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Copepod community dynamics in a highly variable environment : the Strait of Georgia Black, Graeme Robert 1984

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COPEPOD COMMUNITY DYNAMICS IN A HIGHLY VARIABLE ENVIRONMENT - THE STRAIT OF GEORGIA By GRAEME ROBERT BLACK B.Sc, Queen's University, 1980 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in THE FACULTY OF GRADUATE STUDIES (Department of Oceanography) We accept this thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA APRIL 1984 ®'Graeme Robert Black, 1984 In presenting t h i s thesis i n p a r t i a l f u l f i l m e n t of the requirements for an advanced degree at the University of B r i t i s h Columbia, I agree that the Library s h a l l make i t f r e e l y a v a i l a b l e f o r reference and study. I further agree that permission for extensive copying of t h i s thesis for s c h o l a r l y purposes may be granted by the head of my department or by his or her representatives. I t i s understood that copying or pub l i c a t i o n of t h i s thesis for f i n a n c i a l gain s h a l l not be allowed without my written permission. Department of OCEANOGRAPHY The University of B r i t i s h Columbia 2075 Wesbrook Place Vancouver, Canada V6T 1W5 Date May 1 0 , 1 9 8 4 7Q^ Abstract Along the S t r a i t of Georgia's north-south axis there exists a k i n e t i c energy continuum from a high in the Southern S t r a i t which is characterized by strong t i d a l mixing, to a low in the Central S t r a i t where a stable s t r a t i f i e d water column is the dominant physical feature. The copepods for this region were found to arrange themselves along the north-south axis i n an ec o l o g i c a l continuum where small r-selected species dominated the Southern S t r a i t while in the Central S t r a i t the trend was towards larger more K-selected species. The l i f e h i s t o r i e s of the various copepod species are discussed and appear to play an important role in maintaining the "r-K" continuum. Many of the K-selected animals undergo an ontogenetic migration which co n s t r i c t s them to the deeper regions of the Central S t r a i t through f a l l and winter. This migration counterbalances the movement of juvenile copepods which appear in the surface waters through most of the spring. At the surface, the juvenile copepods are exposed to estuarine, t i d a l and wind derived c i r c u l a t i o n s which tend to concentrate them in the boundary regions of the extreme northern Central S t r a i t and in the Southern S t r a i t , away from the overwintering areas. S i m i l a r i t i e s exist between the copepod members of the North Central P a c i f i c Gyre and the S t r a i t of Georgia, yet, those members which are abundant in both environments show l i f e h i s t o r i e s which are markedly d i f f e r e n t between the two regions. Based on their e c o l o g i c a l r o l e , the S t r a i t of Georgia copepods may be divided into five groups: reproductive, growth, reproductive and growth, carnivorous, and r-sele c t e d . The f i r s t four are composed primarily of the more K-selected species. Each group shows unique d i s t r i b u t i o n a l patterns r e f l e c t i n g the d i f f e r e n t forces a f f e c t i n g them, yet none of the groups can be said to be completely independent of the others. - iv -Table of Contents Page ABSTRACT i i TABLE OF CONTENTS iv LIST OF TABLES v i i LIST OF FIGURES v i i i LIST OF APPENDICES x i i i ACKNOWLEDGMENTS xiv INTRODUCTION 1 Objective and Hypothesis 1 B i o l o g i c a l Background 2 Physical Environment 4 Sampling Background 8 METHODS AND MATERIALS 10 Ichthyoplankton Sampling Program 10 Sample C o l l e c t i o n and Preservation 10 Station Selection 11 Sample Processing 12 Predator Wet Weight 13 Copepod Subsampling 14 Copepod I d e n t i f i c a t i o n 15 Limits, Error Estimates and Conversions 15 - v -Table of Contents (cont'd) Page Quantitative Limits 15 Predator: Dry Weight Determination 16 Copepod: Dry Weight Determination 17 Folsom Plankton S p l i t t e r Error 19 Predator Wet Weight Error 19 S t a t i s t i c a l Treatment of Data 19 Raw Data 19 Processed Data 20 Clustering of Stations 20 Clustering of Species 24 RESULTS AND DISCUSSION 26 Limits, Error Estimates and Conversions 26 Quantitative Limits 26 Predator: Dry Weight Determination 27 Copepod: Dry Weight Determination 28 Folsom Plankton S p l i t t e r Error 28 Predator Wet Weight Error 29 Clustering of Stations and Species: An I n i t i a l Perspective . . . 29 Synopsis 30 Detai l s 34 Table of Contents (cont'd) Page Cruise 1 3 ^ Cruise 2 35 Cruise 3 35 Cruise 5 36 Cruise 7 36 Cruise 8 38 Cruise 9 3 8 Cruise 10 39 Cruise 11 40 Community Dynamics 40 Disperal Mechanisms 44 Estuarine Dispersal 45 T i d a l Dispersal 48 L i f e Strategies 51 Predator - Prey 61 Conclusions 63 P r e d i c t i o n 66 Recommendations 67 REFERENCES 68 APPENDICES 128 - v i i -LIST OF TABLES Page 1. Cruise schedule, 1981 77 2. T-test for 351 um vs. 200 um mesh tows 78 3. Copepod wet and dry weights 80 4. Predator wet weight 95% confidence l i m i t s 84 5. Copepod species encountered and abbreviations used . . . . 85 - v i i i -Page LIST OF FIGURES 1. The S t r a i t of Georgia 89 2. Schematized mixing for the S t r a i t of Georgia 90 3. T-S p r o f i l e s for the S t r a i t of Georgia, 1968 91 4. Ichthyoplankton survey g r i d , 1981 92 5. Selected transects and controls 93 6. Predator wet weight vs. dry weight 94 7. Folsom plankton s p l i t t e r : counts vs. confidence l i m i t 95 (a) 1/8 S p l i t (b) 1/16 S p l i t (c) 1/32 S p l i t (d) 1/64 S p l i t 8. Numerical abundance of l i f e cycle stages of Neocalanus plumchrus . 96 9. N. plumchrus: north - south d i s t r i b u t i o n patterns 97 (a) C3 (b) C4 (c) C5 (d) C6 10. N. plumchrus: plume d i s t r i b u t i o n patterns 98 (a) C3 - ix -L i s t of Figures (cont'd) Page (b) C4 (c) C5 (d) C6 11. Clustering of stations 99 (a) Cruise 1 (b) Cruise 2 (c) Cruise 3 (d) Cruise 5 (e) Cruise 7 ( f ) Cruise 8 (g) Cruise 9 (h) Cruise 10 ( i ) Cruise 11 Clustering of s] (a) Cruise 1 (b) Cruise 2 (c) Cruise 3 (d) Cruise 5 (e) Cruise 7 ( f ) Cruise 8 100 - x -L i s t of Figures (cont'd) Page (g) Cruise 9 (h) Cruise 10 ( i ) Cruise 11 13. E q u i t a b i l i t y : north - south pattern . 101 14. E q u i t a b i l i t y : plume pattern 101 15. Pseudocalanus minutus: north - south d i s t r i b u t i o n pattern . . 102 16. P^ . minutus: plume d i s t r i b u t i o n pattern 102 17. A c a r t i a longiremis: north - south d i s t r i b u t i o n pattern . . . . 102 18. Numerical abundance of adult Oithona s p i n i r o s t r i s and S c o l e c i t h r i c e l l a minor 103 19. 0. s p i n i r o s t r i s : north - south d i s t r i b u t i o n pattern 104 20. 0. s p i n i r o s t r i s : plume d i s t r i b u t i o n pattern 104 21. S. minor: north - south d i s t r i b u t i o n pattern 105 22. S. minor: plume d i s t r i b u t i o n pattern 105 23. Numerical abundance of l i f e cycle stages of Calanus marshallae . 106 24. C. marshallae: north - south d i s t r i b u t i o n patterns 107 (a) C5 (b) C6 25. C. marshallae: plume d i s t r i b u t i o n pattern 108 - x i -L i s t of Figures (cont'd) Page (a) C5 (b) C6 26. Numerical abundance of l i f e cycle stages of Metrida lucens . . . 109 27. M. lucens: , 110 (a) C5 (b) C6 28. M. lucens: . I l l (a) C4 (b) MC5 (c) FC5 (d) MC6 (e) FC6 29. M. lucens: . 112 (a) C4 (b) MC5 (c) FC5 (d) MC6 (e) FC6 30. Numerical abundance of l i f e cycle stages of Eucalanus bungii . . 113 31. IE. bungii: 114 - x i i -L i s t of Figures (cont'd) Page (a) Cl (b) C2 (c) C3 (d) C4 (e) C5 ( f ) C6 32. Numerical abundance of l i f e cycle stages of Calanus p a c i f i c u s . . 115 33. C. p a c i f i c u s : north - south d i s t r i b u t i o n patterns 116 (a) C5 (b) C6 34. Numerical abundance of l i f e cycle stages of Ch i r i d i u s g r a c i l i s . 117 35. C. g r a c i l i s : north - south d i s t r i b u t i o n patterns 118 (a) Ju (b) C6 36. Numerical abundance of l i f e cycle stages of Gaidius minutus . . .119 37. G. minutus: north - south d i s t r i b u t i o n patterns 120 (a) Ju (b) C6 38. Numerical abundance of l i f e cycle stages of Euchaeta elongata . 121 - x i i i -L i s t of Figures (cont'd) Page 39. E. elongata: north - south d i s t r i b u t i o n patterns . 122 (a) C2 (b) C3 (c) C4 (d) C5 (e) C6 40. Schematized model for Neocalanus plumchrus d i s t r i b u t i o n . . . . 123 41. T i d a l cycles and coincidental sampling of Southern S t r a i t stations 124 42. Predator - copepod: biomass and r a t i o vs. time 125 43. Copepod biomass: north - south d i s t r i b u t i o n pattern 126 44. Predator biomass: north - south d i s t r i b u t i o n pattern 126 45. Timing of prominent deep-water adult copepods 127 - xiv -LIST OF APPENDICES Page A. Non-reduced, processed copepod data 128 B. Processed predator wet weights 146 C. 351 urn vs. 200 urn mesh tows 148 D. Predator wet weight and dry weight data 151 E. Copepod wet weight and dry weight data 152 F. Predator biomass error estimation 156 - XV ACKNOWLEDGMENTS I would l i k e to thank Dr. Robin LeBrasseur for acting as my advisor at the P a c i f i c B i o l o g i c a l Station i n Nanaimo and for h i s invaluable comments and d i r e c t i o n given both during the proposal and i n the organization of the th e s i s . Many of his ideas have been incorporated into the current thesis and I expect his influence w i l l be with me for many years to come. I am also indebted to Dr. Tim Parsons for acting as my supervisor at the University of B r i t i s h Columbia and for the time he spent i n organizing my thesis committee, as well as, h i s i n s i g h t f u l c r i t i c i s m s of the thesis. A s p e c i a l thanks goes out to John Fulton for h i s constant good humour, patience, and f r u i t f u l discussions. I am gr a t e f u l to Dr. Paul Harrison and Dr. Paul LeBlond who sat on my committee and for reading and c r i t i c i z i n g this manuscript. Much appreciation is extended to Dr. John Mason of the P a c i f i c B i o l o g i c a l Station for making available his very extensive plankton samples from the S t r a i t of Georgia. The s t a f f of the P a c i f i c B i o l o g i c a l Station were very h e l p f u l and provided an excellent setting for research and learning. F i n a l l y , to my Dad, for e d i t i n g the f i r s t draft of the thesis, but most importantly for his encouragement and the confidence he placed in me, I w i l l always be g r a t e f u l . - 1 -COPEPOD COMMUNITY DYNAMICS IN A HIGHLY VARIABLE ENVIRONMENT - THE STRAIT OF GEORGIA Introduction Objective and Hypothesis This study, based upon quantitative zooplankton samples, addresses the concept of community structure. The i n i t i a l hypothesis i s that communities r e f l e c t t h e i r environment. In the present s e t t i n g , the open waters of the S t r a i t of Georgia, the marine environment varies from a region ex h i b i t i n g a well s t r a t i f i e d stable water column i n the Central S t r a i t , to a region of r e l a t i v e l y high turbulence and exchange of water in the Southern S t r a i t (Waldichuk, 1957). In the Central S t r a i t , changes to the physical/chemical properties of the water column occur i n a predictable seasonal fashion over the time scale of weeks to months. In the Southern S t r a i t where t i d a l mixing i s strong, time scales of hours are involved (Samuels, 1979). Thus within the r e l a t i v e l y small scale dimensions (110 km by 22 km) being considered, there exists a dynamic t r a n s i t i o n from a region of low k i n e t i c energy to one of high k i n e t i c energy (LeBrasseur pers. comm.). It may be expected then, that the organisms inhabiting the S t r a i t of Georgia a l i g n themselves within this k i n e t i c continuum according to c e r t a i n features which make them better adapted for l i f e i n one or the other region. The objective of this thesis, based upon the hypothesis put forward, is to describe the copepod communities inhabiting the open waters south of Texada Island i n the S t r a i t of Georgia and i d e n t i f y their c h a r a c t e r i s t i c features. Copepods usually dominate the zooplankton, both i n terms of - 2 -weight and numerical abundance. Being a diverse group of organisms, the order Copepoda is well suited to studies of communities. Williams et a l . (1981) recognized two types of multispecies organization, deterministic and p r o b a b i l i s t i c . This two-fold d i s t i n c t i o n is e s s e n t i a l as the f i r s t implies i n t e r a c t i o n and p r e d i c t a b i l i t y whereas the l a t t e r suggests random ass o c i a t i o n . Thus the following d e f i n i t i o n s : Copepod Assemblage: comprises a l l members of the order Copepoda which are q u a n t i t a t i v e l y retained by a 351 um mesh net, and are found to coexist i n the same water column, at one point i n time. An assemblage may or may not be a community. Copepod Community: an "assemblage" of copepods, whose members share e c o l o g i c a l features enabling them to p e r s i s t and coexist i n a common environment. The oblique tows which are used to c o l l e c t the data for the present research, integrate the catch over the entire water column. As a r e s u l t , several communities are l i k e l y to be c o l l e c t e d i n a single sample since d i f f e r e n t copepod communities may be c h a r a c t e r i s t i c of d i f f e r e n t depth stratums (Harrison et a l . , 1983). Nevertheless, a survey of the l i t e r a t u r e and the observed geographic r e s t r i c t i o n s of c e r t a i n species suggests that the copepods could be sorted into representative groups with varying degrees of success. B i o l o g i c a l Background As early as 1916 (McMurrich, 1916), studies pertaining to - 3 -copepods within the S t r a i t of Georgia were underway. These early investigations concentrated primarily on the taxonomy of the l o c a l species. Quantitative considerations were hampered both by poor taxonomic basis for copepod studies, and the long periods of time required to generate data. In recent years, functional taxonomic keys to the Northwestern P a c i f i c Ocean became available (Brodskii, 1950; Davis, 1955; Gardner and Szabo, 1982). In addition, a v a r i e t y of methods to increase the rate for generating data were developed; sampling stations were lim i t e d , and various automated counters were considered (Parsons and LeBrasseur, 1970; Fulton, 1972; Sheldon et a l . , 1972; Bary, 1980; Haury and Wiebe, 1982). Although several alternate methods to the microscope have been proposed, v i s u a l inspection is s t i l l the only means of assessing the s p e c i f i c composition of a sample which is e s s e n t i a l to the consideration of communities. The commercial success of a f i s h e r i e s has often been linked with a single year class (Parsons, 1975), and s u r v i v a l of early l i f e h i s t o r y stages is believed to be important to the recruitment of cohorts into these commercial stages (LeBrasseur et a l . , 1969; Hourston and Haegele, 1980). While factors a f f e c t i n g the s u r v i v a l of l a r v a l and juvenile fishes are poorly understood ( S a v i l l e and Schnack, 1981; Solemdal, 1981; Lasker and Sherman, 1981), i t is at these early stages that v i r t u a l l y every species of f i s h feed upon copepods (Sherman et a l . , 1981; Paul, 1983). For example, growth rates of juvenile f i s h have been linked to the size and density of available zooplankton (LeBrasseur et a l . , 1969). Parsons and LeBrasseur (1970) found juvenile pink salmon grew faster when offered copepods i n the range of 2 to 4 mm. Prey densities (copepods) were also shown to a f f e c t the growth of A t l a n t i c - 4 -herri n g (Horwood and Cushing, 1978), and l a r v a l anchovy (Lasker and Zweifel, 1978). The coordination between the appearance of seasonally variable prey and the recruitment of l a r v a l f i s h must therefore be of paramount importance to the s u r v i v a l of early l a r v a l f i s h stages. In such sit u a t i o n s success depends on proper timing between the appearance of l a r v a l fishes and th e i r prey items (copepods). Several authors (Steele, 1976; Greve and Parsons, 1978; Matthews and Heimdal, 1980) have noted the trend for stable water columns to maintain f i s h in the upper trophic l e v e l s ; whereas unstable water columns tend to terminate in ctenophores and other gelatinous animals. Cooney and Coyle (1982) in assessing variable copepod grazing e f f i c e n c i e s concluded that low e f f i c i e n c i e s favoured a benthic dominated sink, but e f f i c i e n t grazers favoured pelagic sinks. Physical Environment The S t r a i t of Georgia i s an inland sea separating the mainland of B r i t i s h Columbia and Vancouver Island. It's dimensions are 222 km long, and 28 km wide, with an average depth of 155 m, and a maximum depth of 420 m. Discovery Passage - Johnstone S t r a i t - Queen Charlotte S t r a i t , connect the S t r a i t of Georgia to the P a c i f i c Ocean in the north, while to the south, the S t r a i t is connected v i a various passes to Juan de Fuca S t r a i t and hence, the P a c i f i c Ocean. A prominent feature of the S t r a i t is the Fraser River estuary, one of the world's major r i v e r s (Thomson, 1981). Upon entering the S t r a i t of Georgia, the Fraser River water tends to be directed southwest towards the Gulf Islands (Thomson, 1981). Saltwater entrainment into the brackish surface layer creates a net loss - 5 -of s a l t at depth as the surface layer moves seaward v i a Juan de Fuca S t r a i t (Samuels, 1979). The loss of s a l t to the surface layer i s replaced through a deep inflow of seawater entering from Juan de Fuca S t r a i t . However, owing to the intensive t i d a l mixing over the s i l l s i n the San Juan Archipelago, the r e s u l t i n g deep water flow into the S t r a i t is a mixture of Juan de Fuca S t r a i t and Fraser River water. The s a l i n i t y of this deep inflowing water therefore, varies with changes in the Fraser River discharge. Waldichuk (1957) and Thomson (1981), divided the S t r a i t of Georgia into three p h y s i c a l l y d i s t i n c t regions (Figure 1) based p r i m a r i l y on the s t a b i l i t y of the water columns (Figure 2): North, Central, and South. The Northern S t r a i t extends from the south end of Texada and Lasqueti Islands to the channels at the northern entrance to the S t r a i t . In r e s t r i c t e d channels such as Discovery Passage, t i d a l currents may a t t a i n speeds of 50 cm*S - 1, but such speeds are the exception and for the most part the region is t y p i f i e d by weak t i d a l currents of approximately 10 cm*S~*. Few observations on the surface c i r c u l a t i o n of this region l i m i t most discussions to speculation; Thomson (1981) provides a summary of such d e l i b e r a t i o n s . Freshwater input into this region from the Fraser River or other t e r r e s t r i a l sources is l i m i t e d ; thus the formation of a d i s t i n c t layering system is i n h i b i t e d . In summer however, solar r a d i a t i o n warms the upper few meters creating a shallow but d i s t i n c t surface layer; this enhances the generation of surface currents. Because the northern channels are long and narrow, they possess only limited power to remove the S t r a i t water on the ebb t i d e , thereby r e s t r i c t i n g the exchange of Northern S t r a i t water with more oceanic water (Waldichuk, - 6 -1957). Because of this limited exchange of water, the Northern S t r a i t must r e l y on a slow northward movement of Southern S t r a i t bottom water and winter convective overturning to replenish the oxygen supply of i t s bottom water. Compared to the rest of the S t r a i t , the Northern S t r a i t could be summarized as a s l i g h t l y stagnant region of low k i n e t i c energy. The area south of the Northern region extending to a l i n e running between Point Roberts and Active Pass is termed the Central S t r a i t . Here, the Fraser River water has a pronounced influence on the v e r t i c a l structure and surface flows of the S t r a i t by creating a brackish surface layer of 1-10 m thickness. The extent of this low-density layer varies seasonally with the r i v e r ' s discharge. River discharge i s maximal 3 1 around 11,000 m 'S - which coincides with the period of rapid snow melt i n the higher elevations of the Fraser watershed. Between late June and mid-July, this surface layer may occupy much of the Central S t r a i t . In la t e winter, however, when the discharge i s minimal (1,000 m »S ), the brackish surface layer is r e s t r i c t e d to near-shore areas. In addition to creating a southwest surface flow, the waters of the Fraser River also act to make the surface layer " s l i p p e r y " (Thomson, 1981), favouring the development of wind generated currents. On the flood tide, part of the Fraser River plume appears to be re-directed due north, and, i f aided by favourable winds, may continue north even on the succeeding ebb tide (Giovando and Tabata, 1970). The heavy s i l t load of the Fraser River r e s u l t s i n the plume appearing much l i g h t e r i n colour than the adjacent seawater, and has aided investigators i n following i t s movements (Tabata, 1972). This northward current has been observed to maintain speeds in excess of 50 cm'S - 1. The current may d r i f t towards Burrard'Inlet, or on occassion head for the west side of Howe Sound and the Sechelt Peninsula - 7 -where i t may move p a r a l l e l to the shore or be deflected towards Texada and Lasqueti Islands. The frequency and duration of such currents are unknown. The tides for the region are moderate, entering the Central S t r a i t at about 50 cm*S - 1 and decreasing i n strength as they flood north. Figure 3 shows the c h a r a c t e r i s t i c two-layered system, with the upper layer occupying the top 50 m. This is a very stable system and exis t s year round. The lower layer shows l i t t l e seasonal v a r i a t i o n with temperatures remaining between 8.0 and 10.0°C and s a l i n i t i e s f a l l i n g between 29.5 and 31.0 ppt. Usually an increase of 1°C occurs from summer to winter. The upper layer experiences marked seasonal changes. Winter storms, cold a i r and convective mixing combine to lower the temperature of the upper 50 m. With the commencement of spring, increased solar r a d i a t i o n warms the top 10-20 m. Temperatures are commonly reported to reach i n excess of 15°C and in constricted bays i n excess of 20°C. By mid-May, the Fraser River discharge has maximized and brackish water extends over much of the surface of the S t r a i t recording s a l i n i t i e s of less than 15 ppt. This brackish layer further increases s t a b i l i t y of the water column. From the south of Point Roberts to the northern end of the San Juan Archipelago l i e s a region marked by strong t i d a l mixing, the Southern S t r a i t . T i d a l currents i n the extreme south of the S t r a i t are commonly i n excess of 100 cm*S - 1 and have s u f f i c i e n t turbulent energy to mix the entire water column i n the extreme Southern S t r a i t (Figures 2 and 3). Rock outcrops, boulders and cobbles, t y p i c a l of the bottom substrate of the area, is in d i c a t i v e of strong currents. T i d a l excursions for the surface at this end of the S t r a i t are in the v i c i n i t y of 18 km for flood tides and about 40 km for ebb tides . Because of the i n t e n s i t y of the - 8 -t i d a l currents in the extreme south, surface winds are not expected to have much e f f e c t . As the tides progress north, the rapid increase i n cross - s e c t i o n a l area of the S t r a i t dampens the t i d a l currents increasing the r e l a t i v e importance of wind derived currents. Like the Central S t r a i t , the Southern S t r a i t is influenced by the seasonal changes of the Fraser River, although to a lesser extent. In summer a d i s t i n c t brackish layer can be found at times i n parts of the Southern S t r a i t . Density currents o r i g i n a t i n g i n the Juan de Fuca S t r a i t , augmented by the pumping action of the tides and the estuarine seaward flow, provide continual replacement of the S t r a i t s bottom water (Sammuels, 1979). This renewal process is p e r i o d i c a l l y i n t e n s i f i e d when the density of the Juan de Fuca S t r a i t ' s upper layer greatly exceeds that of the S t r a i t of Georgia's bottom water. In the winter, cold, well oxygenated water having a s a l i n i t y s l i g h t l y less than that of the bottom water displaces the warmer bottom water, a product of the previous summer. During summer, the process of deepwater renewal is believed to be the res u l t of upwelling o ff the continental shelf. The upwelled water which is warmer and of a higher s a l i n i t y than the S t r a i t ' s winter bottom water, then penetrates into the Juan de Fuca S t r a i t and the S t r a i t of Georgia. Sampling Background A single s t a t i o n located i n the deepest part of the S t r a i t of Georgia, was considered to be representative of the S t r a i t for many investigations (Gardner, 1976; Krause and Lewis, 1979). Li m i t i n g plankton sampling to a single s i t e , treated the S t r a i t of Georgia as a homogeneous environment. It is at this point that I wish to further the - 9 -u n d e r s t a n d i n g o f t he S t r a i t o f G e o r g i a copepods t h r o u g h t h e use o f many s a m p l i n g s t a t i o n s , t h e r e b y a c k n o w l e d g i n g the d i v e r s e n a t u r e o f t he S t r a i t o f G e o r g i a . - 10 -Methods and Materials Ichthyoplankton Sampling Program The samples were co l l e c t e d during an ichthyoplankton survey in the S t r a i t of Georgia under the d i r e c t i o n of Dr. J. C. Mason of the P a c i f i c B i o l o g i c a l Station, Nanaimo, B r i t i s h Columbia. The plankton were co l l e c t e d from late winter to spring of the years 1979, 1980 and 1981. A series of oblique bongo net tows, from the surface to the bottom and back to the surface, were used. A bongo net is a dual net with no b r i d l e preceeding the mouth opening; in this i n v e s t i g a t i o n the net has a 351 um mesh. Of the three years under in v e s t i g a t i o n , 1981 was the most intensive survey and the only year considered i n the present research. Eighty stations d i s t r i b u t e d throughout the Central and Southern S t r a i t were sampled on alternate weeks, from mid-Febuary to mid-June (Figure 4, Table 1). This scheme provided nine sampling in t e r v a l s (cruises: 1, 2, 3, 5, 7, 8, 9, 10 and 11), that produced a t o t a l of 720 paired samples. An a d d i t i o n a l , two cruises (4 and 6) sampled 23 s i t e s i n the Northern S t r a i t , these cruises were not considered in the present research. In each of the paired bongo net samples, the l e f t hand side was analysed for hake and pollock eggs and larvae, as well as for euphausiids and coelenterates; the right-hand side was used for copepod analysis. Sample C o l l e c t i o n and Preservation Sample c o l l e c t i o n was r e s t r i c t e d to daylight hours to reduce v a r i a t i o n caused by d i f f e r e n t i a l day and night catches. Flow meters (General Oceanic D i g i t a l Flow Meters), centered at the mouth of each net, - 11 -helped to determine the distance the net t r a v e l l e d . Wire angles, recorded at frequent i n t e r v a l s , helped to determine the maximum depth encountered by the net and ensured proper " f i s h i n g " of the net. The occurrence of mud either i n a cup suspended 10 m below the net's weight, or mud on the weight, or mud in the net i t s e l f , also helped i n estimating the maximum depth encountered by the net. A perfect tow was one when the net fished continually at an angle of 30 degrees plus or minus 10 degrees, and upon recovery, mud would be found i n the cup and on the weight but not in the net. About 90% of the tows belonged to this l a t t e r category. Upon r e t r i e v a l of the sampling gear, the nets were thoroughly washed with sea water and the contents of the net concentrated at the cod end. One l i t r e jars were used to store the samples and a 10% by volume of buffered s o l u t i o n (Borax) of formalin provided the necessary preservation. Station Selection The number of samples were reduced through the se l e c t i o n of transects and control s i t e s (Figure 5). A north-south transect was chosen along the deepest channel of the S t r a i t . This procedure was followed to ensure that both surface and deep water dwelling species, i f present, would be encountered. The transect consisted of 17 stations. Coarse inspection of the north-south transect consisted of analysing every other s t a t i o n commencing in the north with s t a t i o n 8535 and terminating i n the south with 2837. This was followed by a " f i n e " inspection of regions showing sharp changes i n the copepod assemblages as determined from a c l u s t e r i n g process, BMDP Cluster Analysis of Variables, - 12 -which was performed on a VAX 100 computer. This s t a t i s t i c a l technique used untransformed processed data. Stations were amalgamated into s i m i l a r groups using the minimum distance method and based upon the absolute c o r r e l a t i o n c o e f f i c i e n t s between s t a t i o n s . A second shorter transect, s t a r t i n g at the mouth of the Fraser River and running west, was used to examine the influence of the plume on the assemblages. A t h i r d transect was located in the northern part of the Central S t r a i t where i t was believed that v a r i a t i o n in the assemblages could be observed with only minimal perturbations caused by the Fraser River. Four stations from the northern Central S t r a i t provided controls. Features considered in the s e l e c t i o n of these controls were s i m i l a r i t y of depths to stations found i n the Southern S t r a i t , as well as maximum distance from the influence of shores and important estuaries. Sample Processing Processing of a sample involved the separation of the large zooplankton ("predators") and copepod fractions followed by subsampling of the copepod f r a c t i o n and then obtaining the wet weight of the predator f r a c t i o n . These separations were accomplished through the following procedures: 1. A 3350 um sieve was stacked on top of a 250 um sieve. The stacked sieves were submerged i n water to within an inch of the top of the 3350 um sieve. The sample was then poured into the 3350 um sieve. This process reduced the tendency for large - 13 -animals to form clumps and thereby i n h i b i t the passage of smaller animals into the 250 um sieve. The sink plug was pulled to release the water. The sieves were rinsed u n t i l no animals were observed to pass from the 3350 um to the 250 um sieve. 2. Contents of the coarse sieve were placed i n a clear Plexiglas tray mounted on a l i g h t table. Copepods were removed and placed in the fine sieve. A l l non-animal matter was discarded. The animal content remaining i n the 3350 um sieve is defined as the predator f r a c t i o n i n this paper. Thus, size alone was used to describe the predator f r a c t i o n . However, i t is important to note that i t is t y p i c a l l y comprised of siphonophores, polychaetes, amphipods, euphausiids, chaetognaths, juvenile f i s h , etc. The proportions of these animals varied between tows and through the season. No attempt was made to quantify the constituents of this assemblage. Predator Wet Weight 1. The cleaned coarse f r a c t i o n was poured into a p l a s t i c cylinder with a 250 um screen for a bottom. The cylinder and contents was allowed to stand on a damp sponge for about 10 min to remove the i n t e r s t i t i a l water. 2. Contents of the cylinder were placed on f i l t e r paper and patted l i g h t l y with f i l t e r paper u n t i l excess moisture was removed. 3. Contents and f i l t e r paper were weighed on a Mettler P120 balance to the nearest 0.001 g, then the contents were removed and the f i l t e r paper alone was weighed. The difference was determined and rounded to the nearest 0.01 g. This weight was taken as the wet weight of the predator f r a c t i o n . - 14 -Copepod Subsampling 1. Large non-copepod items found in the 250 um sieve were v i s u a l l y removed and returned to the sample container. 2. The contents were then placed i n a modified Folsom Plankton S p l i t t e r with enough water to keep the sample f l u i d . With too l i t t l e water, incomplete mixing was a problem. If too much water was used, r e l a t i v e to the volume of the animals, then keeping the animals in suspension became a concern. 3. Subsampling continued by an alternate sides method to reduce any error due to a bias between sides of the s p l i t t e r . Only enough water was added between subsamples to wash down the sides of the s p l i t t e r . 4. Subsampling was not taken beyond 1/64 of the o r i g i n a l copepod content and was usually terminated when 500 to 1000 animals remained. 5. Contents of the f i n a l subsample were placed i n a sorting tray; the copepods were i d e n t i f i e d and counted under a Wild M5 d i s s e c t i n g microscope. The Folsom Plankton S p l i t t e r was modified to minimize s p l i t t i n g errors by d r i l l i n g four rows of small holes (0.03 mm) through the bottom of the mixing chamber (Longhurst and Seibert, 1967). The outside of the chamber was f i t t e d with an a i r - t i g h t manifold into which compressed a i r was forced. The fine jets of a i r introduced into the mixing chamber of the s p l i t t e r created a vigorous mixing environment, preventing specimens from s e t t l i n g . This adaptation was shown to maintain better s p l i t t i n g consistency among samples than a regular Folsom Plankton S p l i t t e r - 15 -(unpubl. data). Copepod I d e n t i f i c a t i o n Three references provide the primary source for copepod i d e n t i f i c a t i o n (Brodsky, 1950; Fulton, 1968; Gardner and Szabo, 1982;). I d e n t i f i c a t i o n and staging of the various copepods posed few problems. Though i t i s beyond the scope of this research to deal with the taxonomy of these animals; b r i e f mention is made of taxonomic problems common to this i n v e s t i g a t i o n . Pseudocalanus minutus ranges from 1-2 mm in length. Frost (pers. comm.), believes that this animal is not P. minutus, but is a composite of two or three species. U n t i l the various d i s t i n c t i v e features can be incorporated into a key, these animals are c l a s s i f i e d herein as P. minutus. The only other s i g n i f i c a n t taxonomic problem encountered was that of Metridia lucens vs. M. p a c i f i c a . Thorp (1980), investigating specimens from the coastal waters of B r i t i s h Columbia and using a series of detailed measures, concluded that there were three species of Metridia, a l l of si m i l a r s i z e . I endeavoured to apply these measures to d i f f e r e n t i a t e the specimens in the S t r a i t but concluded that the various c r i t e r i a used by Thorp were not applicable to the species found i n the S t r a i t . Consequently, they are referred to as M. lucens, whose description they most c l o s e l y matched. Limits, Error Estimates and Conversions Quantitative Limits Mesh s e l e c t i v i t y was examined to assess what size range of - 16 -copepods was q u a n t i t a t i v e l y retained by a 351 um mesh net. This was done in late July, 1982, using six tows at a fixed location south of Texada Island. Each tow alternated between a 351 um and a 200 um mesh SCOR net. A l l tows started at a depth of 350 m and followed an oblique path to the surface. Flow meters were centered at the mouth of the net; a piece of cord was l a i d across the mouth to prevent the meters from f l i p p i n g and counting on the net's descent. The samples were preserved and processed as previously described, but the predator f r a c t i o n was omitted. Predator: Dry Weight Determination Biomass, as determined by wet weight, is not a true comparison for the weight of predators and copepods since the methodology d i f f e r s between large and small organisms. Biomass however, when taken to a constant dry weight at a set temperature, does provide for a better comparison since the same method can be implemented for both small and large animals. Samples from 23 stations varying both temporally and s p a c i a l l y , were examined. Wet weights were determined as previously described. Once the excess moisture was removed, the following procedure was followed: 1. A l l weights were recorded on a Mettler AE160 balance to an accuracy of 0.1 mg (Damp sponges were packed into the enclosed weighing chamber at least 2 h p r i o r to taking wet weights; whereas, drying-salts were placed in the weighing chamber 24 h p r i o r to dry weight determinations). The balance was set to - 17 -give an integrated weight over a 6 sec period. This procedure reduced fluctuations i n the balances read-out. Fluctuations that occurred were within the accuracy of the balance (0.1 mg) and were infrequent. 2. The animals were removed from the f i l t e r paper, placed i n pre-weighed aluminum containers and then weighed. (The containers were previously dried for 24 h at 60°C, then allowed to cool for 12 h at room temperature i n a desiccator.) 3. The wet animals and the i r containers were placed i n a drying oven set at 60°C. The heavier samples were weighed p e r i o d i c a l l y u n t i l an approximately constant weight was reached. These samples were allowed to dry for an additional 12 h. 4. The samples were removed from the oven and placed i n the desiccator for 24 h. 5. After the 24-h period, the dried samples were weighed. Copepod: Dry Weight Determination When most i n d i v i d u a l copepods, were dried, they could not be weighed on the available balance. Therefore a group of individuals were counted and weighed, and the r e s u l t i n g weight divided by the number of in d i v i d u a l s . This procedure was time consuming and therefore analysis was r e s t r i c t e d to the 22 most common species stages. P. minutus and Calanus marshallae C6, were investigated for geographic v a r i a b i l i t y i n the i r weights by weighing specimens from both the Central and the Southern S t r a i t . Neocalanus plumchrus C5 and C6 was checked for temporal v a r i a b i l i t y by weighing specimens from three d i f f e r e n t cruises. To at t a i n an i n d i c a t i o n of the v a r i a b i l i t y expected from the above weighings - 18 -several stages were re p l i c a t e d fi v e times. Those stages which were not weighed d i r e c t l y were supplemented by weights given i n the l i t e r a t u r e (as referenced). The weighing procedure for the copepods was as follows: 1. Undamaged specimens were removed sequentially and placed i n a labelled v i a l containing 10% buffered formalin. In this way the male to female ra t i o s were preserved. 2. Samples were poured through a small open-ended p l a s t i c tube. The bottom of the tube was held firmly against a 44 ym screen, which lay atop a damp sponge. The sponge absorbed the i n t e r s t i t i a l water. 3. The tube was removed leaving the animals concentrated on the screen. Forceps were used to remove the animals from the screen to pre-weighed containers prepared i n the same manner as for the predator f a c t i o n . The animals were then weighed i n the damp sponge packed weighing chamber, giving the wet weights for the animals and the containers. 4. They were then transferred to a drying oven and dried to a constant weight at 60°C. Once an approximately constant weight was attained, they remained i n the oven for an addit i o n a l 12 h. Cooling followed at room temperature i n a desiccator. 5. After 12 h in a desiccator, the animals were weighed i n a dryin g - s a l t packed chamber and then returned to the desiccator. 6. After 1 h, 10 samples were re-weighed to check on the moisture s t a b i l i t y of the sample. - 19 -Folsom Plankton S p l i t t e r Error Subsampling was a p o t e n t i a l l y major source of error i n the present study. Four subsample f r a c t i o n s , 1/8, 1/16, 1/32 and 1/64, covered a l l the fractions used, and except for the 1/4 s p l i t , which was used only twice, a l l were investigated. To accomplish this task, a sample was chosen for each f r a c t i o n and processed (as previously described). Following the i d e n t i f i c a t i o n and enumeration of the copepods the sample was recombined, then taken to the same s p l i t again and the copepods i d e n t i f i e d and enumerated. This process was repeated fiv e times for each f r a c t i o n . Predator Wet Weight Error To determine the error i n estimating the predator f r a c t i o n , three samples varying i n their recorded predator wet weights were chosen. Once the wet weights were determined (as previously described), the sample fractions (predator and copepod) were recombined and allowed to soak in water for an hour; then again the predator wet weight was determined. This process was repeated five times for each sample. S t a t i s t i c a l Treatment of Data Raw Data The i n i t i a l data derived from a sample may be described as "raw" data. Here, the data was expressed as number of copepods for each subsample or the wet weight of the predator f r a c t i o n for each sample. The f i r s t step i n viewing the data was to convert the raw data to meaningful u n i t s . - 20 -Processed Data The number of copepods per subsample were converted to numbers per square meter by the following formula: numbers*m - 2 = (raw data) X (1/subsample fract i o n ) X (tow depth/(flow meter reading X flow meter constant X surface area of net opening) where tow depth = (wire out)cos(wire angle at maximum wire out) or = distance from bottom on echo sounder based on l o c a t i o n of mud on sampling gear flow meter constant = converts flow meter reading into distance net t r a v e l l e d i n meters. = 0.051 surface area of net opening = (pi) X (0.285) X (0.285) To convert the wet weight of predators per sample to wet weight of predators per square meter, the same formulae was used except there was no subsample f a c t o r . Clustering of Stations Numerical c l a s s i f i c a t i o n as a means of extracting trends and associations from complex data matrices are commonly reported in the l i t e r a t u r e (Williams and Stephenson, 1973; A l l e n and Skagen, 1973; Walker, 1974; Magadza, 1980; Mackas and Sefton, 1982; F i e l d et a l . 1982; - 21 -C o l l i n s and W i l l i ams, 1982j Penas and Gonzalez, 1983). When these a n a l y t i c a l tools are combined with the speed and capacity of a computer, the i n d i v i d u a l investigator is freed to view the data from many d i f f e r e n t angles which previously would have been neglected. By grouping stations into s i m i l a r c l u s t e r s , i t becomes apparent as to which st a t i o n assemblages have the most i n common and i f geographic trends exist for the associations. The pros and cons of various c l a s s i f i c a t i o n methods do play an important role i n the presentation of this work and are discussed within the l i m i t s imposed on this study. F i e l d et a l . (1982), i n a general consideration of analysing multispecies d i s t r i b u t i o n patterns, put forward a scheme which summarizes c l a s s i f i c a t i o n procedures. When applied to the present research, the procedure is as follows: Raw —> Processed —> Reduced —> Transformed — > Data Data Data Data S i m i l a r i t y —> C l a s s i f i c a t i o n Matrix The process of data reduction has been common practice i n most eco l o g i c a l studies. The purpose is to eliminate species which add l i t t l e to the ecolog i c a l understanding of the community, but add greatly to the computational time involved. As pointed out by C l i f f o r d and Stephenson (1975), the only instance where data reduction i s undesirable i s when measures of d i v e r s i t y are being considered. Any interest this study may have had in d i v e r s i t y measures is superceded by the nature of the subsampling routine, whose s e n s i t i v i t y to rarer species varies according - 22 -to the subsampling fractions that are u t i l i z e d . Most cut-off levels are a r b i t r a r y . Unlike a transformation procedure, data reduction does not a l t e r the f i n a l product provided the cut-off leve l is not too severe. In the current study, I have adopted as a basis for data s e l e c t i o n to eliminate any stage whose population does not exceed one percent of the station's population, and appears i n less than h a l f the stations for a given cruise ( C l i f f o r d and Stephenson, 1975; F i e l d et a l . , 1982; Mackas and Anderson, i n press). Transformation of data is intended to reduce the skewness placed on a data set caused by a few very abundant species i n asso c i a t i o n with several less abundant ones. In some investigations the transformation process was eliminated altogether (Stephenson and Williams, 1971), others have used various degrees of root transformations (Stephenson and Burgess, 1980), but the most common is the log transformation ( F i e l d et a l . , 1982; Legendre and Legendre, 1983). In attempting to standardize the l a t t e r process, many workers selected the transformation which would most c l o s e l y normalize their data-set. As pointed out by C l i f f o r d and Stephenson (1975), transformations which normalize data do not necessarily y i e l d the best e c o l o g i c a l explanation. Some workers believed that transformations weaker than those required to normalize the data make the best e c o l o g i c a l sense (Williams and Stephenson, 1973). After c l o s e l y s c r u t i n i z i n g the present data, a need for data transformation was evident. Without such transformations one or two species determine the entire c l u s t e r i n g process. Although community domination is an i n t e g r a l consideration i n this study, the concept does not require a complex process such as cl u s t e r analysis to hi g h l i g h t the - 23 -dominant organisms. Since the data display a wide range of counts'm (10's to 100,000's), a strong transformation is performed: the log(X + 1) transformation (Penas and Gonzalez, 1983). The l i t e r a t u r e contains numerous measures of s i m i l a r i t y , distance and association (Whittaker, 1952; Whittaker and Fairbanks, 1958; Hummon, 1974; Stromgren, 1975; Chester, 1978; Magadza, 1980; Bloom, 1981; Kohn and Riggs, 1982; C o l l i n s and Williams, 1982). In these p a r t i c u l a r instances, the objective of such measures is to summarize the difference or s i m i l a r i t y between two stations, by taking a l l species into account, in a single measure. The "BMDP Cluster Analysis of Variables", the cl u s t e r i n g program available to the author, provides a choice of four such measures: two are based on the product-moment c o r r e l a t i o n c o e f f i c i e n t , and two are based on the angle between two va r i a b l e s . The l a t t e r two measures are se n s i t i v e to the proportion of species i n a sample and to the absolute value for a given species; whereas the former two only take into account the proportion of species. As the analysis moves towards the periphery of a community, the absolute value for the various species is l i k e l y to change, but i f the interactions among the species remain intact then the p r o p o r t i o n a l i t y of the various species should hold. To s a t i s f y this statement, the absolute product-moment c o r r e l a t i o n c o e f f i c i e n t is chosen. The amalgamation of stations into clusters is the f i n a l step i n the process. Three choices are available to the user: minimum distance, maximum distance, or arithmetic average. A l l strategies force the stations into clusters even i f the environment is continuous rather than d i s c r e t e . Minimum distance (or single linkage) is the basic format for - 24 -c l u s t e r i n g . In this choice, a s t a t i o n i s added to an already e x i s t i n g c l u s t e r based on i t s s i m i l a r i t y to i t s nearest neighbour within the e x i s t i n g c l u s t e r . If enough intermediate stations e x i s t , a chain is formed that may cause two very d i f f e r e n t stations being placed i n the same c l u s t e r . Thus, minimum distance tends to produce a representative picture when the clusters are r e l a t i v e l y small, but in a continous environment the larger picture may be d i s t o r t e d . Maximum distance (complete linkage) takes the opposite approach to minimum distance. For a s t a t i o n to j o i n a c l u s t e r i t must l i n k up with the most distant member of the ex i s t i n g c l u s t e r . Therefore, as a cl u s t e r grows, i t is increasingly d i f f i c u l t to add stations to the c l u s t e r . Intermediate to the above two methods i s that of arithmetic averaging. Clusters are amalgamated based on the average s i m i l a r i t y among a l l possible pairings in the two clusters under consideration. This method reduces errors inherent i n the above two methods, pr i m a r i l y that of chain forming, and that of increasing d i f f i c u l t y i n clu s t e r formation with increasing c l u s t e r s i z e . The main drawback to this l a t t e r approach is the loss of an i n d i v i d u a l station's i d e n t i t y once i t has fused with a c l u s t e r . Because this method offers the most representative e c o l o g i c a l pattern (Legendre and Legendre, 1983), the arithmetic average linkage is chosen to assess the data for this research. Clustering of Species As with sta t i o n s , species may be clustered into s i m i l a r groups. The methodology is sim i l a r to that used for cl u s t e r i n g stations, but with the following a l t e r a t i o n s : - 25 -1. Because of the large number of species s t i l l remaining after data reduction, data standardization is necessary to allow r o t a t i o n of the processed data matrix on the computer f i l e . This process is accomplished by expressing each species at a given s t a t i o n as a percent of the t o t a l number for each species over a l l the stations of a given c r u i s e . 2. Owing to the standardization process, a less stringent root transformation is used ( F i e l d et a l . , 1982). - 26 -Results and Discussion Limits, Error Estimates and Conversions Quantitative Limits Table 2 summarizes the t - t e s t for the more numerous copepod members of the 351 um vs. 200 um mesh net comparison. The underlying assumption in this analysis is that a 200 um net catches everything that a 351 um net does plus a measureable portion of smaller sized plankton. Therefore any copepod species caught in s i g n i f i c a n t l y greater numbers i n the 200 um net as opposed to the 351 um net is considered non-quantitative for a 351 um mesh net. _N. plumchrus C5, M. lucens C3, and 0. s p i n i r o s t r i s (>1 mm) showed s i g n i f i c a n t l y greater numbers in the 200 um net. N. plumchrus C5, however, was not expected to show a s i g n i f i c a n t difference as i t is the largest and by far the most abundant copepod member of the samples in question. As subsampling error is not considered in the present analysis and the mean number of N. plumchrus C5's is not too far apart for the two meshes (within 25%), the difference is ignored and the N. plumchrus C5 is considered a part of the 351 Um mesh catch. The means for 0, s p i n i r o s t r i s show a spread of almost three f o l d . However, since 0. s p i n i r o s t r i s is an important component of the S t r a i t of Georgia copepod assemblage, I have decided to maintain 0. s p i n i r o s t r i s with reservation as part of the 351 um net catch. A valuable aspect of the t - t e s t analysis is the difference between M. lucens C4 and M. lucens C3. The means for the C4 copepodites were non-significant having means within 10% of each other; the C3 copepodites however, were s i g n i f i c a n t l y d i f f e r e n t having a f i v e - f o l d - 27 -differ e n c e i n means. The marked difference i n s i g n i f i c a n c e between successive developmental stages enables extrapolation for a cut-off s i z e for the developmental stages of other species of s i m i l a r shape that were not encountered in the July 1982 tows. Fulton (1968), l i s t s the various sizes for d i f f e r e n t copepodite stages of M. lucens. The C4 has a given length of 1.28 mm; the C3 stage has a length of 1.08 mm. For a copepodite stage to be considered q u a n t i t a t i v e l y sampled by the 351 um mesh, the minimum acceptable length cannot be less than 1.28 mm. Other factors such as body width, shape, r i g i d i t y , and p o s i t i o n of appendages may also a f f e c t the retentive q u a l i t i e s of a plankton net. However, body length is the simplest and most r e a d i l y applicable measure to deal with. Predator: Dry Weight Determination The c o r r e l a t i o n between predator wet and dry weight was 0.988. The l i n e a r r e l a t i o n s h i p , the expression for which is given below, is shown i n Figure 6. Dry Weight = 0.1265(Wet Weight) - 0.0209 Sample preservation is not accounted for in the above equation. Omori (1978), i n v e s t i g a t i n g the influence of Borax-buffered formalin on copepod dry weights, found weight losses in excess of 20% a f t e r one week; thereafter, the animal's weight appeared to s t a b i l i z e for the remaining 180 day recording period. Omori found less than a 6% decrease in dry weight when preserved specimens of Calanus sinicus were rinsed with d i s t i l l e d water as opposed to saltwater. He further notes - 28 -that the l i p i d f r a c t i o n of the animal is susceptible to leaching. Therefore, one would conclude that animals such as Neocalanus  plumchrus C5, whose l i p i d content may account for as much as 55% of i t s dry weight (Vidal and Whitledge, 1982), would undergo a greater r e l a t i v e weight loss than many other planktonic organisms. Consequently, weight loss due to method and duration of preservation is l i k e l y to be highly species and stage s p e c i f i c , no single conversion factor is available which could improve the quality of the dry weights attained. Copepod: Dry Weight Determination Weights for the copepod stages which remained after the data reduction procedure are compiled in Table 3. Weights were obtained from the l i t e r a t u r e when present sample numbers were too small to obtain weights d i r e c t l y (Table 3). Weights for N. plumchrus C6 and C5 as well as C. marshallae C6 and P. minutus C6, showed seasonal v a r i a b i l i t y . While other animals l i k e l y undergo seasonal a l t e r a t i o n s to t h e i r dry weight, the present data is not s u f f i c i e n t for a comprehensive consideration of every species weighed. As in the predator f r a c t i o n , no adjustments were made for weight losses r e s u l t i n g from preservation. Folsom Plankton S p l i t t e r Error S t a t i s t i c a l l y , there is l i t t l e difference between a 1/8, 1/16 and 1/32 s p l i t . A l l show a po s i t i v e c o r r e l a t i o n between numbers counted and increasing accuracy (Figures 7a,b,c). The lower l i m i t of the curves appears to be a count of 5 with a 95% confidence l i m i t of about 50% of the mean. Below this l i m i t one is dealing more with "present/absent" data. Decreasing accuracy with the smaller f r a c t i o n s p l i t s is expected as - 29 -each s p l i t should multiply the error by some factor. This expected trend may have been obscured through the use of d i f f e r e n t samples to test each f r a c t i o n , i f the composition of a sample affects the error produced while subsampling. The 1/64 s p l i t also shows a lower l i m i t of about f i v e counts, but the 95% confidence l i m i t is closer to 40% (Figure 6d). A l l four curves approach a 95% confidence l i m i t of 90% at the higher counts. The f i r s t three s p l i t s , 1/8, 1/16 and 1/32, reach this mark at lower counts than did the 1/64 s p l i t . In a l l samples a count of 5 represented less than 1% of the copepod population that was used as part of the cut-off c r i t e r i a to construct the reduced data matrix. The c l u s t e r i n g process however, could not u t i l i z e incomplete species records; therefore some binary data - less than 5 counts used to determine the number of — 2 individuals*m - was included i n the reduced data matrix. This s i t u a t i o n could not be detected u n t i l after a cruise was completely analysed (the point when data reduction occurs). Predator Wet Weight Error Table 4 shows confidence i n t e r v a l s for samples having three d i f f e r e n t weights. From these r e s u l t s i t is apparent that the reported sample wet weights are within 10% of the i r true mean. Clustering of Stations and Species: An I n i t i a l Perspective Abbreviations used for species and stages i d e n t i f i e d from the nine cruises are shown i n Table 5. The la s t two characters of the five-character abbreviation is reserved for a species developmental stage, and is summarized as follows: - 30 -Cl to C6: copepodite stage 1 to 6 Ju: copepodite stages 1 to 5 amalgamated Synops is One of the most r e a d i l y defined communities is that group of copepods whose habitat is determined by depth. An example of such is described by Fulton (1973) for Neocalanus plumchrus in the S t r a i t of Georgia. The animal was found to migrate, s t a r t i n g i n mid-July, from the surface waters to depths i n excess of 300 m, never to return to the surface. Moulting from stage 5 to 6, and the sequential release of eggs and death takes place at depth. Given such a cycle, what is the geographical d i s t r i b u t i o n pattern for this animal? The f i r s t cruise occurred after the majority of N. plumchrus moulted to C6 (Figure 8). The adults show a d e f i n i t e r e s t r i c t i o n to the Central S t r a i t ; a sharp decline i s noted in the numbers occurring between Station 5035 in the north and 4636 to the south (Figure 9d). An additional decline is observed when proceeding from s t a t i o n 5132 to s t a t i o n 4929 (Figure lOd). Other stations displaying reduced populations are the control stations, 7627 and 6727, both of which are in r e l a t i v e l y shallow water (164 m and 79 m, r e s p e c t i v e l y ) . Three Central S t r a i t stations, 5538, 7136 and 7227, having depths in the range of those stations south of 5035, show substantial numbers of _N. plumchrus C6. From these observations depth appears to be a c r i t i c a l factor i n governing adult d i s t r i b u t i o n within the Central S t r a i t . Other parameters must be taken into account however, when the S t r a i t as a whole is being considered. L i f e h i s t o r i e s of other copepods in the S t r a i t of Georgia have - 31 -received scant attention to date. The grouping of animals which overwinter at depth and are r e s t r i c t e d to the more Central S t r a i t water, establishes the f i r s t copepod community of this survey. The data were viewed from three d i f f e r e n t manipulations: c l u s t e r i n g of stations (Figures 11a to i ) , c l u s t e r i n g of species (Figures 12a to i ) , and the e q u i t a b i l i t y of stations (Figures 13 and 14). The l a t t e r measure is expressed as a percent and considers the r e l a t i v e d i s t r i b u t i o n of the 25 most common species at a s t a t i o n ; 100 indicates that the copepods are equally d i s t r i b u t e d among the species stages, while 0 suggests they a l l exist as one species stage. In Figure 11(a to i ) the stations have been clustered into s i m i l a r groups. B a s i c a l l y a two c l u s t e r system exists separating northern and southern stations during the course of the nine cruises. The nothern extent of the Southern c l u s t e r varies from s t a t i o n 3439 to s t a t i o n 5035, a distance of approximately 35 km. Commonly, the extreme east or west plume stations (4929 and 5538 r e s p e c t i v e l y ) , as well, the control stations (7627, 7227, 6727, and 7136) may be added to the southern group. On four occassions (cruises 3, 7, 9 and 10) the Central S t r a i t c l u s t e r could be subdivided into two groups. Figure 12(a to i ) has clustered the species into s i m i l a r groups and aids in describing the members of the deepwater community: N. plumchrus C5 and C6, Eucalanus bungii C5, Euchaeta elongata C2 to C6, Gaidius minutus C6, C h i r i d i u s g r a c i l i s C6, Calanus p a c i f i c u s C5, Metridia lucens C6 female and male, Metridia okhotonsis C6, and Calanus marshallae C5. Eucalanus bungii C5 shows the greatest geographic r e s t r i c t i o n of the deepwater copepods; only at the two deepest stations (during the - 32 -f i r s t cruise) does i t ' s population exceed 2,000'm-2, with a l l other stations showing less than 800*m - 2 (Figure 31e). Metridia lucens female C6's d i s t r i b u t i o n c l o s e l y p a r a l l e l s that of N. plumchrus C5 for the f i r s t cruise (Figure 28e). However, during the second and t h i r d cruises the animal starts to broaden i t ' s d i s t r i b u t i o n into the Southern S t r a i t and from the f i f t h cruise on, i t no longer shows a simple north-south pattern. The juveniles of M. lucens appear to be the only s t r i c t l y surface dwellers during the f i r s t c r u i s e . By the second cruise several of the dominant deepwater species, namely N. plumchrus C6, C. p a c i f i c u s C5, M. lucens female C6, C. marshallae C5, and E. bungii C5, are d e c l i n i n g allowing for a s l i g h t increase in the e q u i t a b i l i t i e s from the mid-seventies of the previous cruise to low e i g h t i e s , for this region. By the f i f t h cruise, most of the deepwater forms are non-existent. The southern c l u s t e r i s dominated by one species, P. minutus (Figure 16). As a r e s u l t the e q u i t a b i l i t y indices for this region r a r e l y exceeds 50 and is found as low as 15. Closely associated with P. minutus is A c a r t i a longiremis (Figure 17), r ranging from 0.83 to 0.94. There are three exceptions, cruise 1 (r = 0.61), cruise 5 (r = 0.49), and cruise 10 (r = 0.69). The lower values r e f l e c t A. longiremis's greater avoidance of the Central S t r a i t . From the t h i r d cruise on C. marshallae C6 joins this group with co r r e l a t i o n s for the group ranging from 0.60 to 0.85. Oithona s p i n i r o s t r i s C6 and Scolecithrice11a minor C6 are c l o s e l y correlated (r = 0.90) during the f i r s t c r u i s e. They show reduced numbers at the shallow control s t a t i o n 6727 and in the Southern S t r a i t , i n d i c a t i n g t h e i r members occur at depth (Figures 19 and 21). - 33 -0. s p i n i r o s t r i s and S. minor, unlike the other deepwater species, show thei r peaks i n abundance to be centered further to the south. However, as the season progresses, 0. s p i n i r o s t r i s ' s numbers in the Southern S t r a i t , s t e a d i l y increase while S. minor's do not, causing the c o r r e l a t i o n between the two to drop to 0.42 in the f i n a l c r u i s e . The appearance of juvenile N. plumchrus i n the t h i r d c r u i s e , mid-March, marks the beginning of spring growth. By early A p r i l (cruise 5) these juveniles show a d i s t r i b t u i o n c h a r a c t e r i s t i c of the o f f s p r i n g of the deepwater community, with their peak abundances occuring i n the northern Central S t r a i t , the Southern S t r a i t and at varying points along the plume transect. Juveniles of other copepods start to appear following the appearance of N. plumchrus j u v e n i l e s . The juveniles of M. lucens for example which had been d e c l i n i n g i n numbers show an increase between the t h i r d and f i f t h c r u i s e . Between the nineth and tenth cruise (mid-May to the end of May) N. plumchrus experiences a reduction in numbers, the f i r s t since i t ' s juveniles were recorded. Accompanying th i s decline, the peak abundance observed in the northern Central S t r a i t has sh i f t e d towards the deeper s t a t i o n s . The decline continues during the f i n a l cruise, with the migration continuing towards the deeper Central S t r a i t , along with the disappearance of the Southern S t r a i t maxima. Thus, i t would appear that N. plumchrus i s completing i t ' s growth phase and returning to depth; most other species have not reached this maturity. Several species of which l i t t l e mention has been made include G. minutus, C. g r a c i l i s and E. elongata. Primarily because of their constantly low populations, any d e f i n i t e statements about t h e i r d i s t r i b u t i o n s is d i f f i c u l t to advance. There is however, a strong - 34 -tendancy for these species to avoid the Southern S t r a i t (Figures 35a and b, 37a and b, 39a to e). Nearly a l l the species encountered for the research period (mid-February to mid-June) were found during the f i r s t c r u i s e . Thus, changes to the copepod assemblages are r e f l e c t e d primarily in th e i r progression through developmental stages. Details Cruise 1 (February 18 to 24) Two b a s i c a l l y d i f f e r e n t c lusters are described i n the f i r s t cruise (Figure 11a). The f i r s t comprises the deep central stations from 5035 north to 8535; the second contains the southern stations south of 5035 to 2837 including the four controls and the extreme east and west plume sta t i o n s . The shallowest s t a t i o n i n the f i r s t c l u s t e r is 232 m; whereas the deepest s t a t i o n for the second c l u s t e r is 223 m. Obviously, the bathymetry of the S t r a i t is important in determining the wintering habitats of many of the copepods. The deep Central S t r a i t stations from 4636 north to 8535, have e q u i t a b i l i t i e s between 71 and 77 i n d i c a t i n g that there are several numerically common species. The e q u i t a b i l i t y decreases from s t a t i o n 4337 to the southern end of the transect. In the extreme south (2837) the e q u i t a b i l i t y index has decreased to 17. C. marshallae C6 forms a loose association with 0^. s p i n i r o s t r i s and S. minor. Although i t occurred in reduced numbers at 6727 and attained maximum numbers over the same geographic range, i t s population in the Southern S t r a i t remains r e l a t i v e l y strong (Figure 24b). - 35 -Cruise 2 (March 2 to 10) The same c l u s t e r i n g of stations is preserved in the second cruise as in the f i r s t with one exception, the extreme west plume sta t i o n , 5538, is added to the Central S t r a i t c l u s t e r . Generally, e q u i t a b i l i t y has increased in the deep Central S t r a i t , with the range being from 76 to 83. The observed decrease in deep water wintering stages of the more common species would account for this increase. Again, e q u i t a b i l i t y decreases r a p i d l y in the Southern S t r a i t to 15 at s t a t i o n 2837. 0. s p i n i r o s t r i s shows increased patchiness, and does not reveal the high c o r r e l a t i o n previously held with S. minor. Cruise 3 (March 16 to 21) The stations no longer c l u s t e r into a simple two-grouping system as before. The Southern S t r a i t , although maintaining previous a t t r i b u t e s , is reduced to the stations south of 4039. Coincidental with P. minutus and A. longiremis, is a pronounced r i s e i n the number of C. marshallae C6, y i e l d i n g a high c o r r e l a t i o n among a l l three' species (0.83). This s i t u a t i o n also increases the e q u i t a b i l i t y i n the Southern S t r a i t to 47 ( s t a t i o n 2837). 0. s p i n i r o s t r i s and S. minor as before are only loosely correlated (0.60). Both species are reduced i n the Southern S t r a i t ; however, 0. s p i n i r o s t r i s increases towards the north. E. bungii C6 and C. p a c i f i c u s C6 appear to have increased at the expense of the C5 stage suggesting that the majority of these animals recently may have undergone a moult to the adult stage (Figure 30 and 32 r e s p e c t i v e l y ) . - 36 -Cruise 5 (March 30 to April 6) In early April, the stations return to the distinct two cluster pattern as was found in the first two cruises. The Southern Strait begins at 4636 and continues south, including both plume stations. Again C^. marshallae C6 and _P. minutus are closely correlated (0.89). Linked to these two species, is the youngest stage of C. marshallae investigated, namely the C3 (0.86). The C3's for N. plumchrus now form distinctive numerical peaks in the northern part of the Central Strait, in the Southern Strait, and in the eastern most plume station (Figure 9a and 10a). N. plumchrus C4 appears, especially in the Southern Strait and 2 9 eastern plume where catches of 12,731'm and 6,841*m were respectively recorded (Figure 9b and 10b). At station 8535, in the extreme north, the high abundance of C3 causes the equitability index at this location to decrease from the high of 82 on the previous cruise to 69. In the south, where the equitability index has been low due to single species domination (P_. minutus), N. plumchrus juveniles (C3 and C4) have increased this index to 51. 0. spinirostris and S. minor were closely correlated, 0.89, as seen in the first cruise. Their close association appears to be the result of increased numbers of 0. spinirostris in the Central Strait which smooths out the northward gradient of earlier cruises. Cruise 7 (April 14 to 20) Three distinct clusters are distinguishable. The first consists of a rather loose association of four northern Central Strait stations: 7829, 7832, 8535 and 6832. This cluster provides an example of one of the problems involved in clustering. Station 7634 shows a much - 37 -higher c o r r e l a t i o n with the f i r s t three members of the cluster than does 6832. Yet, 7634 amalgamates with greater ease into the second c l u s t e r than does 6832, and therefore is placed in the second rather than the f i r s t c l u s t e r . Problems such as this cause d i f f i c u l t i e s i n e s t a b l i s h i n g species patterns which have the greatest influence on c l u s t e r formation. The second cluster consists of the remaining Central S t r a i t stations from 5035 north together with the two extreme east and west plume sta t i o n s . The t h i r d c l u s t e r is composed of the more central plume s t a t i o n , 5132, and the Southern S t r a i t stations from 4636 south. Again, d i f f i c u l t y arises in making sense of the amalgamating process as s t a t i o n 4636 and 5035 are placed in separate c l u s t e r s , yet are c l o s e l y correlated (0.83). Apparently, because 5035 has a higher c o r r e l a t i o n with the extreme plume stations than does 4636, the two stations are separated. E q u i t a b i l i t y for the seventh cruise shows the same pattern as the f i f t h cruise. In the previous cruise high numbers of N. plumchrus C3's supressed e q u i t a b i l i t y i n the northern Central S t r a i t and countered the high _P. minutus numbers in the Southern S t r a i t . During this seventh cruise though, the dominant stage of.N. plumchrus is C4 with the C5 1s r a p i d l y approaching that of the C3's. P. minutus reaches a new peak population in the extreme south i n excess of 100,000 m_2, and not u n t i l s t a t i o n 5931 to the north do i t s numbers appear to l e v e l o f f . As before, both A. longiremis and C. marshallae C6 are c l o s e l y correlated with P. minutus (0.85). The majority of C. p a c i f i c u s have now moulted to the adult stage, making C. p a c i f i c u s the last of the deep water Central S t r a i t winterers to do so. E. bungii is now represented in the Cl and C2 stage. With the exception of a spike at s t a t i o n 5035, these two early - 38 -stages appear to increase numerically towards the south (Figure 31a and b). Cruise 8 ( A p r i l 27 to May 3) Two main clusters are displayed in the eighth c r u i s e . The Central S t r a i t c l u s t e r consists of a poorly correlated (0.59) grouping of stations from 5931 north to 8535, plus the extreme west plume s t a t i o n , 5538. The second c l u s t e r comprises those stations from 5035 south, the two eastern plume stations and the control s t a t i o n , 6727. The recruitment of juvenile N. plumchrus continues to supress the e q u i t a b i l i t y of the Central S t r a i t c l u s t e r ; this measure remains r e l a t i v e l y unchanged in the Southern S t r a i t from the previous cruise. N. plumchrus now predominates in the C5 stage. Although both S. minor and 0. s p i n i r o s t r i s occur at reduced numbers in the Southern S t r a i t , the poor synchrony between the l o c a t i o n of their peak numbers leaves the two copepods poorly correlated (0.55). The number of adult male and female M. lucens are conspicuously reduced i n the Southern S t r a i t ; this s i t u a t i o n i s the opposite to that of the juvenile stages (Figure 27 a to e). The juveniles form loose associations with each other and with C. marshallae C6, P. minutus and A. longiremis (0.72), a l l of which show the i r greatest numbers in the Southern S t r a i t . Cruise 9 (May 11 to 15) The northern Central S t r a i t stations, 8535 and 7634, form an isolated c l u s t e r (r = 0.83). Owing mainly to th e i r high numbers of N. plumchrus C5 (Figure 9c), these two stations are set apart from other - 39 -Central S t r a i t stations. This is also r e f l e c t e d i n the i r r e l a t i v e l y low e q u i t a b i l i t y (53 and 52 r e s p e c t i v e l y ) . A C5 spike of 21,144 m~ , at s t a t i o n 5931 resulted i n a drop i n e q u i t a b i l i t y when this s tation is compared with other l o c a l s t a t i o n s . During this cruise, the Southern S t r a i t c l u s t e r which comprises those stations from 4337 south and the eastern plume s t a t i o n 4929 reach th e i r maximum e q u i t a b i l i t y with only one s t a t i o n f a l l i n g below 50. The remaining stations f a l l into a t h i r d c l u s t e r , whose e q u i t a b i l i t i e s are higher than the previous two cl u s t e r s (60 to79). Cruise 10 (May 25 to 29) Three clusters are described i n this tenth cruise (Figure l l h ) . The Southern S t r a i t group consists of those stations from 4039 south and station 4929 of the eastern plume. A second c l u s t e r is formed from three northern Central S t r a i t s t a t i o n s , 7832, 7934 and 7634, and the western most plume st a t i o n , 5538. The t h i r d c l u s t e r consists of the remaining Central S t r a i t stations from 5035 north and the central most plume st a t i o n 5132. When untransformed data is used, stations 7934 and 6832 show a c o r r e l a t i o n of 0.995. Using a log(X + 1) transformation results in a 0.456 c o r r e l a t i o n . The primary reason for this discrepancy is found at the lower end of the scale where a t o t a l of seven zero values between the two stations are paired with non-zero counterparts. The numbers of N. plumchrus have dropped for the f i r s t time since the C3's were recorded. Accompanying this decline is a short movement of the numerical peak of N. plumchrus from the northern to the deeper Central S t r a i t . This movement is not evident i n the Southern - 40 -S t r a i t , and although the numbers have diminished, the d i s t r i b u t i o n pattern in the Southern S t r a i t is s i m i l a r to that described above. 0^. s p i n i r o s t r i s ' s Southern S t r a i t population has numerically increased to the point where a decrease to the south is d i f f i c u l t to discern. Cruise 11 (June 8 to 14) Several important changes occur in the f i n a l c r u i se. When viewing the north-to-south e q u i t a b i l i t y indices (Figure 13), the sporadic pattern displayed i n the previous two cruises has dissipated, and is replaced by one c l o s e l y resembling the f i r s t c r u i s e . The s t a t i o n c l u s t e r i n g shows two main clusters (Figure H i ) , a southern one from st a t i o n 3739 south, with the remaining stations f a l l i n g into the more loosely knit central c l u s t e r . Station 8535 appears to be an "odd b a l l " ; however, after viewing the data, this oddity appears to be another transformation a r t i f a c t r e s u l t i n g from the mismatch of rarer species. Except for a few trace i n d i v i d u a l s , N. plumchrus exists s o l e l y in the C5 stage. For the f i r s t time since the appearance of juveniles no Southern S t r a i t maxima is displayed. 2 P. minutus reaches i t s peak population at just over 140,000 m-for the season in the extreme Southern S t r a i t . Community Dynamics The physical environment is a s e l e c t i v e force acting on animals which attempt to colonize or maintain themselves within the environment. For example, Pielou (1975) notes the p o s i t i v e r e l a t i o n s h i p between environmental s t a b i l i t y or p r e d i c t a b i l i t y , leading to community s t a b i l i t y - 41 -and a higher d i v e r s i t y . Based on MacArthur and Wilson's (1967) d e s c r i p t i o n of r - and K-selected species, Pielou describes environmental settings which select for r - or K-type organisms. Margalef (1958), also looks at environmental settings and communities, noting that as a community "matures" the d i v e r s i t y of species increases u n t i l competition precludes the less adapted ones. D i v e r s i t y may also be altered through the transport of organisms from neighbouring waters (Margalef, 1958). I believe the views of Pielou (1975) and of Margalef (1958), compliment each other, with the mature community favouring the more K-selected animals, a s i t u a t i o n which r e f l e c t s a stable environment. Both MacArthur and Wilson, as well as Pielou, were considering changes on a much larger scale than the S t r a i t of Georgia, but I believe the basic ideas can be applied to the present s e t t i n g . The c h a r a c t e r i s t i c s of MacArthur and Wilson's r - and K-selected organisms and environments are summarized by Pianka (1970). Pianka compares phyla which are quite di s t a n t , vertebrates versus invertebrates. With modifications to Pianka's c r i t e r i a , the r - and K-selected c h a r a c t e r i s t i c s for copepods and th e i r environments would be as follows: - 42 -r-Selected K-Selected Comments Environment primary production water column - depth - T-S p r o f i l e - freshwater input mixing energies - t i d a l currents - wind Copepod body size developmental time generations/year diapause feeding strategy reproduct ion Community changes i n pop. e q u i t a b i l i t y competition mortality limited unstable - shallow - homogeneous - s t r a t i f i e d - l i t t l e - much excess stable - deep (> 250 m) seasonal strong - strong - strong weak - weak - weak not l i k e l y a factor i n the Str. of Georgia small (<2 mm) large (>4 mm) short (few mn) long (1 yr) several no herbivorous i t e r o p a r i t y rapid low (<50) intense density -independent 1 or 2 yes omnivorous semelparity d i f f i c u l t to assess in many cases seasonal high (>70) lax not assessable density - in the present dependent study - 43 -The above summary shows the extreme c h a r a c t e r i s t i c s with the r - s t r a t e g i s t being the opportunist or ex p l o i t e r and the K-strategist revealing a conservative approach to l i f e designed for a predictable environment. A more r e a l i s t i c picture might be a continuum of adaptations between these two extremes and possibly within a species some of both approaches during d i f f e r e n t parts of the animals l i f e h i s t o r y . Several of the c r i t e r i a l i s t e d are in opposition to Pianka's (1970) summary; however, the reasoning remains consistent. An r-selected copepod must maintain intimate contact with the exploitable environment in order to y i e l d the quickest possible response to environmental changes. An important response would be changes in fecundity, a c h a r a c t e r i s t i c p a r t i a l l y affected by a copepod's food consumption (Heinrich, 1962; Corkett and McLaren, 1978; Landry, 1983). The r - s e l e c t i v e environment should be less predictable than the K-selective environment and at times be food l i m i t e d . The fecundity of the K-selected copepod is determined from the previous seasons growth suggesting a greater time lag in the response of the copepod to environmental changes. Thus i t e r o p a r i t y (repeated reproduction) and not semelparity (single reproduction) should be c h a r a c t e r i s t i c of the r - s t r a t e g i s t . And since I expect the r - s e l e c t i v e environment to be food limited at times, competition should be intense. The remainder of this report w i l l center on assessing the v a l i d i t y of the above statements. This involves a consideration of ind i v i d u a l l i f e h i s t o r i e s of the more common copepods, d i s p e r s a l mechanisms which counter the formation of d i s t i n c t communities, and as well, the dynamics of the communities themselves. - 44 -Dispersal Mechanisms The term "planktonic" is suggestive of organisms completely at the mercy of oceanic currents. No doubt, water movement transports copepods; water quality affects s u r v i v a l ; but the most s a l i e n t factor must be the behavioral and functional biology of the species, y i e l d i n g communities rather than assemblages. The v e r t i c a l d i s t r i b u t i o n of a copepod can be altered through ontogenetic and d i e l v e r t i c a l migrations ( M i l l e r , 1970; Krause and Lewis, 1979; Harrison et a l . , 1983). Horizontal d i s t r i b u t i o n on the other hand, may be affected by turbulence (Omori and Hamner, 1982), although this i s d i f f i c u l t to document. Over the s i l l of an i n l e t , Farmer and Smith (1980) observed a sc a t t e r i n g layer (107 kHz) which they assumed to be plankton, to disappear when strong t i d a l currents produced turbulent waters. Neuston tows c o l l e c t e d on successive nights i n the S t r a i t of Georgia (January, 1983) were found almost void of planktonic animals during rough seas when compared with r e l a t i v e l y calm nights (unpublished observation). Mechanisms other than simple v e r t i c a l movement to avoid an undesirable parameter may be operating. Fulton (1973) observed an increase i n numbers of N. plumchrus C5 at a deep Central S t r a i t s t a t i o n from June through September. This suggests a l a t e r a l movement of animals to this l o c a t i o n . It may or may not be that the animals are r e l y i n g upon random l a t e r a l motion to find t h e i r winter habitat. While the animal is suspended i n the water column, simple v e r t i c a l movement w i l l lead i t to deeper water. Nearer the bottom, however, i t must consider l a t e r a l movement; gradual slopes and anomylous dips and ri s e s i n the seafloor can complicate matters. N. plumchrus may prove to be a valuable test - 45 -organism, when investigating planktonic d i s p e r s a l , to e s t a b l i s h the roles played by the behaviour of the animal and that of the physical oceanography. Estuarine Dispersal The early l i f e h i s t o r y of the copepods may be most important for planktonic d i s p e r s a l . N. plumchrus, has an annual l i f e cycle making i t p a r t i c u l a r l y well suited to follow the animal's seasonal d i s t r i b u t i o n . A model combining estuarine and t i d a l c i r c u l a t i o n can account, in general terms, for the movement and formation of population peaks of the developmental stages into the Southern S t r a i t where the overwintering adults are scarce. A d e s c r i p t i v e model is given (Fi~ure 40), followed by the evidence, pro and con. The model begins with geographically and temporally constricted adults located at depths i n excess of 300 m. Eggs are released from early January u n t i l mid-April (Fulton, 1973). As the developing n a u p l i i r i s e to the surface, at about 26 m a day (Fulton, 1973), they are exposed to r e s i d u a l currents of increasing strength (LeBlond, 1983). Although residual flows are poorly understood i n the Central S t r a i t , there seems to be a d e f i n i t e net progress towards the B r i t i s h Columbia mainland (Yao et a l . , 1982). As the n a u p l i i r i s e to the surface they are carried by these currents, where they become entrained into the Fraser River plume. From the plume, the majority are moved southwest in surface brackish waters towards Haro S t r a i t . Some, by this same brackish layer, are relocated northward when favourable conditions reverse the normally southwest flow of the plume (Giovando and Tabata, 1970). Those animals which are directed to the north, experience few - 46 -problems i n maintaining their geographic p o s i t i o n as surface currents, predominantly t i d a l , are weak i n the Northern S t r a i t . To the south however, a net seaward flow exists at the surface counterbalanced at depth by a net flow into the S t r a i t of Georgia (Waldichuk, 1957). The conti n u i t y and p r e d i c t a b i l i t y of this deep inward flow has yet to be described. As the brackish outflow is confined to the upper 10 m, the deep inflow would occupy a greater cross-sectional area of the Southern S t r a i t ; therefore i t s v e l o c i t y would be reduced. Consequently, t h i s would imply that i f the only b i o l o g i c a l l y reactive movement by N. plumchrus i n the Southern S t r a i t i s a diurnal migration then a loss of N- plumchrus from the Southern S t r a i t to the Juan de Fuca S t r a i t could be expected. However, in turbulent mixing areas l i k e Harrow S t r a i t , plumchrus may descend into slower, less turbulent water flowing northward. Off the coast of Oregon, Peterson et a l . (1979) suggest that during an upwelling event C_. marshallae juveniles remain near-shore and well below the thin surface Ekman layer to prevent offshore advection. The C5's, which d r i f t some 15 to 20 km offshore, upon approaching maturation, descend to depth, and u t i l i z e deeper counter currents to move shoreward. Evidence for the entrainment of JN. plumchrus into the Fraser River plume began with the t h i r d cruise where the C3's make their f i r s t s i g n i f i c a n t appearance. Most of the C3's are found at the eastern plume st a t i o n , 4929, where adults were always scarce. Through to late May, there is a persistent numerical peak of juveniles along the plume transect; usually occurring at st a t i o n 4929 but on several occassions being found further west. The r i v e r i n e fronts of the Fraser River as described by Stronach (1977) would support such a pattern. The location - 47 -of the front and, therefore the peak population, would be dependent on the r i v e r ' s discharge and the point on the t i d a l cycle when the samples were taken. Since the duration of such fronts are sh o r t - l i v e d (LeBlond, 1983), I suspect that much of this production is transported to the south when the concentration mechanism breaks down. This d i s p e r s a l is suggested since the concentration takes place at the surface, and the surface flow is to the south. The concentration of juvenile copepods along the plume transect could also be explained through entrainment processes drawing the animals into the plume coupled with an avoidance of low s a l i n i t y waters by the a n i ~ a l s . Parsons et a l . (1981) predict larger scale (of several kilometers) t i d a l mixing fronts as prominent features of the Southern S t r a i t . The wider based numerical peaks of N. plumchrus extending over several stations were consistent with this p r e d i c t i o n . No overwintering adult N. plumchrus were found i n the Northern S t r a i t during a four year study (from 1965 to 1968, Stephens et a l . , 1969), consequently, the most l i k e l y source for the juveniles i n the northern Central S t r a i t is the Central S t r a i t . The mechanism of estuarine entrainment into the Fraser River plume, followed by periodic displacement to the north, is not as strong an arguement as the proposed southern transport for peak population formation. Movement of intact plume waters to the northern reaches of the Central S t r a i t are rar e l y recorded. Perhaps the residual flows for the northern Central S t r a i t head north rather than to the mainland. Whatever the mechanism that transports juveniles to the northern Central S t r a i t i s , the predicted fronts at the southern end of Texada and Lasqueti Islands (Parsons et - 48 -a l . , 1981), may provide the means for concentration. The picture breaks down on the tenth.cruise (late May). At this time _N. plumchrus exists s o l e l y i n the C5 stage, indicating an end to recruitment from younger stages and a return to depth. An end to emigration from the Central S t r a i t to other reaches of the S t r a i t is also evident. Sizeable populations of juvenile N. plumchrus were found i n the extreme south from cruise fi v e onward, suggesting that some of this production i s lost to Juan de Fuca S t r a i t . It is unclear whether the d i s i n t e g r a t i o n of the southern peak is due to a combination of flushing and reduced recruitment or active migration to deeper waters. A d i f f i c u l t problem comes from tr e a t i n g each cruise as though i t were an instantaneous picture, p a r t i c u l a r l y when some cruises cover a period of seven to eight days. Such a time span may represent between 14 to 16 complete t i d a l cycles, or the time required for a copepod to complete one to several developmental stages (Landry, 1983). In contrasting the Southern and Central S t r a i t population peaks, the prediction may be made that in the northern Central S t r a i t immigration with very l i t t l e emigration is the operative mechanism; whereas, the Southern S t r a i t must r e l y heavily on immigration to counter a p o t e n t i a l l y high emigration. To assess the extent of immigration and emigration, mortality must be considered. Unfortunately, i t is not possible to discern between mortality and emigration in the present research. T i d a l Dispersal In the above model, t i d a l currents are viewed primarily as a means of producing boundary conditions which concentrate organisms. - 49 -Their role regarding transport is acknowledged, but i t is masked by other a c t i v i t i e s i n the S t r a i t . This view may be elaborated by considering those animals whose populations increase continually southward. P. minutus is a prime example of such a d i s t r i b u t i o n . P. minutus is a common member of the copepod assemblage throughout the S t r a i t . In the Southern S t r a i t , i t is the dominant copepod comprising from 46 to 91 percent by number of the copepod assemblage i n the extreme south. In the Central S t r a i t , P. minutus fluctuates between a few hundred and 2,000«m - 2; towards the Southern S t r a i t i t s abundance increases exponentially (southward) and may reach populations i n excess of 100,000«m - 2. The point where rapid increase begins, varies between cruises. I hypothesize that this s i t u a t i o n r e f l e c t s the t i d a l cycle during sampling with P. minutus from Juan de Fuca S t r a i t and the extreme Southern S t r a i t moving i n and out of the Central S t r a i t with the ti d e s . P. minutus is extremely abundant i n Juan de Fuca S t r a i t water and on the Continental Shelf (Chester et a l . , 1980; Mackas and Sefton, 1982). It may be, that inorder to understand the population dynamics of animals inhabiting the Southern S t r a i t , the influence of Juan de Fuca S t r a i t on this region would require consideration. Perhaps, as pointed out by Frost (pers. comm.), two d i s t i n c t species are involved. An attempt was made to discern between the Southern and Central S t r a i t groups of P. minutus on the basis of their mean dry weights for the two regions, but no conclusive evidence was produced. Figure 41 shows the nature of the t i d a l cycle when the sampling occurred, and Figure 15 shows the d i s t r i b u t i o n of. P. minutus i n the Southern S t r a i t . According to the t i d a l hypothesis, high populations are - 50 -expected in conjunction with high flood tides (at the time of sampling) due to the flood tides bringing the copepods further north, and conversely low populations with low tides due to the removal of P. minutus on the ebb t i d e . On three occasions, cruises 2, 8 and 10, anomalous points were observed; a single s t a t i o n was found to have a greater P. minutus population than the immediate more southerly s t a t i o n . On the tenth cruise there was an observed increase of about 1,500'm from s t a t i o n 3439 to 3739; both were sampled on the same ebb tide, but 3739 was sampled f i r s t about 1.0 m above 3439. The eighth cruise yielded a population increase of 3,000'm- moving from stations 3138 to 3739. Again, both stations were sampled on the same ebb tide with 3739 being about 1.5 m higher than 3138. During the second cruise, s t a t i o n 4337 was found to have over 19,000'm more P. minutus than s t a t i o n 3439. As before, both were sampled on the same ebb ti d e ; s tation 4337 was established almost 2.3 m above that of s t a t i o n 3439. These observations are consistent with the i n i t i a l hypothesis. Nevertheless, there were several cruises, such as cruise 9, during which the southern st a t i o n s , sampled on the same ebbing tide, produced a smooth curve reaching i t s maxima in the extreme south, even though the stations were not sampled sequentially from north to south. V e r i f i c a t i o n of the proposed hypothesis cannot be tested from this present paper. To evaluate this topic, several 24-h stations in the Southern S t r a i t , along a north-south axis, and at least one in the Juan de Fuca S t r a i t would be necessary. From this approach, the strength of the c o r r e l a t i o n between population size and t i d a l cycle may be determined, as well as, an i n d i c a t i o n over what distance into the Central - 51 -S t r a i t of Georgia, the b i o l o g i c a l c h a r a c t e r i s t i c s of the Southern S t r a i t may be carried on a given t i d e . Contrary to the P. minutus r e s u l t s , those animals which migrate to the Southern S t r a i t from the Central S t r a i t could be expected to provide a d i f f e r e n t picture. As long as there exists recruitment to the Southern S t r a i t and f r o n t a l a c t i v i t y N. plumchrus would be expected to peak in the Southern S t r a i t . If the P. minutus populations are assumed to move with the tides, then the N. plumchrus population could be expected to peak further north of the P. minutus peak. This s i t u a t i o n seems to be the case. L i f e Strategies Communities whose composition is influenced purely by physical parameters, p a r t i c u l a r l y depth, are r e l a t i v e l y easy to recognize. When i n t e r s p e c i f i c interactions are involved the communities become more d i f f i c u l t to define as the interactions themselves elude v e r i f i c a t i o n . The wide array of l i f e strategies found i n the S t r a i t of Georgia provide a testimony to the many i n t e r s p e c i f i c encounters that may occur. C l a s s i c ecology states through the "Competitive Exclusion P r i n c i p l e " , that two species in competition for the same l i m i t i n g resource, given s u f f i c i e n t time, cannot coexist (Lack, 1944). Whether i t is niche "overlap" or niche "narrowness" which allows for coexistence is not immediately evident, but no doubt, competition does cause the " r e a l i z e d " niche (functional niche) to d i f f e r from the optimum niche (Pielou, 1975). Thus, through competitive i n t e r a c t i o n s , various l i f e stategies a r i s e . Consistent with such l o g i c , is the lack of extensive temporal or spacial d u p l i c a t i o n by any two copepod species i n - 52 -the S t r a i t of Georgia. Neocalanus plumchrus, as described by Fulton (1973) is an example of a K - s t r a t e g i s t . It overwinters and releases i t s eggs at depth. The timing of these events coordinates the appearance of the juvenile stages inconjuction with the spring bloom. This approach may be considered preplanned as no stimulus is r e a d i l y apparent which might t r i g g e r spawning. For N. plumchrus, reproduction is an " a l l - o u t " e f f o r t r e s u l t i n g i n the death of the adult. During the f i r s t 42 days of sampling, when egg release is maximal, the average dry weight of adults dropped over three-fold, from 0.376 to 0.114 mg (Table 3). Growth and l i p i d storage takes place during the period of maximal primary production followed by the C5's return to depth. The temporally unpredictable summer production (Mackas et a l . , 1980) occurs after N. plumchrus returns to depth. During the C5's growth phase the average dry weight increases almost 70% from 0.219 mg to 0.704 mg over a 55-day period (Table 3). By reducing the time spent i n surface waters, N. plumchrus may be l i m i t i n g i t s exposure to predators and seaward-flowing surface currents. L i t t l e yearly v a r i a t i o n i n the timing of these events is shown (compared with Fulton, 1973). Such preciseness may be a result of a highly predictable, stable environment. At Ocean Station P, in the North Central P a c i f i c Gyre, the phytoplankton standing stock shows l i t t l e seasonal v a r i a b i l i t y and seldo— exceeds 2 mg chlorophyll a*m- despite a rapid increase i n primary produ c t i v i t y during the spring ( M c A l l i s t e r , 1962; Stephens, 1964, 1966, 1968). This unique s i t u a t i o n has been attributed to the e f f i c i e n t herbivores of this region (Parsons et a l . , 1977), of which N. plumchrus is the primary species. That is N. plumchrus's grazing rate is greater - 53 -than the rate of primary production. In contrast, the S t r a i t of Georgia combines greater and e a r l i e r water column s t a b i l i t y increasing the exposure of primary producers to the radiant energy of the sun, and d i f f e r e n t phytoplankton species, leading to a greater rate of primary production, a rate which exceeds N. plumchrus's capacity to control them. Hence, in the oceanic environment where primary production and low standing stocks of phytoplankton evidently have lead to food l i m i t a t i o n ( M i l l e r et a l . , i n press), reproduction i n N. plumchrus is nearly continuous (eight months of the year). In the S t r a i t of Georgia where primary production and standing stocks of phytoplankton are much greater, there is a synchronous reproduction ( l a s t i n g approximately two months) peaking i n late February and leading to maximum copepod biomass coincidental with peak spring production. Upon descent to depth, the C5's at Station P immediately commence to mature and reproduce. Another difference found i n the oceanic population i s the apparent a b i l i t y of N. plumchrus to prolong the time spent i n a stage p r i o r to the C5 ( M i l l e r et a l . , i n press). Although i n spring, chlorophyll is i n excess i n the S t r a i t of Georgia, the developmental times for the C5 stage appears to be sim i l a r between the two environments. Consequently, d i f f e r e n t i a l fecundities between the two locals would be suspected, since the adult stage i n both situations i s a non-feeding one. Frost et a l . (1983), l i s t three c h a r a c t e r i s t i c s of N. plumchrus's feeding that adapt i t to the North Central P a c i f i c environment. These include: rapid and accelerated response to changes i n food concentrations, a b i l i t y to feed at below previously suspected threshold concentrations (levels common to the North Central P a c i f i c ) , and a b i l i t y to feed on a wide array of p a r t i c l e s i z e s . - 54 -These a d a p t a t i o n s seem u n i m p o r t a n t i n c o a s t a l h a b i t a t s so t h a t any s e l e c t i v e f o r c e t h a t wou ld m a i n t a i n such a b i l i t i e s wou ld be d i f f i c u l t t o p r e d i c t . R e p e t i t i o n o f F r o s t e t a l . ' s e x p e r i m e n t s i n t h e S t r a i t o f G e o r g i a c o u l d i n d i c a t e why t h e N o r t h C e n t r a l P a c i f i c Ocean ( F r o s t e t a l . , 1983) and t h e S t r a i t o f G e o r g i a (Parsons e t a l . , 1969) a n i m a l s have d i f f e r e n t f e e d i n g t h r e s h o l d s (< 10 ug C l - 1 , and 62 ug C * l - 1 r e s p e c t i v e l y ) . Two s y m p a t r i c c a l a n o i d s p e c i e s , Calanus m a r s h a l l a e and C. p a c i f i c u s , a re s e p a r a t e d t e m p o r a l l y i n t h e i r b r e e d i n g seasons i n t h e S t r a i t o f G e o r g i a . C. m a r s h a l l a e a d u l t s a re d e c l i n i n g f r o m a peak i n abundance a t t h e s t a r t o f t h e F e b r u a r y s a m p l i n g . The appearance o f younger c o p e p o d i t e s i n s i g n i f i c a n t numbers a t t he end o f March s u g g e s t s t h a t t h e d e c l i n i n g C6 p o p u l a t i o n i s a b r e e d i n g p o p u l a t i o n . C. p a c i f i c u s h o w e v e r , s t a r t s t h e season w i t h t h e C5 's f a l l i n g f r o m a s e a s o n a l h i g h and t h e C6 1 s a t a min imum. The a d u l t C. p a c i f i c u s peaks i n m i d - A p r i l . These o b s e r v a t i o n s c o r r e s p o n d w e l l w i t h t h o s e o f Woodhouse ( 1 9 7 1 ) . Woodhouse n o t e s , t h a t a d u l t male C. m a r s h a l l a e a re common o n l y i n t h e e a r l y p a r t o f t h e y e a r s u g g e s t i n g a c o n s e r v a t i v e , one g e n e r a t i o n a y e a r l i f e c y c l e ; whereas C_. p a c i f i c u s shows a second peak around September . J u d g i n g f r o m s p e c i e s c l u s t e r i n g b o t h s p e c i e s C5 s t a g e s appear t o o v e r w i n t e r a t d e p t h . U n l i k e N. p l u m c h r u s h o w e v e r , t he d i s t r i b u t i o n o f t h e C6 's more c l o s e l y resemb les s u r f a c e d w e l l i n g a n i m a l s . Perhaps b o t h Calanus s p e c i e s must w a i t f o r a s t i m u l u s such as c h l o r o p h y l l b e f o r e r e l e a s i n g t h e i r eggs ( M a r s h a l l and O r r , 1955; H e i n r i c h , 1962; Conover , 1 9 6 7 ) . C e r t a i n l y a more o p p o r t u n i s t i c app roach t h a n t h a t o f N. p l u m c h r u s . - 55 -At the time of the f i r s t cruise, adult C. marshallae were 17% heavier (dry weight) than the C5's, that i s , the adults were feeding. By early A p r i l , the adult population had reached i t ' s lowest leve l of abundance, suggesting an end of the reproductive population. Adult weight at this time is s i m i l a r to that of the C5 stage at the time of the i n i t i a l cruise; therefore energetic requirements for reproduction appear to be met during adult feeding. A l l C. marshallae stages, except C5, reach their maximum populations in the extreme south i n the l a t t e r c r u i s e s . This pattern being s i m i l a r to P. minutus would suggest input from Juan de Fuca S t r a i t or surrounding waters. Intere s t i n g l y enough, the C5's followed this pattern in only one cruise (cruise 8). One possible explanation is that the C5 stage i s primarily a deep-water one, or at least a sta~e which avoids turbulent waters. Eucalanus bungii provides one of the more curious l i f e h i s t o r i e s . M i l l e r , et a l . ( i n press) at Ocean Station P uncovered strong evidence suggesting that oceanic E. bungii has a two- or even a three-year l i f e cycle. The f i r s t year of growth appears to take most of the animals to the C3 or C4 stage; the second year i s for maturation and reproduction. By examining the development of the gonads, to determine the spawning condition of adult female E. bungii, about 20% of the spent females were estimated to l i v e to reproduce in t h e i r t h i r d year. The appearance of juveniles, in the surface waters at Station P showed two peaks: the f i r s t occurred in early May - a product of those which had spawned the previous year - and the second more productive spawn came i n early July from the newly matured females. M i l l e r et a l . suggest that such a l i f e cycle arises when there exists yearly v a r i a t i o n s in the p o t e n t i a l of an environment to support the development of _E. bungii. - 56 -This multi-year l i f e cycle does not seem to apply to the S t r a i t of Georgia, at least not in the fashion as described by M i l l e r et a l . ( i n press). While the juveniles did show two peaks in abundance over the course of the study period, the only possible stage which could carry E. bungii onto a second year would be the C6 as there is a clear developmental ascent to this stage. However, adults were absent from the water column in September of 1970 (Krause and Lewis, 1979), and may indicate a one-year l i f e cycle i n the S t r a i t of Georgia. M i l l e r et a l . ( i n press) suggest that the l i f e h i s t o r y of E. bungii at Station P r e f l e c t s an adult stage where i t i s r e l a t i v e l y safe compared to the juvenile stage. Given the abundant alga l resources of the S t r a i t of Georgia then, i t is not s u r p r i s i n g to find more mature overwintering stages in the S t r a i t . This concept of reduced mortality in the adult stages c o n f l i c t s with the opinion expressed by Parsons and LeBrasseur (1970) for N. plumchrus. However, the predatory pressures in the oceanic environment are l i k e l y d i f f e r e n t from that of the S t r a i t of Georgia, and may select for a d i f f e r e n t size range of copepods. Or perhaps, es t a b l i s h i n g i t s stringent overwintering depth requirements sets precedence over predation. It is also possible that E. bungii's transparent body is a d i f f i c u l t image for v i s u a l predators to seek. To pursue the gonadal development of E. bungii in the S t r a i t of Georgia as M i l l e r et a l . ( i n press) have done at Station P, could be valuable; e s p e c i a l l y in conjuntion with v e r t i c a l l y discrete samples. This proposed study could determine whether or not spent females of E. bungii return to depth for a second spawning in the S t r a i t of Georgia. Metridia lucens has received l i t t l e attention i n the l i t e r a t u r e as a member of the S t r a i t of Georgia copepod assemblage, despite - 57 -M. lucens being the most numerically common copepod in the Central S t r a i t during the winter u n t i l the appearance of the juveniles of N. plumchrus1. Strong d a i l y v e r t i c a l migrations of 100 m or more by the adult females (Stone, 1977; Thomas, 1982), deep-water dwelling adult males, and surface inhabiting juveniles, produce a complex picture. Evidence i n the current research for deep-water dwelling adult males stems from the geographic r e s t r i c t i o n s of the males to the Central S t r a i t (Figure 28d), and the lack of any concentrated population along the plume transect (Figure 29d), c h a r a c t e r i s t i c of surface dwelling copepods which I believe accumulate i n t i d a l f r o n t s . The adult male i s more s h o r t - l i v e d than the adult female since the male to female r a t i o drops between the C5 and C6 stage (Figure 27a and b). The f i r s t few cruises correlate _M. lucens adult females with the deep-water species. Consequently, i f the males are located at depth, then mating and possibly release of eggs would also take place at depth. Therefore, for mating to take place, the adult females must locate i n that part of the S t r a i t favouring male populations. At the star t of the sampling for the current research, M. lucens appears to be near the end of i t s reproductive phase; the adult females are ending a decline, the adult males are almost non-existent, and the juvenile stages are on the increase (Figure 26). The dry weights of the adult females are almost invariant suggesting that they may be feeding during the reproductive phase, therefore implying some v e r t i c a l migration (assuming they are herbivorous and t h e i r food is r e s t r i c t e d to the surface layer, M i l l e r , pers. comm.). This pattern breaks down upon termination of reproduction, and is l i k e l y caused by the increased influence of surface currents on non-reproducing females. Towards the end of the sampling season the - 58 -juvenile populations have moved into adult ranks and again adult males and females are d e c l i n i n g . This trend would most l i k e l y indicate the i n i t i a t i o n of a second generation, however, the females did not show the same geographic r e s t i c t i o n as i n the e a r l i e r spawning. It is d i f f i c u l t to say i f K. lucens becomes more opportunistic in the summer, as l i t t l e is recorded with regards to i t s sum-er production. In late winter however, i t ' s reproductive e f f o r t does seem to be more K-selected, although i t i s not clear i f i t s reproductive timing i s predetermined or i f a stimulus is required. Several small but numerically important species including P. minutus, A. longiremis, S. minor, and 0. s p i n i r o s t r i s can only be outlined b r i e f l y . Owing to th e i r small s i z e , only the adults are considered quantitative; therefore the inferences concerning t h e i r l i f e h i s t o r i e s are l i m i t e d . The peak of abundance of P. minutus in the Central S t r a i t occurs during the f i r s t cruise, mid-February. The population decline following this cruise is accompanied by a f o u r - f o l d reduction in the animal's dry weight. A plausible i n t e r p r e t a t i o n of these events would be the synchronous reproduction of overwintering P_. minutus. An aspect of the overwintering strategy is the accumulation of l i p i d s , but to assess the animals v e r t i c a l d i s t r i b u t i o n is d i f f i c u l t . A Pseudocalanus species has been s i t e d as member of the winter surface community (Koeller et a l . , 1979). Here, d i f f e r e n t species may be implied; the taxonomy of this genus is s t i l l being studied. Both S. minor and 0. s p i n i r o s t r i s , on the last cruise, reach th e i r peak i n abundance (Figure 18) which is in agreement with Stephens et a l . (1969). The summer zooplankton assemblage is t y p i c a l l y composed - 59 -of such small herbivorous copepods. The number of generations per year that Pseudocalanus and Aca r t i a species have i s related to the water temperature and food a v a i l a b i l i t y (Corkett and McLaren, 1978; Landry, 1983), and this may be considered an r-selected strategy. In t r o p i c a l waters Pseudocalanus species may have as many as nine generations.(Corkett and McLaren, 1978), l i k e l y i t has fewer i n the colder B r i t i s h Columbian waters. I would speculate, based on t h e i r r e l a t i v e l y small size (<2 mm), that S. minor and s p i n i r o s t r i s are also multi-generation per year species. The copepods discussed up to this point are either herbivorous or omnivorous. There is a small but p o t e n t i a l l y important group of carnivorous adult copepods, including: C h i r i d i u s g r a c i l i s , Gaidius  minutus, and Euchaeta elongata. Other such copepods exist in the S t r a i t , but invariably are found only i n trace numbers. Of the three carnivorous species mentioned, only the l i f e . h i s t o r y of E. elongata has been investigated (Pandyan, 1971; Evans, 1973). Generally, no contradictory evidence was found between this current research and the e a r l i e r work of Evans (1973). Evans found naupliar and f i r s t copepodite stages located at depths below 100 m. This early depth s t r a t i f i c a t i o n is consistent with the v e r t i c a l preference of egg-bearing females (Evans, 1973; unpubl. data). Also found i n this same stratum are hake and pollock larvae (Mason, pers. comm.); Bailey and Yen (1983) and Yen (1982) have found traces of hake larvae i n the guts of E. elongata adults. Evans found the t h i r d to sixth copepodites throughout the water column; these stages had d e f i n i t e diurnal migrations. With such d i e l patterns and using the same logic as for M. lucens, E. elongata could be expected to show a very patchy d i s t r i b u t i o n from north-to-south in the S t r a i t . Such - 60 -a d i s t r i b u t i o n is not borne out, in fact, for a l l observed stages, C2 to C6, the Southern S t r a i t was almost completely void (Figures 39a to e). Evans supports LeBrasseur's (1955) estuarine return flow idea as the probable mechanism for maintaining such a d i s t r i b u t i o n . If this i s true, then E. elongata would necessarily spend a longer part of i t s time at depth as the return flow i s slower than that of the opposing seaward surface flow. Therefore, on any given day a part of the population may be undergoing d i e l migrations, while another part would be r i d i n g the return flow. The estuarine return flow however, is primarily towards the Fraser River plume and not the deeper Central S t r a i t . The lack of any concentration along the plume transect suggests that an a l t e r n a t i v e explanation is necessary. The C4, C5 and C6 populations of E. elongata fluctuate, whereas the C2 and C3 populations reveal increasing numbers as the season progresses (Figure 38). Evans found that the C4's also increased in the summer. Although reproduction maximizes in the spring and summer, some reproduction occurs year round, suggesting year-round feeding. This is further supported by the r e l a t i v e l y constant dry weight shown for E. elongata C6 from mid-February to mid-April (Table 3). Possibly, the C5's and the C6 females would show reduced d i e l migration in the winter and more time spent at depths where the prey, copepods, are located. Based on the feeding appendages of the adult females, C. g r a c i l i s and G. minutus are considered to have carnivorous adult stages (Fulton, pers. comm.). In fact, G. minutus specimens have been taken from the S t r a i t with pollock larvae in t h e i r mouths (Mason, pers. comm.). Both the adult populations of these two species may exceed that of adult E. elongata, and both species show an avoidance of the Southern - 61 -S t r a i t s i m i l a r to E. elongata (Figures 35a to e, 37a and b), but their seasonal population levels d i f f e r . C. g r a c i l i s is s i m i l a r to E. elongata in that fluctuations i n the adult population do not show a seasonal trend (Figure 34), but the juveniles of C. g r a c i l i s are most abundant between mid-April and early May. As the d i s t i n c t i o n between the various copepodite stages of C. g r a c i l i s have not been described taxonomically one cannot state with any certainty i f this varies markedly from E. elongata. G. minutus's l i f e h i s t o r y resembles that of an annual herbivore (Figure 36). During the f i r s t three cruises the adults drop r a p i d l y from a peak that is followed by a modest decline. A few juveniles that may be C5's exist during the f i r s t c r u ise, but these have molted to the adult stage as almost no juveniles are found on the second and t h i r d c ruises. Starting at the end of March, the juveniles increase markedly, implying that after reproduction the adults usually die. Adult G. minutus are e a s i l y recognized as their l i p i d stores are a bright orange or yellow. Whether or not the adults overwinter at depth is not clear, nor can one determine from the present study whether mature adults are feeding. As there appears to be a major decline in the adult population following reproduction, the mature adults are l i k e l y i n a non-feeding phase. Predator - Prey Interactions between predator and prey involve many complex actions which mostly l i e beyond the means of this thesis. In this current work the predator and copepod biomass are b r i e f l y assessed. Neither the predator nor the prey fractions are complete, for example, no account is taken of l a r v a l f i s h , the small-non-copepod f r a c t i o n , and - 62 -animals that are not retained by a 351 um mesh net - a l l are p o t e n t i a l prey items. Besides many pote n t i a l predators, such as juvenile f i s h , are capable of evading the sampling net. Another aspect of this association that is not considered is the v e r t i c a l d i s t r i b u t i o n of the predator biomass; however, the predator mass is assumed to be d i s t r i b u t e d p r o p o r t i o n a l l y in the water column to the copepod mass, an assumption whose v a l i d i t y is questionable. It is also assumed that because the prey biomass show d i s t i n c t seasonal trends, the predator biomass should do the same. This would largely depend on the various l i f e h i s t o r i e s of the d i f f e r e n t predator species. Seasonally, the r a t i o of copepod to predator biomass is almost t o t a l l y dependent on the copepod biomass since the predator biomass shows l i t t l e v a r i a t i o n (Figure 4-?). Geographically, the r e l a t i o n s h i p between copepod and predator may also vary with decreases i n the predator's weight (Figure 43 and 44). This l a t t e r condition i s a feature of the Southern S t r a i t , perhaps the turbulent waters of this region hinder v i s u a l or v i b r a t i o n - s e n s i t i v e predators, or, this may be a response by the predators to the decreased prey biomass i n the Southern S t r a i t . By concentrating their reproductive e f f o r t over a r e l a t i v e l y short time span, not only are copepods able to take advantage of the abundant spring bloom, but may also provide a means of "fl o o d i n g " the predator for a short period with more prey than i t can e f f i c i e n t l y handle (Holling, 1959). Cushing (1971) noted the r e l a t i o n s h i p between transfer e f f i c i e n c y (the r a t i o between production at one trophic leve l to the next), from the primary to the secondary producers, and primary production. For the S t r a i t of Georgia, primary production levels are - 63 -reported to range seasonally from 0.2 to 1.2 gC*m- «d (Parsons et a l . , 1970), and on occasion as high as 8.0 gC*m~ 2 ,d~ 1 (Stockner et a l . , 1979). Usin~ the former range and Cushing's r e l a t i o n , a transfer e f f i c i e n c y of between 15% for the lower production and 8% corresponding to the spring production maxima is obtained. From the current data, an estimate of the transfer e f f i c i e n c y between the secondary and t e r t i a r y producers can be made. Since I have moved up one trophic l e v e l from that used by Cushing, I would expect the transfer e f f i c i e n c y to be lower than that predicted by Cushing. However, a maximum transfer e f f i c i e n c y of 50% is attained, approximating the seasonal production low and a minimum transfer e f f i c i e n c y of 11.4% associated with the peak of the spring bloom. Although the general pattern shown here appears consistent with Cushing (1971), the values yielded are high. Besides the problems outlined e a r l i e r regarding the underestimation of the prey f r a c t i o n , the trophic l e v e l immediately above the herbivores may be overestimated by the addition of c e r t a i n large omnivores, such as euphausiids. Both situa t i o n s result i n high transfer e f f i c i e n c i e s , and act further to outline the gap that exists i n the knowledge of predator to prey interactions in the Georgia S t r a i t . Conclusions In conclusion, the S t r a i t of Georgia copepods may be c l a s s i f i e d into fi v e e c o l o g i c a l l y d i s t i n c t groups; each group showing some s p a t i a l and temporal p a r t i t i o n i n g from the other groups. 1. Diapause and Reproductive: The overwintering herbivorous copepods which exist in the deeper sections of the Central S t r a i t , subsist on l i p i d s accumulated during the previous - 6 4 -spring and summer. They may reproduce i n a n t i c i p a t i o n of the spring bloom without feeding, placing these copepods at the more K-selected end of the spectrum. These animals are believed to be non-feeding stages. From this group arises the second group in which competition for food could be very a c t i v e . Members include: C. mashallae C5; C. p a c i f i c u s C5; E. bungii C5; M. lucens female C6 ? and male C6; N. plumchrus C5 and C6; P. minutus? 2. Growth: Juvenile herbivores concentrate i n the surface waters of the northern reaches of the Central S t r a i t and in the Southern S t r a i t . This is an e x p l o i t a t i v e part of t h e i r l i f e cycle and would l i k e l y have the greatest impact on chlorophyll standing stock. This same group is also l i k e l y to be the most suseptable to mortality i n f l i c t e d by l a r v a l and some juvenile fishes and other predators, as well as, seaward advection by surface waters. Members include: C. marshallae C3 and C4; C. p a c i f i c u s C3 and C4; _E. bungii C l , C2, C3 and C4; M. lucens C4 and C5; N. plumchrus C3, C4 and C5. 3. Reproductive and Growth: This group consists of copepods whose l i f e cycle shows members i n the f i r s t group, but whose adult stage is a feeding stage and l i k e l y r e l i e s on stimulus i n the surface waters such as food to commence reproduction. They are not confined to the deeper Central S t r a i t . - 65 -Members include: C_. marshallae C6; C. pa c i f i c u s C6; E. bungii C6; M. lucens female C6; JP. minutus?. 4. Carnivorous: A l l numerically prominent carnivorous copepods appear to be confined to the Central S t r a i t . Within this group, reproduction may be seasonal or an ongoing process, however, in a l l cases there does appear to be a spring maxima of juv e n i l e s . In some cases the maxima follows the herbivore maxima while i n others the carnivore and herbivore maximas coincide. The juveniles may be herbivorous. Members include: C. g r a c i l i s ; E. elongata; G. minutus. 5. r-selec t e d : These copepods can be subdivided into two groups: a) The f i r s t consists of S. minor and 0. s p i n i r o s t r i s . During late winter their geographic range is s l i g h t l y broader than the deepwater members of the f i r s t group; they may at th i s time be mid-water. b) The second is comprised of A. longiremis and P. minutus. Their populations fluctuate markedly between cruises and may be the res u l t of these animals having r e l a t i v e l y short generation times as suggested by Corkett and McLaren (1978) or t i d a l advection into the Southern S t r a i t from - 66 -neighboring waters. Their populations always peak at the extreme southern end of the Southern S t r a i t . This supports the hypothesis of high k i n e t i c energy systems, l i k e the Southern S t r a i t , favour more r-selected copepods. P r e d i c t i o n Given a physical d e s c r i p t i o n of a coastal body of water based on the k i n e t i c energy in the system, I expect one would find K-selected copepods dominating the low k i n e t i c energy system and r-selected copepods predominating in the higher k i n e t i c energy system. Further, i f a general species l i s t for a region is a v a i l a b l e , and accompanied by an understanding of the ecology of the species on the l i s t , then s p e c i f i c species may be suggested to dominate the environments i n question. As an i l l u s t r a t i o n , I w i l l use Johnstone S t r a i t , a region of high k i n e t i c energy. Characterized by narrow channels and strong t i d a l surges, the waters of Johnstone S t r a i t are continually agitated top to bottom despite depths to almost 500 m (Thomson, 1981). Even in July at the peak of summer heating, the surface waters ra r e l y exceed 10.0°C. Certainly, the oxygen levels i n this S t r a i t appear s u f f i c i e n t at a l l depths to support copepods (minimum 3.9 ml 02*1 _ 1, Thomson, 1981). I would expect this region to be dominated by small r - s t r a t e g i s t s such as 0. s p i n i r o s t r i s , A. longiremis or P. minutus, or even by copepods too small to consider in the present research such as Microcalanus, Clausocalanus or A c a r t i a c l a u s i . Come spring and summer large numbers of those copepods l i s t e d in group two may be added by the surface currents (predominantly t i d a l ) . Because of the cold temperatures and the lack of s t r a t i f i c a t i o n - 67 -in the water column, i t is l i k e l y that primary production for this region would be low and therefore copepod biomass should also be small. The possible exception may come in the spring and summer through the advection of copepods from neighbouring waters. The predictive power of this r e l a t i o n s h i p may not hold under conditions of prolonged s t a b i l i t y where the primary production i s expected to be dominated by f l a g e l l a t e s ; a s i t u a t i o n c l o s e l y resembling that of the instable system (Wyatt, 1980). The moderately stable system is usually characterized by diatoms and the nature of the primary production l i k e l y a f f e c t s that of the secondary producers (Wyatt, 1980). Recommendat ions 1. Increased attention to i n d i v i d u a l species l i f e h i s t o r i e s through the c o l l e c t i o n of v e r t i c a l l y discrete samples over time scales of one to two weeks, coordinated with a look at the ph y s i o l o g i c a l state of the animal, i n p a r t i c u l a r , the development of the gonads and the extent of energy reserves the animal accumulates. 2. 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Cruise schedule, 1981 Cruise Name Date No. of Stations Analysed SG81-1 February 18 - 24 20 SG81-2 March 2 - 1 0 17 SG81-3 March 16 - 21 17 SG81-5 March 30 - A p r i l 6 15 SG81-7 A p r i l 14 - 20 17 SG81-8 A p r i l 27 - May 3 16 SG81-9 May 11 - 15 18 SG81-10 May 25 - 29 18 SG81-11 June 8 - 1 4 17 - 78 -Table 2: T-test for 351 um vs. 200 Um mesh tows Location: 49 27.8 N 124 03.9 W Station depth: 395 m 351 u 200 u Species Stage Mean Mean Neocalanus (No. ni ) plumchrus C5 39433 52322 Calanus t sig.(0.05) 4.230 yes marshallae C5 3483 4671 0.836 no pa c i f i c u s C5 Metridia lucens , FC6 MC6 FC5 MC5 2397 2585 0.181 no 1243 2039 879 1439 1138 2608 2036 2073 0.245 no 1.156 no 2.238 marginal 1.787 no - 79 -Table 2 (cont'd) C4 7175 C3 1593 Pseudocalanus minutus C6 4728 C5 9934 Oithona s p i n i r o s t r i s (>lmm) 2878* Gaidius minutus 2552 Scolecithrice11a minor 1621 6469 0.501 no 8660 24.152 yes 5942 1.116 no 13225 2.023 marginal 8249 6.846 yes 2473 0.109 no 2181 1.205 no * = only two tows considered - 80 -Table 3: Copepod wet and dry weights (i n mg) Species Stage Tr i p wt.wt. Dry Wt. Dry/Wet Percent N. plumchrus C6 81-1 3.468 0.376 10.8 81-3 2.926 0.173 5.9 81-5 2.998 0.114 3.8 C5 81-7 2.661 0.219 8.2 81-9 3.489 0.382 11.0 81-11 3.840 0.704 18.3 C4 81-7 1.029 0.065 6.3 C3 81-7 0.241 0.017 7.1 C. marshallae C6 81-1 1.448 0.204 14.1 81-2 1.396 0.184 13.2 81-3 1.356 0.172 12.7 81-5 1.436 0.164 11.4 81-9(12)1.660 0.190 11.5 C5 81-1/9 1.038 0.213 20.5 C4 0.45(3) 0.030(10) C3 0.20(3) 0.013(10) C. p a c i f i c u s C6 81-5 0.752 0.100 13.3 C5 81-1/2/ 0.561 0.112 '20.0 C3 0.12(3) 0.008(10) - 81 -Table 3 (cont'd) M. lucens FC6 81-1/2/ 0.662 3/7 0.064 9.7 MC6 81-3 0.160 0.017 10.6 FC5 81-3 0.152 0.012 7.9 MC5 81-2/3 0.128 0.013 10.2 C4 81-2/3 0.036 0.004 11.1 M. okhotonsis C6 2.65(la) 0.260(7) Ju 0.37(lb) 0.037(7) E. bungii C6 81-3/5 7.562 0.278 3.7 C5 81-1/2/ 3.175 0.196 6.2 3 C4 1.44(lc) 0.089(11) C3 0.61(lc) 0.038(11) C2 0.23(lc) 0.014(11) Cl 0.10(lc) 0.006(11) E. elongata C6 81-1/2/ 5.595 0.838 15.0 3/5/7 C5 81-1/2/ 3.386 0.557 16.5 3/5/7 C4 2.23(5) 0.345(8) C3 1.00(5) 0.155(8) C2 0.70(5) 0.108(8) - 82 -Table 3 (cont'd) P. minutus C6 81-1 81-2 81-3 81-7 0.135 0.119 0.052 0.109 81-9(12)0.134 0.020 0.017 0.005 0.014 0.013 14.8 14.3 9.6 12.8 9.7 S. minor C6 81-1/2 0.130 0.017 13.1 0. s p i n i r o s t r i s C6 C. g r a c i l i s C6 Ju 81-1/2/ 0.022 0.002 3/7 1.35(6a) 0.123(9) 0.70(6b) 0.064(9) 9.1 G. minutus C6 81-1/2 1.614 0.146 Ju 0.84(6d) 0.076(9) 9.1 C. columbiae Ju 1.05(6c) 0.096(9) A. longiremis C6 81-9 0.066 0.006 9.1 S. brevicaudatus C6 0.14(4) 0.020(4) - 83 -Table 3 (cont'd) (la) Bogorov, 1959 (lb) Bogorov, 1959: stages 2 to 5 averaged ( l c ) Bogorov, 1959: weights adjusted according to measured C5 (2) based on length to weight r a t i o of C5/C4/C3 (3) based on length to weight r a t i o of N. plumchrus C4/C3 (4) based on winter population of P. minutus (5) based on lenght to weight r a t i o of C6/C5 (6a) used E. elongata curve and adjusted to G. minutus C6 (6b) averaged lengths of C3 to C5 (6c) used length r a t i o of C. Columbia C6 to (5. minutus C6 times the estimated weight of G. minutus Ju (6d) used weight r a t i o of C. g r a c i l i s C6 to G. minutus C6 r a t i o times the estimated weight of C. g r a c i l i s Ju (7) used average wet to dry weight r a t i o for a l l M. lucens stages (8) used average wet to dry weight r a t i o for C6 and C5 (9) used average wet to dry weight r a t i o for G. minutus C6 (10) used average wet to dry weight r a t i o for N. plumchrus C4 and C3 (11) used wet to dry weight r a t i o for C5 (12) Southern S t r a i t s t a t i o n used - 84 -Table 4: Predator wet weight 95% confidence l i m i t s Mean Std.Dev. Confid %Conf 3.04 8.14 17.88 0.21 0.27 0.22 0.26 0.34 0.27 91.53 95.88 98.49 - 85 -Table 5: Copepod species encountered and abbreviations used Species Abbreviation Order: Harpacticoida Aegisthus mucronatus Giesbrecht, 1892* AMu Order: M o n s t r i l l i d a e M o n s t r i l l a longiremis Giesbrecht, 1892 MLo M. spinosa Park, 1967 MSp M. wandelii Stephensen, 1913 MWa Order: Cyclopoida Lubbockia wilsonae Heron and Damkaer, 1969 LWi Oithona s p i n i r o s t r i s Claus, 1863 OSp Order: Calanoida A c a r t i a c f . c l a u s i Giesbrecht, 1889 AC1 —' longiremis ( L i l l j e b o r g , 1853) ALo Aetideus divergens Bradford, 1971 ADi A. p a c i f i c u s Brodsky, 1950 APa Bradyidius saanichi Park, 1966 BSa - 86 -Table 5 (cont'd) Calanus marshallae Frost, 1974 CMa — P a c : * - f i c u s s.l.Brodsky, 1948 CPa Candacia columbiae (Giesbrecht, 1889) CCo Centropages abdominalis Sato, 1913 CAb C h i r i d i u s g r a c i l i s Farran, 1908 CGr Epilabidocera longipedata (Sato, 1913) ELo Eucalanus bungii Giesbrecht, 1892 EBu Euchaeta elongata E s t e r l y , 1913 EE1 Eurytemora americana Williams, 1906 EAm Gaetanus intermedius Campbell, 1930 Gin Gaidius minutus Sars, 1907 GMi G. pungens Giesbrecht, 1895 GPu £. v a r i a b i l i s Brodsky, 1950 GVa Heterostylites longicornis (Giesbrecht, 1889) HLo L u c i c u t i a f l a v i c o r n i s (Claus, 1863) LF1 Metridia lucens s.l.Boeck, 1864 MLu (the u maybe replaced by either F or M: Female or Male respectively) M. okhotensis Brodsky, 1950 - MOk Mesocalanus tenuicornis (Dana, 1849) MTe Neocalanus c r i s t a t u s Kroyer, 1848 NCr N. plumchrus (Marukawa, 1921) NP1 Pseudocalanus c f . minutus (Kroyer, 1848) PMi Racovitzanus antarcticus Giesbrecht, 1902 RAn Scaphocalanus brevicornis Sars, 1900 ScB S. euchinatus Farran, 1905 ScE - 87 -Table 5 (cont'd) S c o l e c i t h r i c e l l a minor (Brady, 1883) SMi o v a t a (farran, 1905) SOv Spinocalanus brevicaudatus Brodsky, 1950 SpB Tharybis f u i t o n i Park, 1966 TFu Tort-anus discaudatus Thompson and Scott, 1897 TDi * new to the S t r a i t of Georgia - 88 -Figures Note: Many of the figures use the north-south axis of the S t r a i t of Georgia running from s t a t i o n 2837 to 8535. Not every s t a t i o n was analysed along this transect so i t was necessary to integrate between analysed stations to improve the computer graphics. A l l north-south figures contain 17 stations as opposed to the 16 stations i n c l u s i v e between stations 2837 and 8535. This resulted from the use of a s t a t i o n 6530 in the f i r s t cruise which was extrapolated onto a l i n e between stations 6231 and 6832. A l l numerical abundance of l i f e cycle stages as well as, biomass vs. time figures, use values averaged over the entire north-south transect. - 89 -Figure 1. The S t r a i t of Georgia (From Thomson, 1981) - 8 9 a -Figure 2. Schematized mixing for the S t r a i t of Georgia (From Waldichuk, 1957) - 91 -Figure 3. T-S p r o f i l e s for the S t r a i t of Georgia, 1968 (From Crean and Ages, 1971) C FLATTERY C. FLATTERY C. FLATTERY C. FLATTERY BOUNDARY CMUDQ! - 92 -Figure 4. Ichthyoplankton survey g r i d , 1981 (Courtesy J.C. Mason) - 93 -gure 5. Selected transects and controls - 94_-Figure 6. Predator wet weight vs. dry weight Dry weight = 0.1265 (wet weight) - 0.0209 r = 0.988 - 9 5 -Figure 7. Folsom Plankton Splitter: (a) 1/8 Split (b) 1/16 Split (c) 1/32 Split (d) 1/64 Split counts vs. confidence limit - 95a -200-95% C.I. as a % - 95b -- 95c -2 0 0 100 8 0 6 0 4 0 Mean Counts per Subsample 20-10 8 6-o o o o 8 90 70 50 30 10" I 9 5 % C.I. as a % - 95d -- 96. -Figure 8. Numerical abundance of l i f e cycle stages of Neocalanus  plumchrus N e o c a l a n u s p l u m c h r u s - 97 ..-Figure 9 . JN. plumchrus: north - south d i s t r i b u t i o n patterns (a) C3 (b) C4 (c) C5 (d) C6 - 97a -8 5 3 5 5 , 4 6 5 / m z 2 8 3 7 ^ - ^ 2 plumchrus C6 8 5 3 5 l2 ,000/m2 2 8 3 7 plumchrus C3 8535 3 4 , 0 0 o / m 2 2 8 3 7 plumchrus CA - 98 -F i g u r e 10. N. p l u m c h r u s : plume d i s t r i b u t i o n p a t t e r n s ( a ) C3 ( b ) C4 ( c ) C5 ( d ) C6 - 98a -N. plumchrus C6 4929< 3 7 3 5 3 8 | 1 2 , 3 6 4 / m 2 N. plumchrus C3 N. plumchrus C 4 to 4929 < 5 S 3 8 J N. plumchrus C5 38^13 /m2 - 9 9 -Figure 11. Clustering of stations (a) Cruise 1 (b) Cruise 2 (c) Cruise 3 (d) Cruise 5 (e) Cruise 7 ( f ) Cruise 8 (g) Cruise 9 (h) Cruise 10 ( i ) Cruise 11 60 - 99i -Figure 12. Clustering of species (a) Cruise 1 (b) Cruise 2 (c) Cruise 3 (d) Cruise 5 (e) Cruise 7 ( f ) Cruise 8 (g) Cruise 9 (h) Cruise 10 ( i ) Cruise 11 o o NPIC6 ' -CPaC5 -MLFC6 -M0kC6 -EEIC4 -CMaC5 -NPIC5 -EBuC5 -EEIC6 -EEIC5 -EEIC3 -GMiC6 -EEIC2 -GMiJu -CMaC6-SMiC6 -0SpC6 -CGrC6 -CGrJu -CCoJu -CPaC6 -ML.MC6-PMIC6 -AL0C6 -MLFC5 -MLMC5-MLuC4 -00 o Similarity o -1^  o - a Similarity o o NPIC6 ' 1 CPaC5 ' EBuC6 1 GMIC6 I L EBuC5 1 1 EEIC3 ' EEIC6 1 NPIC5 1 EEIC2 1 SMIC6 = MLFC6 1 M0kC6 1 CGrC6 1 " CGrJu 1 EEIC5 EEIC4 , CCoJu CPaC6 PM1C6 1 AL0C6 MLMC5 j 0SpC6 MLuC4 CMaC6 CMaC5 MLMC6 MLFC5 5 a Similarity o o NPIC6 ' • EBuC6 1 GMIC6 1 . CPaC5 1 1 EBuC5 1 SpBC6 1 EEIC6 1 EEIC2 1 ' I 0SpC6 MLFC6 = 1 SMiC6 EEIC5 1 EEIC3 NPIC3 CPaC6 CGrC6 CMaC6 1 PMIC6 ' AL0C6 CGrJu M0kC6 EEIC4 MLMC6 MLFC5 1 MLMC5 MLuC4 o NPIC6 °-CPaC5 -GMiC6 -NPIC4 -CMaC4-CGrJu — CGrC6 -EEIC4 -EEIC3 -EEIC2 -SMiC6 -0SpC6 -NPIC5 -MLFC5 -MLMC5-NPIC3 -MLuC4 -C M a C 6 -PMiC6 -C M a C 3 -CPaC6 -EEIC6 -EBuC6 -MLMC6-CMaC5 -EEIC5 -MLFC6 -AL0C6 -00 o Similarity CD o o g Similarity NPIC5 5 = 1 CPaC5 ' MLFC6 NPIC4 1 NPIC3 CMaC5 1__ CMaC4 CMaC6 1 PMIC6 1 I AL0C6 1 EEIC5 0SpC6 1 EEIC4 — CMaC3 1 EEIC3 1 CGrC6 CPaC6 1 EEIC6 1 CGrJu ' EEIC2 j I GMIC6 1 EBuC6 MLMC6 1 SMIC6 EBuC2 EBuCI MLFC5 1 MLMC5 1 MLuC4 NPIC5 -NPIC4 -NPIC3 -EBuC2 • CPaC5 -CMaC6-MLMC5-MLuC4 -0SpC6 -PMiC6 -AL0C6 -MLMC6-EEIC6 -CGrC6 • GMiC6 • CGrJu • EEIC3 -EEIC2 -GMIJu -MLFC5 -EEIC5 • MOkJu -EEIC4 • MLFC6 -SMIC6 • CMaC5• CMaC4 -CPaC6 -EBuC3 -CD o Similarity 0) o o ro o 5 oo Similarity o o NPIC5 / = MOkJu NPIC4 1 NPIC3 EBuC3 CMaC6 MLFC5 1 MLMC5 ' MLuC4 1 PMIC6 1 1 AL0C6 1 MLFC6 1 CGrJu EEIC6 EEIC2 1  GMiJu ' 0SpC6 j CGrC6 1 I GMIC6 MLMC6 j SMIC6 EEIC5 -1 EEIC3 EEIC4 CPaC6 CMaC5 CPaC5 o NPIC5 -CMaC5 -EBuC4 -NPIC4 -C6rC6 -CMaC6 -MLFC5 -0SpC6 -MLMC6-MLuC4 -EEIC6 -MLMC5-CGrJu -EEIC3 -PMiC6 -SMiC6 -AL0C6 -EEIC4 -EBuC6 -EEIC2 -6MIC6 -GMiJu — MLFC6 -EEIC5 -CPaC6 -CPaC5 -EBuC3 -00 o Similarity cn o •A o o o 00 o Similarity NPIC5 CMaC5 • CMaC6 CPaC6• EEIC4 • PMiC6 A L 0 C 6 • C P a C 5 C P a C 3 • SM1C6 CGrJu MLMC6 EEIC6 • EEIC2 GMUu EBuC6 EEIC3 • EEIC5 GMiC6 • MLuC4 CMaC4 EBuCI • 0SpC6 • EBuC5 MLFC6 MLFC5• MLMC5-O O - 101 -Figure 13. E q u i t a b i l i t y : north - south pattern Figure 14. E q u i t a b i l i t y : plume pattern - 1i01a -4929 E q u i t a b i l i t y - 102 -Figure 15. Pseudocalanus minutus: north - south d i s t r i b u t i o n pattern Figure 16. P. minutus: plume d i s t r i b u t i o n pattern Figure 17. A c a r t i a longiremis: north - south d i s t r i b u t i o n pattern - 103 -Figure 18. Numerical abundance of adult Oithona s p i n i r o s t r i s and S c o l e c i t h r i c e l l a minor - 103a -- 104 -Figure 19. 0. s p i n i r o s t r i s : north - south d i s t r i b u t i o n pattern Figure 20. CJ. s p i n i r o s t r i s : plume d i s t r i b u t i o n pattern - 104a -8535 0. s p i n i r o s t r i s 0. s p i n i r o s t r i s - 105 -Figure 21. S. minor: north - south d i s t r i b u t i o n pattern Figure 22. S. minor: plume d i s t r i b u t i o n pattern - 105a -8 5 3 5 7 , 8 4 l / m 2 m i n o r 4929 "5 ^ 7 5538 6 , 5 5 6 / m 2 s. m i n o r - 106 -Figure 23. Numerical abundance of l i f e cycle stages of Calanus marshallae 2-x l 0 3 / m 2 2 3 5 7 8 9 10 II Cruise - 107 -Figure 24. C. marshallae: (a) C5 (b) C6 north - south d i s t r i b u t i o n patterns - 108 -Figure 25. C. marshallae: plume d i s t r i b u t i o n pattern (a) C5 (b) C6 - 108a -- 109 -gure 26. Numerical abundance of l i f e cycle stages of Metrida lucens - 110 -Figure 27. M. lucens: sex r a t i o s (a) C5 (b) C6 - 110a -Figure (a) (b) (c) (d) (e) M. lucens: north - south d i s t r i b u t i o n patterns C4 MC5 FC5 MC6 FC6 1111a -8535 u c e n s C 4 8533 M. l u c e n s C5 Ma i e - 112 -Figure 29. (a) (b) (c) (d) (e) M. lucens: plume d i s t r i b u t i o n patterns C4 MC5 FC5 MC6 FC6 - 112a -- 113 - -Figure 30. Numerical abundance of l i f e cycle stages of Eucalanus bungii - 113a -- 114 -Figure 31. E. bungii: north - south d i s t r i b u t i o n patterns (a) Cl (b) C2 (c) C3 (d) C4 (e) C5 ( f ) C6 8 S 3 5 8333 8335 5 t ^ v 8 E. b u n g i i C3 264 / m2 E. b u n g i i C4 - 115 -Figure 32. Numerical abundance of l i f e cycle stages of Calanus p a c i f i c u s 2 x l 0 3 / m 2 2 3 5 7 8 9 1*0 II C r u i s e - 116 -Figure 33. C. p a c i f i c u s : north - south d i s t r i b u t i o n patterns (a) C5 (b) C6 - 116a -- 117. -Figure 34. Numerical abundance of l i f e cycle stages of C h i r i d i u s  g r a c i l i s - 117a -2 3 5 7 8 9 lb i'i Cruise - 118 -Figure 35. £. g r a c i l i s : north - south d i s t r i b u t i o n patterns (a) Ju (b) C6 - 118a -8535 g r a c i l i s 8535 g r a c i - 119- -Figure 36. Numerical abundance of l i f e c y c le stages of Gaidius minutus - 119a -- 120 -gure 37. J3. minutus: north - south d i s t r i b u t i o n patterns (a) Ju (b) C6 - 120a -- 121 -Figure 38. Numerical abundance of l i f e cycle stages of Euchaeta  elongata E u c h a e t a e l o n g a t a - 122 -Figure 39. E_. elongata: north - south d i s t r i b u t i o n patterns (a) C2 (b) C3 (c) C4 (d) C5 (e) C6 - 122a -8535 8535 8535 - 123 -Figure 40. Schematized model for Neocalanus plumchrus d i s t r i b u t i o n - 124 -Figure 41. Tidal cycles and coincidental sampling of Southern S t r a i t stations - 124a -Cruise I T 5035 T 4636 4039 4337 . * 2837 3439 February 22 2 3 2 4 I000 2 0 0 0 Cruise 2 0 5 0 0 . t • 4337 I500 t •4636 5035 , t'2837 34394 • March E 2 I000 . r 4636 2 0 0 0 Cruise 3 0500 I500 f 4039 March 2 0 3I38 -21 I500 4 0 3 9 C r u l s e 5 4636 OIOO I000 April .t . 3 4 3 9 2 6 3 8 * t ' -3138 1000 Cruise 7 2 0 0 0 "t4337-•4636 . April t 5 0 3 5 18 3138 u 3739 . • t 2837 19 1000 2 0 0 0 Time of Day 0 5 0 0 OIOO 1000 Cruise 8 •" 1 4 0 3 9 . ' 3 7 3 9 ' 4 3 3 7 i f ' ' . . t -2837 31381 5035 May 2 0 5 0 0 1500 OIOO 1000 Cruise 9 f 4 0 3 9 ^ • t ' • •4337 3 4 3 9 - 2 8 3 7 3138 • • ' May 1 5 0 5 0 0 I50p Cruise 10 f 4 0 3 9 • 4 6 3 6 ' t. • t 3 7 3 9 - 2837-3 4 3 9 May 28 29 1500 OIOO 1000 Cruise II . ' ' t 1 . t 2638 • •3138 t '4337 T 5 0 3 5 . , - 3 7 3 9 June 13 0 5 0 0 1500 - 125. -Figure 42. Predator - copepod: biomass and r a t i o vs. time - 125a -j 2 3 5 7 8 9 IO II Cruise - 126 -Figure 43. Copepod biomass: north - south d i s t r i b u t i o n pattern Figure 44. Predator biomass: north - south d i s t r i b u t i o n pattern - 1 . 2 6 a -8 5 3 5 C o p e p o d B i o m a s s 8 5 3 5 P r e d a t o r B i o m a s s ... - 127. -Figure 45. Timing of prominent deep-water adult copepods APPENDICES Appendix A: Non-reduced, processed copepod data (Values i n N/m2) C r u i s e : SG81-1 Scat i o n : 7829 7832 8535 7934 7634 6530 5931 5035 4636 4337 4039 3439 2837 7136 7627 7227 6727 4929 5132 5538 Depth (in meters): 311 251 287 385 393 352 324 280 223 181 174 170 209 200 164 185 79 127 232 181 NP1C6 4661 3963 3845 5811 6953 3654 2918 3090 758 148 90 38 38 1801 972 1587 4 152 1848 1295 NP1C5 541 118 410 708 501 68 37 32 16 16 8 0 0 29 13 16 0 0 0 0 CMaC6 2496 2691 1324 1751 2600 2131 1933 2447 2368 1789 1681 1395 1226 2334 1294 2237 693 2000 2978 1538 CMaC5 571 296 536 1006 595 406 465 354 647 328 218 154 115 648 263 587 122 380 342 308 CPaC6 30 0 32 37 .125 34 19 97 16 0 8 26 0 58 20 0 0 0 68 32 CPaC5 2135 828 1670 3465 3320 1725 1933 1642 789 197 188 64 38 692 387 587 30 304 856 599 MLFC6 4510 4791 5736 6780 9490 6597 6320 4153 3647 1412 931 704 249 3746 3251 7997 11 2304 3423 4404 MLMC6 60 59 32 75 94 68 37 64 111 82 105 90 96 14 20 16 0 127 34 49 ML PC 5 90 89 95 37 31 101 74 0 U l 16 53 205 77 58 20 32 34 25 68 49 MLMC5 782 680 126 261 345 271 353 258 237 181 98 346 460 72 99 651 366 51 205 243 MLuC4 601 2691 378 671 407 575 539 612 916 1182 210 1613 1245 317 204 841 1169 279 240 372 M0kC6 120 118 63 112 251 203 93 161 79 82 0 0 0 14 26 48 0 0 171 81 MOkJu 0 0 32 37 0 34 19 0 0 0 0 0 0 0 0 0 0 0 0 0 EBuC6 0 0 32 112 157 169 93 64 0 16 0 13 0 0 0 0 0 0 0 0 EBuC5 301 148 315 2086 2881 778 409 193 79 0 8 0 0 0 0 32 0 0 34 32 EBuC4 0 0 0 37 0 0 19 32 0 0 0 0 0 0 0 0 0 0 0 0 EBuC3 0 0 0 37 0 0 0 0 0 0 0 0 0 0 0 0. 0 0 0 0 EE1C6 150 0 63 75 282 237 112 32 32 0 0 0 0 14 0 63 4 13 0 0 EE1C5 60 30 63 186 125 135 112 193 79 66 15 0 19 14 7 ' 0 0 25 34 16 EE1C4 271 59 158 75 125 169 130 225 126 49 8 0 19 58 53 79 0 25 34 65 EE1C3 90 89 63 186 94 237 167 161 47 16 23 0 0 14 13 63 0 13 103 16 EE1C2 90 30 0 75 313 304 167 97 16 16 0 0 0 0 0 16 0 13 34 0 PMiC6 4090 5086 2301 3241 4259 3349 3271 4604 5226 5204 6124 12787 37861 3876 2390 6093 1671 2329 3697 2882 SMiC6 421 621 - 536 894 501 1725 1134 1062 1152 673 270 294 172 836 381 603 126 532 1574 745 0SpC6 1203 2188 1859 2123 2255 6597 3290 3058 2352 968 368 589 19 3962 611 2190 495 1266 7256 1328 CGrC6 120 30 63 112 63 169 149 129 142 98 23 26 0 43 39 127 0 13 171 97 CGrJu 210 89 32 261 376 406 335 161 316 427 75 102 38 173 92 317 0 25 205 372 GMiC6 692 887 599 1676 1315 2470 1208 612 111 16 8 0 0 14 13 111 0 25 308 16 GMiJu 30 30 0 75 31 744 186 32 0 16 0 0 0 0 0 0 0 0 34 0 Cruise: SG81-1 (cont'd) Stat ion: 7829 7832 8535 7934 7634 6530 5931 5035 4636 4337 4039 3439 2837 7136 7627 7227 6727 4929 5132 5538 Depth (in meters): 311 251 287 385 393 352 324 280 223 181 174 170' 209 200 164 185 79 127 232 181 A L 0 C 6 60 59 0 37 0 34 0 0 189 279 308 333 402 29 13 0 19 76 0 16 CC0C6 0 0 63 0 63 34 0 64 126 0 15 13 0 14 0 0 0 25 34 32 CCoJu 60 0 32 112 63 101 19 129 126 49 23 0 0 14 7 32 0 51 0 65 ADiC6 0 0 32 0 0 0 19 0 L6 0 8 26 0 14 0 0 0 13 0 0 ADiJu 0 0 0 0 31 0 19 0 0 0 0 38 0 0 0 16 0 0 34 0 SpBC6 0 0 0 37 63 0 37 0 0 0 0 0 0 0 0 0 0 0 0 0 ScBC6 0 0 32 75 94 0 37 64 0 0 0 26 19 0 0 0 0 0 0 0 KAnC6 30 0 0 0 0 0 0 32 16 0 15 13 0 0 0 0 0 0 0 0 TDiC6 0 0 0 0 0 0 0 0 0' 0 0 13 96 0 7 0 0 13 0 0 I ro 10 1 C r u i s e : SG81-2 S t a t i o n : 7829 7832 8535 7934 6832 5931 5035 4636 Depth ( i n m e t e r s ) : 335 282 316 382 390 343 296 232 NP1C6 3239 2164 3246 5173 3676 2502 949 505 NP1C5 67 69 33 133 37 55 17 0 NP1C3 0 0 0 0 0 0 0 • 0 NP1C2 0 0 0 0 0 18 17 0 CMaC6 1518 1575 1410 1691 724 846 1254 3380 CMaC5 304 277 295 99 167 74 102 122 CPaC6 34 35 98 33 37 37 34 61 CPaC5 .1552 866 1082 2818 2544 1785 1271 566 CPaC4 0 0 0 0 0 0 17 0 CPaC3 0 0 0 0 0 0 0 0 CPaC2 0 0 0 0 0 0 0 15 MLFC6 5364 4795 4295 7030 4679 2355 4949 1178 MLMC6 304 104 98 199 167 294 441 260 ML PC 5 405 L38 492 497 260 74 492 459 MLMC5 L282 692 1311 2122 854 239 949 581 MLuC4 1754 2701 984 2222 2340 331 1797 2630 M0kC6 169 69 131 232 130 74 119 15 EBuC6 236 69 164 298 371 386 . 339 107 EBuC5 810 156 852 1658 1021 938 576 153 EBuC4 0 0 33 33 0 0 0 0 EBuC3 0 17 0 0 0 0 0 0 EE1C6 202 104 164 232 130 147 169 107 EE1C5 67 173 131 232 56 92 0 31 EE1C4 34 35 66 99 149 166 51 46 EE1C3 135 69 66 232 371 239 220 61 EE1C2 202 52 197 232 186 184 • 203 260 PMiC6 2496 1298 1902 5040 2748 1343 3407 12892 SMiC6 2294 1523 918 1227 1263 1361 1339 627 OS pC6 2294 2233 2656 4244 1393 294 1661 1285 CGrC6 169 104 197 166 241 350 136 46 CGrJu 135 260 66 332 260 276 186 76 GMiC6 708 415 590 928 724 865 966 245 GMiJu 0 0 0 0 0 0 51 0 AL0C6 0 0 33 232 0 55 17 321 4337 3439 2837 4929 5132 5538 7136 6727 7227 178 171 190 138' 245 188 209 183 138 236 69 57 69 904 651 2104 469 109 0 0 0 0 17 17 0 0 4 0 14 0 0 0 0 0 0 0 15 0 0 14 0 0 0 0 0 1165 894 2669 1516 1071 1185 945 2068, 621 15 55 227 62 84 67 46 226 60 0 14 0 21 50 17 46 129 23 398 69 114 206 535 618 732 663 169 0 0 57 0 0 0 0 0 0 0 14 57 0 0 0 0 0 0 0 41 114 14 17 0 0 0 0 2581 701 398 1619 2192 9013 4208 3264 542 236 179 -1-7-0-— 48 218 417 198 194 72 ' 811 248 341 144 268 684 366 646 188 1136 825 398 316 1155 801 732 663 1273 2802 2324 1760 563 1071 1419 808 953 1363 0 0 0 48 33 317 122 210 4 0 0 . 0 0 117 67 30 0 0 0 0 0 L4 184 17 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 28 0 7 100 117 46 48 11 0 55 0 7 33 267 122 16 4 0 14 57 14 134 0 30 16 15 15 0 0 14 151 67 15 . 32 19 0 28 57 21 184 100 0 32 8 33096 13929 81435 714 2242 2754 2272 4024 877 619 413 227 494 987 684 1250 856 425 2050' 536 568 480 1506 818 1159 1664 764 74 0 0 69 134 167 ' 46 178 11 59 28 0 75 234 217 168 436 87 59 14 0 34 535 83 15 16 4 0 0 0 0 17 0 0 0 0 324 275 738 21 33 0 0 16 8 Cruise: SG81-2 (cont'd) Stat ion: 7 8 2 9 7832 8535 7934 6832 5 9 3 1 5035 4 6 3 6 4337 3439 2837 4929 5132 5 5 3 8 7136 6 7 2 7 7227 Depth (in meters): 335 282 3 1 6 382 390 343 296 232 178 171 190 138 ' 245 188 209 183 138 CC0C6 0 17 0 33 37 37 0 15 L5 0 0 7 33 100 0 48 0 CCoJu 34 0 66 66 93 55 17 76 0 14 0 21 50 0 30 16 23 ADiC6 34 17 0 0 56 0 34 15 29 0 0 0 17 0 0 16 0 ADiJu 0 0 0 33 0 18 0 15 0 41 0 0 0 17 15 0 8 SpBC6 101 156 131 199 74 55 17 0 0 0 0 0 . 0 17 0 0 0 ScBC6 0 35 66 66 0 37 119 15 15 14 0 0 17 0 0 0 0 TDiC6 0 0 0 0 0 0 0 46 0 28 0 0 0 0 0 0 0 HL0C6 0 0 33 33 0 0 17 0 0 0 0 0 0 0 0 0 0 KLoJu 0 0 0 0 0 0 0 0 0 14 0 0 0 0 0 0 0 CAbC6 0 0 0 0 0 0 0 0 0 41 0 0 0 0 0 0 0 C r u i s e : SG81-3 Scat i o n : 7829 7832 Depth ( i n meter 313 25 NP1C6 1561 1239 NP1C5 0 0 NP1C4 0 0 NP1C3 49 33 CMaC6 455 1071 CMaC5 0 33 CMaC4 0 0 CMaC3 0 0 CPaC6 98 167 CPaC5 748 736 CPaC4 0 0 MLFC6 2570 6478 MLMC6 927 105 MLFC5 862 1975 MLMC5• 1252 3214 MLuC4 4651 348 MOkC6 114 167 EBuC6 618 636 EBuC5 114 167 E1C6 81 117 E1C5 8L 67 E1C4 114 84 E1C3 211 117 E1C2 276 251 P M i C 6 1561 2712 S M i C 6 976 125  OSpC6 3708 279 CGrC6 163 84 CGrJu 98 151 G M i C 6 553 335 GMiJu 16 0 A L 0 C 6 49 84 CC0C6 0 33 8535 7934 7634 ): 326 372 373 1567 2164 2008 0 0 18 0 0 0 31 34 0 3259 1454 705 0 68 0 0 0 0 0 34 0 188 169 211 1003 2130 1620 0 0 0 3384 3517 2184 1003 812 845 1034 575 845 1003 1082 1444 1786 2299 969 0 0 35 877 1488 881 407 406 634 219 135 88 125 203 88 0 34 123 63 101 106 564 406 159 2601 1995 1004 1222 1488 705 5170 2604 1022 157 101 229 94 169 123 721 845 493 0 0 0 63 135 0 0 34 53 6231 5532 4636 353 345 208 1088 858 301 0 0 0 0 0 0 45 130 105 417 820 662 75 19 15 0 0 0 0 0 0 238 112 135 924 839 391 0 0 0 5619 1622 3221 1490 969 1806 954 1174 4906 1431 1100 3371 2146 2498 3431 60 37 120 358 130 105 104 56 0 149 75 60 238 37 105 224 37 15 358 93 90 477 205 45 2325 1957 1972 1401 1249 1219 2400 1286 1430 238 149 150 134 37 120 462 317 60 30 0 0 30 19 30 60 112 15 4039 3439 3138 187 192 184 50 64 0 0 0 0 0 16 33 99 128 163 812 1184 6214 33 0 0 17 0 0 0 0 0 33 80 33 83 80 228 17 0 0 1674 656 944 1757 1344 1887 1691 1408 1724 1094 1504 1594 2222 2671 2115 50 0 0 17 0 0 0 0 0 33 16 0 50 32 0 50 0 33 17 16 0 50 32 0 2354 2767 7191 962 368 390 978 576 456 66 96 65 83 96 0 0 16 0 0 0 0 33 96 163 0 0 0 2837 4929 5132 206 ' 132 239 0 68 354 0 7 0 0 20 0 62 258 97 9999 536 1546 0 7 32 . 0 0 0 0 7 0 62 109 97 31 149 354 0 0 0 966 1854 2448 1090 903 1578 1838 407 1224 2305 306 1159 1215 469 2158 0 41 129 0 14 97 31 0 0 0 14 129 31 20 64 0 0 161 31 20 97 31 7 290 21773 1100 1514 716 1331 1192 93 1433 2158 93 0 97 0 14 322 0 0 483 0 0 32 467 20 32 93 0 0 5538 7227 6727 186 185 168 259 421 237 0 0 15 0 0 0 49 16 0 470 681 769 0 0 15 0 0 0 16 16 0 243 194 163 680 859 296 0 0 0 2235 1605 2914 988 1605 636 583 989 2071 664 956 1804 1377 1070 1139 130 16 30 16 146 118 0 16 15 32 49 30 49 16 59 16 146. 89 49 81 30 97 113 0 2576 1669 1671 680 989 1316 1863 1653 1420 194 113 89 292 276 222 65 259 222 0 0 0 32 97 59 0 0 15 Cruise: SG81-3 (cont'd) S t a t ion: 7829 7832 8535 7934 7634 6231 5532 4636 D e p t h ( i n m e t e r s ) : 313 225 326 372 373 353 345 208 C C o J u 33 17 31 0 0 60 75 45 ADiC6 . 0 0 0 0 0 0 0 15 A D i J u 0 0 0 0 ' 0 0 0 0 APaC6 0 0 0 34 0 0 0 15 SpBC6 49 17 188 101 106 15 19 0 ScBC6 0 0 31 34 0 30 75 0 TDiC6 0 0 0 0 0 0 0 0 HL0C6 0 0 0 0 18 0 0 0 U L o J u 0 0 0 34 0 0 0 0 CAbC6 0 0 0 0 0 0 0 0 CLaC6 0 0 0 0 0 0 0 0 GInC6 0 0 0 0 0 0 0 0 E L o J u 0 0 0 0 0 0 0 0 ML0C6 0 0 0 0 0 0 0 0 MWaC6 0 0 0 0 0 0 0 0 MSpC6 0 0 0 0 0 0 0 0 )39 3439 3138 2837 4929 5132 5538 7227 6727 .87 192 184 206' 132 239 186 185 168 0 0 0 0 0 0 0 0 15 0 0 33 0 7 0 16 0 15 0 0 0 0 7 32 16 0 ' 0 0 0 0 0 0 0 0 0 0 33 16 0 0 0 32 16 32 0 0 16 0 31 0 0 0 0 0 0 0 65 31 0 0 32 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 31 0 0 0 0 0 0 0 0 31 0 0 0 0 0 0 0 0 31 0 0 0 0 0 0 0 33 0 0 0 16 0 0 0 16 0 0 0 0 0 0 0 0 16 0 0 0 0 0 0 0 0 0 0 0 7 0 0 0 0 Cruise: SG81-5 Station: 7829 7832 8535 Depth (in meters): 330 250 335 NP1C6 864 305 958 NP1C5 36 0 0 NPLC4 468 1157 548 NPLC3 6808 10902 17863 CMaC6 144 274 548 CMaC5 72 61 0 CMaC4 0 0 68 CMaC3 36 61 137 CPaC6 468 396 548 CPaC5 612 579 684 CPaC4 0 0 0 CPaC3 0 0 0 MLFC6 4322 5238 4928 MLMC6 2233 974 1163 MLFC5 1909 1401 1643 MLMC5 2341 2010 2943 MLuC4 1693 2619 2943 M0kC6 72 30 0 EBuC6 864 152 342 EBuC5 0 0 0 EBuC4 0 0 0 EE1C6 108 . 152 68 EE1C5 72 30 137 EE1C4 144 122 205 EE1C3 360 426 137 EE1C2 792 822 274 PMiC6 1117 1279 2190 SMiC6 1225 1127 1574 0SpC6 2846 2802 4175 CGrC6 288 30 137 CGrJu 216 183 137 GMiC6 180 61 411 GMiJu 0 0 0 7634 6832 5931 4636 394 385 321 223 1241 360 138 0 73 36 0 193 584 288 932 3803 7886 3208 4867 4898 146 252 69 97 0 72 35 32 0 0 35 161 146 36 0 97 292 180 207 258 803 901 552 193 0 36 0 0 0 36 0 0 3286 757 5385 4866 1972 1730 1070 2900 1898 1622 1622 4544 2191 2560 1208 4254 1898 1839 2002 2610 0 0 0 0 803 144 587 32 0 0 0 32 0 0 0 0 73 36 69 97 146 36 207 161 146 108 104 32 365 216 552 97 511 685 690 129 2410 2091 1312 3255 1898 1298 2416 806 4454 3857 4453 1740 365 144 104 32 365 216 380 161 876 865 311 0 0 0 35 0 4039 3439 3138 2638 165 192 176 199 0 0 0 0 86 202 253 64 4894 12731 7530 3493 8186 20679 19934 11684 114 943 1266 2286 57 67 127 254 172 337 759 445 29 337 443 1016 229 202 127 191 143 67 63 64 0 0 127 127 0 0 127 64 9245 3503 4430 1778 1746 3099 2595 1524 7299 4311 4240 2159 5982 4580 7720 3747 3692 3839 4240 3874 0 0 0 0 286 337 127 0 29 0 0 0 0 0 0 0 0 67 0 0 0 0 0 64 0 0 63 0 0 0 0 0 0 0 0 0 3921 9498 44993 60644 544 808 633 572 2118 2425 1582 1588 57 0 0 0 114 0 0 0 0 67 0 0 0 67 0 0 4929 5538 '• 6727 7136 139 179 . 180 197 0 29 61 186 129 29 122 8 6841 2679 608 97 13552 6800 9730 194 516 177 122 56 0 29 0 16 323 29 0 16 387 177 122 8 65 235 426 331 258 265 365 363 0 29 0 0 65 88 61 8 4969 4975 7723 476 4324 2767 2007 2388 4711 2384 2919 524 2388 1648 3953 678 4130 2237 2797 1331 0 59 122 0 0 118 487 40 0 0 0 8 0 0 61 0 0 59 122 24 129 59 122 ' 24 0 147 122 65 0 147 182 145 0 177 182 234 2710 4327 2493 1840 1226 1884 1520 2396 2710 3503 6264 6479 0 88 304 266 0 412 608 355 0 0 182 468 0 0 0 0 Cruise: SG81 -5 (cont'd) Station: 7829 7832 8535 7634 Depth (in meters): 330 250 335 394 A L 0 C 6 0 0 68 219 CC0C6 0 61 0 0 CCoJu 0 0 68 73 A D i C 6 0 30 0 0 ADiJu 0 0 0 0 APaC6 0 0 0 0 SpBC6 108 30 205 146 ScBC6 0 0 0 0 ScaC6 0 0 0 0 TDiC6 0 0 0 0 HLoJu 36 0 "0 0 CAbC6 0 0 0 0 RAnC6 36 0 0 0 C I n C 6 0 0 0 0 GaeJu 0 0 0 0 EAmC6 0 0 0 0 C l a C 6 0 0 0 0 E L 0 C 6 0 0 0 0 ELoJu 0 0 0 0 BSaC6 0 0 0 0 BSaJu 0 0 0 0 )832 . 5931 4636 4039 3439 3138 2638 4929 5538 . 6727 7136 385 321 223 165 192 176 199 139' 179 180 197 288 0 32 29 135 127 318 129 147 122 56 0 0 32 0 0 0 0 0 59 61 8 0 104 0 0 0 0 0 0 0 0 0 0 0 0 0 0 63 0 0 0 0 8 36 0 32 0 0 0 0 0 29 0 0 0 35 0 0 0 0 0 0 0 0 0 36 0 64 0 0 0 0 0 29 0 24 72 69 0 0 0 0 64 0 0 0 0 0 0 0 0 0 0 0 65 0 0 0 0 0 0 0 0 127 191 0 59 0 0 72 0 0 0 0 0 0 0 0 0 0 36 0 0 0 0 0 318 0 353 61 65 36 0 0 0 0 0 0 0 0 0 0 0 35 0 0 0 0 0 0 0 0 . 0 0 0 0 29 0 0 0 0 0 0 0 0 0 32 0 0 0 0 0 0 0 0 0 0 0 0 67 0 0 0 0 0 0 0 0 0 0 0 63 64 0 0 0 0 0 0 0 0 0 127 127 0 0 0 0 0 0 0 0 0 0 0 65 0 0 0 0 0 0 0 0 0 0 65 0 0 0 C r u i s e : SG81-7 Station: 7829 7832 8535 7634 6832 5931 5035 4636 Depth ( i n m e t e r s ) : 283 232 302 397 390 338 292 217 NP1C6 66 0 0 74 69 39 37 0 NP1C5 5403 6363 12726 4482 2746 2781 3727 3418 NP1C4 5139 6303 39682 9704 4050 3583 10211 10130 NP1C3 2833 2881 13889 4889 824 979 6298 4039 CMaC6 66 120 342 148 34 98 335 311 CMaC5 198 120 479 185 34 137 224 249 CMaC4 132 0 684 37 69 176 708 186 CMaC3 659 120 205 U l 0 59 410 124 CPaC6 395 600 616 407 481 294 224 311 CPaC5 66 300 0 333 412 98 75 124 CPaC4 132 0 " o 0 0 39 37 0 CPaC3 66 0 137 74 69 39 37 0 MLFC6 2438 13746 4379 2593 5286 1664 8460 3356 MLMC6 3228 2401 7184 2556 5664 3779 3056 2237 MLFC5 922 3301 2053 3148 961 1410 5031 4537 MLMC5 1977 4082 4721 3852 2197 2017 4584 4350 MLuC4 3755 5402 5473 3296 4256 6247 6373 6277 M0kC6 0 60 0 0 0 0 0 0 MOkJu 0 0 0 37 0 0 224 186 EIiuC6 198 120 137 222 309 59 37 0 EBuC5 0 0 0 0 0 0 37 0 EBuC4 0 0 0 0 0 0 37 0 EBuC3 0 0 0 U l 69 78 335 0 EBuC2 66 0 0 444 34 392 1416 311 EBuCl 66 360 274 704 515 372 1193 249 EE1C6 132 240 342 185 275 137 37 62 EE1C5 66 360 137 74 103 59 149 124 EEIC4 66 240 137 74 378 215 112 0 EE1C3 0 240 274 407 892 509 224 62 EELC2 132 360 684 778 789 450 261 62 PMiC6 3163 1501 2326 2556 1407 1684 7230 7085 SMiC6 1515 1681 3010 1445 1854 2389 2311 1989 0SpC6 5534 6843 7800 4371 5904 8126 5441 7147 4337 3739 3138 2837 4929 5132 5538 7136 6727 184 186 178 191* 138 221 185 175 162 0 0 0 0 0 0 0 0 0 3495 9482 4019 1890 7915 10380 4355 9885 6180 8387 21149 14569 10248 6596 17512 6786 11136 11023 5553 12931 11492 10539 2895 8050 2093 2502 3441 272 517 1256 2180 147 636 101 63 64 39 345 314 581 220 494 338 250 191 272 345 377 2471 73 636 439 375 127 155 517 377 581 73 424 236 63 191 39 0 314 145 110 71 135 125 382 155 57 314 291 0 71 135 63 191 0 57 126 0 0 0 34 0 64 0 0 63 0 0 141 0 0 0 1359 2299 4459 654 5900 9533 4085 11011 7582 1010 2299 188 2035 1173 3389 3646 2753 1274 2912 3161 2512 800 1429 7838 2296 2815 2166 3728 4253 5777 4143 2052 9039 4152 3879 1975 7028 7471 4773 5451 3555 7344 5942 4692 5989 0 0 0 ' 0 73 0 0 0 0 39 0 0 0 147 282 34 63 64 0 57 0 0 0 141 34 63 0 0 0 0 0 0 0 0 0 0 0 57 0 0 0 0 0 0 0 0 460 63 73 0 141 68 125 0 311 575 565 654 183 353 540 313 191 388 747 691 800 147 424 473 125 64 39 0 0 0 37 0 135 0 64 0 0 0 0 73 71 34 125 255 0 57 0 0 110 0 203 438 510 39 0 0 0 110 141 101 188 127 0 0 0 0 73 0 135 188 0 10135 33447 46030111205 2162 7697 3376 1752 2612 1709 747 126 218 1136 2401 1148 2190 1784 2524 2471 1382 509 4031 5014 5570 6068 7773 Cru ise : SG81-7 ( c o n t ' d ) S C a t 7 8 2 9 7832 8535 7634 6832 5931 5035 4636 4337 3739 3138 2837 4929 5132 5538 7136 6727 Depth ( i n m e t e r s ) : , , , „ 283 232 302 397 390 338 292 217 184 186 178 191 138 221 185 175 162 CGrC6 0 0 137 74 309 157 37 124 0 0 0 0 37 71 169 125 64 CGrJu 593 660 753 370 1133 294 186 373 39 0 0 0 0 141 270 250 573 GMiC6 132 480 821 556 584 196 149 0 0 0 0 0 73 0 101 188 64 GMiJu 0 0 0 37 103 59 0 0 0 0 0 0 0 0 0 0 0 AL0C6 0 0 205 222 34 39 522 373 660 1092 1068 1454 73 424 101 0 191 CC0C6 132 60 0 74 0 39 75 0 0 0 0 0 37 0 68 125 0 CCoJu 0 60 68 37 0 59 37 0 0 0 0 0 0 0 0 0 0 ADiC6 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 63 0 ADiJu 0 0 0 37 0 0 37 0 0 0 0 0 0 0 0 0 0 SpBC6 66 60 68 74 137 98 75 0 0 0 0 0 0 0 68 0 64 ScBC6 0 0 0 111 0 20 37 62 0 0 0 0 0 0 0 0 0 ScaC6 0 0 0 0 0 0 37 0 0 0 0 0 0 0 0 0 0 TD iC6 0 0 0 0 0 20 0 0 0 57 0 145 0 0 0 0 0 HLoJu 0 0 137 148 69 0 0 0 0 0 0 0 0 0 0 0 0 CAbC6 0 60 0 0 0 0 0 0 0 115 0 0 0 0 68 0 0 KAnC6 0 60 0 0 0 20 37 0 0 57 0 0 0 0 0 0 0 LFLC6 0 60 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 EL0C6 0 0 0 0 0 0 0 0 0 0 0 218 0 0 0 0 0 ELoJu 0 0 0 0 0 20 0 0 39 57 0 145 0 0 0 0 0 CaeJu 0 0 0 0 0 0 37 0 0 0 0 0 0 0 0 0 0 LuWJu 0 0 0 0 0 0 37 0 0 0 0 0 0 0 0 0 0 ClaC6 0 0 0 0 0 0 0 0 39 0 0 0 0 0 0 0. 0 BSaC6 0 0 0 0 0 0 0 62 0 0 0 73 0 0 0 0 0 BSaJu 0 0 0 0 0 0 0 0 0 0 0 0 110 0 0 0 0 Cruise: SG81-8 Station: 7829 8535 7133 6832 5931 5035 4337 4039 3739 3138 2837 4929 5132 5538 7136 6727 Depth (in i meters): f 229 340 396 390 350 310 181 188 193 185 190 151 247 197 192 150 NP1C6 70 0 0 0 70 0 0 0 0 0 0 0 0 0 0 0 NP1C5 L5138 50750 13704 6540 6703 11568 7544 10919 19020 6677 6167 28381 14154 12032 27145 27441 NP1C4 14858 40922 17110 2770 3002 4273 4348 6739 9597 5664 8409 9569 8351 4117 11625 8059 NP1C3 3855 12083 4622 577 559 1695 1647 1322 2139 1789 2313 3212 1486 511 3655 2997 CMaC6 280 564 0 154 209 295 198 256 752 954 1402 524 849 223 240 333 CMaC5 911 1692 243 885 1327 1253 560 384 578 656 1051 787 425 287 1079 3730 CMaC4 70 322 0 77 140 516 99 128 0 119 771 0 212 255 120 799 CMaC3 0 0 0 0 0 0 33 0 0 0 140 0 0 32 0 133 CPaC6 210 322 324 308 279 147 165 256 231 179 70 131 495 128 180 67 CPaC5 140 161 243 115 70 221 . 99 43 173 119 280 0 71 0 60 0 CPaC4 0 0 0 0 0 0 0 0 0 119 210 131 0 32 120 67 CPaC3 0 0 0 0 0 0 0 0 58 0 0 0 0 0 0 0 MLFC6 3574 1692 4541 3885 4888 5600 10739 6270 1619 537 561 3015 6086 6064 8209 6461 MLMC6 4205 3867 3406 6424 4050 3979 2602 938 1734 954 491 3277 4388 2489 4794 1798 MLFC5 1892 1933 973 1039 1327 3095 3492 3455 1272 4054 2523 3015 4671 2011 1438 2464 MLMC5 1822 1611 1784 1308 1466 2652 4315 2900 3237 4114 8059 3670 3963 1947 1079 1865 MLuG4 1121 2094 1379 2193 1606 2726 6786 2986 5550 6260 12965 5899 4175 1787 899 2398 M0kC6 0 0 0 0 70 0 0 0 0 0 0 0 0 0 0 0 MOkJu 140 0 81 77 140 74 132 43 58 0 0 6  0 96 0 0 EBuC6 0 161 81 269 349 74 0 0 0 0 0 0 71 0 60 0 EBuC5 0 0 0 0 0 0 0 0 0 0 0 0 0 32 0 0 EBuC4 70 161 0 0 0 0 198 171 173 0 0 131 71 191 120 0 EBuC3 70 81 405 0 140 589 264 256 405 298 140 328 425 383 360 133 EBuC2 210 81 81 154 349 221 231 256 289 119 350 459 142- 64 240 ' 0 EBuCl 0 0 0 0 0 0 33 43 116 0 70 66 0 32 0 0 EE1C6 280 403 243 154 70 0 0 0 58 0 0 0 0 32 60 133 E1C5 140 0 81 154 419 74 33 85 173 0 0 0 0 96 60 200 EE1C4 631 0 162 77 559 74 198 0 0 0 0 66 0 32 419 67 E1C3 350 161 568 231 349 74 33 341 0 0 0 66 142 191 180 0 EE1C2 350 403 811 385 628 147 33 128 0 60 0 0 142 128 0 0 PMiC6 1542 725 649 1231 1885 6484 8071 8658 38850 35891 74003 3605 3609 3032 839 4196 SMiC6 1262 564 1054 1962 1327 2137 1186 1109 867 417 140 721 1628 862 479 1199 0SpC6 6448 5156 5676 8694 9845 4126 4250 3284 1908 1073 491 1835 3539 4404 6831 2531 Cruise: SG81-8 (cont'd) Station: 7829 8535 7133 6832 5931 5035 4337 4039 Depth (in meters): 188 229 340 396 390 350 310 181 CGrC6 280 242 649 539 349 74 0 0 CGrJu 421 1208 892 692 140 147 0 0 GMiC6 140 644 405 423 279 147 0 0 GMiJu 140 161 649 77 349 74 0 0 A L 0 C 6 0 161 0 308 140 295 296 299 CC0C6 0 0 0 77 70 0 0 0 CCoJu 0 0 81 38 140 0 0 0 ADiC6 70 0 0 38 0 0 0 0 ADiJu 0 0 0 0 0 0 33 0 SpBC6 140 161 243 192 0 74 0 43 ScBC6 0 0 0 0 70 0 0 0 TDiC6 0 0 0 0 0 0 0 0 IIL0C6 0 0 0 0 0 0 0 0 H L o J u 70 0 0 0 0 0 33 0 CAbC6 0 0 0 0 0 0 0 43 RAnC6 0 0 0 0 0 0 33 0 GaeC6 0 0 0 0 0 0 0 0 GaeJu 0 0 0 0 0 74 0 0 E L 0 C 6 0 0 0 0 0 0 0 85 ELoJu 0 0 0 0 0 0 0 0 3739 3138 2837 4929 5132 5538 7136 . 6727 193 185 190 151 247 197 192 150 0 0 0 0 71 160 300 0 58 0 0 0 142 223 899 266 0 0 0 0 71 64 180 0 0 0 0 0 0 0 60 0 1098 894 1962 262 71 0 0 533 0 0 0 0 0 32 60 0 0 0 0 0 71 0 0 67 0 0 0 0 0 32 60 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 58 0 0 0 0 0 0 0 0 0 210 0 0 32 0 0 58 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 60 210 0 0 • 0 0 0 0 0 70 0 0 0 0 0 58 0 0 0 0 0 0 0 0 60 0 0 0 0 0 0 0 238 491 0 0 0 0 0 0 60 280 0 71 0 0 0 Cruise: SG81-9 Station: 7829 7832 8535 Depth (in meters): 316 245 347 NP1C6 0 0 0 NP1C5 17168 24409 •42907 NP1C4 2120 6186 2924 NP1C3 71 336 711 NCrC5 0 0 0 CMaC6 283 134 158 CMaC5 1554 403 1185 CMaC4 71 0 79 CMaC3 0 , 0 0 CPaC6 141 471 237 CPaC5 71 134 237 CPaC4 0 0 0 CPaC3 0 0 0 MLFC6 10951 14861 3240 MLMC6 3744 4774 3082 MLFC5 565 672 711 MLMC5 495 336 1027 MLuC4 918 740 1185 M0kC6 0 0 0 MOkJu 141 336 79 EBuC6 0 134 0 EBuC5 0 0 0 EBuC4 0 67 0 EBuC3 141 202 237 EBuC2 0 134 79 EBuCl 0 67 0 EE1C6 353 134 316 EE1C5 283 538 158 EE1C4 283 202 0 EE1C3 424 471 316 EE1C2 353 269 553 PMiC6 424 1009 869 SMiC6 1484 2085 1185 7634 6832 6231 397 387 356 0 0 0 51849 7063 9453 8125 1144 983 2848 276 227 0 0 0 335 276 0 335 888 681 0 0 0 0 0 0 84 178 151 335 0 227 0 0 0 0 0 0 10051 3689 5521 3350 4952 4916 670' 986 529 586 769 378 921 1401 1134 0 39 0 0 158 227 84 39 0 0 0 0 0 79 303 84 395 378 251 99 0 0 20 0 84 158 303 0 118 151 0 335 151 168 178 151 1424 750 454 503 947 1059 754 2249 2042 5931 5035 4337 352 295 187 83 69 0 24144 7299 26215 1499 757 3155 83 0 451 83 0 0 167 207 526 1249 757 676 0 0 0 0 0 0 250 275 451 83 138 75 0 69 0 0 69 0 9574 7919 5483 4079 4889 2329 1249 1721 5634 250 1928 2329 1082 2135 2554 83 0 0 333 344 0 167 138 0 0 69 0 83 69 75 83 551 0 0 0 75 83 0 0 167 69 75 333 413 75 83 413 75 500 482 75 416 413 0 250 1997 7587 1998 2892 826 4039 3439 3138 185 172 190* 0 0 0 49564 25825 30401 1877 3084 5387 526 470 1033 0 0 0 826 209 590 2779 732 664 0 0 0 0 0 0 0 52 0 0 105 0 0 52 0 0 0 0 3079 1150 590 2103 1411 1992 2028 1568 3025 2478 1568 2730 2178 2562 5608 0 0 0 0 0 74 0 0 0 0 0 0 0 0 74 225 209 74 0 52 0 0 0 0 0 0 0 75 0 0 75 0 0 451 0 74 75 105 0 7134 9253 15274 676 209 590 2837 4929 5132 207 135 255 0 0 0 12277 40393 23530 3508 2372 1927 .1012 271 80 0 0 0 1079 542 482 337 407 562 0 0 0 67 0 0 67 136 80 270 0 . 80 135 0 0 135 68 0 2361 1762 6987 1417 1288 4658 8095 1830 1285 7555 813 964 5801 1288 1446 0 0 0 202 0 161 0 0 80 0 68 0 0 0 161 0 136 321 202 271 161 67 0 0 0 0 0 0 0 402 0 0 642 0 0 161 67 68 482 40878 813 1526 67 474 1446 5538 7227 6727 188 193 133 0 0 0 7906 13924 13189 228 2904 1374 98 132 550 0 0 0 163 330 275 423 396 343 0 0 0 33 0 0 0 198 206 130 0 0 65 0 ' 69 33 0 0 2050 8315 15249 2245 2640 481 716 1386 756 1269 1254 343 1464 528 618 33 66 0 65 330 206 33 0 0 33 0 0 98 198 0 390 198 137 130 0 0 ' 33 0 0 33 198 0 0 0 206 65 198 206 130 66 137 130 66 0 1074 792 206 1659 1386 687 Cruise: SG81-9 (cont'd) Stat ion: 7829 7832 8535 7634 6832 6231 5931 5035 4337 4039 3439 3138 2837 4929 5132 5538 7227 6727 Depth (in meters): '190 316 245 347 397 387 356 352 295 187 185 172 207 135 255 188 193 133 0SpC6 6147 8607 7823 6785 5583 4765 4746 5233 2178 1953 2666 1623 1282 2779 6023 4978 11020 6732 CGrC6 212 336 711 503 296 76 333 69 0 0 0 0 0 0 482 163 264 69 CGrJu 424 807 553 251 197 303 333 69 75 0 0 0 0 0 241 130 462 893 GMiC6 636 67 553 335 197 0 83 757 0 0 0 0 0 0 241 130 396 0 GMiJu 353 269 79 670 355 76 167 275 0 0 0 0 0 0 321 33 132 0 GPuC6 0 0 0 0 0 0 83 0 0 0 0 0 0 0 0 0 0 0 AL0C6 0 202 158 84 39 76 0 69 526 526 680 1033 2091 203 80 0 0 0 CC0C6 0 0 0 168 20 76 0 0 75 0 52 0 0 0 241 33 0 0 CCoJu 0 0 0 84 79 76 83 69 0 0 0 0 0 0 80 33 66 0 ADiC6 0 0 0 84 0 0 0 207 0 0 0 0 0 0 0 0 0 0 ADiJu 0 0 0 84 0 0 0 0 0 75 0 0 0 0 0 33 0 0 APaC6 0 0 0 0 20 0 0 0 0 0 0 0 0 0 0 0 0 0 SpBC6 71 67 316 0 59 151 0 • 0 0 0 0 0 0 0 0 0 0 0 ScBC6 0 0 79 84 20 76 0 69 75 0 52 0 0 0 80 0 0 0 TDiC6 0 0 0 0 0 0 0 0 75 0 0 0 67 0 0 0 0 0 CAbC6 0 0 0 0 0 0 0 0 0 0 0 74 67 0 0 0 0 0 RAnC6 0 0 79 0 79 0 83 69 0 0 0 0 67 0 0 0 0 0 ClaC6 0 0 0 0 20 0 0 0 0 0 0 0 0 0 0 0 0 0 HL0C6 0 0 0 0 39 0 0 0 0 0 0 0 0 0 0 0 0 0 HLoJu 0 0 0 84 39 76 83 0 0 0 0 0 0 0 80 0 0 0 BSaC6 0 0 0 0 20 0 0 0 0 0 0 0 0 0 0 0 0 0 BSaJu 0 0 0 0 20 0 0 0 0 0 0 0 0 0 0 0 0 0 GaeC6 0 0 0 0 20 0 0 . 0 0 0 0 0 67 0 0 0 0 0 EL0C6 0 0 0 0 0 0 0 0 0 0 0 74 607 0 0 ' 0 0 0 ELoJu 0 0 0 0 0 0 0 0 0 0 0 0 67 0 0 0 0 0 C r u i s e : SG81-10 S t a t i o n : 7829 7832 8535 7934 7634 6832 5931 5532 4636 Depth ( i n meters) : 328 270 340 391 410 388 331 328 221 NP1C6 0 0 0 0 0 0 0 56 0 NP1C5 1981 2876 9463 10880 37267 10746 10487 8053 14334 NP1C4 108 105 80 0 0 76 211 222 155 NPLC3 0 0 0 0 0 0 0 0 0 CMaC6 216 0 239 0 . 0 152 70 278 155 CMaC5 684 386 557 919 1040 533 1548 444 232 CMaC4 0 0 0 0 0 0 0 0 0 CMaC3 0 0 0 0 0 0 70 56 77 CPaC6 108 105 398 0 400 152 0 167 0 CPaC5 108 105 80 77 240 0 70 111 0 CPaC4 0 0 0 0 0 0 70 0 0 CPaC3 36 0 0 0 0 0 0 0 0 MLFC6 4646 5541 13041 2222 2959 L905 5349 8831 9763 MLMC6 3061 2104 1670 3984 2799 4801 3871 3943 2015 MLFC5 108 245 318 0 400 152 633 1722 697 MLMC5 144 386 398 153 160 76 141 389 1007 MLuC4 252 175 239 153 400 229 422 611 465 M0kC6 108 0 0 0 0 76 0 111 0 MOkJu 0 140 80 0 0 0 0 278 155 EBuC6 108 386 239 383 320 229 70 56 0 EBuC5 36 0 0 0 0 76 0 0 0 EBuC4 72 70 318 306 160 229 211 56 155 EBuC3 360 351 239 0 160 76 70 167 77 EBuC2 , o 35 0 0 0 0 70 0 0 EE1C6 396 140 159 230 80 152 141 56 0 EE1C5 108 0 159 77 80 76 352 278 77 EE1C4 144 35 159 77 80 305 141 167 77 EE1C3 180 140 159 230 ' 240 152 141 111 155 EE1C2 1044 947 1034 690 960 610 352 778 232 PMiC6 936 1157 875 383 480 152 3097 2721 1937 SMiC6 1116 1052 1113 1916 1200 2134 3941 1277 1162 OSpC6 7131 6102 6521 5900 5278 6173 4645 3666 3719 CGrC6 144 140 239 230 480 305 282 111 0 4039 3739 3439 2837 4929 5335 5538 7627 7227 181 203 169 202 141 292 184 181 217 0 77 0 0 0 0 0 0 0 4178 12656 12225 11886 4372 29194 2292 7288 5310 159 384 310 457 207 307 0 69 72 40 0 0 0 59 0 0 0 0 199 384 248 152 384 307 0 34 179 478 384 683 229 30 1076 239 172 215 0 0 0 0 0 0 34 0 72 40 0 372 0 30 0 34 103 0 119 307 62 76 59 77 68 69 72 119 77 0 152 30 0 68 0 36 0 77 0 76 0 0 34 0 0 40 0 62 0 0 0 0 0 0 6884 3528 5026 1448 3929 5992 3079 9454 5668 1313 537 496 990 561 1613 1745 3644 3731 517 1918 993 1295 1182 461 308 241 144 398 920 1241 1448 1034 230 205 344 72 796 1304 1800 1829 1477 691 513 138 718 0 0 0 0 0 154 0 0 0 80 0 0 0 59 230 137 138 36 119 0 62 0 0 154 34 206 0 0 0 0 76 0 0 0 0 0 239 ' 77 186 0 118 384 171 103 108 0 153 62 152 30 230 0 172 395 0 0 0 0 30 0 0 103 72 40 0 0 0 0 77 68 206 36 40 0 186 0 89 77 68 378 215 40 77 0 0 30 77 205 481 72 159 77 62 0 59 230 239 138 179 40 153 62 0 89 768 239 309 287 4218 23318 21843 36648 2925 3227 1539 550 969 1552 844 186 229 679 1537 1608 1409 1543 5730 3758 3351 5029 3456 4379 6226 4813 4628 119 0 0 0 0 0 239 206 215 Cruise: SG81-10 (cont'd) Station: 7829 7832 8535 7934 7634 Depth (in meters): 328 270 340 391 410 CGrJu 144 105 318 153 240 G M i C 6 396 105 398 460 240 GMiJu 612 456 716 460 320 GPuC6 72 0 0 0 0 A L 0 C 6 180 70 318 77 80 CC0C6 36 0 0 . 77 80 CCoJu 108 70 . 0 77 0 A D i C 6 0 0 80 0 0 ADiJu 0 0 80 0 0 APaC6 0 0 0 0 0 SpBC6 72 70 239 230 400 ScBC6 108 35 159 230 240 TD i C 6 0 0 0 0 0 HL0C6 0 0 0 0 0 HLoJu 36 0 159 0 80 CAbC6 0 0 159 0 160 RAnC6 0 0 . 0 0 0 E L 0 C 6 0 0 0 0 0 ELoJu 0 0 0 0 0 BSaC6 0 0 0 0 0 AMuC6 0 0 0 0 0 >832 5931 5532 4636 4039 3739 388 331 328 221 181 203 152 282 i l l 0 279 77 0 141 167 387 40 0 152 70 222 465 80 0 0 0 0 0 0 0 76 0 611 465 279 844 0 0 56 0 0 0 0 70 111 0 40 0 0 0 0 0 0 0 0 0 0 0 40 0 76 0 0 0 0 0 76 0 0 77 0 0 0 0 167 77 0 0 0 0 0 0 0 77 76 0 0 0 0 0 229 70 56 0 0 0 0 0 56 0 119 153 76 70 56 0 0 0 0 0 0 0 0 0 0 0 0 0 0 77 0 0 0 0 0 0 0 0 56 0 0 0 3439 2837 4929 5335 5538 7627 7227 169 202 141 292 184 181 217 t 0 0 0 307 239 275 323 0 0 0 0 137 69 251 0 0 0 461 . 137 138 574 0 0 0 0 0 0 0 1303 533 266 538 137 206 395 0 0 0 77 34 0 0 0 0 0 0 0 0 36 0 0 30 0 0 0 72 0 0 30 77 0 34 0 0 0 0 0 0 0 0 0 0 0 0 34 34 0 0 0 0 0 34 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 154 0 0 0 0 0 0 230 0 0 0 0 0 30 0 0 0 0 0 76 0 0 0 0 0 62 0 0 77 34 0 0 0 0 0 0 34 0 0 0 0 0 0 0 0 0 Cruise : SG81-U Stat ion: 7829 7832 8535 7634 6832 5931 5532 5035 Depth (in meters): 320 248 326 408 392 347 319 285 NP1C6 0 38 0 0 79 0 0 0 NP1C5 6090 3851 1034.1 10150 9996 20338 14263 9116 NP1C4 0 0 73 41 79 0 0 0 NCrC6 0 0 0 0 0 77 0 0 CMaC6 145 381 73 247 157 77 228 656 CMaC5 1595 343 583 1726 1417 2707 835 590 CMaC4 145 114 73 164 79 77 76 0 CMaC3 0 0 0 82 0 77 0 0 CPaC6 0 38 0 164 79 155 76 328 CPaC5 218 76 218 205 0 309 303 131 CPaC4 0 191 218 0 0 0 0 66 CPaC3 73 38 218 82 79 0 0 0 MLFC6 6018 3699 3641 3863' 787 1315 2428 7149 MLMC6 3335 4576 1238 2835 2597 2243 3490 1508 MLFC5 218 114 655 123 551 232 228 984 MLMC5 290 153 510 205 394 309 303 787 MLuC4 145 343 655 616 157 541 759 525 MOkC6 73 0 0 41 79 0 0 0 MOkJu 0 76 73 0 0 0 76 131 EBuC6 145 114 73 164 157 155 76 0 EBuC5 0 153 291 863 394 155 ' 0 394 EBuC4 0 38 0 41 0 0 0 0 EBuC3 0 38 0 41 0 0 0 0 EBuC2 0 0 0 0 . 0 0 228 197 EBuCl 73 76 73 0 315 309 303 197 EE1C6 73 38 146 205 472 77 303 66 EE1C5 435 191 364 247 79 0 0 197 EE1C4 218 191 73 164 315 155 152 131 EE1C3 725 534 0 452 787 619 531 328 EE1C2 870 572 437 657 1417 619 379 394 PMiC6 1740 1487 655 1151 1495 1547 2504 12527 SMiC6 3045 4194 1165 1931 2991 3480 4248 9116 OSpC6 3625 4576 2694 863 5746 7269 9787 12133 4337 3739 3138 2638 4929 5132 5538 6727 7136 183 196 183 270 120 247 190 178 196 0 0 0 0 0 0 0 0 0 3924 2594 2664 2135 5127 5001 2594 4659 3444 0 62 0 0 30 0 0 0 0 0 0 118 76 0 0 0 0 0 129 618 592 839 90 286 329 436 270 322 309 237 457 271 286 219 201 180 0 247 118 229 0 71 73 67 60 0 0 118 152 0 71 0 34 30 386 185 770 915 181 143 183 168 270 193 494 592 1296 0 214 402 101 90 64 0 296 457 0 0 183 67 180 0 247 533 610 30 71 37 0 60 13188 5929 592 2058 1930 5358 2777 5362 1587 450 309 237 457 573 1643 1096 3150 1467 515 1359 59 229. 422 643 402 704 569 772 1050 118 534 543 500 475 436 359 515 556 770 457 814 786 292 670 689 64 0 0 0 0 0 0 0 0 0 62 0 0 30 0 0 34 30 64 0 59 0 0 71 37 168 150 129 62 355 0 211 143 329 335 329 0 0 0 0 0 71 0 34 0 0 62 0 0 0 0 0 34 0 0 247 178 229 0 " 71 183 0 0 64 1173 474 381 0 429 146 ,"0 0 0 0 0 0 0 286 110 168 60 64 185 0 0 121 214 219 268 180 64 124 0 0 121 214 219 201 120 257 124 0 76 211 786 365 134 419 129 0 0 0 30 286 146 268 629 15311 40329 59018140351 5730 2358 4238 3217 2036 1866 1297 178 76 1749 6287 2229 2715 1258 8427 8832 6512 3354 8294 9359 10340 3385 3504 Cruise: SG81-11 (cont'd) Stat ion: 7829 7832 8535 7634 6832 5931 5532 5035 4337 3739 3138 2638 4929 5132 5538 6727 7136 Depth (in meters): 320 248 326 408 .392 347 319 285 183 196 183 270 120 247 190 178 196 CGrC6 290 153 0 288 157 0 303 197 0 0 0 0 0 143 146 101 90 CGrJu 508 343 218 123 157 387 76 525 257 0 0 0 30 286 183 168 120 GMiC6 870 229 874 205 157 0 0 131 64 0 0 0 0 143 0 101 30 GMiJu 725 534 364 1027 866 387 607 459 64 0 0 0 0 929 37 168 60 GPuC6 0 0 0 0 0 0 0 0 0 0 59 0 0 0 0 0 0 AL0C6 0 38 218 41 79 0 0 66 257 371 1066 2287 30 0 37 34 90 CC0C6 0 0 73 0 79 77 228 66 0 0 0 0 0 71 0 0 0 CCoJu 0 114 73 82 79 0 0 131 64 0 0 0 0 0 0 101 30 ADiC6 0 38 0 0 0 0 0 66 0 124 0 0 0 0 0 34 0 ADiJu 0 0 0 0 0 77 0 0 0 62 0 0 0 0 0 34 0 SpBC6 73 38 0 82 157 77 0 0 64 0 0 0 0 0 0 0 0 ScBC6 73 0 73 82 236 77 76 0 0 0 0 0 0 0 0 0 30 TDiC6 0 ' 0 0 0 0 0 0 66 0 0 0 229 0 0 0 0 0 HL0C6 73 0 0 0 0 0 0 0 0 0 0 76 0 0 0 0 30 HLoJu 145 0 73 0 0 0 0 0 0 0 0 0 0 71 0 0 30 CAbC6 0 0 0 41 79 0 0 0 0 0 59 152 0 0 0 0 30 RAnC6 0 0 0 82 79- 0 0 0 0 0 0 0 0 0 0 0 0 EL0C6 0 38 0 0 0 0 0 0 0 62 710 1144 0 0 0 0 0 ELoJu 73 76 0 0 0 0 0 0 0 0 59 76 0 0 0 0 0 BSaJu 73 0 0 0 0 0 0 0 0 0 0 0 0 0 0 34 30 GaeC6 0 0 0 0 0 0 - 0 0 0 0 0 76 0 0 0 0 0 GaeJu 0 0 0 0 79 0 0 0 0 62 59 0 0 0 0 0 0 MSpC6 0 0 0 0 79 77 0 0 0 0 0 0 0 0 0 0 0 Appendix B: Processed predator wet weights (values in grams) Cruise: SG81-1 Station: 7829 7832 8535 1.738 L.546 1.960 7934 7634 6530 2.380 2.765 1.410 5931 5035 4636 4337 4039 3439 2837 7136 7627 7227 6727 4929 5132 5538 1.684 1.646 0.878 0.615 0.228 0.262 0.342 0.879 0.966 1.196 0.148 0.660 1.116 1.208 Cruise: SG81-2 Station: 7829 7832 8535 1.646 1.485 1.444 7934 6832 5931 1.840 1.917 1.317 5035 4636 4337 3439 2837 4929 5132 5538 7136 6727 7227 1.601 0.777 0.737 0.262 0.192 0.272 0.627 0.745 0.678 1.346 0.741 Cruise: SG81-3 Stat ion: 7829 7832 8535 1.406 1.077 2.828 7934 7634 6231 1.543 0.870 1.535 5532 4636 4039 3439 3138 2837 4929 5132 5538 7227 6727 1.335 0.657 0.661 0.068 0.069 0.188 0.795 1.227 0.735 0.492 1.148 Cruise: SG81-5 Stat ion: 7829 7832 8535 1.716 2.387 2.739 7634 6832 5931 2.072 1.162 0.948 4636 4039 3439 3138 2638 4929 5538 6727 7136 1.673 1.122 1.350 0.554 0.757 0.795 0.701 1.181 0.987 -P» cn Cruise: SG81-7 Stat ion: 7829 7832 8535 0.747 0.964 1.588 7634 6832 5931 1.883 2.422 1.574 5035 4636 4337 3739 3138 2837 4929 5132 5538 7136 6727 1.659 2.497 0.394 0.695 0.414 0.712 1.109 1.849 0.847 1.933 1.453 Cru ise : SG81-8 S t a t i o n : 7829 8535 7133 1.891 2.527 2.560 6832 5931 5035 1.941 2.077 1.581 4337 4039 3739 3138 2837 4929 5132 5538 7136 6727 1.633 1.024 0.425 0.368 0.353 0.731 1.140 1.342 1.471 0.940 Cruise: SG81-9 Station: 7829 7832 8535 1.431 1.574 1.416 7634 6832 6231 1.717 1.586 1.818 5931 5035 4337 4039 3439 3138 2837 4929 5132 5538 7227 6727 1.509 1.177 1.345 1.404 0.981 0.358 0.992 0.528 1.392 0.892 2.183 1.092 A p p e n d i x B ( c o n t ' d ) Cru ise : SG81-10 S t a t i o n : 7829 7832 8535 7934 7634 6832 5931 5532 4636 4039 3739 3439 2837 4929 5335 5538 7627 7227 1.531'1.102 1.551 1.812 1.354 1.244 1.328 0.945 1.756 1.033 0.586 1.046 0.973 1.708 1.133 0.628 0.686 0.830 Cru ise : SG81-11 Stat i o n : 7829 7832 8535 7634 6832 5931 5532 5035 4337 3739 3138 2638 4929 5132 5538 6727 7136 0.982 1.682 1.534 1.604 1.068 1.538 1.089 1.133 1.327 1.627 1.290 2.375 1.183 1.208 1.149 0.840 0.618 Appendix C: 351 um vs. 200 um mesh Cows, July 1982 (values in n/m2) Location: 49 27.8 N 124 03.9 W Station Depth: 395 m Tow 351 um 200um Species Stage 1 3 5 2 4 Neocalanus plumchrus C5 37216 39973 41111 50304 57890 Calanus marshallae C6 0 0 109 0 210 C5 2828 4668 2952 5786 2097 pacificus C6 792 718 2077 590 210 C5 2715 1197 3280 2126 4195 C4 566 957 219 472 1678 C3 226 479 109 0 419 Euca Lanus C6 113 0 0 236 0 C5 339 359 328 354 419 C4 113 0 0 0 0 C3 0 0 109 0 0 C2 0 120 109 0 0 Metridia fC6 1131 957 1640 1299 419 mC6 1697 1795 2624 2126 2307 fC5 905 1077 656 1063 2307 mC5 1471 1316 1531 1535 2727 C4 5430 6582 9512 5314 6922 C3 1584 1556 1640 8266 9229 okhontens is Euchaeta elongata 113 0 219 118 0 C6 113 0 109 354 0 C5 0 0 0 0 419 C4 339 479 765 1417 839 C3 226 . 598 875 354 210 C2 679 359 328 472 210 Cl 113 0 219 0 0 48771 0 6129 1043 1434 391 0 130 782 0 0 0 1695 3390 2738 1956 7172 8484 0 0 130 130 522 261 0 Appendix C (cont'd) Tow Species Stage Pseudqcalanus minutus C6 C5 Scolecithricella C6 C5 minor Oithona spinirostris (>lmm Our idius grac i L is Gaid ius minutus Aet ideus divergens  Spinocalanus brevicaudatus Aeartla longiremis Candac ia columbiae Scaphocalanus brevicornis Racovitzanus antarct icus 5882 10294 1131 : 1018 )4638* 1244 2262 113 452 339 0 113 113 351 Um 3 . 3710 10652 1436 0 3351 718 1676 0 120 239 239 0 0 4592 8856 2296 219 2405 875 3717 219 765 109 0 109 0 6731 10392 2126 354 9447 709 1771 0 236 118 0 0 0 200 pm 4 6922 13634 1678 210 8390 629 2517 210 1049 0 419 0 0 6 4173 15648 2738 522 6911 782 3130 0 652 130 130 0 0 Appendix C (cont'd) Tow Species Stage Tharyb is fultoni Tortanus d iscaudatus 351 um 3 0 0 0 0 0 0 200 um 4 0 0 130 130 Tow Summary Station Depth: Wire Angle: Tow Time: Vol. Filtered: Fract ion sub.: 388 m 395 m 30 15 10 min 7 min 304 m3 288 nv 1/32 1/32 402 m 395 m 40 20 6 min 8 min 315 m3 292 m3 1/32 1/32 395 m 402 m 30 25 8 min 7 min 328 m3 264 m3 1/64 1/32 * = size limitation not considered. Appendix D: Predator wet weight and dry weight data Cruise; Wt.Wt. Dry Wt. Station & Tin & Tin 81--7 4636 18 2379 3.4052 81--7 7634 12.4147 2.8752 81--I 8535 16 .0550 3.2222 81--1 6530 13 .9291 2.9865 81--1 2837 3 .6933 1.7529 81--9 7227 15 .3782 2.8949 81--1 7634 20 8171 3.9280 81--3 6231 12 .6178 2.7293 81--7 4929 7 .9506 2.2051 81--1 5132 9 .2696 2.2519 8110 7934 13 .2721 2.8203 81--9 3138 3 7499 1.7098 81--7 4337 2 .9962 1.5726 81--9 2837 8 .6467 2.3113 81 -9 7832 11 .3133 2.5010 81 -9 6231 13 .6148 3.0218 81 -2 5931 ,9 .5529 2.5204 81 -7 ,6832 20.3273 3.9347 81 -2 4929 3 .8981 1.7139 81 -1 ,5538 10 .6058 2.4594 81 -3 ,8535 12 .5964 2.5930 81 -3 ,3439 2 .0054 1.4342 81 -3 ,7832 8 .8686 2.1746 Tin Wet Dry Weight Weight 1 .3743 16.8636 2.0309 1 .3774 11.0373 1.4978 1 .3730 14.6820 L.8492 1 .3761 12.5530 1.6104 1 .3774 2.3159 0.3755 1 .3715 14.0067 1.5234 1 .3759 19.4412 2.5521 1 .3795 11.2383 1.3498 1 .3855 6.5651 0.8196 1 .3687 7.9009 0.8832 1 .3804 11.8917 1.4399 I .3780 2.3719 0.3318 I .3814 1.6145 0.1912 1 .3755 7.2712 0.9358 1 .3756 9.9377 1.1254 1 .3707 12.2441 1.6511 1 .3748 8.1781 1.1456 1 .3809 18.9464 2.5538 1 .3745 2.5236 0.3394 1 .3728 9.2330 1.0866 1 .3775 11.2189 1.2155 1 .3728 0.6326 0.0614 1 .3734 7.4952 0.8012 Appendix E: Copepod wet weight and dry weight data (values in mg) Spec. Cruise No. T i n Wt.Wt. Dry Wt. Wt.Wt. Dry Wt. Animal & T i n & Tin /Animal /Animal Southern S t r a i t PMiC6 81-9 200 70.1 96.9 72.7 0.134 0.013 CMaC6 50 69.7 152.7 79.2 1.660 0.190 AL0C6 100 69.5 76.1 70.1 0.066 0.006 Central S t r a i t NP1C6 81-1 25 69.1 153.3 78.0 3.368 0.356 69.4 157.6 79.3 3.528 0.396 69.9 155.9 78.5 3.440 0.344 69.4 157.2 79.7 3.512 0.412 70.2 157.5 79.5 3.492 0.372 81-3 70.0 143.8 74.7 2.952 0.188 69.6 140.0 73.4 2.816 0.152 68.8 154.1 76.0 3.412 0.288 69.7 153.6 74.0 3.356 0.172 69.8 122.1 71.4 2.092 0.064 81-5 69.9 145.7 72.3 3.032 0.096 69.6 151.0 72.9 3.256 0.132 69.7 129.6 71.3 2.396 0.064 69.4 147.8 72.2 3.136 0.112 . 69.8 149.1 74.0 3.172 0.168 ro A p p e n d i x E ( c o n t ' d ) Spec. Cruise No. Tin Animal Central Strait NPLC5 81-7 25 69.9 69.6 69.4 69.3 70.1 81-9 70.1 70.0 69.3 69.7 69.5 81-11 69.4 70.1 69.7 69.3 69.6 NP1C4 81-7 100 69.9 NP1C3 81-7 100 69.7 CMaC6 81-1 25 69.6 81-2 69.7 70.0 81-3 69.8 81-5 69.6 Wt.Wt. Dry Wt. Wt.Wt. Dry Wt. & T i n & T i n /Animal /Animal 130.6 75.3 2.428 0.216 134.4 75.3 2.592 0.228 135.4 74.2 2.640 0.192 140.0 74.9 2.828 0.224 140.5 76.0 2.816 0.236 158.2 79.9 3.524 0.392 152.8 79.8 3.312 0.392 155.7 78.9 3.456 0.384 165.0 79.6 3.812 0.396 153.0 78.2 3.340 0.348 164.8 85.5 3.816 0.644 167.5 87.9 3.896 0.712 171.4 88.5 4.068 0.752 165.9 88.4 3.864 0.764 158.5 85.8 3.556 0.648 172.8 76.4 1.029 0.065 93.8 71.4 0.241 0.017 105.8 74.7 1.448 0.204 103.8 74.3 1.364 0.184 105.7 74.6 1.428 0.184 103.7 74.1 1.356 0.172 105.5 73.7 1.436 0.164 Appendix E (cont *d) Spec. Cruise No. Tin Wt.Wt. Dry Wt. Wt.Wt. Dry Wt. Animal & Tin & Tin /Animal /Animal Central Strait CMaC5 81-1 25 69.5 95.6 73.9 1.004 0.176 69.3 92.4 73.4 0.924 0.164 81-9 69.0 96.7 75.3 1.108 0.252 69.2 96.5 75.3 1.092 0.244 69.9 96.4 75.6 1.060 0.228 EBuC6 81-3 25 70.1 254.6 76.3 7.380 0.248 70.0 261.4 75.8 7.656 0.232 81-5 69.5 264.3 77.2 7.792 0.308 69.9 258.9 77.5 7.560 0.304 '70.1 255.7 77.5 7.424 0.296 EBuC5 81-1 25 69.5 138.0 74.3 2.740 0.192 69.5 144.9 74.7 3.016 0.208 81-2 70.6 153.8 75.4 3.328 0.192 69.5 153.1 74.7 3.344 0.208 81-3 70.0 155.2 74.5 3.408 0.180 CPaC6 81-5 50 .70.1 107.7 75.1 0.752 0.100 CPaC5 81-1 25 69.4 82.0 71.8 0.504 0.096 70.1 84.2 72.8 0.564 0.108 81-2 69.8 84.5 73.0 0.588 0.128 69.8 84.6 72.8 0.592 0.120 81-3 69.5 83.4 72.2 0.556 0.108 MLFC6 81-1 50 69.4 103.4 72.6 0.680 0.064 81-2 69.2 100.3 72.4 0.622 0.064 81-3 69.7 103.9 73.0 0.684 0.066 81-7 69.5 100.8 72.7 0.626 0.064 69.9 104.8 73.0 0.698 0.062 Appendix E (cont'd) Spec. Cruise No. Tin Wt.Wt. Dry Wt. Wt.Wt. Dry Wt. Animal & Tin & Tin /Animal /Animal Central Strait MLMC6 81-3 100 69.8 85.8 71.5 0.160 0.017 MLFC5 81-3 100 70.2 85.4 71.4 0.152 0.012 MLMC5 81-2/3 MLuC4 81-2/3 PMiC6 81-1 81-2 81-3 81-7 SMiC6 81-1/2 0SpC6 81-1/2 /3/3/7 EE1C6 81-1 81-2 81-3 81-5 81-7 EE1C5 81-1 81-2 81-3 81-5 81-7 100 69.3 200 69.4 100 69.4 69.2 70.2 69.1 70.2 200 70.0 750 69.2 25 69.7 69.4 70.3 70.5 69.9 25 69.8 69.4 69.5 69.7 69.8 82.1 70.6 76.6 70.2 82.9 71.4 81.1 70.9 75.2 70.6 74.4 69.6 81.1 71.6 95.9 73.4 85.4 70.6 221.0 89.7 217.9 91.8 199.6 91.9 198.2 89.7 212.5 91.4 149.4 80.6 156.0 86.3 151.6 84.6 160.0 84.6 154.5 81.7 0.128 0.013 0.036 0.004 0.135 0.020 0.119 0.017 ^  0.050 0.004 0.053 0.005 0.109 0.014 0.1295 0l017 0.0216 0.00187 6.052 0.800 5.940 0.896 5.172 0.864 5.108 0.768 5.704 0.860 3.184 0.432 3.464 0.676 3.284 0.604 3.612 0.596 3.388 0.476 GMiC6 81-1/2 50 69.2 149.9 76.5 1.614 0.146 Appendix F: Predator biomass error estimation Cruise; Station 1 81-1;4929 3.2 81-2;5931 8.3 81-7;6832 17.9 Tri a l 2 3 4 5 3.0 3.1 3.2 2.7 8.2 7.7 8.4 8.1 17.6 18.2 17.8 17.9 

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