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Some factors influencing the distribution of pelagic copepods in the Queen Charlotte Islands area Cameron, Frances Ella 1955

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SOME FACTORS IHFHJEHOINO THE DISTRIPUTIOI OF PELAGIC COPEPODS IH THE qJEEI CHARLOTTE ISLANDS AREA by Prances B l l a Cameron  A T h e s i s Submitted i n P a r t i a l F u l f i l l m e n t of the Requirements f o r the Degree of . Master of A r t s In the Department of Zoology  We accept t h i s t h e s i s as conforming t o th© standard r e q u i r e d from candidates f o r the degree of Master of A r t s  Members of the Department of Zoology THE UNIVERSITY 0P BRITISH COLUMBIA April  1955  -.1  Abstract  The d i s t r i b u t i o n of c e r t a i n copepod species o f the north coast of B r i t i s h Columbia suggests that  breeding  i s r e s t r i c t e d t o l i m i t e d regions of w e l l ~ d e f i n e d temperature and  salinity characteristics,  the spread description  Currents ar® r e s p o n s i b l e f o r  of j u v e n i l e s and a d u l t s from these areaa*  A  of the probable c i r c u l a t i o n p a t t e r n i n the  v i c i n i t y of the Queen C h a r l o t t e Islands as I n f e r r e d from plankton c o l l e c t i o n s  1® submitted*  TABLE OF CONTENTS PAGE INTRQDUCTION «#•«•*»«•••»•»#»•••#»»•»•••»*«*»«*»»•»»*»  X  METHODS AID MATERIALS «#*,..»•...•»*«.•••«•»  ,. #.  4  V e r t i c a l D i s t r i b u t i o n ••*•••*•»*•••••••••«•<.•••••••«  5  H o r i z o n t a l D i s t r i b u t i o n »**••.•#**«*•»•»•*»«*••••.»*  8  RESULTS •»...,»•..=»»««*«•«»»»»•••»«•«••*»••»«••*•••<•  11  DISCUSSXON •••»•••••#«••»••••••«••»••••«*«••»••••«»•«»  18  The distribution®/Of the more common species  18  Currents suggested Py the d i s p e r s a l of eopepeds .**«  3®  Tasu I n l e t »•••«»»»••*#»•»•••«»«»••»••«•»»•»•«•«»•  39  Dixon Entranoe ««»»##**«**»#»**%•«*»*«**•**»##**«*  45  Masset I n l e t •»•««••»•»••».•«•»•*»••••••«••••••«••  49  SUMMARY AND 0OK0KJSI0HS »»«••...•,*•..,•.»•*.,»,,,.••*  57  ACI10WL1DG1EITS  59  . . . . f t . , . , * . . . . . . . . . * . . . * , . . . . . . . . . .  BIBLIOGRAPHY «*••«•»»«»«»»**•*•»«•••. *»•»•*»,•*«**••**** 1  APPE'MDIl  60  • L l S f 0? TABLES TASK I.  PAGE The Occurrence and Eelativ® Abundance of th©  Gopepod Species II*  i n A l l th© Plankton Sample®  «... Appendix  S t a t i o n s Compared i n Determining the 'Upper V e r t i c a l L i m i t s o f Three Specie® #••..».»...«».«  III.  S i z e Eangea i n maw o f Several Spool©a of Copepods •Found i n Dixon Intrance *.....*.*.».*.....,•....  IV*  IS  IT  The V e r t l e a I / D i s t r i b u t i o n af Centropegea .momurrlohi aa I n d i c a t e d by i t s Oeourrenee i n Supplementary and Deep Hauls »»..•..«.»*.•,.#.•#  V,  The V e r t i c a l D i s t r i b u t i o n o f Tortanna  23  diaoaudafcaa.  aa Indicated by i t s Occurrence In Supplementary and Deep Haul a *»*».•««#*»»•.**•»*»*»**•*.«»»***.» VI* VII.  2*7  Mas set I n l e t S t a t i o n 89 *• .#*«.»#..#..#*».«»..»**• * Appendix The Age Proportions of the Oopepodid Stages of M e t r l d l a laeens and Calanui flnmerohleua  Appendix  LIIT O P FIOtfHES  FIGURE It  PAGE  Plankton s t a t i o n s taken adjacent t o the Queen Charlotte Islands by the Canoollm I t during th® summer of 1953 »..*.».«;».»»,».«,»•,•„.,..,.,»,,  2.  3  The occurrence of Gepepod species i n samples taken from d i f f e r e n t depths to the surface  ...  14  3#  Tli® d i s t r i b u t i o n of Faraea&anua, parvus ........».,.,»  18  4,  The d i s t r i b u t i o n of A o a r t i a e l a u s i .**.*..,.,.».  20  5,  a) The s i z e variation of  aaartia c l a u a i i n  Dixon Entrance b) Currents suggested by t h i s v a r i a t i o n #«**•*»*«•*«*  21  @t  The d i s t r i b u t i o n of Oentrppagea acmurrlehi ..........  22  7»  a) The s i z e v a r i a t i o n of Centropages momayriehi i n Dixon Entrance b) Currents suggested by t h i s v a r i a t i o n ,•»»,.».»»,»-.  8* ' The d i s t r i b u t i o n of Tertanua dlaoaudatus #•.«*..*..»* 9.  2S 28  a) The viae v a r i a t i o n of Tort anus dis.oaudatua i n Dixon Entrance b) Currents suggested by t h i s v a r i a t i o n ... *  29  10. Th© r e l a t i o n of temperature to th© d i s t r i b u t i o n of Oncaea oonifera • • •»•  »•«  •. •..  31  11. The d i s t r i b u t i o n of .Oneaea o o n i f e r a .................  32  12. The d i s t r i b u t i o n of Corye&eus a f f i n i s .*#,«....•.*...  33  13. The v e r t i c a l d i s t r i b u t i o n of Ooryoaeus a f f i n i s  34  14. The d i s t r i b u t i o n of Calanus f Inmarohloua ............  36  FICRJHS 15.  PAGE  The v e r t i c a l d i s t r i b u t i o n of Galanus flnmarchlcua o f f laden i n l e t i n Dixon Entrance * •  ....*  36  16.  The d i s t r i b u t i o n of M e t r l d l a lueena  37  17 •  The d i s t r i b u t i o n of Eucalanus b u n g i i ,  38  18.  The v e r t i c a l d i s t r i b u t i o n of three species o f f laden I n l e t i n Dixoa Entrance ***...«*.#*•.#•*»•#*•  It*  A proposed c i r c u l a t i o n of the surface water above 14 meters i n Tasu I n l e t .  20.  38  *...  42  A proposed c i r c u l a t i o n of the water between about /  14 and 30 meters i n Tasu I n l e t 21.  ...................  A proposed c i r c u l a t i o n of the deep water of Tasu I n l e t <«.<•»..•••.•..*»«*.•«..*•.«•.•.•«,.••.•••..  22»  47  A proposed c i r c u l a t i o n of water between about 40 and 100 meters l a Dixon Entrance •«#*«•.»**»,»*  26.  47  A proposed c i r c u l a t i o n of water between about 15 and 30 meters i n Dixon Entrance .,....»..,.,.....,  25.  43  A proposed c i r c u l a t i o n of th© surface water In Dixon Entrance  24*  43  The d i s t r i b u t i o n o f the deep water oopepods i n Tasu I n l e t ant immediately outside ..•.*«..*.».».#  23*  42  48  The r e l a t i v e abundance and v e r t i c a l d i s t r i b u t i o n of oopepods at s t a t i o n 80 In Masset I n l e t over a 24 hour p e r i o d .........  27.  The r e l a t i v e abundance and h o r i z o n t a l d i s t r i b u t i o n of Gentropages memurrlohl i n Masset I n l e t  28.  51  52  The r e l a t i v e abundance and d i s t r i b u t i o n of A c a r t j a o l a u s l i n Masset I n l e t  53  ill FIGURE 29.  PAGE The r e l a t i v e abundance and d i s t r i b u t i o n of Oncaea eonIfera In Masset I n l e t  30*  .......................... 54  The population- s t r u c t u r e s of l e t r l d l a lucena at the s t a t i o n s i n Masset I n l e t  31.  54  The h o r i z o n t a l ranges of s e v e r a l copenodid stages of M e t r i d l a luoens i n Masset I n l e t  /  ...........  55  ijwjtoarcrxon Daring th© summer of 195$, the waters adjacent t o the Queen C h a r l o t t e Islands were surveyed I n the O.G.M.v*. Csneolim I I under' the sponsorship of the Defence Be search Board' and the s c i e n t i f i c d i r e c t i o n o f Dr* ft*F, Seagel and Mr. F . G . Barber. Plankton samples were taken a t each s t a t i o n ( F i g , I ) w i t h p h y s i c a l d a t a , i n order t o permit a study o f the r e l a t i o n s h i p of plankton t o ©eeanbgraphlQ conditions, i n the a r e a . One of the most s t r i k i n g f a c t o r s became evident, when the deeper hauls were seen t o contain d i f f e r e n t copepod species tn&n were ©aught i n hauls from shallow water.  This situation  suggested t h a t the species had c e r t a i n v e r t i c a l ranges some of which were i n deeper water*  f u r t h e r study determined  t i c a l d i s t r i b u t i o n s of many eopepod speeles.  the v e r -  With t h i s inform-  a t i o n , i t was p o s s i b l e t o study the f a c t o r s e f f a s t i n g h o r i z o n t a l distribution* Th© occurrence as w e l l as the abundance of various' species appeared t o be a s s o c i a t e d with great d i f f e r e n c e s i n s a l i n i t y and temperature*  The regions where species were  abundant and present i n a l l stages were assumed t o be breeding areas*-  Inere the eopepods were rare and represented by a few  stages only,, they were considered, i n many eases to be I n d i v i d * n a l s s c a t t e r e d from an adjacent breeding population. T h i s s c a t t e r i n g i n turn was th© r e s u l t of another physical factor.  The currents although not known at the time  of study were suggested as r e s p o n s i b l e f o r the d i s p e r s a l of  individuals from breeding areas.  Aa scattering seemed to be  prevalent i n several l o c a l i t i e s , i t became possible to suggest the currents i n these areas.  3 E N T R A J C  D I X O N  E  o  8< *  7/ J  .  86  96  •  ^>  90 '  93  5 4°  J  99  •  »  n  .  ( *~-r^  /  I j j  0?  INLET  L 106  105V  9  1  .102  m  >N  z' • . ^ Y  Z^T**  /  BASSET INLET  /i  t \  *  107  ^  ^»  t—^^^^  S T R ;M T  *  *  \-<  5 3°  H E C A T E  «  THETIS INL£T^^  TASU INLET  „;  V  52°  1 3 3'  j  13 2 * "  13 1°  Figure 1, Plankton atationa taken adjacent t o the Queen Charlotte Xslanda by the Cancolim I I during the summer of 1953.  4 METHODS AND  MATERIALS  Plankton samples were taken between J u l y 18th and  August  8th i n 1953, i n the Queen C h a r l o t t e Islands area i n c l u d i n g Dixon Entrance and Hecate S t r a i t . shown i n F i g u r e 1.  The  s t a t i o n p o s i t i o n s are  A t a l l s t a t i o n s i n d i c a t e d , v e r t i c a l hauls  were taken from near the bottom to the  surface with a t h i r t y  i n c h diameter net having a number t e n mesh*  At about on©  half  of the s t a t i o n s , a d d i t i o n a l hauls were made from one, three or f o u r t e e n meters, to the  surf see* • A ten Inch diameter net  used f o r these hauls*  At s t a t i o n 89 i n Masset I n l e t , samples  was  were obtained from water pumped through a hose from a d i s c r e t e depth*  The samples were preserved i n f o r m a l i n . the samples were considered o n l y q u a l i t a t i v e l y as no  d i r e s t accurate measurements were made of the volumes of water filtered*  An  i n d i c a t i o n of f a u l t y f i l t r a t i o n was  noted a t some  s t a t i o n s (69, 72, 73, 86, 87, 112, 114, 115, 116, 128) where ifi c r e s t e l l s aorvsfiioa and supplementary h a u l but not that the  A c a r t l a c l s n a i were captured i n i n the deep h a u l .  the  T h i s suggests  l a r g e net used f o r the deep haul was clogged by the  time I t reached the surface and f a i l e d t o capture species l i v i n g near the s u r f a c e . A l s o , no attempt was made t o take d u p l i c a t e  samples*  Many workers (Barnes and M a r s h a l l , 1951, and Hardy and  Gunther,  1935) have found that v a r i a b i l i t y between d u p l i c a t e plankton h a u l s can he l a r g e , due to the patchy d i s t r i b u t i o n of the plankton.  Hence, In c o n s i d e r i n g numbers of plankton animals i n the  samples s t u d i e d , only gross d i f f e r e n c e s ar© considered s i g n i f i c a n t .  s The  cop©pods were determined t o species except f o r a  few genera found In th© deep water.  Most wer© i d e n t i f i e d by  referring t o Davis (1949) although Giesbrecht {1903) and B s t e r l y (1905, 1911,  1924)  (1892), Sara  were a l s o c o n s u l t e d .  D i f f i c u l t i e s arose i n the determination of a few species which were neither common or widespread.  In a few cases, i t seemed  evident that f a r t h e r systematic work i s necessary i n t h i s r e g ion,'  For example, a species o f Gaetanns Was  found*  According  to Jesperson and Russel #(1952) v e r y few males of t h i s genus have been d e s c r i b e d . of i n v e s t i g a t i o n .  T h e r e f o r e , these specimen* are i n need .•'  ••As'the method of sampling d i d not J u s t i f y exact q u a n t i t a t i v e c a l c u l a t i o n s o n l y approximate numbers of each species were determined. fomr terms. thousand  8  Sare  cubic  The numbers at each s t a t i o n were denoted by tt  r e f e r s t o approximately one animal i n a  centimeters, "few* to t e n , ^common® to twenty*  f i v e and "abundant  11  to . f i f t y *  As no d e f i n i t e numbers d i v i d e d  one category from another, i t was d i f f i c u l t i n tea©  caaas to  decide which term to use, but wherever the r e s u l t s wer©  crucial  the samples were reeheeked. Vertical  BsM^Mm*  I t was more speeies.  immediately apparent that the deep hauls contained This strongly suggested that different species  were l i v i n g a t d i f f e r e n t depths.  Evidence i n d i c a t i n g that  speeies occur within c e r t a i n v e r t i c a l ranges can be seen i n  * D i r e c t o r s of P u b l i c a t i o n . S e c t i o n r e f e r r e d to i a written by W. Verwoort.  many systematic and e c o l o g i c a l p u b l i c a t i o n s on the copepods. Depth d i s t r i b u t i o n s are*given i n general, hy Davis (1949), S l e s b r e c h t (1893) and Wilson (I9©0), while 5uruhaal (1953) has s t u d i e d In considerable d e t a i l the v e r t i c a l d i s t r i b u t i o n s of  copepods i n Japanese waters*  In an attempt to'deduce t h i s  d i s t r i b u t i o n "from''the ©eeurrsnee Of speeies i n the" hauls,'the samples were arranged In order of i n c r e a s i n g depth,  fhe appear-  ance ©f speeies i n the deeper samples'gave  an i n d i c a t i o n of the  approximate depth at wfaieh they ©ocurred.'  In order t o present  t h i s data g r a p h i e a l l y the samples were considered i n groups of twenty-five*  F o r example the f i r s t group i n c l u d e d  samples  taken from th© surface to a depth of 10 meters while the ' second group i n c l u d e d samples taken from s l i g h t l y g r e a t e r depths t o the surface.  For eaeh species the percentage of  samples i n which I t occurred i n each group was considered.  In  Figure 2. where a speeies occurred i n a group, i n more than f i f t y percent of' the samples i t was represented by a heavy l i n e , I f less, by a f i n e l i n e *  I t ©an be seen i n Figure 2 that  Olauaocalanus apuicornis' occurred i n only the deepest group of samples and In l e s s than f i f t y percent of them* represented by a t h i n l i n e i n the l a s t group.  I t i s then Pauedoealanug  ainatu8# on th® other hand, occurred i n more than f i f t y pereent of th© samples i n a l l groups.  Sense, I t i s represented by a  heavy l i n e through a l l groups* The upper l i m i t s to the v e r t i c a l d i s t r i b u t i o n s of the Speeies l i v i n g near the surface could be f a i r l y w e l l determined by comparing th© r e s u l t s of the supplementary haul with those of  the deep haul*"- The  analyses of the species and t h e i r approx-  imate numbers In the two hauls y i e l d e d important to how  r e s t r i c t e d a species was  I t moved down and how  the age  Information  as  to th© upper l a y e r , i f and when  groups were arranged  vertically.  I f a species were common i n the surface haul and i n the deeper h a u l , i t was  assumed to be l i v i n g almost e n t i r e l y at the  f a c e , while-atspecles absent from the surface h a u l .but i n the  deep-haul  sur-  present  presumably occurred somewhere below the depth  of the surface haul* ' iome s p e c i e s , however, l i v e d well below the depth® of the supplementary h a u l s .  In some l o c a l i t i e s where deeper s t a -  t i o n s were close together, i t was  p o s s i b l e to estimate  the  approximate depth «hero the uppermost i n d i v i d u a l s occurred comparing adjacent  s t a t i o n s , one with and on® without  species.  The depth of th©  found was  l e s s than that of an adjacent  by  th©  s t a t i o n where the species was  not  s t a t i o n where i t  occurred, and t h l a depth was t a k e n as tho uppermost l i m i t to the v e r t i c a l d i s t r i b u t i o n of th®  species at the second s t a t i o n .  For example, i f Met.rid.ia lucena was a depth of 100 50  meters and absent at an adjacent  aaefcera, then i t was  low 50  metere.  common at s t a t i o n % j' with s t a t i o n "b "at  assumed to be l i v i n g at atation'V'be*  Although the r e s u l t s appear adequate, i t i s  r e a l i z e d t h a t e r r o r may  he present because of the dependency  of a p a r t i c u l a r upper l i m i t on the depth of th© adjacent  However,  haul.  they seem workable and have y i e l d e d some valuable  Information.  Horizontal Distribution* In tho  d i s c u s s i o n * h o r i z o n t a l d i s t r i b u t i o n s have been  depicted on small maps on which the occurrence  of the  and i t s r e l a t i v e abundance are Indicated by c i r c l e s .  speciea The  dis-  t r i b u t i o n s are considered i n the l i g h t of temperature and a a l i n i t y data as w e l l aa geographical l o c a t i o n and i n a eases, bottom topography* p a r t i c u l a r point was th©  The  occurrence  taken to mean one  i n d i v i d u a l s were abundant, present  few  of a species at a  of s e v e r a l things t  if  i n a l l stages and  pro*  duclng eggs, the c o n d i t i o n s i n t h i s l o c a l i t y have been in  the  recent past and.are favourable f o r the reproduction of t h a t s p e c i e s , i f the i n d i v i d u a l s were r a r e and represented hy a few  only  stages then they were considered In some instances to be  s c a t t e r e d from a l a r g e r population.  In the l a t t e r case i t i s  d i f f i c u l t t o d i s t i n g u i s h between a population which Is i n c r e a s i n g and a group of I n d i v i d u a l s merely s u r v i v i n g .  However, each  case i s considered s e p a r a t e l y and ao o v e r a l l i n t e r p r e t a t i o n i s used. With a c l e a r e r idea of c o n d i t i o n s which a r e optimum and suboptimum f o r each species,' i t becomes p o s s i b l e t o v i s u a l i z e centers of reproduction and th® movements of i n d i v i d u a l s away from these l o c a l i t i e s , by the currents prevalent i n th© D i s p e r s a l was widespread nor r a r e .  area.  best s t u d i e d in spec!©® which were n e i t h e r Feeudcoalanua mlnutua and Qlthona  helgolandlca were too common everywhere.  The  l i m i t s f o r sur-  v i v a l of these two are probably w e l l outside the c o n d i t i o n s i n t h i s area.  E p j l a b i d o c e r a amphltrltes and Aetidema armatua on  the other hand were so few and s c a t t e r e d as to be almost  u a a l e a o f o r a atudy of  diaporeal*  Variation in tfet a l i a of aopapoda btafuw a vary neeful  ofittrlan fw dl»tla§«lahlag populfttioaa. Tha absolute slsa adalt «©p«p©da haw bean found to vary with l o c a l i t y baaing 1958)  and aeaaon im&y  1060)•  of  In tho literature there la no  Mieattotifenatthe else of the adult d«panda en t h a length of tlma i t h«« be«n ais a d u l t * Cotkalderins tbt aiaea of the d i f f erent; davelojpiaenfcal stage a of pavaral oopopoda (Jonaaon 1@M a and Stfbnir XttM) i t l a ardent that alee inereaaea oeosar dur-lng m® moult tafcatwn atagea« en attwapt to e x p l a i n the variation of the alaea of •Suite* 0*»tng ©©italderafcragfeti©Haerwtloa %im% m iaereaee in teaperatare w i l l prottoo* an ineremae i n the rate o f 4*«a&* opraent. 1« r a marks i);wt I f a otipapod deireleped in cold water, i t would e a t ware than a ©opened developing i n warm water under the sane food oonditlons. Ia the a i a a of any one stag® Oapettda on tha aitse of the pm&Lma atage aa well as e» food o a n d i t l o n * in the ye**nt peat* B a a i n g auggeeta that temperature, isflxjena* Ing tha langtt* «f ftndtag tine, and f@ed ewaaentration* a f f a s t * l a g t h efttnaantoonsuaad could &mm the e l s e variations that h® ©neerved near Greenland* III  In one area at one tine* those f a a t a r a would a f f e o t th® • growth o f a l l oopapada the mm*  afeath*? or not tal.® effect v a r i e d , the net reault would be apparent in the aiae variation o f the- a d u l t * o f t h a t a r e a * JSeoo* a wry ;nmrrow else range would be **pa*tad« ffhia was fatal t o be th® oaae, first i n Even i f neither the past conditions are known nor their exaot effect on the slue o f the ©opapod, Metrldla luoana.  tha  10 narrow size ran^e l a one l o c a l i t y i s i a I t s e l f u s e f u l , as eopepods d i e p e r a e d from one art® mj  im reeognltsed by their  else* l a Dixon Entrance the else ranges of s e v e r a l a p e o l s a were determined In order t o find the extent end direotlon o f dispersal*,  that* ipeeiee were measured from the t i p of the  head to the end of the sandal faunae, using an oenlar mlero-  mUr* nnrted o f f i n ,05 mm* taterwle* l a the site ranges were apparent*  Slgaifi«*nt  differences  These eould be used to  d e t s e t direction o f *@abt®i»Ing* Harked d i f f a r t n a a a i n th® proportions @f the age else see Of a f e w apt**** was aoaaldvvwA i n one o r two esses to i n d l o a t w population 4tatlnatt%*n«eja v but only when the water seas i n that i d e a l i t y was ahown t® be d i s t i n c t by else differenees In ©tfaer epeelee o r t h a preaenee o f different speeiae. This Method waa mum e f f e e t i v e i n considering the diaperaad of the sub» smrfaa® spealea i n whloh the age group* have a dafiiiite v e r t l e a l d l i t r i b m t i o a t tha young being n e a r e r the surfsee•  fhe preeenee  o f young only, at a station iadleeted that they had drifted away from above the adult® which l i v e d i n deeper water*  &a  this teehnifie waa <n*anlltabi*e only i n Masset Inlet and waa not uaed throughout tha study the results are presented i n the discussion. The tern population refers In this study to a group of animals of one speeies whieh has developed i n one area at one time.  I t i s also uaed i n this sense by Barnes and Marshall  (1961).  U mmhm  The water flowing  of the Rorth F a o l f l o Drift Cswdrmp «J| aj. l i W ) # This broad diffuse c u r r e n t divides Just south of the l o l a a d * the northern, arm moving up the eeeeb has a f a i r l y uniform temperature whioh fluctuates seasonally to. about 2 0 0 aefeara hut rarely wanna t© above 15 degrees oentlgrs.de • with thaae ta^eratur® etearaeteriatle® this are® mm be ©onalderad I n one of tiea^reaht1!.faunlatle sonoo (©ieebreeht 1898). la hag divided the plankton fauna into three geographical zonest the southern sold water, tha eantral war® water and the northern eold water* the last la Halted to the south by the 15 degree aantlgrato laothem* Seaee, th® waters being studied belong in this tone* OoMMa^eatly* the eopepods are mostly northern c o l d water epeeles* fhe almost complete absence of war® water apeoleo aaaea th® • l a a a l f l o a t U n d i f f i c u l t t o a @©naiderably» Thirty two ayodioa o f oopooodi ware determined* These are Hated below* ooaea  from  past th© %t©©n Charlotte Islands  the northern p o r t Ion  Acartia  ol«aai  A*loiigirwodf Aetidoua  aieabrficht  (tiUjoMg)  aratabue (Boeck)  Brwdyidiua a i m l l i e  0  (Sara)  Salawe erletatua Kr^yer C. f i a a a i r o h i o i u i (tunaerna) 6* tonsua Brady  12 #Candacla Columbia© Campbell Centropages raomurri c h l W l l l e y Chlridius g r a c i l i s Clausoealanua  0  J&pran  a c u l c o r n l a (Dana)  -*Epilabidocera amphitrites (Momurrich) Eucalanus  b u n g i l OlesbrecLt  luehaeta Japonic*. Marukawa Eurytemora hlrundoides  (Nordqviat)  Grasbairas sp« Heterorbabdus proximus l e t r i d l a longa  0  Davis  (Lubbock)  1. luoena Boeek -Faraealanus parvus (Glaus) Pleur©mamma ap. Pseudeeal&nu© a i m bus ( I r ^ y e r ) Ihinealenus nasutus® Siesbreeht - one specimen seen S o o l e o l t h r l e e l l a minor (Brady) S* eubdentata. (Esterly) Tertaaus diseaudatsua  (Thompson and Scott)  aObryeaeua a f f i n i s If ©Mar r i c h Oithona helgolandlea Claus 0* plumifera B a i r d Oncaea c o n i f e r a Giesbrecht  0g.B*OHDE-R B4HPA0TXOQ3CDB4 M l c r o a t e l l a norveglca (Boeck) M. rosea Dana o probably  13 A few other harpactacoids were present In the samples hut as these were not common or widespread,  they were not determined.  As mentioned before a l l these species are northern c o l d water forms, although a few are a l s o found In warm water. are p e c u l i a r to the l o r t h P a c i f i c . asterisk.  Figure 2.  These are marked with an  The d i s t r i b u t i o n of a l l the species a t the s t a t i o n s  can be seen i n Table I The  Several  (Appendix).  v e r t i c a l d i s t r i b u t i o n of these species i s shown In A l i n e i n the f i g u r e i n d i c a t e s t h e occurrence of a  species at the s t a t i o n s having depths as i n d i c a t e d .  As was  mentioned before the s t a t i o n s were grouped according t o depth. The dark l i n e I n d i c a t e s that a species was found i n more than f i f t y percent of the s t a t i o n s o f the group i n d i c a t e d . galanug finmarchlous was caught  Although  i n a few cases near the s u r f a c e ,  because i t was more common i n deeper water i t was considered as a subsurface form.  Of the species which were caught i n hauls  of 22 meters or more, one, Bradyldius a l m i l i s , I s benthic and w i l l not be considered a s p a r t of the plankton.  Aetidiua  armatua and E p i l a b i d o c e r a amphitrltea are too sparce f o r f u r t h e r consideration. The upper l i m i t s t o the v e r t i c a l d i s t r i b u t i o n s of the subsurface species could be roughly determined by comparisons of adjacent s t a t i o n s , i n one of which the species Is e i t h e r absent or r a r e .  The compared s t a t i o n s are shown In Table I I .  The depth of th© shallow s t a t i o n s are the depths of the upper l i m i t s to th® species' d i s t r i b u t i o n . quite v a r i a b l e .  These are seen to be  T h i s v a r i a b i l i t y prevents the d e t a i l e d use of  14  Deep forma  X* longa 0* a e a i t o r s i a aaetamia H*fe**oitafe0a« Blm®mmmm 0. c^XoatolM S« jspontoa  C* arletava*  $n»*» Surfaee  Ponaa  A. H M t B I o#. p l u n i f e r a v. tumm §m fttmrnmhim® c* t f f i a u  — .  M* F©gft F# p » » » a 1» nirundoides Snrfae® Forms  0* e o a i f e v * i» n o r v t g i a * f« dlaeftudatae  0* wtm#*Uto& A* longtraait 4*. aXanai •  ; •  tm miia&ttts 0« beXgoXan&loa  Bepfeh Rangaa,**  figara  1*X0  ,  14*20  8* Tha ooeurrenoe of oopopo4  83*48  apooloa d i f f e r e n t depth* t o the a u r f a o e ,  _ _ _ _ _ _  44-10 ©*?-64l§  la awttpXat taken froa  15  TABLE I I STATIONS COMPARED I I DETERMINING THE UPPER VERTICAL LIMITS OP THREE SPECIES  l e t r l d i a lucena  Calamis finmarchicus Present Sta* Dep* M 64 104 09 97 86 111 112  44 75 91 65 91 119 . 91  i  Euoalanus b u n g i i  Absent S t a , Dep* M  Present Sta. Dep. M  Absent S t a . Dep. M  Present Absent Sta. Dep. S t a . Dep. M M  68 103 72 90 85 108 113  97 83  366  90 86  36 91  119 110  175 33  122 109  104 24  81 83 114 82  25 36 53. 36 55 100 45  35  135 366 164 206  90 86 108 96  36 91 100 46  average depth d i s t r i b u t i o n s f o r any one area. upper l i m i t s have been used, except f o r one  Instead l o c a l  case  (page H)  for  p a r t i c u l a r areas. F i n a l l y , eleven species were caught only In th© very deep water, 87  meters or more.  s h e l f on the Entrance.  These s t a t i o n s are mostly o f f the  west coast and i n th© deeper regions o f Dixon  The  species found were not represented by great  numbers and wer© not widespread, h a l f of them wer© caught only i n two  or three of th© very de©p h a u l s .  These species ar©  pre-  sumably c h a r a c t e r i s t i c of th© deep oceanic water which occurs close to the  west coast and In Dixon Entrance.  ature (Jespersen and Hussel 1962)  In the l i t e r -  these species ar©  oceanic, while th© others mentioned above are mostly The Entrance  termed nerltio.  r e s u l t s of measurements of s e v e r a l species i n Dixon ar© recorded i n th® f o l l o w i n g Table I I I .  17 TABLE I I I SIZE R&IGES I I MILLIMETERS OP SEVERAL SPECIES OF COPEPODS FOUND IN DIXOH ENTRANCE  Adult Males Sta, 99 98 97 71 75 75 96 88 90 91  A, o l a u a i  0*70*1*00  A. l o n g i r e m i s 0*95-1*01 •1,^0*1,06  0.72*0*75 0*72-0#78 0*91-1.00  0*96-1*01 0,90* 0*9©# 0*92-1,00  T, dlscaudatus  G. mcmurrichi  2  1*53-1,80  1,43  1,73*  1.41*  p 1.86-2,10* 1,52*1*77  1.27-1*60 1.30-1.44 1.44-1.78  1.30-1*50  1# 25*1,33 1*24*1.55  Adult Females 8t*'« 99 98 97 71 73 75 96 88 90 91  *1  A« o l a u s l  0*85-1.04 0.92-0*97 0*90*0.93 0*88-1*00 0.95*1,12  A, l o n g i r e m i s 1*17-1*31  T. dlscaudatus 1,87*1.94s X. 62*  1,20-1*40 1.14*1.35 • 1,20*1.25 - 1*15*1*25 1*09*1.27  specimen measured  2-2 specimens measured  0. mcraurrichl  1,48-1.52  1.70* 1.63*2.40  1.52-1.83  1.35-1.57  1.45*1.69^ 1.40-1.72  O  18 DISCUSSION fhe h o r i z o n t a l d i s t r i b u t i o n s of the more common species can he considered i n d e t a i l ,  Each i s considered s e p a r a t e l y ,  because as a s p e c i e s , each has i t s own p h y s i o l o g i c a l l i m i t s , seasonal f l u c t u a t i o n and v e r t i c a l  distribution.  Spceiea l i v i n g near .the surfaces Faraealanus parvus: P. garyus was found i n the upper 14 meters of water i n seven out of nine s t a t i o n s where supplementary hauls were taken (Table 1 Appendix).  At f i v e of these s t a t i o n s the numbers caught  i n th® deep haul approximated those i n th© shallow h a u l , so i t was assumed that here P. parvus was l i v i n g almost e n t i r e l y a t the surface.  At two station®  i t occurred s l i g h t l y below 14 meters as  w e l l as a t th© s u r f a c e , while at th© remaining two s t a t i o n s i t was rare and was caught e n t i r e l y below the surface l a y e r . species therefor©  This  can b© considered  as a surface form r a r e l y occurring  P. parvus was found i n l a r g e numbers a t only three l o c a l i t i e s (Figure 3 . ) .  In T h e t i s and  Tasu I n l e t s I t seems l i k e l y that P. parvus has Increased l o c a l l y as these are i s o l a t e d populations, the absence of t h i s species at the s t a t l o n s being i n d i c a t e d by the arrows. P i g . 3.  Th© d i s t r i b u t i o n of Paraoalanua parvus  19 In Dixon Entrance, however, the p o p u l a t i o n may have come from elsewhere,  As there were no P. parvus In laden o r Masset  Inlets,  i t i s p o s s i b l e that i t could have d r i f t e d i n t o Dixon Entrance from the north or th© e a s t , T h e t i s and Tasu I n l e t s have th© highest surface of the area sampled  (Table I ) ,  temperatures  As P. parvus i s l i v i n g mostly a t  the s u r f a c e here, these temperatures may be c o r r e l a t e d with I t s abundance* Glestorecht ( 1 8 9 2 ) c l a s s e s t h i s species as c h a r a c t e r i s t i c Of a l l three f a u n i s t i o zones.  However, he has stated that i t s  northern l i m i t i a at the 10°C. Isotherm,  Although t h i s l i m i t a t i o n  i s an approximation, with a c o n s i d e r a t i o n of winter temperatures, s t a t e d by Barber and Tatoata (1954) t o be about 7*7°0.  i n Dixon  Entrance, I t i s r e a l i z e d that the water temperatures i n t h i s area i n d i c a t e that P. parvus i s l i v i n g here close to i t s northern I f t h i s I® the case, then temperature as a l i m i t i n g f a c t o r  limit,  would toe expected t o i n f l u e n c e the abundance of P. parvus. I t i s not suggested that jP*_ parvus cannot survive during the winter but i t probably does not breed then, as the temperatures are below S l e s b r e e h t ' s l i m i t * warms, breeding could commence*  However, In the spring as the water I f waters In d i f f e r e n t  localities  warm sooner than In o t h e r s , then these w i l l a f t e r a time be popul a t e d with more P. parvus* A c a r t l a clau.s.li A. c l a u s l l i v e s immediately below the surface i n most areas and does not appear t o extend much deeper than about 15 or 20 meters. meters.  Only at two s t a t i o n s was i t found completely below 14  The  20 speeies Is most common  i n three l o c a l i t i e s ( F i g . 4.). Th© greatest numbers were found i n Dixon Entrance and laden I n l e t *  For  some unknown reason I t i s absent or rare from th© surface waters of deep s t a t i o n s *  As t h i s species i s  e u r y h a l i n e , I t may p r e f e r low s a l i n i t y water found at the surface near land where runoff supplies f r e s h water. Temperature, however, may a l s o have an important i n f l u e n c e on th©  seasonal f l u c t u a t i o n s of t h i s  eopepod, s i m i l a r t o P. parvus* Figure 4* Th© d i s t r i b u t i o n . of Acartia c l a u s i .  Giesbrecht  (1892) has noted that  A , clausi. i s c h a r a c t e r i s t i c of warm and northern c o l d water hut I s  l i m i t e d t o the north by the 11°0*  Isotherm.  I t s p r o f u s i o n , then,  may b@ du© a t the time o f sampling t o th© previous as w e l l as present existence of low s a l i n i t y warm (11°C.) water. In a d d i t i o n t o the above f a c t o r s , the e f f e c t of a s c a t t e r i n g from population center's Is evident i n Dixon Entrance* c l a u s i was stmdi©d here i n more d e t a i l .  By determining  A.  th© s i z e  ranges of th® animals a t d i f f e r e n t s t a t i o n s I t was p o s s i b l e t o d i s t i n g u i s h two separate Th©  populations  and t h e i r d i s t r i b u t i o n s ,  r e s u l t s may be seen In th© f o l l o w i n g i l l u s t r a t i o n (Figure 5 ) . Th©  d i s t i n c t separation between th© size ranges of the  21  Figure 5,  a , Size V a r i a t i o n of A c a r t i a c l a u s l i n Dixon Entrance b. Currents suggested by t h i s v a r i a t i o n  22. copepods at s t a t i o n s 96 and 91 i s s t r i k i n g .  T h i s i s taken to  mean that A, e l a u a l at both these stations have grown up i n d i f f e r e n t environments temperature  probably i n s l i g h t l y d i f f e r e n t food and  c o n d i t i o n s (see page 9 ) and thus are considered as  separate p o p u l a t i o n s . At s t a t i o n 90, tha males belong w i t h the population a t s t a t i o n 96 while the females are a mixture of copepods from s t a t i o n s 96 and 91.  S t a t i o n 98 contains a mixture  of animals of both sexes from s t a t i o n s 96 and 91,  The mixing  and spreading of these populations suggests the currents ( P i g . 5) which may be r e s p o n s i b l e .  These w i l l be d i s c u s s e d  l a t e r In the account of Dixon Entrance as a whole. Oentropages  memurriehi.s £ . mciaurriehl was  found  at 26 s t a t i o n s mostly i n Dixon Entrance and Hecate  Strait*  have supplementary h a u l s , the r e s u l t s of which give a f a i r l y good I n d i c a t i o n of I t s v e r t i c a l distribution.  The  chart (fable IV)  following  summarizes  the r e s u l t s . Although there are too few s t a t i o n s at n i g h t , at these, C, mcmarriohj e s p e c i a l l y the Figure 6,  The  distribution  of Oentropages m o m u r r l c h l  young stages Is found up to the surface.  During the day, there  23 fmm i v fBB  VERTICAL' OISTKlSJTlOfl OF QhMTROF&ggS MGgJBftlOHl AS INDICATED m ITS OCGOSHBHOB I I WPPLHMEST&HY AND DEBP~ItAULS  Station  Depth Meters  Above  ' iwtature Moo* o n l y  Depth Meters  Below Koa*  immature only  Tim©  £ * Few r » lure Is tome tendency f o r I t t o remain  below*  it I s anall popula*  A t t h i s time, when  at the aujtif**** i t I s represented hy the whole of a  t l o n o r by the upward extension o f a large population* teem,' then, that £ . »e«irrlohi tha  younger stages npperaoet*  I t would  Uvea Just below the surface with Although a t times i t occurs above  14 metere* I t shows a tendency t o move down during t h * daytiKte. Moat o f the population., however,  Uvea  s l i g h t l y below 14 meters.  24 '  £• mcmurrlohl I s most common In Hecate S t r a i t .  In Dixon  Entrance i t i s common only a t on© s t a t i o n while on th© west coast i t i s rar© and s c a t t e r e d , represented' by only a few Immature specimens.  T h i s s i t u a t i o n suggests that seasonal f l u c t -  u a t i o n s ar© responsible f o r th© differ©no© i n numbers. Th© small separate populations on th© west coast and i n laden I n l e t would appear t o he populations i n c r e a s i n g under present c o n d i t i o n s , which may have been present i n Keeate S t r a i t p r e v i o u s l y .  However,  the data does not i n d i c a t e anything d e f i n i t e . S c a t t e r i n g by currents does seem responsible f o r some o f the r a r e occurrences i n Dixon Entrance* s t a t i o n s were measured.  Copepods from s e v e r a l  The r e s u l t s can be seen i n the f o l l o w -  i n g diagram ( f i g u r e ?)# I t has been suggested that 0 . atoaarriehl i s l i v i n g s l i g h t l y below the surfaee, extending i n t o deeper water than does A . c l a u s i * thus being subject to d i f f e r e n t water movements. In the western part of Dixon Entrance, r e l a t i v e l y large specimens of £* maiaurrichl ar© common..  These, however do not  move past s t a t i o n §0* as the eopepode a t t h i s l o c a l i t y are w e l l below the sis© range of those a t s t a t i o n 96*  Although i t was  very spars© a t s t a t i o n WI the one specimen was c l o s e r In sia© to those f a r t h e r west*  The few copepods a t s t a t i o n 90 have  l i k e l y corn© out of laden I n l e t .  These a r e small and can b©  traced westward i n t o Masset Sound ( s t a t i o n 73). The ©astern part o f Dixon Entrance contained very few C. momurrichi, too few f o r measurements.  Because of the pre-  dominance of young stages here, i t does not seem l i k e l y that they have com© from the west.  Figur© 7*  a) Th© s i z e v a r i a t i o n of Oantropage® moimrriohi i n Dixon Entrance b) Currents suggested by t h i s v a r i a t i o n  Th©  26 s i z e rang© of th© male© In Masset Sound i s f a i r l y  wid© I n c l u d i n g s i z e s comparable to those i n Masset and Haden I n l e t s and at s t a t i o n 96.  I t i s evident that th©  small cope-  pods at s t a t i o n 73 could have come from s t a t i o n 9 0 , west, while th© medium s i z e d i n d i v i d u a l s may i n s i d e Masset I n l e t .  directly  hav© com© from  Th© l a r g e r copepods her© are only com-  parable i n s i z e to those at s t a t i o n 96.  Because t h i s species  i s so common at s t a t i o n 73, i t i s not conceivable that i t has spread from s t a t i o n 96, through regions where i t l a scarce, unless perhaps, the i n t e r v e n i n g area has i n th© recent past contained more £• memurriohi. Although  the problem of seasonal f l u c t u a t i o n s i s  present a l s o i n t h i s speeies, some I n d i c a t i o n of s c a t t e r i n g b y water movements i a suggested  by th©  size rang© v a r i a t i o n  and population s t r u c t u r e s of t h i s speeies i n Dixon Entrance. The c u r r e n t s I l l u s t r a t e d w i l l b© discussed i n r e l a t i o n to others i n the  account of Dixon Intranee.  Tortanus diseaudatusi. Th©  v e r t i c a l d i s t r i b u t i o n of t h i s species can b©  reasonably w e l l deduced.from the twenty-flv© s t a t i o n s which have supplementary hauls*  In Table ¥ on page 27 the s t a t i o n s  are arranged i n a time sequence.  The depth of th©  supple-  mentary haul i s given with th© approximate numbers of copepods.  Below the depth of th©  shallow h a u l , th© numbers are  based on th© r e s u l t s of th© deep hauls. The  species tends to remain .below 14 meters although  It ean survive i n the surf's©© water*  The eggs ar© l a i d  both  27  TABLE V THE  VERTICAL DISTRIBUTION OP TORTAMOS PISCAPPATHS A3 INDICATED BY ITS OOOOBREHCE IH StlPPLiSMEllTARY Alt)* BEE*' H A U L S  Tim®  Station  72 126 73 64  0642" oaoo 0800 0810 94 0845 71 0900 ' 69 1055 127 1130 101 1215 ©s 1315 114 1400 128 1415 87 1500 129 1515 Off Adam Ek. 1630 121 1630 82 1645 115 1710 1715 8S< 130 1730 111 1800 83 1900 131 ' 1930 116 1945 68 '201$ 70 0145 110 0225 ©  m  eggg  © ~ common  Above Depth Hot*  3  1  14 14  14 14  Below Dopth Bos.  ©  3 14 3 1 14 1 r (imm) 1 14 14 • 3 14 f 14 3 o 14 14 14 10 • 14 3 14 14 10 r , 1mm, 14 imm. 14 1 1 14  f - few  r  *  r 4 e r  j. 2*  f e © © a l l stages f f r  f r  A f r  f f © r imm. * * e f t  V  e  r , ItBBU  f c c r  pare  above and below 14 meters but th© young seem to occur just above th© a d u l t s . Johnson (1934a) has Indicated that t h i s speeies i s very widespread although r a r e l y becoming abundant.  I t s p r o f u s i o n In  28 th« area.sampled could not be r e l a t e d to the p h y s i c a l characteri s t i c s of water at twenty meters*  As t h i s species i s a primary  c a r n i v o r e , the presence of s u i t a b l e animal food may toe responsi b l e f o r i t s abundance, e s p e c i a l l y when the s u r v i v a l of the very young n a u p l i i depends on the presence of small protozoans i n the surrounding  water.  As the food c o n d i t i o n s could not  be  estimated# the reason f o r i t s abundance In c e r t a i n areas i s still  questionable* The  s p e c i e s , however, i s very widespread (Figure 8 ) , a r e s u l t perhaps of I t s v e r t i c a l distribution*  Occurrence mostly  below 14 meters enables  the a d u l t s  to remain Inside i n l e t s with  shall-  ow s i l l s f o r example, laden I n l e t as w e l l as In the slower moving subsurface waters outside the i n lets*  On the other hand i t s  o c c a s i o n a l presence i n the surface l a y e r brings I t under the Influence of f a s t e r currents* The  water below 14 meters however,  must toe responsible f o r most of the d i s p e r s a l of t h i s species. Figure ©* The d i s t r i b u t i o n of Tortanus dlscaudatus "• caudatus was 9 a*  ' ""  measured*  This d i s p e r s a l was In Dixon Entrance  where T*  evident dls-  The r e s u l t s of t h i s are seen I n Figure  Although scarce In Dixon Entrance  l a r g e r her< than In laden I n l e t .  i t is distinctly  Because of t h e i r s i m i l a r s i z e  ranges, the copepoda In Dixon Entrance  are considered to toe one  Figure 9  a) Tto.© sis© v a r i a t i o n of Tprtanua digCaudatua i n  Dixon Entrance b) Th© currents suggested by t h i s  variation  c o n t i n u o u s population i n which the a d u l t s have been subject to th©.same past c o n d i t i o n s *  There i s only a s l i g h t suggestion  i n the s i z e of the female at s t a t i o n 72 that a few Naden I n l e t cop®pods ar© spreading out i n t o the open water.  On© of th© two  cop©pods measured i n Masset Sound was, however, u n u s u a l l y large.  I t could not have o r i g i n a t e d from i n s i d e the i n l e t  because her© T. diaeaudatua i s extremely r a r e .  Besides, th©  one specimen t h a t was measured from her© was much smaller than that i n th© sound*  The other copepod f e l l w i t h i n th©  range of the Dixon Entrance cop©pods*  sis©  I t could have o r i g i n a t e d  from s t a t i o n 96 or l e s s l i k e l y from s t a t i o n 99.  An  the sisse  v a r i a t i o n of Q. mopurrlehl a l s o i n d i c a t e d a subsurface inflow of Dixon Entrance water i n t o Masset Sound i t l a quit® p o s s i b l e that t h i s i s a l s o suggested by the d i s t r i b u t i o n of T. dlaoaudatug* Oneaea. conifer*, t 0. c o n l f e r a l i v e s i n a f a i r l y wide range of depths,  ^rom  an a n a l y s i s of the r e s u l t s from s t a t i o n 69 i n Masset I n l e t (Table ¥1 appendix) i t i s evident that t h i s species Inhabits water from 75 meters to th® surface* SO  Th© presence of eggs at  or 60 meters i a s t r o n g l y i n d i c a t i v e that conditions only  her© ar© favourable f o r breeding*  In Tasu I n l e t i t also occurs  i n th© subsurface as w e l l as th© surface water* ar© present i n hauls taken from 14 meters*  Th© eggs here  Proa th©  tempera-  ture data of both l o c a l i t i e s exemplified i n Figure 11, I t Is evident that th© eggs ar© being l a i d at a temperature of about 11°0.  31 A l l th® s t a t i o n s where 0, e o n l f e r a occurs have temperat u r e s a l l o w i n g breeding.  I n the f o l l o w i n g Figure 10 are  arranged the temperature curves f o r s t a t i o n s where 0, e o n l f e r a was  A) common or fewj B) rare and C) absent.  The l a s t group  Is made up o f s t a t i o n s taken i n west coast i n l e t s .  These were  chosen on the grounds that 0, e o n l f e r a seems t o t y p i f y Inshore waters (Figure 11),  The s t a t i o n s where 0, e o n l f e r a i s , p r e s e n t  are a l l c h a r a c t e r i s e d by a,high surface temperature, above 13*5°0V*  Of the s t a t i o n s where 0, e o n l f e r a i a absent but hav-  i n g temperatures a t which i t can breed (127, 128, 129, 10§* 108, 109| 110| 117) only three have h i g h surface temperatures and these (117,  110, 109) are shallow, having very l i t t l e bottom  water below 11°0*»  I t would seem* then* that £» e o n l f e r a he-  comes -common In l o c a l i t i e s where there i s a f a i r l y deep l a y e r T A S U  I N L E T  MASSET  4  V  on  ABSENT  I 00  M  TEMPERATURE  S C A L E  Figure 10. The r e l a t i o n o f 0. e o n l f e r a  10  c  VERTICAL  temperature  LINES  AT  II  C  t o the d i s t r i b u t i o n of  32 of water below 11°C.  as w e l l as warm surface water.  Perhaps  the development of the young r e q u i r e s a high temperature.  This  apparent dependency on warm surface water Is s u f f i c i e n t to l i m i t 0» e o n l f e r a to i n l e t s In t h i s area,  Nowhere In the  out-  s i d e water does the temperature r i s e above 13.5°C.. 0, conjfe.ra i s represented a l s o by a few I n d i v i d u a l s at other s t a t i o n s (Figure 11),  nohbreedlng  On the  f o u r of these s t a t i o n s where eggs or both eggs and  west eoagfc adults  were absent are seen to tee s l i g h t l y north of the main population centers of Tasu and T h e t i s I n l e t s *  The d i s t r i b u t i o n suggests  s c a t t e r i n g by a northward moving current i n t h i s area* Tasu I n l e t  f  Froa  0, e o n l f e r a Is presumably swept out i n th©  surface  water and c a r r i e d northward. Only a nonbreedlng part of  the  population moves out her® as e  no  egg sacs were found at s t a t i o n 124, may  I t s r a r i t y f a r t h e r north Indicate that i t Is dying or  being widely dispersed,  In three  I n l e t s to the north, however, £* eonlfera. Is represented by a few I n d i v i d u a l s i n the deeper water below 14 meters*  I f the  c i r c u l a t i o n of water In these i n l e t s i a . t y p i c a l , then the Figure 11. Distribution of Oncaea e o n l f e r a  surface  water i s moving out and being r e placed by an inwardly flowing  33  bottom water,  fhe unusual ooourrence  d©©p water her© perhaps  of 0, c o n l f e r a i n th©  suggests that a t the time o f sampling  i t was being c a r r i e d i n th© deeper water i n t o these i n l e t s from outside. S c a t t e r e d i n d i v i d u a l s a l s o occur i n Dixon Entrance*  As  these have not d r i f t e d out of l a s s e t or laden I n l e t s , i t would  seem  l i k e l y that they come from north of the sampled area.  Coryoaeus a f f i n l e t jC. a f f i n i a i s d i s t r i b u t e d much l i k e 0 . c o n i f e r s . Breedi n g p o p u l a t i o n ar© present i n Tasu and T h e t i s I n l e t (Figure 1 2 ) although they ar© not as obviousl y breeding as 0 . c o n i f e r a *  How-  ever, th© r e l a t i v e l y large number of t h i s species i n T h e t i s I n l e t i a assumed t o have . a r i s e n there. In Tasu I n l e t a female an egg sac was seen*  carrying 'Th© r a r i t y  ©f ®gg sacs may merely mean t h a t the copepods are j u s t beginning to mat© and that th© breeding cycle i s attuned t o seasonal fluctuations.  C . a f f i n i a Is more  common i n Puget Sound where popu l a t i o n s with many females Figure 1 2 .  Th© d i s t r i b u t i o n  of Coryoaeus a f f i n i a  carry-  i n g egg sacs have been noted,  34 Whether or not _0*  a f f i n l g breeds In the same c o n d i t i o n  as £• e o n l f e r a , I t Is d i f f i c u l t to say,  I t l a however, common  where the warmest surface waters of the area are found. may  This  be s i g n i f i c a n t t o the development of the young stages,  although t h i s s p e c i e s i s not always found i n the surface l a y e r . Another f a c t o r I n f l u e n c i n g i t s d i s t r i b u t i o n becomes evident with a c o n s i d e r a t i o n of Tasu I n l e t ,  C, a f f I n i a seems  to be f l o u r i s h i n g i n the long, arms hut does not spread i n t o the main part of the i n l e t *  0.  r i g h t out of the i n l e t ,  A reason f o r t h i s l a c k of d i s p e r s a l  may  e o n l f e r a on the other hand, moved  be seen In the v e r t i c a l d i s t r i b u t i o n of 0,  affinis.  I t Is  absent or at times r a r e , i n the surface water, while £* e o n l f e r a i s always present  there ( f i g u r e  13),  119  Time  S  X  •F  S  a  1  w  a  F  0  1330  1630  1030  1045  Figure 13*  The  1815  v e r t i c a l d i s t r i b u t i o n of Ooryoaeus a f f i n i s  I t might be suggested that C,  a f f i n i s i n occurring i n the deep  l a y e r o n l y f o r a time i s kept i n the arms by an inwardly i n g bottom current*  flow-  I t s mere l a c k of d i s p e r s a l at any rate  would suggest t h a t t h i s species might be able to Increase  In  other l o c a l i t i e s but has f a i l e d to reach them* There are some apparently scattered £« a f f i n i s at f i v e other s t a t i o n s *  The  l o c a t i o n s of these, however, give no clues  35 ' t o th© o r i g i n of the i n d i v i d u a l s * side the sampled Subsurface  'I'hey may have come from out-  area*  apealeat  fhe f o l l o w i n g three speeies, £ . fInmarohlcus* M. luoens a  a  d  £ • h u n g i i , ar© l i m i t e d to th© deeper water as seen In  F i g u r e 2 and i n Table I(Appendix).  Th® v e r t i c a l d i s t r i b u t i o n  of th© age groups of the f i r s t two species was determined from the data of s t a t i o n 89 i n Masset I n l e t , ( F i g u r e 26, page-5 Table f t , -Appendix),  7  ,  I n both, the young live, c l o s e r to the  surface than the a d u l t s *  Both sexes o f M. lucens l i v e a t  approximately the same depth*  The male C. finmarchieus, how-  ever, appears to occur' In deeper water than th© female.  Th©  absolute depths are not considered t y p i c a l , as M, luoens adult© are c e r t a i n l y l i m i t e d to water deeper than 30 meters In Dixon Entrance*  Because o f the s c a r c i t y of E . bung|i,, i t was not  p o s s i b l e to determine th© d i s t r i b u t i o n o f th© eg© groups* S t a t i o n 89 data a l s o i n d i c a t e s t h a t if* lucens and C* finmarchlQUs perform a d i u r n a l v e r t i c a l m i g r a t i o n , the l a t t e r moving up from.deeper  water, c l o s e t o th© surface a t n i g h t .  Other authors {Clark, 1933 and M i e h o l l s , 1933) have found t h i s same behavior i n these two sps©!©a. (1911b), however, Buealaa.ua  According to B s t e r l y  elongatua, a-close  r e l a t i v e of E,  b u n g j j , does not migrate v e r t i c a l l y In San Diego Bay.  Whether  or not t h i s i s t y p i c a l a l s o of E . b u n g l i In th© Queen Charlotte area i s not known.  T h i s s p e c i e s , however, i s not found i n th©  shallower hauls which capture 0. fjnmarehjoua and M* luoens*  30 ' Oalanua finmarohiousi The h o r i z o n t a l d i s t r i b u t i o n of t h i s species can b© Figure 14#  seen i n  fhe r e s t r i c t i o n of the  a d u l t s to the deeper water i s very n o t i c e a b l e , as w e l l as the wider d i s t r i b u t i o n of th© younger stages. Although i t can not be the young l i v e i n th©  shown that shallower  water i n Bixon Entrance, i t i s quite p o s s i b l e that during t h e i r v e r t i c a l migration they come up i n t o the surface waters long.enough to be F i g u r e 14. Th® D i s t r i b u t i o n - o f galenas f inmarchious the  transported onto shallow  areas*  T h i s process may be v i s u a l i z e d i n  f o l l o w i n g diagram. (Figure 15) I l l u s t r a t i n g the v e r t i c a l d i s -  t r i b u t i o n of £ . flnaarchicu® at th© l o c a l i t y i n d i c a t e d by the l i n e i n f i g u r e 14*  Figure 15,  The v e r t i c a l d i s t r i b u t i o n of Oalanus o f f Naden I n l e t In Dixon Entrance  finmarehloua  37 T h i s e f f e c t i a a l s o n o t i c e a b l e i n Tasu I n l e t , where the young move i n t o the shallow arms away from th© main population i n the deep part o f th© i n l e t .  In both these p l a c e s , s c a t t e r i n g Is  probably  the r e s u l t of currents just above SO meters.  Metriala  luoenst  The d i s t r i b u t i o n of M» luoens (Figure IS) suggests that i t i s l i m i t e d t o deeper water than i s 0 . flnmgrchleua*  T  h e upper  l i m i t s t o i t s v e r t i c a l rang© i n different areas (Table II) a l s o indicates this*  I t s r e s t r i c t i o n t o th® deeper water i s very  l i k e l y r e s p o n s i b l e f o r the l a c k of d i s p e r s a l of the young stages. In o n l y two l o c a l i t i e s , o f f Baden Inlet and In Tasu I n l a t , i s  this effect evident* Euoalanua b u n g l i *  As this species was so rare, no conclusions could toe drawn from th© age distribution®.  s  tag© V"  copcpodids and adult females wer© present  i n n e a r l y a l l samples i n  which I t was found*  'Ihere i s th©  s u s p i c i o n , however, that t h i s species i s more common than the plankton samples i n d i c a t e * Johnson (1943) has found that i t i s quit© capable o f evading plankFigure 18* Th® d i s tribution of M e t r l d l a lueena  ton nets.  58  Th© upper l i m i t s to i t s v e r t i c a l d i s t r i b u t i o n (Table I i ) i s very s i m i l a r to that of M. luoens.  I n spit© of t h i s , t h i s  species does not spread onto shallow areas the v e r t i c a l d i s t r i b u t i o n s together  (Figure 17).  of the three species are  When  considered  (Figure 18) o f f Kaden I n l e t , t h i s e f f e c t Is n o t i c e a b l e .  The f a c t that i t probably does not migrate upward i n t o an onshore c u r r e n t , evident I n the d i s t r i b u t i o n of th© other s p e c i e s , very l i k e l y accounts f o r i t s l i m i t e d  two  distribution.  /  ST. 91  92  90  97  81  DEPTH METERS  h50  C-F I N M A R C H I C U S M L U C E N S -  Figure 17. Th© d i s t r i b u t i o n of Buoalanus b u n g i l  MOO  E . BUN6II  Figure 18. Th© v e r t i c a l d i s t r i b u t i o n of three species o f f Naden I n l e t i n Dixon Entrance  39 Part 2 j  Currents suggested: by the d i s p e r s a l of Copeooda  The d i s t r i b u t i o n s of a l l the species i n c l u d i n g those which are rare are considered now i n several areas, with suggestions o f p o s s i b l e c u r r e n t patterns*  These areas a r e i n order;  Tasu i n l e t , Dixon Entrance, Masset I n l e t * Tasu I n l e t I Tasu I n l e t I s f a i r l y l a r g e and h i g h l y productive body of water on the west coast ©f Moresby Ialand(Figure ! • ) •  At the  t i n e o f sampling i t s plankton c o n s i s t e d l a r g e l y of copepods* The composition of I t s fuana, however, r e f l e c t s the i n f l u e n c e o f open ocean water, by the presence o f salps and siphonophores, as w e l l as water from shallow areas c h a r a c t e r i z e d by pelycepod larvae.  In a d d i t i o n to t h i s the presence of three long bays  make t h i s i n l e t a complex body of water*  The r e s u l t s of seven  deep plankton h a u l s as w e l l as f i v e supplementary hauls along with c o n s i d e r a t i o n s of the p h y s i c a l data and topography do, however, give some idea of I t s c i r c u l a t i o n . The main part of Tasu I n l e t i s l a r g e and r e l a t i v e l y deep (Figure 21)• I t I s separated from the open ocean by a narrow s i l l , with a depth o f 30 meters*  Towards the heads of the three  bays, th® depth decreases t o about 30 or 40 meters*  bright I n -  l e t , however, has a deeper pocket (maximum depth - 175 meters) behind a s i l l of 50 meters depth. Surface temperatures are high, up t o 19°C. i n Botany Bay,  T h i s warm water i s a l s o present In the main part of the  i n l e t but not outside the mouth,  f h e temperature, however, de-  creases sharply with depth, water a t 20 meters being around  40 1Q G.,  fhe deeper water a t s t a t i o n 118 Is near 7 ° C , The  0  s a l i n i t y a l s o Increases  sharply with depth from about 28^oo  to 3 2 ° / oo a t 20 a s t e r s becoming as high as 33 In the deeper water  a t s t a t i o n 118.  The conditions at th© surface suggest  that there has been an unmixed slowly moving l a y e r of f r e s h e r water there f o r some time, during which I t has warmed considerably. The d i s t r i b u t i o n o f surface plankton, although does suggest some outward flow of surface water.  scarce,  Two rather  r e s t r i c t e d species common i n Tasu I n l e t , 0. e o n l f e r a and J?» parvnf, are found In a l l surface samples although the majority occur below 14 meters.  Only two other species, 0. helgolandlca  and P. alnutus were a l s o c o n s i s t e n t l y found at the surface, but as these were so widespread I t was not p o s s i b l e to determine the centers o f population Increase.  0, e o n l f e r a and P. parvus  seem t o breed In the three arms (Figure 19). e r numbers, 0. e o n l f e r a without  They occur In l e s s -  i t s eggs, sacs at s t a t i o n 118.  0. e o n l f e r a only reaches s t a t i o n 124 presumably v i a a surface current*  I s the temperature i a low and the s a l i n i t y high here,  considerable mixing must have occurred between s t a t i o n s 118 and 124.  The presence of 0. c o n i f e r a suggests that some of the  mixed water has o r i g i n a t e d from the surface a t s t a t i o n s 118* A surface outflow from Wright I n l e t and Botany Bay i s a l s o suggested by the d i s t r i b u t i o n of £ . a f f i n i s , represented here by a few i n d i v i d u a l s (Figure 20). Species l i v i n g below 14 meters as well as i n shallow water are apparently l i m i t e d t o the bays (Figure 20)* Although these copepods are represented by few i n d i v i d u a l s , t h e i r l a c k of  41 d i s p e r s a l suggests  that  water between 14 and 30 meters i s not  moving out of the bays.  Alao^the abundance of 0,  conifers  and P. parvus* which l i v e mostly below 14 meters i m p l i e s that very l i t t l e  water a t these depths i s moving out*  The bottom water of the arms may be t y p i f i e d by the presence of G, fjnmarohlcue. the  In Figure 2 1 , the d i s t r i b u t i o n of  a d u l t s and l a t e copepodld stages i s seen to be i n deeper  water only.  The young, however, l i v i n g above th® adults appear  to  have spread i n t o the shallow bays i n d i c a t i n g an inward flow  of  water j u s t above 50 meters and p o s s i b l y up th® 25 meters.  M. luoens seems t o be r e s t r i c t e d t o the deeper water at s t a t i o n 118* at  The presence o f a few young i n d i v i d u a l s i n the deep pocket  s t a t i o n i n Wright I n l e t , however, suggests that strays ore  being c a r r i e d t o a s l i g h t extent over the s i l l et about 50  meters*  The remaining speeies i n the deep part do not seem to scatter*  T h i s suggests very l i t t l e  v e r t i c a l movement o f the deep  water* The r e l a t i o n s h i p of the deep water i n s i d e the s i l l outside I s i l l u s t r a t e d by (Figure 22}*  a comparison of s t a t i o n s 118 and 124  F i v e species l i v e Inside below 100 meters, M*  luoens and S. minor b e i n g uppermost. are  to that  Except f o r M* luoens a l l  represented by fewer i n d i v i d u a l s her© than outside.  At  s t a t i o n 124, an a d d i t i o n a l f i v e species occur presumably belonging  to the deeper water.  The presence of stage V oopepodids and  female a d u l t s of E* b u n g i i i s of considerable importance.  By a  comparison o f adjacent s t a t i o n s In other p a r t s , th© upper l i m i t s to  i t s v e r t i c a l d i s t r i b u t i o n I s near 38 to 100 meters.  This  sp©ci©s then c h a r a c t e r i z e s th© water about 6 to 70 meters below  42  Figure 19,  Figure 20,  A proposed c i r c u l a t i o n of the surface water above 14 meters i n Tasu I n l e t  A proposed c i r c u l a t i o n of water between 14 and about 50 meters i n Tasu I n l e t  43  C A L A N U S OoCO  Fl N M A R C H I C U S A O " L T S C O P E P O D I D STAGES  METRIDIA ADULT  l-V  L-UCENS DISTRIBUTION  C0PFP0D1D I- I I I DISTRIBUTION  •o- o-oo -c^r;- • v  •oo- o-o -o-o  •o-— —o-o-o-o-c -  -o o-ooo-o-\o  ofo  ,  •o -o-o-o-o-o-o-©\o-o-o-Oi  o---o, oo-o-o-o-o* -o-o-o -o" 1  Figure 21*  4 proposed c i r c u l a t i o n of thedeep water of Tasu Inlet __,  50-1 175- Y DEPTH  METERS 340  600  Figure 2 2 .  The d i s t r i b u t i o n of the deep water oopepods i n Tasu i n l e t and immediately outside  s i l l depth.  As S, b u n g l l does nob occur Inside the I n l e t , i t  may be s a i d the water ;Jusfc below s i l l depth l a not e n t e r i n g the i n l e t a t the present time nor has l l k o l y done ao in  the recent  past. Further evidence f o r t h i s I s o l a t i n g e f f e c t of the may be seen i n the population structure of S. japonioa.  sill The  r a r e i n d i v i d u a l s a t s t a t i o n 118 were a l l oopopodids at stage V* Outside stages XV, V as w e l l as female adults and eggs were found.  The e x i s t e n c e of rare I n d i v i d u a l s of only one stage I n *  side the i n l e t suggests that thee© have perhaps been c a r r i e d i n t o the i n l e t at some time but at present are separated from those o u t s i d e . In  According to Campbell  th® deep water at 200  develop*  t o 400  (1934) E. Japonica breeds  meters*  Here a l s o , the young  I t cannot be assumed then ,the copepodlds i n s i d e the  i n l e t a r r i v e d there a t a previous time v i a an Inwardly f l o w i n g current which c a r r i e d a few young o f f the top of the 1* population outside*  Japonica  Although i t seems l i k e l y that water 'below  the  s i l l depth has at some time i n the past moved up and over  the  s i l l , I t - l a not doing so a t the time of samplIn(S*  Any water  moving i n t o the i n l e t must be from a depth of above 20 t o 36 meters* In  t h i s a n a l y s i s , i t l a suggested that a slow surface  current moves out of the arms i n t o the main part of the i n l e t some of I t f l o w i n g over the s i l l .  There Is some movement of  subsurface water i n t o the arms at 20  to 50 meters.  The deep  water i n the center of th® i n l e t Is not mixing v e r t i c a l and at the  time of sampling Is d i s t i n c t from the deep water outside  the  inlet*  45 Dixon Entrance1 The water Immediately o f f s h o r e , north of Graham Island i a s u f f i c i e n t l y well sampled, to be Investigated i n d e t a i l . The presence of large areas of unsampl.. d water to the north and west make t h i s a n a l y s i s vary d i f f i c u l t .  The currents proposed  a r e , thus, only suggestions but they do I l l u s t r a t e the  possibil-  i t i e s f o r th© us© of plankton animal d i s t r i b u t i o n . Dixon Bntranc® has a narrow, shallow shelf ( f i g u r e 25). About 20 m i l e s from the shore o f f linden I n l e t the depth increases to 300 meters*  Only on®  sample was taken i n water t h i s deep  about 5 miles n o r t h of Lang&ra  Island*  The water i s c o l d , about 11° to 13°G, at th© surface dropping t o about 6°C. near the bottom a t th© deeper s t a t i o n s . The s a l i n i t y ranges from about 30foo t o 35°bo s t the surface* The deep water has a f a i r l y uniform s a l i n i t y of about 33?oo« Th© currents as estimated from th© d i s t r i b u t i o n of copepods w i l l be considered In three groupsS surface to 10 or 15 meters ( f i g u r e 23), between 15 and 30 meters (Figure 24) and below 30 meters (Figure 25)• The  surface movements a r e suggested by the d i s t r i b u t i o n  of A, c l a u s i , P. parvus and 1. norvegjca (Figur© 23)* Th© sis© v a r i a t i o n of A , c l a u s i i s the best i n d i c a t o r her© of an eastward movement.  I t s s c a r c i t y i n th© east suggests that i t i s  a l s o being c a r r i e d northward*  Th© occupation of tha eastern  water i n d i c a t e s that t h i s Is a somewhat separate water mass and receive s a few i n d i v i d u a l s of A. c l a u s i from th© west and possi b l y from Masset I n l e t ,  Th© flow of surface water from Haden  I n l e t as i n d i c a t e d by A * c l a u s i , i s a l s o suggested by th© d i s t r i b u t i o n of M» norvesica. This species a l s o occurring offshore  46  from laden I n l e t may p o s s i b l y i n d i c a t e an onshore surface current here. An eastward movement of water a t 15 t o 30 meters i a a l s o evident i n the s i z e v a r i a t i o n s of £ . mcmurriohi and f« dlgoauda* tug (Figure 24}, the lengths of these copepods being comparable r i g h t across the coast*  Some flow of water out of Maden i n l e t  i s a l s o i n d i c a t e d but t h i s seems to be much l e s s than th® outward flow o f surface water*  Because these species are so  scarce I t i s impossible to Judge whether or not they move north. However, a separate water mass In the east Is a l s o implied by the presence of l a r g e A» l o n g l r e m i a , above the s i z e rang© of those found In the western part*  A l s o the population of £» a e a u r r i o h l ,  composed almost e n t i r e l y o f young animals may be d i f f e r e n t t o that i n the west* fhe s c a r c i t y of a d u l t s i n t h i s area i n d i c a t e s that i n d i v i d u a l s from the l a r g e population i n Masset I n l e t are not moving out and eastward* since i t i s made up of a l l stages i n c l u d i n g many a d u l t s *  ®n the contrary, the large s i z e v a r i a t i o n  of £ . mcmurriehi I n Masset Sound Implies that some at l e a s t , , have come f r o a f a r t h e r west ( s t a t i o n 06), and the entrance t o Baden I n l e t * and a few p o s s i b l y from i n s i d e Masset Inlet*. The i n f l o w of s l i g h t l y subsurface  water i n t o Masset Sound Is a l s o  Indicated by T. dlscaudatus, a species extremely rare i n Masset Inlet. Movements of the deeper water (Figure 25) can be v i s u a l i z e d as responsible f o r the spreading of the copepodid stages of C. finmarchicus and 1* lucens onto the shallow  areas.  47  Figure 23*  Figure 24.  A proposed c i r c u l a t i o n of surface water in Dixon Entrance  A proposed c i r c u l a t i o n of water between 15 and 30 meters i n Dixon Entrance  48  Q O  C  O M M O N  H A R E  $*S#tj»» 25. A pro-pooed aireulation o f m t t r between about 40 ' and 100 maters In Dixon Intranae  &• t^mr^m  eee» t© be l i v i n g s l i g h t l y above M* .^OOQI aa  i t s young a r e spread onto mat o f the shallow areas* An example of tfel* was aoea In Figure 13* This aaattaring  pmmmbtj  ln-  dioaiea an onshore isovament of water at aboat 40 ©r even 80 meters,  f h * spread of II* Ittoena young onto t h e eholf in only 1  one plaee, opposite laden i n l e t i n d i c a t e * an exoeptional onateore ©xtrrent news at abont 50 a * t o r s *  UpwaUimg i n this region  i a ale© suggested by the appearanee at station 81 of £ . orlstafcns* a apaoiea otherwise occurring i n atieh. deeper water (station 83) from 364 staters*  T h i s specie a does net occur a t station 88 at  a depth of 806 meters. Of spools 1 note l a tho spread of very young C» f i n y r e h * Icaa i n t o laaaet Sound (eopepodid stages 11 and 1 1 1 ) . The o r i g i n o f these oould o n l y he the open w a t e r , as o n l y stage 7  being found Inside Masset I n l e t .  The presence of these young  stages here along with specimens of 0. acaturrlohl and T* d l s caudatus confirms the idea that water s l i g h t l y below the surface was moving i n t o Masset Sound from th® outside at the time of sampling* Masset  Inlett Masset i s the l a r g e s t of the I n l e t s studied (Figure 1 ) *  I t 1® about twenty miles long and f i v e miles wide*  The western  part s i t u a t e d elose to high mountains i s the deepest, about 80 meters as sounded from s t a t i o n 78 ( f i g u r e 27)*  Towards the  e a s t e r n r e g i o n the water shallows t o twenty meters.  The narrow  sound opening Into the eastern p a r t , extends f o r about twenty m i l e s w i t h a depth of twenty meters before opening i n t o Dixon Entrance* At the time of examination, th© surface water was warmer i n Masaet I n l e t than i n ©ixon Intranee*  I t decreased f r o a about  14°C. a t th© surface t o 8®0* In the deeper water* In the deep water was unusually low*  The s a l i n i t y  At the surface, I t was  22foo* Increasing s l i g h t l y a t a i d depth but decreasing again towards the bottom*  Although there are many r i v e r s entering  the I n l e t from a l l d i r e c t i o n s , the Yakun River entering the southeastern corner from a l a r g e drainage area to the south I s thought t o be the main c o n t r i b u t o r of f r e s h water,  The low  s a l i n i t y near the bottom as w e l l as the length of the shallow sound make t h l a i n l e t very i n t e r e s t i n g . The i n l e t was surveyed on th® 20th of July*  Both a deep  haul and a supplementary haul were taken at each s t a t i o n . S t a t i o n 89, however, was occupied f o r 24 hours on J u l y 24th.  50 A supplementary h a u l , pumped hose samples and a deep haul were taken four times during that period* Ten C«  species of copepods were found i n Masset I n l e t s  rocmurrlehi  P* minutua  A. longlremla,  T. discaudatus  A* c l a u s i  C. finmarchious  S*  amphltrites  0.  0,  M*  luoens  conlfera  0. h e l g e l a n d i e a and ?* mjnutus ar® Only rare specimens of E .  Inlet*  were captured.  Th©  s i x remaining  helgolandlca  ooaaaon and widespread i n th© a a p h i t r i t e s and T. diaoaudatua. species ar©  sufficiently  numerous and v a r i e d i n concentration to i n d i c a t e th© p h y s i c a l factors  influencing their distribution, Th© v e r t i c a l d i s t r i b u t i o n of th© s i x species at s t a t i o n  89 ©an be seen i n Figure 26 (Table VI Appendix).  T h i s diagram  Shows the r e l a t i v e abundance of species at d i f f e r e n t depths over a 24 hour p e r i o d .  Three species £ . mcmurriohi, A*  and A. p l a u g l are l i v i n g near the surface * to occur near three meters here*  lonqlremls,  A. lon^irernla- seems  At other s t a t i o n s I t occurs  below t h i s but i t i s assumed to b© l i v i n g j u s t below three meters*  C, mcaairrlehl occurs fro® th©  surface to about 10  meters* C. momurriohi Is common and probably breeding upper 14 meters as young wer© a l s o found here.  in th©  Its horizontal  d i s t r i b u t i o n and r e l a t i v e abundance Is i l l u s t r a t e d In the f o l l o w ing diagram ( f i g u r e The  27).  low s a l i n i t y surface water, I f Massot Inlet i s t y p i c a l  of most I n l e t s with a f r e s h water inflow, would tend to move out and over th©  sound.  £. mcmurrlohl I s , however, l i v i n g  and  51  T I M E 0015-  Figure 26.  034  5  300  -1536  The r e l a t i v e abundance and V e r t i c a l d i s t r i b u t i o n of copepods at S t a t i o n 8® In Masset I n l e t over a 24 hour p e r i o d .  52  Cj  C O M M O N  O  PEW R A R E  Fifty*  17*  The  relative abiandaaee and h o r U o n t s i i d i s t r i b u t i o n  and reprodueing i n t h i s l a y e r *  I t ©euld not become abundant  where tea aurfeae water was teeing rapidly replaced.  .g.ttOMai'glaM  would aero U M g r hoeoiae ooonon i n s l o w l y moving water* I t s abundante suggests a slow movement of aurfaoa water and the  possible eacietonee of an eddy i n the i n l e t * £• olaufli. unlike tho aforementioned speolea l i v e s entirely i n t h a s u r f a e e three l i t e r s * I t ean be seen from F i f u * 28, t h a t i t occurs n o s t oetawoillf opposite the mouths of large bays* AS aeon i n previous r e s u l t s , 4* claual i s often eowon i n b*ya. t y p i f i e d by Saden Inlet* f  A l s o the v o r t i c a l distribution  stakes i t s u b j e c t to tho outward flowing surfaee water*  It Is  possible then, that A# olaual i s more abundant i n the bays opening into iaaset Inlet* If ao, those oepepods would quite  move out into t h e mtm part and aeaouat f o r t h e l a r g e numbers s t Station* 77 and 78* fhe other three spoelea are most abundant i n the deeper  they do occur near tha surface. 0* eonlfera iaoena oeour in fewor numbers near the surfaee but  water a l t h o u g h and JM.  F i g u r e 28*  The d i s t r i b u t i o n o f A e a r t i a c l a u s i i n ttaaeet I n l e t .  C. f inma.rob.ieua approaches  the  surf so® only at night*  may i n d i c a t e th® movements of the deeper  These speeies  water*  Although. j>« c o n i f e r s l i v e s i n the whole water column, females carrying, egg sacs wer© seen only i n pumped samples taken near SO meters and i n the deep v e r t i c a l h a u l , were a l s o noted i n these samples*  Free ®gg sacs  As t h i s s p e c i e s , character-  i s t i c of th© superfamily th© Oyolepoidea,. c a r r i e s I t s eggs unt i l they hatch, i t can be assumed that th© f r e e egg sacs had f a l l e n o f f females a f t e r the sample was c o l l e c t e d .  T h i s evidence  suggests' that £# c o n i f e r s i s reproducing i n the deeper water at ahout §0 o r ©0 a s t e r s and i f so could only be producing young In the western part of th© i n l e t where these depths are found. 0, c o n l f e r a * however, i a found i n l e s s e r numbers and without egg  sees f a r t h e r ©sat (Figure 20), Considering that production  occurs only i n the west, th© "individuals l i v i n g near th© surface must d r i f t westward away frost t h e i r o r i g i n .  54  Figure 29.  fhe d i s t r i b u t i o n of Onoaea e o n l f e r a  The v e r t i c a l d i s t r i b u t i o n o f M. 3,uoen,s i s s i m i l a r to that of 0. e o n l f e r a .  The a d u l t s l i v e a t 30 to 50 meters, while the  young l i v e above t h i s and «re c o n t i n u a l l y present even I f scarce In the surface l a y e r ,  f i g u r e 30 (Table VII Appendix) Indicates  th© population s t r u c t u r e s of t h i s species at d i f f e r e n t s t a t i o n s , S T A T I O N S  7 7  7 6  7 8  noiW ungift ittuffig DEVELOPMENTAL  Figure 30. °  7 4  TTLMJSYM  7 5  m m g Y v i  STAGES  The population structures of Metrldia luoens In Masset I n l e t  55 T h i s data i s presented a l s o i n Figure 51.  Th© extents of th©  d i s t r i b u t i o n of a d u l t s and copepodlds i s Indicated.  I t can  seen that th© a d u l t s ar© present only In the western area  be  while  copepodlds i n stags v" range f a r t h e r east and copepodlds stage I cover th© whole sampled area.  A® only the young ar©  th© c u r r e n t s responsible f o r t h i s must be  Figure 31.  dispersed,  present above 30 meters.  The h o r i z o n t a l d i s t r i b u t i o n of s e v e r a l oopepodld stages of i s t r l d i a luoens In Masset I n l e t  Oalanus f l n p a r c h i e u a i s represented  In Masset I n l e t  by  copepodlds i n stag© ¥ although a few adult females were seen as w e l l as an o c c a s i o n a l oopepodld stag© IV. s t a t i o n s 77, 70, 75, 89 (Tabl©  TO  I t occurs e n t i r e l y at  Appendix).  Although I t does  move up to the surface at night, I t l a not dispersed, as Is M. luoens, over the r e s t of th® i n l e t . spreads the young of M. as I t does not  Hence the current which  luoens must not be very f a s t or  steady  cause C. fInmarchlous to spread while the species  56 Is near the surface* Th® three species l i v i n g i n the deeper water appear to he i s o l a t e d i n Masset I n l e t by th© long shallow sound.  Two  M * luoens and jO. c o n i f e r s are reproducing at the time of but i t i s d o u b t f u l that G* finmarohious i s doing so.  of them, sampling  T h i s specie a  i s mostly represented by stage ? copepodlds, a few adults and o c c a s i o n a l stag© IV oopepodld.  an  I t Is very d i f f i c u l t to say  whether these were washed i n t o Masset r e c e n t l y or whether they have been there f o r some time.  I t seems p o s s i b l e that th©  very young stages of £ . finmarohious. may the i n l e t .  At th© time of sampling  have been c a r r i e d Into  I t I® present i n i t s very  young oopepodld stages l a Masset sound*  I t i s not impossible  that a strong Inflow i n t o the sound could b r i n g a f a i r number of t h i s species i n t o the i n l e t * In sumnary, I t seems that there are no d e f i n i t e r a p i d moving currents i n Slasset I n l e t *  As species l i v i n g near the  surface are able to reach l a r g e numbers by reproducing, i t i s reasonable to suggest that very l i t t l e  surface water Is moving  out over the sound at the time of sampling*  There i s some  i n d i c a t i o n , however, that surface water i s moving out of the bays.  A l s o there seems to be a s l i g h t eastward movement of water  above 50 meters*  Apparently very l i t t l e  water i s moving i n t o  th© I n l e t over the sound, judging from the absence of young £* *inmarohious i n s i d e th©  Inlet*  57 SUMMARY. AID COHCttTSIOHS Thirty-two  species of copepods .were i d e n t i f i e d .  c h a r a c t e r i s t i c of northern c o l d water.  A l l were  These species were found  to have c e r t a i n v e r t i c a l ranges, some o c c u r r i n g only In the deep water*  Hence, fewer species were encountered from hauls at  shallow s t a t i o n s * With a c o n s i d e r a t i o n of v e r t i c a l d i s t r i b u t i o n s , the r e l a t i o n o f copepod d i s t r i b u t i o n and abundance to p h y s i c a l conditions was s t u d i e d .  Temperature was  found to Influence th® abundance of  several species l i v i n g near the surface.  Aa these species were  l i m i t e d to the north by temperatures present i n th© area during the winter, I t was  suggested t h a t they became abundant only i n  th© summer and f i r s t i n l o c a l i t i e s which warmed e a r l y .  It  was  a l s o evident that several species l i v i n g In deeper water could breed only In the low temperature® there, thus not o c c u r r i n g In l a r g e numbers i n the shallower water.  In a d d i t i o n to temperature  low s a l i n i t y seemed to c o r r e l a t e with the presence of one known to p r e f e r water of low s a l i n i t y .  species  I t l a apparent, then,  from t h i s study that p h y s i c a l c o n d i t i o n s have some influence on the presence and abundance of copepod® In the Queen Charlotte area. D i s p e r s a l was  a l s o seen to influence d i s t r i b u t i o n .  s c a t t e r i n g of a species from i t s breeding area could be and currents responsible f o r t h i s are suggested. the extent of surface outflow and  subsurface  The  traced  In lasu I n l e t ,  Inflow are Indicated  as w e l l as th© I s o l a t i o n of the deep water Inside th©  shallow  , 58 s i l l from that outside* was  In Masset I n l e t , th© surface outflow  not outstanding, i n s t e a d , th© existence of slowly moving  addles seemed evident. appeared  Species l i v i n g In th© deep water her© a l s o  to be i s o l a t e d by the long shallow s i l l .  Although  no  i n f l o w from Dixon Entrance Is evident at th® time of sampling, i t Is suggested that p e r i o d i c a l l y a f a i r l y l a r g e amount of water moves Into Masset I n l e t from Dixon Entrance* A study of Dixon Bntranc© i n d i c a t e d an eastward flow of surface water as w e l l as a movement of deeper water onto the s h e l f p a r t i c u l a r l y o f f Naden I n l e t * T h i s study although e x p l o r a t o r y has i n d i c a t e d some of the f a c t o r s which need to be considered l a future plankton studies* It cal  has i l l u s t r a t e d the b i o l o g i s t ' s ne©^ f o r knowledge of p h y s i c o n d i t i o n s , ©specially the movements of water which have  been shown to have a considerable e f f e c t on  th® d i s t r i b u t i o n of  copepods. A l s o many problems concerning the plankton animals themselves ar© evident. fluctuations, t h i s area*  A more b a s i c knowledge of t h e i r seasonal  v e r t i c a l migrations and physiology Is r e q u i r e d f o r  I t i s f e l t that l a order to solve these problems,  th© plankton should be s t u d i e d as such and not as supplements t o physical studies*  A© t h i s study has shown, copepods have poss-  i b i l i t i e s i n current determinations but these w i l l not b© f u l l y u n t i l more I s known about th© animals  themselves*  realized  59  ACKNOWLEDGMENTS The author wishes t o thank those who c o l l e c t e d the samples,©specially Dr. R, P. Scagel and Mi*. P.G. Barber and the o r g a n i s a t i o n s which made th© c r u i s e p o s s i b l e , th© P a c i f i c 0c©apograph!e r o u p and th© Defence Research Board. &  Thanks  a l s o go t o f e l l o w students i n c l u d i n g s e v e r a l at th© U n i v e r s i t y of Washington. E s p e c i a l l y appreciated i s the encouragement and c a r e f u l guidance given throughout th© study by Dr. l.M. Cameron.  eo BIBLIOGRAPHY. Barber, P. and S. Tabata, 1954. The Hecate S t r a i t oceanographie p r o j e c t . Prog Rep. Pac. Coast S t a t . , F i s h . Res, Bd. Canada no. 101:20-22. Barnes, H. and S.M, M a r s h a l l , 1951, On the v a r i a b i l i t y of r e p l i c a t e plankton samples and some a p p l i c a t i o n s of "contagious" s e r i e s to the s t a t i s t i c a l d i s t r i b u t i o n of catches of r e s t r i c t e d periods. Journ. Mar. B i o l , Assn. 308 233-263. Campbell, M.I., 1934, The l i f e h i s t o r y and post embryonic development of the copepods, Calaaag tonaua, Brady and Buohaeta Japonica Marukawiu #ourn." Canada B i o l .  -M. n p i r  C l a r k e , G.L., 1933. D i u r n a l migration of plankton i n the Gulf of Maine and i t s c o r r e l a t i o n with changes i n submarine i r r a d i a t i o n . B i o l , B u l l , 65:402-436. Dahl, F., 1893. Plueromamma, ©In Xrebs mit Leuchtorgan* Zool* AM. 16:104-109, Davis, C.C., 1949, The pelagic Copepods of the northeastern P a c i f i c . Univ. Wash. Pub, i n Biology. 14:1-118. Digby, P.B.S., 1950. at Plymouth.  The b i o l o g y of some planktoule copepods Journ. Mar. B i o l , Assn. 29:393-438.  Ekman, S., 1953. Zoogeography of the sea. son L t d . , London*  Sedgwick and Jack-  E s t e r l y , CO., 1905. The p e l a g i c Copepods of tha San X>i©ge r e g i o n . Univ. C a l i f * Pub. Z o o l . 2:113-233. E s t e r l y , CO., 1911a, T h i r d report on th© Copepoda of the Diego region* Univ. C a l i f . Pub. Z o o l . 6:313-352.  San  E s t e r l y , 0,0., 1911b. The v e r t i c a l d i s t r i b u t i o n of Supaligns eiongatus i n th© San Diego region during 1909* "oniv. C a l i f , P u b . Zool* 8*1-7* E s t e r l y , CO., 1924, The f r e e swimming Gopepoda of San Francisco Bay. Univ. C a l i f * Pub. Sool. 26:81-119, P i s h , C L . , 1936 a. The b i o l o g y o f Calanus f inmarchicus i n th© Gulf o f Maine and Bay o f M n d y T " B i o l . B u l l . 70: 118-141. P i s h , C.L., 1936 b . The b i o l o g y o f Psuedocalanua minutus In th© Gulf of Maine and Bay o f Pundy. B T o l T B u l l . "70s 193-216,  61 Furuhaai, K., 1953. On th© v e r t i c a l d i s t r i b u t i o n of animal plankton In the Sea of Japan o f f San'in d i s t r i c t i n th© summer of 1953. Pub. 2©to Mar, B i o l . Lab. 38 81-74. Giesbreeht, f , , 1892. Syatemetik and F u a n i s t i k der p e l a g i s chen Copepoden des Golfes von Seapel und der angrenzenden Meerabsenltts, Fuana und F l o r a des Golfes von leap©!, monog. 19, B e r l i n . Hardy, A.C, and E.R. Gunther, 1935, fhe plankton of the South Georgia whaling grounds and adjacent water 1926-7. D i s c . Rep, no, 11. Jesperson, P. and F.S. Hussel, 1952, ( D i r e c t o r s ) . Flehea d" i d e n t i f i c a t i o n du zooplankton* Cons, E x p l , Mer», not, 1*17, 30-49. Johnson, 1»W,, 1934a. The l i f e . h i s t o r y of th© copepod Tortanus discaudatus (Thompson and S c o t t ) . B i o l , B u l l .  #MM-20C Johnson, M.S., 1934b. The developmental stages of the ccpepod g g i l a b l d o c e r a a m p h i t r i t e s Memurrioh, B i o l , m i l . 67*466483.' Lebour, M.V., 1916, Stages l a the l i f e h i s t o r y of Galanua flnaaarchloua (Gunnarus) experimentally reared by I r , L.R. Grawshay In th® Plymouth Laboratory, Journ. Mar. B i o l . Assn. 1 1 i l S l o h o l l s , A*G., 1933. The b i o l o g y o f Galanua flnmarohjoue* I . Reproduction and seasonal d i s t r i b u t i o n i n the Clyde Sea area during 1932. Journ. Mar.Biol,Assn. 19t 85*101. Sara, G.O., 1903. The Crustacea of Norway. Museum.  V o l , IV.  Bergen  Sverdrup, H.U., Johnson and R.H. Fleming, 1952. The oceans. P r e n t i c e - H a l l i n c . , Hew York* U s s i n g , H.H., 1938. The Biology of some important plankton animals i n the f i o r d s of East Greenland, Medd,  oWnland, Bd. 10081-108.  Wilson, C.B,, 1950* Oopepoda gathered by the United States steamer " A l b a t r o s s " from 1887 to 1909, c h i e f l y i n th© P a c i f i c Ocean* U.S.Iat.Mus.Bull. 100, v o l , 14, p t . 4.  Afturaoc  S - rare F - faw  (3 * ammam A «• atouftdkant  a * «SS® 1 » iKtmafcar© o n l y -* » a l l i l t g t i t o  T ABLE I  THE OCCURRENCE AND RELATIVE ABUNDANCE OF TEE COPSPOD SPECIES I I ALL THE PLANKTON SAMPLES Hecate s t r a i t  Locality 69  68  Station  23 1 22 1 44 1 6  Saul Depth  104  102  101  18 1  36  75  69  27  u.e R  C P p F C P P P F c F " c" 9 ' 1 "¥'A""" ~ c' RR ft ? p T "o"T'Ttl P aF IR C R "inr""« T~¥~ " '"'"C " f ' '0  0 f  c  ,  — — — ~  fi-  .  ..  ,... -....,„.„ ,, ,„,, tr  , ,  _ -  w  C 0 c  p R  0  R R  .—-« ii  ———— P  01  Ri  „,,• — . ,  C  Si  ' ;  .  s  R  C C F C  if  _ ,  ' R  91  ^ — _ — « — — . — — ~ ~ — — • —  :  R  C 0  0  L  s  100  12.6  /  0  0.  R * rare e - eggs  103  14.5  S u r f t o t Temp/c 14.8 CopepodsJ 0. Helgoland!ea P. mfnutu© A. c l a u s i A, longer imls C. racmurrlchl T. d i s T a u d a ' t u s , nor vesica eon I f era E. hlrundoidos" P. Darvus 1. r o s e a • C. a f f i n i a ' " C,' f Inmarchicua M. lucens E. nmohitrltea 0. pluwifera .. A . araatus 1» h u n g l l slafilFl C. erlatatua" 6. tonsua S. minor 1 . japohiaa 0. oolumb-Ia© . Pleuromamma Betordrhabdua .. Saetanus C« a o u l c b r n l a M. lonft® Rhlncalanus Chiridlus S. aubdeptata  70  65  64  — —  —.—.——  ;  ;  ~ — —  — . —  — ——•  • **—  f  -  ~—,—>—.—• •— — —— —.—••——— ~— _ — — . - „,„,.,,..,. » . , ,——.—.—,—— ———•—-— , , . ,—,—~— ————.———, , . — — „— , -—,—:—:————-•— ;  , . ,„„,,.....  f> . few 0 - common 1 - Immature o n l y  ,—-—•  —  — * —  A - abundant -> * a l l stages t o  TABES I  (continued)  Dixon &ntrance  Locality 71  Station Haul Depth _ _ _ _ _  _  127 1 91 _ _ 12.7  S u r f a c e Temp.  Cooepods: 0. h e l g o l a n d l o a / P F l rnlnutus 0  A. clausl  A. l o n s e r l r n l a C Fl C. racmurriehiF"  "T» discauoTatua 1.' nb'rve^lca" 0. eonlfera  99  e  R  X  TT  X  72  90  73  91  92  90  97  81  88  47 54 5 15 3 27 8 35 65 »i-5 i o 42 3 45 • — " ' H 9 — — 124 13.4 12.0 11.9 t  __ 1 c e o  x  ~  x  T  TTX  G  X  F 0 C CCA  C t~  C C F C  C C C C  X  xx~x  - f — i ! — m r  TT  T l IT F  X~l  T—rr—X F  TTX  hiruHd*oTdes  T He  J  affinlt  T  ^^ 'F£hS7cM^u?"^?* M. lucens  E. amphitritgi  0. p l u m i f a r a A. armatus E. hu  K slSilaV  X  R. T  *UT~cri s t a t u s C.  tonsus  S. rainor  ponaea .. "C~. co lum'biae  Pleuromamma i. j " » .' " f . ' g Beterorhabdug, Gaetanua C. a o u i c o r n i s M. l o>na< nga  r a m  1  fthincalanus "Chiridiua ~S. a u b d e n t a t a "  X  X  A  T Rc  x x —  x—c~x~""xr  X  OA F* C  —TR  "M. rosea C.  96  "I  TABLE I (continued) "Dixon Entrance 87 8 4 T !  Locality  BT,I,94 93 "95  83 Station Haul Depth, 20S 10 364 10 42 42 5 10 5 91 3 55 5 meters Surface Temp. 12.9 11.8 °C 11.8 Copepodss C C GA C C 0 C O.helgolandloa C G C P.minutua" ' C 5" 5"  R _ rare  P- few  C - common  A - abundant  e -  1 55 25 22  If© st ^Goas t I n l e t a 0.6106 110^ 108 109 107 i l l 14  35 14 100 12.7  24 219 119 13.7  A C  eggs  I - Immature only  all  stage to  T A B L E I (continued) Locality" —TTZ U S Station f>ep'tn" Surface Temp* ''I4»'8 1S»1 Copepods* O.helRolandlca C 0 R~X F.minutus A.clausl A , longe Hmra'~~~^X X O.mcmurrlcM  ¥ . d i s cauda' tu a" M.norvegica" '"'" O.conTfera E.hiruDdQidea  "X  ITS——JTW~ TBTo c  X~ X X  1  X  "0  -  s  urnr  X  F  X  T S s  TiT TBTo"  G  A  X  Z  X  "IT X  TO  X  X X  X X  Ii  7  F  X  XT  F" X  X  Gaetanui" tr.aeuieornla M.longa Rhincalanus Chiridius S«aubdentata R - rare  F - few C - common  A - abundant  e - eggs  1 - immature only  - - a l l stages t o  it  a  o  jctjc  0 •  1  If  ill  38  r u  1  i  si  IP  n  He  o  I  a  1  4 erf SHI o  erf  8  to  o  3  HI  a  lo  o  o  a  o ow  H»  ©  s> o  I  I  I  I  °5 I f I i M  |  1  a  0  ST  §  3  |  w t  I  * 2  p  o  II  imsi  piHHICtHH •13 O <  o  El  I  Ho  o  fl  I  &5 o o  ml  o  M  92s  o  81 so mm  o  a  - J o  o  O  13  H H  O  4 o  6»  w  3 -a o o c4  a* w o o ojo  o  g  8  ft  01  oi PR  S a m «n  of?  IK  a  4  3 da  S3  I O CJ »  O *  *  8 a  TABLE ¥11  T H E  ^gg  F10P0RTI0IS OF OOP1POD1T1 STAGES OP M. lusena AMD £.  ,  .  II  station  fatrloli^wM — w r  ^  Calanus  tinaareaions.  W  flnmarehl©as  III  iv  — g ^ ^ Z Z I I Z Z —  15"  

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