis. Tasto, R- N. 1975. Aspects of rhe biology of the Pacific sr^g}.lotn sc]u]pin,Leptocottus armatus Gj:(rrd, in Anaieim Bay. .f,?E. Lane afld C. W. Hill (eds.), The marin€ resources of Anaheim Bay. Cal. Dept. Fish came, Fish Bull. 165. Pp. 123 135. weiss, E. F., Jr. 1969. The age and growrh of rhe merine cottid Leptocottus armatr6. proc. Montana A.c d,. Sc|. 29163-f1. wiebe,J. P. 1968. The reproducrive cycle ofthe vivaparons seaperch, CJ)matogaster agregata. Can. J. Zool. 4G1221 1234. Wilson, D. C., and R. E. Milleman. 1969. Relarionships of femxle ege afld size to embryo number and size of the shiner perch, Clmatogaster aggregata. J. Fish. Res. Bd. Can. 262339-2344. Receiued 23 Marcb 1984 Acceptud for publication 14 August 1984

240 Bayer