in · 2013. 7. 8. · 1,. up to 330 m.in length when matureo 2. large, widely flaring oral lobeso...

FARY LAND DEPARnGiT OF FBSEARCW AND EDUCATION hesapeake Biological Lzboratorg, Solomons, Md. Ref. No. 58-55 . Eugene Cronin, Director

THE DISTRISUTIGN OF CTENOFlSORA IN TKE PATUXENT ESTUARY DURI_Nt; THE SUY?t3EB OF 1958 - - I--..-..- ---I-

f2knophcrr-d. have received perhaps less at%ention and study than any other phylum. There have been several extensive works concerning Cijenophores, such q s MayerTs (1912) Ctenophores -- of the Atlantic Coast of North Alherica; how-

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ever, publications a re relat ively few and far between, Nothing could be located i n the l i t e ra tu re dealing with the subject of this stucty, namely, t h e distri- bution and abundance of Chnophores i n an estuary. Therefore, any investigation tha t is carr ied out on t h i s phylum will be of value,

One possible reason fo r the Lack of study is t h a t t h e i r biological im- portance has not been and i s not being fully realized. They constitute a major consumer of zooplankton and phytoplankton, I n fact, according t o Nelson (19251, the rela%ive amounts of same forms of plankton are "to some extent correlated" with the abundance of the Ctenophores i n the plaslkton. An examp1.e of t h e i r economic imparhnce is the i r predation upon oyster larvae, which is of special 9nteL*est to the Chesapeake Bay region,

'khe en t i r e investigation of Ctenophores i n the Chesapeake Bay ems, of which this study is a part, will include same aspects of t h e i r life history, growth, reproduction, feeding and f w d habits, abundance and df stribution, plus other aspects of t h e i r biology which w i l l psobably be added t o the program la ter . The purpose of the ent ire project i s t o supplement and add t o the bialo@cal knowledge and understanding of Ctenophores as a group and of the several individ-

I na3 species found i n the area t o be studied. That pa r t or phase of the pro Sect with which # i s report is concerned deals with the abundance, distributcion and size range of Ctenophores i n the Pa.twcentilives estuary of the Chesapeake Bay, the possible fac tors involved, and implications which can be drawn from the cjbsema%ioas r

t According t o Nayer (l9l2), .three species of Ctenophores a re found in the Chesapeake Bay: -- Beroe ovata, Ymemiopsis gardeni and Ymmiopsis Icidgf,.

, Thus far i n the collections of t h i s smmer, no specimens of Beroe ova- have -- been absemed, Separation of the two species of Plnenriopsis was very difficult partly because the specimens are very del icate and easily damaged and pa r t ly because of t h e apparently unsatisfactory systematics i n the genus, Eayer f 1912)

' gZves the foUowing distinguishing features between the two species:

I/ Conducted f i e l d work, summarized data, and presented summary upon - W c h t h i s paper i s based,

Suggested problen;, provided general supervision of field work, analysis, and presentation. , .

1,. Up to 330 m. in length when matureo

2. Large, widely flaring oral lobeso

3. fink=tsh hWo 4, So discoidal w a r t s on

OX'&. lobes*

1. 35-40 me in length when mature*

2, O r a l lobes small, om- sj.x%h to one-fifth of the length of the body.

3. B1uris.h hue. 4* BiscoidaL warts on oral

I~b&sa

A large amber of the specimens which w z z observed either did not confom~ to the above charac.i;eristics or displayed an iz?;ermdia.te condilzion, F>or exa.qple, m%ny inr3i05duals possessed large flaring a d lobes and a bluish or purplish hue, OCner~ wM.& also had tihe la rge o r a l la3es were covered w i t h discoidal warts on the entire body* including khe oral lo'bes,, S t i l l others which w e r e large 533 s ize (SO& mm,) possessed smll oral loSes* Only -0 specimens were seen which pos- sessed alL of the aboae-zamed features 2ar He garded, Ths maSority conformed 21o %he description of He Xe5dyi, heref fore; there appears ta be consLderable variat-iun ernd o v e r ' l a p p r n g n e WO s p d e s and these charsc%eristics 620 mt s e a to p v i O f 3 a satisfactory d i s W t b n between &e~r, The above-enkioned obsemations w e r e merel;y f i e l d obsemXons and Rr,zher study nus% be made .l;o deterraS.ne -tihe vaJ.i&ty of Mayex-1 s se-parz%i,.on. Payer st;aba i+arther %hat "P9. l d d y i is a creature of the p u , ~ sea m%r along the outer shores, whi3.e Id:

W v e s in protected bays and k x d d s h water@,, This is definitefy fn erfirTs fes s E. Zeidfi i s concernmi. Thmughout thi.s report only %he term W m i . dll 6e -;yed which viU ~efer to He - leir&+

Ia refereace to previous rep+& of the clfsWbnt%on and 8bd-e of l4md8 &a, Mayer (1912) s k k s *at i t s rrtnge extends fxom Tinyard Slmnd to &&?ha. Nelson (1925) dtes seraral acccnri%s l a which it was reported t;o haw occurred 5x1 peat raft;s* or nzzpiads" off %he coast of Hew Jersey, Rho& Idand and Mods Hole in sorat? yezl.sl In other years it was very rare.

* P r o c ~ ~

SaxpXng was out by weehly f'ield trips up the Fakwent RLper, which is characterized by a &creasing sa3Azi.w gradsen%. Seven s&ist&ans %era es%abM.&ed at w h i c h sapZes were Aakm. 'fhese stations we-:

1. The Chesapeake BioXogid L&ora-d;ow jlder. 2. Town Point, 3- Broomels Island. 41 The Patumnt River BTidge .at bnediat . 5 The mouth of H ~ l ~ l k b g Creek. 6. Deep Landing, 7, !The mouth of Black Swaarp meek. 8* Lower Marlborn (if ~ e e c ~ ] ,

Collect2ons were ma6e two staSons begondi where the last C%emphore was taken. At each station, fif ken r?rinr,t;FI bO%tom and surface tows were taken. The to* the was later decreased to -ran minutes because of the very Zarge numbers of anjlnals tha t were being c-. Lwge meshed planMan nets were

eap3to;$ed in s~unpIfng. Several weightis were attached t o the rope about a foot above one of the nets so tihati it would remain on %e bottom while being towed* The neb were thirty c~!ntimzters in diameter and the estlsabd flow of w a t e r through t;he net daring a fifteen minu.t;o k w was not snore approxbate3y

./ 44 c&ic meters, During a .ken m t e tow the flaw was ab0u.t; 29 cubic meters.

Bcfr specimen thak was coflected was masused to &8 nearest half cent&- for total, lely~ehr lfieasulwu~nts were made in a w h i t e enamel pan. ifhe ten-

peratwe was noted at each atat ion w i t h a reversible Lhemmeter for %he bottom and surface, The gensrdl. mather conditions, Kind direc%Lon and velocifg, and the turb9d3.w and roughness of the watef w e r e a l so recorded f o r each statioa, Bat- %urn anll sarface xater sap- were kken a d t;he sakizlity w a s detejr-d in the Bbr,rakmy by the hydK,meM method.

Of the greatest isprtance in any abrmdance and dlisMbukLon study is tibe mliabiXty of %he sanq?Zjlag t echigw; that is, is it %akbg a represenktim smp1e &OBI %he pop&l.af;ion? A special tr"tp w a s made to the Severn aver f o r the purpose of ZRsting the method being used. This was carded o a t as follows:

I B i n s sfa%&ons were chosen on what w a s originally p-ed to be on the basis of a

I sa3inity gradient, %e sallnittes were determined iri t;he field by the hyrbmmetier. TZlis s h m d &at there was so much variation that %e ori@na2 plan had to be d5sregarded. Botiiorn tows w e r e made a& each sta-n the form of *me cbcles

: i of approx5.nately the sarae &ze. One taw coasis%ed of three circles and three

wepe made a t eadn sta.tbn for the purpose of rep3icat;ion. The &ads f r a m eaeh were nemured to ihe nea.resl half centsmeter and recorded. The %ern- p ra twe was also taken at each station. The results were graphed and statistical t e s t s =re cet3csiLated.. Figure I shows the mean 1engtE.t and stmidarb dewAation of %he catch of each tow and station. 'ff~ere does not appear to be a sigfilfiemt pax4.ation between any of € 2 ~ mew, dl3ihonlgh tMs has mt been tested mathe- maticaUg. Tho mans U e between %NO and %wee cenfimebxs* Those arrislals ool- , lected in the downstream stat;ions seem to be slightCLy larger than those upstrearri. The ;initicathn derived f m W s is that the saw popUaaorr was being smp1ed a t all sta%iolzs. Eie results of the repUcation of ibe hws suggests that the eamplAng method proxLdes a reasonably accurate picture of the siae of azx5md.s fn t;he population from w h i c h it is mpl iag , Asswring &at the total +ambier of an3maLs obtained at each staP,5.on represents t&e $rue d m frequency m g e of the propu2atiLon at that strat;ion, It can be seen %at %he mans of each tow are @te close to that of: the station. T h i s 5s espcia33.y app-nt & e ~ e +JIB sample froa a dngX.8 tas oonsfsted of one h m as mre inrEviduals, A farther i&cati(oa of W s 0'235ervation is Figure XIs wh5& shows the size Itistribntion of all %he azrislPils collected in 1&he Severn aver and that of each t o w in &Ah the -re nmbwed one hundred or greater*

It is not knm shebher or mt f;hs saxpJing technique gave an amirate I p5cfare of the abundance of Hmmio &s The tous made i n the Patuxent ESves were made in a w e s t r & & f i G e r h a p s a series of short Gous or tow8 mada Sn a circle over the sanu3 amount of %me would be better.

There are several, persons wfio deserve credit and fhanks for the* part in t3Xi.s s1udyS including 3Sss Charlotte Epfa;ztg~~, Fx. Curtis Allen, 2-55, W i a m E.

I IBogers, Dr. 8- J- Ktll icyl and especia3ly Dr. Rut21 Griffith, who provided 5. mral.rtable \ assisbnee i n organizing the field work and regulas3.y participated irr the field I operation siurhg most of %he summer, 1

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/ The follb~rdng a m the resa3ta produced by eleven weeks of samgling during the s-er of 3958:

The itiijtrribu.trion an8 abundance of Blnemiopds were qnib irregular. Thjts c o ~ 3 . t ~ n was especWjr evjdent in the Severr, River sap-. The animals a p p w d fo be carrcentrakd in some areas; and s p s e in other areas only a f e w hw,drerf yards a~my. Figure 3: shows t;he cakh by fim and by sAttLon. The nm.&rs of spet5ntens obtaiaec! in the three taws at each station were, for the most past, qn5.t~ U?l;iI"~rn* However, there was a large difference between s%a- %ions. A t the most downs%mm stattion (11, a large catch was Wmn. Con- siderably fewer animals were ob-d from the n e d &ee sla%Lons (=, In and IV), ~ t h a Large amomti of variation among these three. Smpling a% the next; sl;aS;ion (TI) resulted 3a a Urge ca%ch+ The s ize distdbutf on, on the other hand, was xemrWl$ un-ifon;i in the Severn aver, as pmviously dna- cated. This was c@ke ~Ufere~nt fmm ttre si%ut;ion found fn %he Patwsent liives, ~ M c h will be &.scassed later in W report.

Thls 5rregul;rrity In a-ce and distribabion canld be seen in %he Sat-t 52i-v-e~ also (~igures I11 and XIL). !!!here paa8 considerable variat50n at %he same sta%ion f m ~ week %o week even tho@ the changes i n the sdl2~Lty and teqgmature appeared to be &nor,

Amt;Pler observation of W s ineguh5t;y w a s made i r r fWmt a2 the Chesapeake B.o;lo@ca3 I;Ebora%ory where specfroelas were being col leckd for a s a m t y .taXerance exprAmm.t;4 The tide was flooding and there was a fairly rapLd ~ O F T 02 water. A great aaarry Fmmiop&s were observed approxi- makly %eo -to f5fteen feet fro1 shore in about two and feet of water.. They appeared in a strip five .t;o ten feet w5de, mw5ng with We currezt. Farf;"ner out 2mrn &ore, only ;an occasional. o m could be seen*

GemTa13yt %he larger k4nemiapsis were found %n higher saliai%y Wer and the smXLer ones in lower salinity asakr. This is srh0'1.m on Figare ZlI. The s i z e range was arbitrar5J.y divided on the basis of early s q 1 e s inb %wa parts, mall, up to Wee centimeters in aength, and large, three and one- half centcimetexs and larger. Figure XIB shows 8 mtypicar2g cakh at an up- .strew & domtraa~ s%atbn during J-. Berg. f e w of the large IsdiuidMLs were %&en below a ~ U r z i t y of 7.0 parts per thousand, whereas the sdU.er tmbnale -ere observed dbwn to a saUx&ty of 4*3 parts per t b m d . The W e r mnermiopsfs were found at khe dow~strem a%at;ions, ;ia&ading the Laboratory L i f and Tcli~n Soh%, in considerable rrumbers rin two succeeding weeks, Emever, they appeared to be co~centrated f n the area of .the Pe%wo~nt ELtrer Bridge at Genedict where the salidty ranged between 5.8 svld 7*0 parts per thousand throughout most of the swmer. The smaller animals were by fir mre ~ m u s than the Zarger ones*

Bo paftem of abw&mce as related tx, temperature could be foMd. Tfie cmplete taperatme range 5a which FSrteaio sSs was observed during t?xis Swim extended from 21*7" C. to 28.80 6. wi TT-- a general gradual r ise Ulmughou.tmost of the s\ursner, There was a rather sharp decrease 5x1 temperature noticed "he lasL week of s=pling. Other than this, there was lit$le weekly va5at;ion. As can be obaemd in FSgures IIL and VII* apparently no variation in abundance can be attr%buted to -temperature.

I; tb With regard to salinity, no pattern of abundance of FheIrJo sfs could. be _%_ f o d as reh.ted t;o varia.t;Zon i n salinity except the coinci ence of low

s W t y and %he Lack of &ls. As previ~wly stated, the lowest s W C y in which Mnemiopsis was found was k.3 paTts per thousand. In tdze Severn Bvert spec5xsns were taken i n 3 8 parts per thousmd. Belar these salinities none were found Zn the respective estaa~ie8~ Therefore, #at s%a%ion, or area, atw"riich t h e sal inity fZuctnates around 4.0 parts per thousznd perhaps could be called %he u f m e area1' of occurrence. T h i s

1 area appeared .i;o be loceted at the mouth of Wnt&ng Creek (r'igur~es XC and

f m) during the ent-ire sur;rmer, except for one week when the sdlixxity showed a pmnomed increase a% aL3, sta.e;ions. A t this station there 2s a weak coirzcidence 'mtrseen s&nj.t;y enc! abundance. On dars when the saki&.* was re&~tivel.y h w , E.inemicrpsis were present, and on &w when the salinity was

I ! m2atively UV very f e w or none were found. At t;he Zocatims where the salinity was above the apparent lower limits of tolerance, no re2ationsMp between 4 . h i i i y and abundance was suggesf~d. !

I I . , s4 A t the Ugher d M t y statbns domstrearn, there were ctiffirences beween

I I the abmdance of Ctempllores at the surface and at 'the bottom. However, th5.s was very ir'pegulas - stme days there were gpeat;er n e r a at the sw-

E f a arid on other dtyrs the sibason was reversed. There is no apparent exphu&ion. A t firs% it was t b @ C that the roughness at the surface was responsible for the greater numbers at the bottom. Bowever, Wsr probabxy was not the case since often greahr n ~ ~ s were taken a% the surface wben 4ihe n b r was quite choppy* The pattern of abundanoe throughout -the mmer

L was p e r a l l y similar for the snrface and botton. r ! ( I

me tak ing of u a y b . t ; t o m surface tows does not give a clear pSc%ure of the verthal distriibution. The use of one or prefembly mre

I nets at i n f e w t e depths and perhaps at ten-foot btemaX~l would be < t , better. f * "r ' i ! 6 , kt the upstream skWns where the saEx&ty was re1ati.U 3att namely Ule f

flsrlaxent Wdge a% PRnedict an3 HmWng Greek, more indixidnals were obiabed f m .the boLtom tow Bas fmm the surface, exeep% for me case when the sal5nikies of fb 9Urfaee snd bottom were %he satae. A t aU. other .t;iEles, sewenken in a;ll, the bottom saXi.r&t&es were Ugher -khan &e surface, sng- i': gestLng %bat sdLirbty &&-b be reLzted to %he control3ing factor. There was o m case where t&e bottom saLln5ty was higher than a t of the &ace. This

i ap-ed at Seep hnding at the time when a e sll'r.nsties a* a U stations &wed a narkmi increase. The water at this s k t i o n was not mry deep, w?xLch m y OX may not have had aometh5ng .to do with the observation. Barever, there mas a pnounced difference &%ween the bottom (6-2 u/m) and surface (4.5 o);oo) s w a e s *

f , h The abWame and distribution over the period of sang,ling ean be seen cm

Filgures S1V gind XVT- An influx of very snra33. individuals was observed at the beem3ing of J-. The vast maJority of W s e were in the one-ha centimeter size cLass, wb2ch included all specinens f r o m *75 ceatimter ~~ to the srsa5lt-

b . est that could be seen with dAe W e d eye. The peak of abundance appeared in the middle or tio~ard the end of July, By the f*st of August, a sharp de- crease in d e r s was noted aad the srnaUest size class (.s centimeter) had -appeared by the middle of the month. 'ithe peat inELux appeared f i r s t at

i the upstrean stst;ions and persistee there a greater leogth of SloaU. &&ivj_duals, bat not as ~ m a U . as FBXV~OUSIY~ are striU. f m d there but in much reduced n&rss

/ 8. mrn .tha m e ;of the peak of ab~W.ae of the sna3.l anima;ls in ddJuly , ~0

individuals tsfren in iikre samples became progressively larger, 'I'fhis is ;in-

d i c a W on Fig~ss Xa and XTI. The increase in sim was noted f o r both upst~eam aJld downs- sbt&cme

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#%ough t M s stac;Y has covered a relat5vel.y &or% perLod of %be, setr- t k j n g s have been learned abcu% F ~ o sis* Xt seems rezsonably certain Wt

---+Ti they are not uniformLy dfsWibuted, that is eir occurmace is spotZy and -uneven. The fftcf; .I;&% mexicr sis As a planlrt;onic Tom and 5 % ~ moaemen.1;~ are l;lrgely, if

__I;%_ mt fi03ls1 dekrmine by water currents -which are in turn cox~%rolled by the tZdBf 3 movements and kinds m y be one factor concerned Perhaps ~ m i o p s i s possess more

r=on"crol over their mvwneni;s than is realized. ff this is the case, they w o l d haw %he power to concentrate theaselves in an area where con&tions are more favoraba, such as an area of a b m h t food.

St; appears as fd P " I . o sis spawns k.om la& June unt i l raid-August, I

~5,431 the peak appea3.islg &d- ----?- ex, a k June, This finding is based an the presencn of .rrerj satall aniraals, most of whlch a-ppared to be passag thmugh the em17 stages shorW,y foIfow3.q hakhing as described by -a'dapr (1912). He states that ha%chhg occnrs dbouf 30 h m s a fh r fertiL3imtion of the egg. Therefore, -tjle W m of spamxhg ieaa be fasirly w e l l deternrined. T h i s was an at;Jrpical summer as f a as saUr&ty, which has been 10wer than n o d , and temperat-, which has also shown a %owes .than average trend, a r r concerned, Therefore, W s obsemed

. %reeding period also nay mPI 5e tmcal wi%h regard to time and dwation*

The presence sf tihe larger bdi83.dwds downstream in higher s a U n i t i e s and the slza;ller i t~dividuab upstream in the lower saUaL%5es was rebt5,vew con- s t One possible exp1a~tti,on far t M s observatbn 5s thai; the eggs are de- posited dmmtream by the Wger adult arxinals and -&tat either the eggs or the yaung larvae were carried upstream where they developed. The larger anbds m a d have greabr control over their mvenents the ~~e and the eggs and could prrsslbly resfst the tendency to drift upsfream. Rnothex p o s ~ 5 ~ 2 S t y is %!at the pund individuals were d e m d by enemies and/or by ~ b a 3 i s ~ c adul.ta, The srsX2. d i d appear fn consf eerable numbers at m e downstrean stak5ons for two successive kxeks and then. conplete3y disapgeared. It does seem adUely %hat the ;pow would undergo complete mrtall%y.

TnrztSng to the question of abgration, it appears +.&at E1Inemicpsi.s has . X t t l e control over borhonW mipratixlin. T h i s is probablg delemined brwater

. currents Wch are i n tunn dependant upon numy factors such as t i d a l movements zrnd a id+ Vertical migration, on the other hand, would appear to be ntore wlm- tam since B e forces Zx, be overcome are mt so strong. No pattern of vertical mipation could be deten;rined, It could be said that Paeniopsis can regulate the5r hori,%ox).tal mmment Zn the sense that they can move vertically up or down to a holoizontally moving curren* and be Lramported mme distance.

I There is s t i l l much to be &ne with regard to the abundance and &slrS- A .. n A AL- a_-- t1-> .- . . - -

. . , - .' ; . .

&a&-&*..-:.,& k4P L* G x >d

- 7 - beginning. Only if this study is carried an over extended period of tim

b

caa the abunda3lw and distribu%ion be b o r n aszd anderslood. These a m several. factors whLch hat% not yet been included in this study and wEch are suggested for the f i b :

1. The ~est ical distribntion of Ctemghores.

2, The m u e n c e of tides upon hopil;ontdl. and vertical d.istribution, if i I

3. The difference 'betweerr n i g h t t W end cia* ~ s t ~ b u ~ o n , '

1 . 4. The U f e m c e b e k e n summer a3ld winker distr&but&on.

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1 I ;i

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Agassiz;, Be 8 6 f ~ X ~ r b h bTgcan Aczlephae.it 132. Cat. plPus. Cump, Zool., No. X I , Haward.

Bauer, P. 1910. ttUber den aascheimad nervose IlegrCLation der FUmerbewegung bei den Rimqutitlen.l Zkchs* VergXeich Phgs i .01~ ;to+

3Lgelow, He 3. 1912. '!A A,vi.ew of Hayex's Ckaophores .of the AXLarrtic C o s t of 1 g ~ r t h . Ante~9.ca.a Science, Aug. 23, 19l.2, Po 2k3.

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f91E;. wEhq$lorat%c+n of the Coastal Wakrs between Nova Scotia a d Wsapeake Bag, July atEd August 39U1 by the Oe S. &shed.es Schoomr, Grampuso Oceanography a;nd Planktones Rill., f4useum of Conzp. Zaol., " W G e , Tol. UX, go, 4.

1 B o s s r n , J • 19570 *A Pre-rg S m y of tbe Small Fifatant Fauna $21 the ! .I_ _ VLc3nit.b of %he %a-Floor off: B Q t b , Eorthumberbnd.tt Jam. AlaimaX

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J E ; c o ~ ~ , 26(2) : 3!3.-366.

I . 1898. %Die Ctenqphoren der Slankbn-Expedit;ion.n Ergeb* P3mk. Eqpd. 2 Ka.

k * ~onf3eJds B. Re 1934, mCooKtZwai;ion an& KovemnG of the Swimbg P la te s sf PInerrtS~p~isrn BUU. 66.

++ - ' . 3936. Tte&nera%&on in F!iaen~Lop&s.~ BLcl., B U . 70.

U36. n A p i c a l Dominance and Polari%y 5.n ~ned.opsis.* Biol. Bull* 70. .

and A. G o l d h e 3937. #PhpioLngicdt Gradbat &n HnmrLoplstis duriag Regenerati~n.~~ BTxrL. B u U . 73.

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'53awy&ff, G. L933. flPIoTpholog5.e et Biologic des Ciwnoplana.* Arch. iWo1. 33xpte Gen. 75.

., 1936, nbes CtenopMdes des eaux de l'lnda&mcln Bul. Bio2. - Fraace Belg. 10.

. 1937. %es Gastrodes.e Acad* Sci. Paris , Coaapte Bend* 2&

. L938. Wu3.tipUca2;lon asextxee chez les C.tenaplana.* Acad. S a r paris, Conpt, Rend. 206.

. 29b8. contribution a la Co1~wiseance des Cbno@oms felagiws des twux de 1tIndocMrie Ball. Biol. France Belg. SO, S46, pp* 233-370.

t ?

I B -3h;llaningt;op, E. and R. WameiS.* 9 *SchooL of Harveslifish Feeds on Sea Wa&mts.n Hary1~~1d Tjcjewatfer News, Oct. 1955, Bole 12. ;j

4 . i

1 11881. nOn ehe AcdLeopae 09 t b East b a s t of Hew England.@ ii ~ b ~ , d e , V 0 3 . r . m r fi

~ ~ y , E. 3. and 3, H. 3 2 ~ ~ . 1938. %mnbescence in Absence of O~rggen in tihe C%nophore Hne~~Lqsis .~ J m e Cell0 Colnpilrr Physlol. l2.

H&&rt Ke 1927. a V ~ m 'iSemmpttm der Ctem@oma" ~tschr. Mrpb. OkoL -Tiere , 9. I .

HerWg, a, X88O. Wber den Bau tier CPRnophoma*" JeaaS.sdbe ~-f;schr, Wat-. a.

I KatushiXov, X* Em 1955. W,.I;a9cl Orebneirka Beroe cu~ximis Fabe# (The &ad. Nauk

Bncaid, To 1915. %3y~ZRr &%me Ln Washington*tl T w o Second AM0 Heeting Paciric Pisheries, Saa Francisco, p. 4.

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3 .* r

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'4

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