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Aggregation and the energetics of the barnacle Balanus glandula Darwin Wu, Shiu-sun 1977

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AGGREGATION AND THE ENERGETICS OF THE BARNACLE B.Sc, The Chinese University of Hong Kong, 1971 B.Sc. Spec. Hon., The University of Hong Kong, 1972 M. P h i l . , The University of Hong Kong, 1974 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENT FOR THE DEGREE OF DOCTOR OF PHILOSPHY i n THE FACULTY OF GRADUATE STUDIES DEPARTMENT OF ZOOLOGY We accept this thesis as conforming to the regaired standard THE UNIVERSITY OF BRITISH COLUMBIA B<tlanus glandula DARWIN SHIU-SUN,WU December, 1977 Shiu-sun Wu, 1977 In present ing th is thes is in p a r t i a l fu l f i lment of the reguirements for an advanced degree at the Univers i ty of B r i t i s h Columbia, I agree that the L ibrary shal l make it f ree l y a v a i l a b l e for reference and study. I fur ther agree that permission for extensive copying of th i s thes is for scho la r l y purposes may be granted by the Head of my Department or by his representat ives . It is understood that copying or p u b l i c a t i o n of th i s thes is fo r f i n a n c i a l gain sha l l not be allowed without my wr i t ten permiss ion. 6 Department of 3 £ The Univers i ty of B r i t i s h Columbia 2075 Wesbrook Place Vancouver, Canada V6T 1W5 i i 1 B S T B A G T Barnacles ( Balanus ^ l a n d u l a ) were allowed to s e t t l e i n 5 pre-determined p a t t e r n s with an i n c r e a s i n g degree o f aggregation { i . e. i s o l a t e d i n d i v i d u a l s , p a i r e d i n d i v i d u a l s , 9 i n d i v i d u a l s i n a group, 81 i n d i v i d u a l s i n a group and 'crowded* i n d i v i d u a l s with number > 12295 ra-2). The energy budget :-C = P.BT. + P.E. * H.AQ. * R.AER. + M + F Where :-C = Consumption P.BT. ..= Pr o d u c t i o n of body t i s s u e E.E. = Production o f egg R.AQ. = Aquatic r e s p i r a t i o n B.AER. = A e r i a l r e s p i r a t i o n H = Molting F = F a e c a l p r o d u c t i o n was c o n s t r u c t e d f o r b a r n a c l e i n d i v i d u a l s f o r the f i r s t year settlement of each of the above 5 p a t t e r n s . A l l the budget items on the r i g h t hand s i d e of the equation were measured and i i i consumption was found by t h e i r summation. Laboratory f e e d i n g experiments showed that consumption values d e r i v e d frcm the summation method balanced c l o s e l y with those obtained from feeding experiments. J LL a i a n d u l a had a very high a s s i m i l a t i o n e f f i c i e n c y ( 91.8 to 99.4 % ) but a low gross production e f f i c i e n c y ( 22.8 t o 26.4 % ) and net production e f f i c i e n c y { 24.7 t o 28.6 % ).. A l a r g e p r o p o r t i o n cf energy i n t a k e { 64.8 to 67.7 J ) was l o s t i n r e s p i r a t i o n . The second most important budget item was egg production ( 12.3 to 15.3 % ) ; f o l l o w e d i n decreasing order by :- s h e l l production ( 6.1 to 7.2 % ) > body t i s s u e p r o d u c t i o n ( 3.9 to 4.6 % ) > molting { 1.1 to 2.3 ? ). Aggregation s i g n i f i c a n t l y reduces the consumption, a s s i m i l a t i o n , production , as w e l l as the energy standing crop of body t i s s u e , egg and s h e l l o f an i n d i v i d u a l barnacle. Among these, egg pro d u c t i o n appears to be most s e n s i t i v e to crowding. When comparing the egg : body t i s s u e r a t i o and s h e l l : t i s s u e r a t i o between crowded and uncrowded i n d i v i d u a l b a r n a c l e s , a decrease i n degree of crowding was a s s o c i a t e d with : (1) an i n c r e a s e i n the egg : body t i s s u e r a t i o and {2) a decrease i n s h e l l : t i s s u e r a t i o of the i n d i v i d u a l s . T h i s appears to suggest a d i f f e r e n c e i n energy p a r t i t i o n i n the b a r n a c l e s with d i f f e r e n t degrees of crowding. Uncrowded i n d i v i d u a l s a c q u i r i n g an adequate amount o f energy , could a f f o r d t o channel more energy i n t o egg i v output and hence produce more progeny. On the other hand, crowded i n d i v i d u a l s a c q u i r e a H a l t e d amount cf energy and apparently conserve a l a r g e r p r o p o r t i o n of t h e i r energy i n (1) b u i l d i n g body t i s s u e r a t h e r than egg and (2) b u i l d i n g a t a l l e r s h e l l so as t o i n c r e a s e the general feeding area of the p o p u l a t i o n . F i n a l l y , the e n e r g e t i c s of a n a t u r a l b a r n a c l e ( B. g l a n d u l a ) p o p u l a t i o n was s t u d i e d . The consumption, energy flow { a s s i m i l a t i o n ) , p r o d u c t i o n and m o r t a l i t y were found to be :- 6844.9, 6666.6 , 2897. 1 and 2518.1 Kcal w~ z yr-»> r e s p e c t i v e l y . The energy flow and production values are among the highest when compared with t h a t of other animal p o p u l a t i o n s and t h e r e f o r e s t r o n g l y suggest the f u n c t i o n a l importance of B± g l a n d u l a i n the l i t t o r a l system. The young age groups ( < 1 year s e t t l e m e n t s ) were most important i n c o n t r i b u t i n g to the p o p u l a t i o n energy flow and production . V TAELE OF CONTENTS MSTRfiCT .\ .... i i TJBLE OF CONTENTS V LIST OF TABLES .............. ............ ...... . . . . . . . . v i i LIST OF FIGURES AND ILLUSTRATIONS v i i i JCJNOWLETDGEJIENTS . . . . v . . . . ........................... . . X (1) INTRODUCTION ............................. ..... .. . 1 (2) T E J R H I N 2 L O G Y 11 (3) MATERIALS AND METHODS ............. 14 3.1 The Experimental P o p u l a t i o n s 3.2 Determination Of Dry weight, Ash-free Height And C a l o r i f i c Values 3.3 Production { P.BT., P.E. AND?.s.) 3.4 R e s p i r a t i o n { R.AER. AND B.AQ. ) 3.4.1 A e r i a l R e s p i r a t i o n 3.4.2 Aguatic R e s p i r a t i o n 3.5 F a e c a l P r o d u c t i o n And Mo l t i n g Rate (M And F) v i 3.6 Laboratory Energy Eudgets 3.7 P o p u l a t i o n Dynamics And Energy Flow Of The N a t u r a l Barnacle P o p u l a t i o n 3.8 E s t i m a t i o n Of E r r o r Involved In Using Long Term Mean Temperatures In C a l c u l a t i n g R e s p i r a t i o n E n e r g e t i c Cost RESULTS . . .... 33 4.1 C a l o r i f i c Values And Energy Standing Crop 4.2 Production 4.3 R e s p i r a t i o n 4.4 F a e c a l P r o d u c t i o n And M o l t i n g Rate 4.5 Annual Energy Budgets 4.6 Laboratory Energy Budgets 4.7 P o p u l a t i o n Dynamics And Energy Flow Of The N a t u r a l Barnacle Population 4.8 E s t i m a t i o n Of E r r o r Involved In Using Long Term Mean Temperatures In C a l c u l a t i n g R e s p i r a t i o n E n e r g e t i c Cost (5) DISCUSSION . . .. . .. ....... . 95 (6) l I 2 I J ! 2 2 ! i CITED 110 (7) APPENDICES 120 v i i LIST OF TABLES Table 1. Proximate a n a l y s i s and c a l o r i f i c value of b a r n a c l e ( B_. qlandula ) s h e l l Table 2. Annual energy budgets f o r i n d i v i d u a l b a r n a c l e s < B. glandula ) from the v a r i o u s aggregation p a t t e r n s over the p e r i o d May, 1975 to A p r i l , 1976. Data i n c a l o r i e s and the percentages of energy c h a n e l i e d t o each items are shewn i n b r a c k e t s by assuming that consumption eguals 100 % (For meanings of a b b r e v i a t i o n s , see Terminology i n text) Table 3. Laboratory energy budgets f o r t h r e e groups cf B._ 2l3£JalS (n=40), with independent measurement f o r each budget item f i n c a l o r i e s ) . * Cs = Consumption, d e r i v e d from summation method, * C F = Consumption, d e r i v e d from f e e d i n g experiments. (For meanings of other a b b r e v i a t i o n s , see Terminology i n t e s t ) Table M. Annual energy budgets f o r each i n d i v i d u a l s i z e c l a s s of b a r n a c l e s on the experimental shore over the p e r i o d May, 1976 to A p r i l , 1977. Data expressed i n K c a l 0.08m - 2 y e a r - 1 . Consumption was found by the summation method. The percentages of energy cha n n e l l e d t o each item were shown i n brackets by assuming that consumption eguals 100 % Table 5. The annual energy budget f o r t h e b a r n a c l e p o p u l a t i o n on the experimental shore over the p e r i o d Way, 1976 to A p r i l , 1977. Data expressed i n K c a l m~2 y e a r - 1 . Consumption was found by the summation method. The r e l a t i v e percentages of energy c h a n e l i e d to each budget iterm were shown i n brackets , by assuming t h a t consumption eguals 100 % Table 6. The annual m o r t a l i t y l o s s f o r each i n d i v i d u a l b a r n a c l e s i z e c l a s s on the e x p e r i m e n t a l shore over the p e r i o d May, 1976 t o A p r i l , 1977 . D a t a expressed i n K c a l 0.08 m - 2 y e a r - 1 Table 7. Comparison between the use of a c t u a l f l u c t u a t i n g a i r / water temperatures and *long term' mean a i r / water temperatures i n c a l c u l a t i n g r e s p i r a t i o n e n e r g e t i c c o s t f o r two h y p o t h e t i c a l b a r n a c l e i n d i v i d u a l s v i i i LISl Q.1 IIGJIilJI MP IliilSTeATIONS F i g . 1 The v a r i o u s experimental aggregation p a t t e r n s on the s e t t l i n g p l a t e s :-(a) ' I s o l a t e d » (b) 1 P a i r • (c) * 9 ' <d) * 81 ' (e) ' Crowded * F i g . 2. The experimental rocky shore near the P a c i f i c Environment I n s t i t u t e , West Vancouver, B r i t i s h Columbia F i g . 3. Seasonal changes i n the c a l o r i f i c values of v a r i o u s t i s s u e s of Bj_ g l a n d u l a , from June, 1975 to May, 1976 F i g . 4. The monthly mean energy s t a n d i n g crop of body t i s s u e of i n d i v i d u a l b a r n a c l e s ( gl a n d u l a ) from the v a r i o u s aggregation p a t t e r n s F i g . -5* The monthly mean energy s t a n d i n g crop of eggs of i n d i v i d u a l b a r n a c l e s ( B^ . glandula ) from the v a r i o u s aggregation p a t t e r n s F i g . 6. The monthly mean energy s t a n d i n g crop of s h e l l s of i n d i v i d u a l b a rnacles ( jB A glandula ) fron> the v a r i o u s aggregation p a t t e r n s F i g . 7. The monthly mean t o t a l energy s t a n d i n g crop o f i n d i v i d u a l b a r n a c l e s { JL«. glandula ) from the va r i o u s aggregation p a t t e r n s F i g . 8. D i f f e r e n c e i n energy p a r t i t i o n between i n d i v i d u a l s ( B. glandula ) from the v a r i o u s aggregation p a t t e r n s . Egg : Body t i s s u e r a t i o F i g . 9. D i f f e r e n c e i n energy p a r t i t i o n between i n d i v i d u a l s | JB. g l a n d u l a ) from the v a r i o u s aggregation p a t t e r n s . S h e l l : T i s s u e r a t i o F i g . 10. The monthly mean production o f body t i s s u e c f i n d i v i d u a l b a r n a c l e s ( B^ glandula ) from the v a r i o u s aggregation p a t t e r n s F i g . 11. The monthly mean production of egg of i n d i v i d u a l b a r n a c l e s ( B*_ g l a n d u l a ) from the v a r i o u s aggregation p a t t e r n s F i g . 12. The monthly mean pr o d u c t i o n of s h e l l of i n d i v i d u a l barnacles ( 13. glandula ) from the v a r i o u s aggregation p a t t e r n s ix Fig. 13. 02 consumption rate of egg masses, ovarian tissue and naupliar embryos cf B t glajQjula under dif f e r e n t temperatures. Fig. 14. A e r i a l 0X consumption rate of body tissue of B. glandula in r e l a t i o n to body weight and temperature Fig. 15. Aguatic Oj. consumption rate of body tissue of B. gland.ula in r e l a t i o n to body weight and temperature Fig. 16. Holting frequency cf B A qlandula over the period June, 1975 to Hay, 1976. 500 individuals were observed for each date Fig. 17. Faecal production rate of B± glandula over the period June, 1975 to May, 1976. 500 individuals were observed for each date Fig. 18. The si z e fregUency d i s t r i b u t i o n (in mg dry wt.) of the barnacles sampled from an area of 0.08 m2 from the experimental shore during the period Hay, 1976 to A p r i l , 1977. Each class i s represented by separated l i n e s on the probability paper and the number of each class i s given with the l i n e . The modal value of each class can be found at the 50 % intercept X ACKNOWLEDGMENTS I am most g r a t e f u l t o my wife, I r e n e , f o r much of her s p i r i t u a l support and t e c h n i c a l a s s i s t a n c e throughout my Ph. D. Study i n Canada. Thanks are p a r t i c u l a r l y due to Drs. C. Levings and D. R a n d a l l , f o r t h e i r h e l p f u l d i s c u s s i o n s and c r i t i c i s m s i n the past three years, and a l s o f o r reading the d r a f t of t h i s t h e s i s . I have b e n e f i t e d from the d i s c u s s i o n s with Drs. T. Ca r e f o o t , T. Parsons and R. Foreman, and I would a l s o l i k e t o thank them f o r r e a d i n g t h i s t h e s i s . I am indebted t o Drs. P. Breen and F. Bernard o f the P a c i f i c B i o l o g i c a l s t a t i o n , f o r readin g parts of my t h e s i s ; and Dr. F. T a y l o r of the U n i v e r s i t y o f B r i t i s h Columbia f o r s u p p l y i n g the diatom c u l t u r e . F i n a l l y , I would l i k e to express my g r a t i t u d e t o the Canadian gcverrirent f o r a l l o w i n g me to use the f a c i l i t i e s a t the P a c i f i c Environment I n s t i t u t e and a l s o f o r the tenure o f a Canadian Commonwealth s c h o l a r s h i p . 1 {1) INTRODUCTION E n e r g e t i c s t u d i e s can provide v a l u a b l e i n f o r m a t i o n p e r t a i n i n g t o a v a r i e t y o f e c o l o g i c a l l y i n t e r e s t i n g q u e s t i o n s . For example, the r a t e of energy flow has been found to be u s e f u l i n a s s e s s i n g the r e l a t i v e importance of the c o n t r i b u t i o n of po p u l a t i o n s t o the f u n c t i o n i n g of a community; d i s r e g a r d i n g the s i z e , metabolic r a t e and d e n s i t y of the animals ( S l o b c d k i n , 1962; Macfadyen, 1964; Odum, 1968, 1971). The v a r i o u s e f f i c i e n c i e s i n energy t r a n f e r (e. g. e c o l o g i c a l e f f i c i e n c y , a s s i m i l a t i o n e f f i c i e n c y and pro d u c t i o n e f f i c i e n c y ) are o b v i o u s l y p e r t i n e n t to the understanding of the ecology of the animal; e s p e c i a l l y i n resource u t i l i z a t i o n and trophodynamics. Margalef (1963; 1968) has r e l a t e d the s u c e s s i o n a l patterns and s t a b i l i t y of communities to energy flew and e f f i c i e n c y of energy u t i l i z a t i o n . H i r s h f i e l d and T i n k l e (1975) have s t r e s s e d the importance of the use of energy budgets i n a n a l y s i n g animal r e p r o d u c t i v e s t r a t e g i e s . fit the ecosystem l e v e l , energy flow s t u d i e s have been c a r r i e d out mostly on simple n a t u r a l a q u a t i c systems. For example, i n lake Mendota (Juday, 1940) ; Cedar Bog lake (Lindeman, 1942) ; S i l v e r s p r i n g (Odum, 1957) ; Root s p r i n g ( T e a l , 1957) ; s a l t marsh system (Odum and Smalley, 1959 ; T e a l , 1962); Cone s p r i n g ( T i l l y , 1968); Marion l a k e ( I f f o r d , 1969) and 2 on an abandoned beaver pond system (Hodkinscn, 1975). More r e c e n t l y , Perkins (1975) has a l s o attempted to c o n s t r u c t an energy flow model of an i n t e r t i d a l a s s o c i a t i o n . Most of these s t u d i e s have been c r i t i c a l l y reviewed by Mann (1969). Fewer s t u d i e s have been c a r r i e d out i n t e r r e s t i a l ecosystems, except i n an e l d f i e l d (Odum et a l , 1962) and more r e c e n t l y cn a shor t grass p r a i r i e (Andrew et a l , 1974) and on a suburban lawn system ( F a l k , 1976). In view of the d i f f i c u l t i e s of working on complicated n a t u r a l j ecosystems , seme other workers have t r i e d t o approach the problem with l a b o r a t o r y s t u d i e s on a microcosm. Such an approach was e x t e n s i v e l y used by Hichman (1958) and Slobodkin (1959; 1960; 1961; 1962) on micro-crustaceans. However, i t i s obvious t h a t , while working on the whole n a t u r a l ecosystem i s never easy, s t u d i e s of l a b o r a t o r y microcosm may be u n - r e a l i s t i c . Even working on the p r e v i o u s l y c i t e d examples of simple ecosystems must i n v o l v e much time and manpower. For understanding ecosystems, e n e r g e t i c s t u d i e s cn n a t u r a l p o p u l a t i o n s are t h e r e f o r e badly needed (Mann, 1S69). According to IBP terminology (Petrusewicz and Macfadyen, 1970), the general energy budget f o r a h e t e r o t r o p h i c organism can be represented by the f o l l o w i n g eguaticn :-C = P + B * U + F 3 W h e r e : -C = Consumption P = Production B = Sespiration U = Excretion F = Faecal production The energy budget of a population can be extrapolated from the energy budget of an i n d i v i d u a l provided that certain population parameters (e. g. density, population structure and mortality , etc. ) are known. Each item i n the above eguation i s measured i n energy units {calories) and are additive. The value of each item i s estimated independently. any item which i s p a r t i c u l a r l y d i f f i c u l t to measure can be found by difference since, t h e o r e t i c a l l y , the energy budget eguation should balance. However, measuring a l l the items i n the equation i s desirable since this provides a double check on the accuracy cf the equation (Hughes, 1970; Crisp, 1971). The measurement of each item per se usually does not involve great d i f f i c u l t i e s . The most intractable problem i s to extrapolate laboratory data (e. g. consumption and respiration) to the natural condition . For example, the metabolic rate may be double (Sinberg, 1956) or as high as 12.8 times <Tucker, 1968} i n comparison with that measured under constrained conditions i n the laboratory. It i s always d i f f i c u l t to estimate to what extent the metabolic rate of the animal i n i t s natural condition has been altered during the measurement period i n the laboratory. Some attempts have been made tc measure the respiration i n the f i e l d . For example, the b e l l jar method has been used by a number of workers to measure the oxygen uptake of benthic communities {Teal, 1957; Odum, 1957; Patmatmat, 1968) although the conditions within the b e l l jar may also have a large deviations from the natural conditions. LeFebre (1964) has t r i e d to measure the metabolic rate of pigeons by the excretion of D 2 0 1 8. He found that the metabolic rate of the pigeons i n the natural condition may be 7 or 8 times higher than that measured i n the laboratory. The application of radiotracer technique in measuring 0 2* 8 uptake in the f i e l d i s rewarding, although i t i s s t i l l i n a rudimentary stage. Three approaches are generally employed i n energetic studies. The f i r s t i s to keep the animals (or the population) e n t i r e l y under laboratory condition; thus a l l the budget items can te measured with precision ( e . g . Conover, 1966; Carefoot, 1967a; 1967b; 1970; Ooohan, 1973; Conover and L a l l i , 1974; Cosper and Beeve, 1975). Some other workers have t r i e d to estimate production i n the f i e l d and other budget items ( e . g . consumption, respiration and faecal production) in the laboratory (e. g. Kuenzler, 1961; Paine, 1971; Hughes, 1970; 5 1971a; 1971b). Yet another approach which involves d i r e c t measurement of consumption in natural conditions i s i l l u s t r a t e d by the work of Paine (1965) and Haynes (1974) . The majority of the reported energy budgets do not involve the measurement of a l l items (e. g. Gdum and Smalley, 1959; Kuenzler, 1961; Teal, 1961; Paine, 1965; 1971; lawton, 1970; 1971; Hughes, 1971a; 1971b; Otto, 1974). In those with independent measurement of a l l the items, few have been cl o s e l y balanced. Carefoot (1967a) found that i t was necessary to double the re s p i r a t i o n data in his energy budget of ftplysia .punctata ••, in order to balance the eguation. M i l l e r and Mann (1973) found that t h e i r energy budget for the sea urchin Stionqylocentictus 3roebachiensis was largely unbalanced, and concluded t h i s was the res u l t of dissolved organic matter l o s t . Similar results were obtained in the study of L i l l y (1975) cn the t r o p i c a l urchin Tjrijaneustes ventricosus . Hughes (1970) worked on the energy budget of the bivalve Scrobicularia plana and found that the consumption value derived from the summation method and feeding experiments d i f f e r e d by 13 to 14 SS. It i s not uncommon to f i n d that the budget eguation i s l a r g e l y unbalanced even i f a l l the budget items are precisely measured under r i g i d l y controlled laboratory conditions (Carefoot, personal communication). As Slobodkin (1962) has pointed cut : « Precise measurement of energetic r e s u l t s seems almost impossible and even rather crude measurements are time consuming and expensive 6 Moreover, t h i s indicates the necessity of double checking the accuracy of the equation by measuring a l l the items i n order to provide an estimation of error. A number of of studies have been reported on the energetics of animals of diverse phyla and habitats at the individual or population l e v e l s . Due to the vast amount of l i t e r a t u r e , i t i s therefore, only appropiate to review those on marine invertebrates. Relatively few studies have been carried out on the subtidal benthos; except for the sea urchins St ronqylocentiotus drgebachiensis (Miller and Mann, 1973) and Tripneustes ventricosus ( L i l l y , 1975) ; and the opisthobranchs Aplysia punctata and Dendronotus frondosus (Carefoot, 1967a; 1967b; 1970) . For pelagic animals, energetic studies have been carried out on the r o t i f e r Brachionus p l i c a t i l i s (Doohan, 1S7.3) ; the copepod Calanus hyperboreus (Conover, 1966); the chaetcgnath Sacjitta higpisda (cosper and Reeve, 1975); the mysid (Lasenby and Langford, 1973) and the pteropod Cliona limacina (Conover and L a l l i , 197U). A number of energetic studies, however, have been carried out on a variety of l i t t o r a l animals. For example, i n 7 the gastropods L i t t c r i n a i r r c n a t a (Odum and Smalley, 1959), Tegula funebralis (Paine, 1971), three species of Merita (Hughes, 1971b) the opisthobranchs Navanax inerffljs (Paine, 1965) and archidoris pseudoarqus (Carefoot, 1967b); the limpets l i s s u r e l l a barbadens (Hughes, 1971a), acmaea seabra (Sutherland, 1972); the isopods Tylos punctatus (Haynes, 1974) , Circlana harfordi (Johnson, 1976); the bivalves Modiolus demissus (Kuen2ler, 1961) , Scrobicularla plana (Hughes, 1970) , the oyster Crassostrea y i r g i n i c a (Dame, 1976) and the pclychaete Neanthes vireus (Kay and B r a f i e l d , 1973). Barnacles are one of the universal species upon the rocky shores in temperate waters (Stephenson and Stephenson, 1949; Lewis, 1964). Along the P a c i f i c coast from the aleutian islands to the northern border of Mexico, Balanus glahdula Darwin i s the major barnacle species (Barnes and Barnes, 1956). The mean number and biomass i n barnacle beds i n Monterey bay, C a l i f o r n i a , have been estimated as 11820 m-z and 2100 g dry wt. m-2, respectively (Glynn, 1965). Connell (1970) alsc reported that the population density of B A glandula at Point Roberts near the mouth of Fraser r i v e r ranged frcm between 3000 and 32000 m-2. Based on resettlement on several denuded quadrats, Glynn (1965) estimated that the production of this barnacle was 3.2 g dry wt. m - 2 month - 1, although i t i s obvious that only the new settlement was accounted for i n his study. With such a large number and biomass, the barnacles may channel 8 a large amount of energy from the pelagic environment to the l i t t o r a l community and, in turn, support a large number and variety of t h e i r predators [ e. g. the gastropods Thais la me 11 os a , T.. emarginata and Xs. cana l i e u lata (Connell, 1S70) ; the s t a r f i s h Pisaster pchraceus and the nemertean Jm_plectcjnema a i a c i l e (Kozloff, 1973) ] . The barnacles must, therefore, play an important r o l e in the food web dynamics of the l i t t c r a l system. However, suprisingly, the energetics cf barnacles have not been studied; except by Perkins (1975) who gave a very crude estimation on the energy flow of the B,. gla.nd.jula population i n his i n t e r t i d a l association - energy flow model. It was, therefore, the purpose of t h i s thesis to attempt a detailed study on the energetics of the barnacle Bj. glandula . Aggregation i s a commonly observed feature of natural barnacle populations, and thus was taken into account as a factor i n the present study of barnacle energetics. Aggregation leading to over-crowding i n barnacles has been shown to retard the growth rate of Balanus balancides (Crisp, 1960; Meadow, 1969), and cause a delay i n the onset of sexual maturation of Elminus modestus (Crisp and Patel, 1S61). Crisp and Davies (1955) also noted that individuals at the edge of a clump have a greater growth rate and concluded t h i s was a resu l t of a lesser competition for food. However, Crisp (1960) suggested that the reduction i n growth caused by high population 9 density was small ; he also suggested (Crisp, 1964) that i n t r a -s p e c i f i c competition at high population density i s important only in l i m i t i n g i n d i v i d u a l size and fecundity and has l i t t l e e ffect on t o t a l bicmass production per unit area. Unfortunately, only the basal diameters (Crisp and Davies, 1955; Meadow, 1969) or the t o t a l weight (Crisp, 1960; Crisp and Patel, 1961) of the animal, including the s h e l l , were used as c r i t e r i a in a l l these growth studies. The basal diameter i s unlikely to be a good index for growth since (1) the barnacles can grow in height i n compensation when crowded (Barnes and Powell, 1950) and (2) t h i s did not provide any information about the tissue of the animal. The measurement of to t a l weight including the s h e l l would place a very large emphasis on the s h e l l rather than the body tissue and egg (which stores the majority of energy) since the dry weight of the former i s some 40 to 50 times larger than the l a t t e r . It i s therefore not cle a r , from previous studies, whether the growth rate of the tissue i s affected or not. This study approached the problem by analysing the energy p a r t i t i o n and the energy budget of i n d i v i d u a l barnacles with d i f f e r e n t degrees of aggregation. The main purposes of t h i s thesis were therefore to (1) construct and analyse the energy budget of ind i v i d u a l specimens °f B. glandula under d i f f e r e n t degrees of aggregation; and (2) to estimate the energy flow, production and mortality of a 10 natural barnacle |B. glandula) population and hence to evaluate the functional Importance of the population in the l i t t o r a l community. Experiments were designed to answer the following questions :-(1) How do barnacles budget t h e i r energy ? Do the animals have a different budget under d i f f e r e n t degrees of aggregation ? To what extent does crowding aff e c t the energetics of the animals ? (2) How much energy i s being channelled from the pelagic into the l i t t o r a l system through the barnacle populations ? How much energy i s elaborated i n the form of barnacle production and becomes available to the next trophic l e v e l ? How e f f i c i e n t are barnacles i n assimilating energy and transfering i t into production ? How important i s B^ . glandula , then, in the l i t t o r a l community i n a functional sense ? 1 1 (2) TERMINOLOGY Following IBP terminology (Petruswicz and Macfadyen, 1970), the energy budget f o r an in d i v i d u a l barnacle can be represented by the following eguation :-C = P.BT. + P. E. + P.S. • B.AER. .• R.AQ. ,+ M • F 8 h er e: -. C = Consumption P.BT. = Production of tody t i s s u e 1 P.E. = Production of egg 2 P.S. = Production of s h e l l E.AER. = A e r i a l r e s p i r a t i o n B.AQ. .= Aguatic re s p i r a t i o n H = Melting F .= Faecal production ,* The testes were d i f f i c u l t to i s o l a t e and were incorporated into body tissue in t h i s study. 2 In this study, *eggf i s defined as the t o t a l of ovarian tissue, egg masses and naupliar embryos . 12 And : -A = C - P F = P.BT. + P.E. P.S* E = R. A EE. + R. AQ. Assimilation efficiency= A/C Gross production e f f i c i e n c y = P/C Net production e f f i c i e n c y = P/A Where : -A = Assimilation P = Production E = Respiration Preliminary determination of urea and NB^ (Strickland and Parsons, 1S68) indicated that NH^ rather than urea was the major excretory product for B.. glandula . The p o s s i b i l i t y of excreting ur i c acid by £}_. glandula has not been investigated. NH3 with low c a l o r i f i c value i s often considered to be unimportant i n energetic studies (Crisp, 1S71). Based on determinations i n a preliminary experiment, i t was estimated that 1 mg of barnacle tissue would excrete 53.5 x 1 0 - 6 c a l . of NH3 per month and i t was therefore considered negligible i n this study. 13 The loss of dissolved organic matter (DOM), which may be important i n some animals (e, g. 42% of consumption and 68% of assimilation in the sea urchins according to M i l l e r and Mann, 1973); as well as the d i r e c t uptake of DOM from the sea water (Stephens, 1968) were not considered i n the present study because quantitative measurement of DOM i s always d i f f i c u l t . S p e c i f i c dynamic action (SDA) was considered important in the energetic studies of copepods (Conover, 1966) and sea urchins ( L i l l y , 1975). The e f f e c t of SDA could vary largely with the guality and guantity of diet, as well as the period and intensity of starvation prior to feeding ( L i l l y , 1975). The role of SDA was not taken into account in th i s study since i t i s d i f f i c u l t to determine these parameters for glandula . 14 (3) MATERIALS AND METHODS 3. 1 The experimental lobulations Barnacle cyprids have a strong tendency to orientate towards p i t s and grooves when s e t t l i n g . Based on this •rugophilic* {Crisp and Earnes, 1954) behaviour of the cyprids, barnacles can be established in a pre-determined pattern by d r i l l i n g small p i t s i n any desired pattern on a smooth surface (Crisp and Patel, 1961). Applying t h i s technique, the following aggregation patterns were developed on 0.61 m - 2 (78 x 78 cm) plexiglass plates {Fig. 1) :-(1) Isolated i n d i v i d u a l s (designated as * Isolated • hereafter) (2) Paired individuals (designated as * Pair ' hereafter) (3) Individuals i n a group of nine (designated as * 9* hereafter) (4) Individuals i n a group of eighty one (designated as *81» hereafter) The number of individuals on each of these plates was equal (n=1458). The distance between indivi d u a l s when they f i r s t s e t t l e d was 1.75 cm f o r *Isolated' and 0.75 cm for a l l other barnacle populations. 1 5 At the end of March, 1975, three replicates of each of the above patterned plexiglass panels, together with two roughened plexiglass panels f to harvest barnacles s e t t l i n g at t h e i r •natural' density (n = 200000-m-2 for the newly s e t t l e d spat which had decreased to 12295 m_2 in late October) and designated as 'Crowded* hereafter ] were put at the same t i d a l l e v e l (2.6 to 3.3 m above chart datum) at random on the p i l i n g s of the wharf of the P a c i f i c Environment I n s t i t u t e , West Vancouver, B r i t i s h Columbia (123° 15»SJ, 49° 20»N). It was anticipated that the degree of crowding would decrease following the order 'Crowded' > »81» > '9» > 'Pair' > 'Isolated'. The plexiglass plates were f i r s t put against a black wooden panel since a dark color encourages cyprids to s e t t l e (Gregg, 1945). The black wooden panels were removed when the cyprids had f i l l e d up the p i t s on the panels. The barnacle spat settled in late May, grew and touched each other (except indiv i d u a l s i n 'Isolated') in July and attained t h e i r maximum size i n late October. The panels were checked p e r i o d i c a l l y and any undesired additional s e t t l e r s (e. g. other specimens of Bj. ijlandula , the mussel Mvtilus edulis and the green algae Bnterorcorpha sp. ) were removed. An analysis of variance showed no s i g n i f i c a n t difference (p = 0.05) i n the growth rate of the barnacles cn a l l 16 the p i l i n g s a f t e r they had s e t t l e d for 1 month. Twenty individuals were removed at random for each pattern each month : twenty single i n d i v i d u a l s were removed from the 'Isolated* panel, ten pairs were removed from the 'Pair' panel, two whole groups of nine plus two indivi d u a l s were removed from '9*; a proportion of a whole group of 81 were removed from *81» and further sampling from t h i s group was avoided in the subsequent samplings. F i n a l l y , twenty i n d i v i d u a l s were removed at random from the 'Crowded' panel every month. 3.2 JDetermination of dry weight, ash- free weight and c a l o r i f i c value In t h i s study, the dry weight of a l l the barnacle material <i. e. body tissue, ovarian tissue, egg masses, naupliar embryos, molted exoskeletons, faecal p e l l e t s and shell) was obtained by weighing with a CAHN G2 electrobalance after drying i n an oven at 100° C for 48 hours. The percentage of ash-free weight of a l l these barnacle materials, except the s h e l l , was determined by ashing a known weight of dried sample in a furnace at 500° C for 4 hours. The dried body tis s u e , ovarian tissue, egg masses, naupliar embryos, molted exoskeletons and faecal p e l l e t s were homogenized with a mortar and pestle, and pelleted. The 17 c a l o r i f i c value of the materials sere determined by burning the p e l l e t s in a P h i l l i p s o n micro-bomb calorimeter. In determining the c a l o r i f i c value of the molted exoskeleton and faecal p e l l e t s , the materials were mixed with benzoic acid to burn since the combustabilities were low. No correction was made for acid production, or the burning of the f i r i n g wire, since these were considered to introduce n e g l i g i b l e error (Paine, 1964). Barnacle s h e l l i s mainly composed of CaC0 3 , c h i t i n and protein (Barnes et a l , 1976). In th i s study, the percentages of the l a t t e r two components in the barnacle s h e l l s were determined, using the methods described by Barnes et a l (1976). The t o t a l energy content of the s h e l l was then found by the summation of the energy content of the c h i t i n and protein f r a c t i o n s , assigning a c a l o r i f i c value of 5.65 and 4.10 for protein and c h i t i n , respectively (Crisp, 1971). 3.3 Production (P.BT., P.E., P.S.) Twenty i n d i v i d u a l samples of barnacles were taken at random from each of the patterns each month. They were cleaned in running sea water with a brush before dissecting. The egg and the body tissue were dissected from the barnacles. The dry weight, ash-free weight and c a l o r i f i c value cf the various components of the barnacle were determined, using the methods already described. The energy standing crop of the various 1 8 components for each i n d i v i d u a l barnacle was then found by multiplying t h e i r dry weight and percentage of ash-free weight by the c a l o r i f i c values. The monthly mean energy production of the body tissue (P.BT.), egg (P.E.) and s h e l l (P.S.) was found by the differences i n the energy standing crop of each item between consecutive samplings. In estimating egg production, the summation of the difference i n the energy standing crop of ovarian tissue and egg masses, as well as naupliar embryos was considered. Negative production of egg masses or naupliar embryos was neglected and assigned a zero value since t h i s merely indicated the discharge of larvae between sampling periods. Negative production of ovarian tissue was accounted for because t h i s would indicate reabsorption (Barnes and Archituv, 1976) . 3.4 Bespiration (B.AEB. And B.AQ.) 3.4.1 Aerial respiration {8. A E B.) The ovarian tissue, egg masses and naupliar embryos were dissected from B. glandula and the 0 2 consumption measured by a Gilson d i f f e r e n t i a l respirometer at 5° , 10° , 15° and 20° C , after acclimatizing the tissues at the experimental temperature for one hour. The tissues were then dried and the 19 0 2 consumption rates were expressed as ^ ul 0 Z mg dry t i s s u e - 1 hour - 1 . The a e r i a l 0^  consumption rate of the barnacle tody tissue was measured in a f a c t o r i a l designed experiment with 4 weight groups and 4 temperatures. Barnacles on mussel s h e l l s were co l l e c t e d near P a c i f i c Environment Ins t i t u t e during low tides, after cleaning i n running sea water, they were sorted into 4 size groups with mean body tissue weight of 0.95 ± 0 . 1 2 , 2.98 ± 0.13, 4.86 ± 0.25 and 8.27 ± 0.60 mg (mean ± S.D.). The animals were acclimatized i n an incubator at the experimental temperature for at l e a s t 12 hours before the experiment. Pneumatostcmes indicating a e r i a l r e spiration (Grainger and Newell, 1965) were observed i n nearly a l l the experimental animals. The a e r i a l r e spiration of the whole animal was measured by a Gilson d i f f e r e n t i a l respirometer at 5° , 10° , 15° and 20° C. Barnacle and mussel s h e l l were used as a control in the experiments. The body tissue, ovarian tissue, egg masses and naupliar embryos of the barnacle were dissected and dried separately at the end of the experiment. The o 2 consumption of the body tissue was then found by subtracting the 0 2 consumption of the associated ovarian tissue, egg masses or naupliar embryos from that of the whole animal at the same temperature. The 0 2 consumption rate of the body tissue was then calculated and expressed as jil 0 2 mg dry t i s s u e - 1 hour - 1 . 20 3,4.2 Agnatic respiration {B. A Q.) Preliminary experiments showed that s a l i n i t y within the range of 13 to 23 So had no s i g n i f i c a n t e f f e c t upon the aguatic respiration of B. glandula (Hu, unpublished data). Since the s a l i n i t i e s near the present study area nearly always f a l l within such a range (Stock'ner and C l i f f , 1976), s a l i n i t y was considered to be unimportant in the present study. Subsguently, sea water of s a l i n i t y 23 %& was used in a l l the experiments to measure the r e s p i r a t i o n of barnacle tissues. The <32 consumption of the dissected ovarian tissue and egg masses were measured at 5° , 10° , 15° and 20° C in a respiration v i a l (YSI Hodel 53) with 0 2 saturated sea water. Occasional s t i r r i n g using a magnetic s t i r r e r was provided every 15 minutes; measurements were obtained over a 1 hour period. The 0 2 concentration was determined by an 0 2 electrode with a Radiometer PHH 72 unit. The 0 2 consumption cf the tissue was then found by the difference i n 0 2 concentration before and after the experiment and expressed as pi 0 2 mg dry t i s s u e - 1 hour - 1 . The aguatic r e s p i r a t i o n of the naupliar embryos was not measured because s t i r r i n g caused a release cf the swimming na u p l i i which increased the 0 2 consumption. The aguatic 0 2 r e s p i r a t i o n of the body tissue of B. glandula was measured i n a f a c t o r i a l designed experiment with 4 21 different water temperatures and 4 d i f f e r e n t weight groups. Barnacles on mussel s h e l l s were collected at low tides, they were cleaned i n running sea water and sorted i n t o 4 groups with mean body tissue weight of 0.44 ± 0.01, 2.43 ± 0.30, 4.06 ± 0.29 and 9,98 ± 0.26 mg. The animals were allowed to acclimatize at the experimental temperature for 24 hours before the experiments. Continous aeration was supplied during the period of acclimatization. The 0 2 consumption of the various size groups at water temperatures of 5° , 10° , 15° and 20° C was measured. Immediately before the experiment, the barnacles were transfered to the resp i r a t i o n chamber. The sea water i n the respi r a t i o n chamber had the same s a l i n i t y and temperature as the acclimatising sea water, except that i t had been f i l t e r e d through a 30 ji plankton mesh to remove large size plankton. The respiration chambers were placed i n a constant water temperature bath ( ± 0.2° G). a current was provided by a magnetic s t i r r e r placed under the mesh-gauze {which separates the animal and the st i r r e r ) i n the resp i r a t i o n chamber. The measured Oz consumption was accepted only i f over half of the barnacles were observed to have active c i r r a l beat during the period of measurement. The 0 2 saturation of the sea water in the res p i r a t i o n chamber was never allowed to f a l l below 60 %. {According to Prasada Rao and Ganapati (1969), 0 2 saturation above t h i s l e v e l has no ef f e c t upon the respiration of 22 barnacles). 0^ concentration of the sea water before and after a one hour experiment was measured by a pclarographic 0 2 electrode and a Radiometer PHM 72 unit. The 0 2 consumption was found by the difference of 0 2 concentration before and after the experiment. As in the a e r i a l r e s p i r a t i o n experiment, the 0 2 consumption of the body tissue was found by subtracting the 0 Z consumption of the associated ovarian ti s s u e , egg masses and naupliar embryos from the 0 2 consumption of the whole active animal. The 0Z consumption was expressed in pi 0 2 mg dry body t i s s u e - 1 hour - 1 . The a e r i a l and aguatic 0 Z consumption of the body tissue, ovarian tissue egg masses and naupliar embryos were converted into c a l o r i e s using an o x y c a l o r i f i c value of 4.8 cal/ml 0 2 (Crisp, 1971) after correcting the 0 2 consumption to N. T. P. Such an o x y c a l o r i f i c value assumes a mixed diet of protein, carbohydrate and fat for the animal (Crisp, 1971) which would be appropiate for barnacles, as they normally feed on plant (phytoplankton) and animal (zcoplankton) materials. Energy l o s t to a e r i a l and aguatic r e s p i r a t i o n for the month was then calculated for an in d i v i d u a l barnacle, taking into account (1) the mean body weight of the animal (2) the associated ovarian tissue, egg masses and naupliar embryos (3) the t o t a l hours of submergence and emergence for the month at 3 2 3 ni t i d a l l e v e l calculated from a l o c a l tide table and (4 long term a i r and water temperature of the month (Envirc Canada / Atmospheric Environment, 1973; I n s t i t u t e Oceanography, the University of B r i t i s h Columbia, 1973). Appendix 1 and 2), 3.5 Faecal production and molting rate (H and F) Both faecal production and molting rate sere determined every month in the same experiment, Barnacles on mussel s h e l l s of approximately the same size as the indivi d u a l s from * 81*' were co l l e c t e d from the p i l i n g s one day before the experiment. They were cleaned with a brush in sea water and submerged from the wharf overnight in a cage. The faecal production and molting rate of the animals were then measured i n an experimental period of 24 hours. The barnacles were taken out of water at the s t a r t cf the experiment. Five r e p l i c a t e s with about 100 animals per tray were exposed to a i r at the mean temperature of the month for a certain period of time, then into running sea water for the rest of the 24 hour period. The r e l a t i v e time for the animals exposed to a i r and submerged i n running sea water within the 24 hours experimental period, was determined accordingly by the t o t a l submergence / emergence ratio of the month at 3 m t i d a l l e v e l . At the end of the experiment, the barnacles were ) the craent of (see 24 dissected and the mean dry weight of th e i r body tissue determined. The molted exoskeletons and faecal p e l l e t s were collected with a micro-pipette and t h e i r dry weights , ash-free weights and c a l o r i f i c values determined , using the methods already described. The faecal p e l l e t s of B.. glandula are large and sheathed, and hence can be recovered e a s i l y . The molting frequency of a single i n d i v i d u a l within the month was estimated from the percentage of molting in the experiment and expressd as number of molts i n d i v i d u a l - 1 month-1 .The faecal production rate was expressed i n og dry faeces mg dry body t i s s u e - 1 month - 1 . The energy l o s t in faecal production for each month was then calculated from the c a l o r i f i c value, the ash-free weight content of the faecal p e l l e t s , and the faecal production rate. The energy l o s t in molting for the month for an i n d i v i d u a l was calculated from the c a l o r i f i c value, the ash-free weight percentage of the molted exoskeleton, the mean weight of the molted exoskeleton and the molting frequency of an individual in the month. An annual energy budget was then constructed for an in d i v i d u a l barnacle from each of the aggregation patterns, by summing up the values for each item. Consumption was then found by summing a l l the budget items on the right hand side of the 25 equation. 3.6 laboratory energy budget To estimate the error involved i n deriving the consumption values by the summation method, a short term (36 days) experiment was performed in the laboratory i n August, 1976. Energy budgets were constructed for three small groups of JLi ^Jandula i n which a l l the budget items, including consumption, were measured independently, J*. glandula on mussels s h e l l s were collected during low tides. Individuals with mean dry body tissue weight 6.1 ± 0.4 mg were sorted for the experiments. The standing crop energy of the body tissue and ovarian tissue i n 30 individuals were determined before the experiment, using the methods already described. Another 120 sorted individuals were equally divided into three groups. They were put i n separated trays in a f i l t e r e d sea water system (in which fresh sea water was f i l t e r e d sequentially through : commercial sponge - glass wool - 100 ji mesh - 3 0 ^ 1 mesh - 10 jx mesh - Whatman No. 1 f i l t e r paper). The sea water of the system was checked daily with an inverted microscope to ensure the absence of algae and d i n o f l a g e l l a t e s . The water temperature was continously recorded by a thermograph (Hyan model F), The fa e c a l materials and molted exoskeletons 26 produced by each group were collected daily, and energy loss in faecal production and molting was estimated by methods described above. Each group of barnacles was fed and exposed tc a i r once or twice d a i l y . The barnacles were fed i n a plexiglass chamber {same as the respiration chamber described before) vhich was incubated in a constant temperature (± 0.2°C) water tath. 400 ml of log phase growing diatom { Skeletoneroa costatum ) was fed to the barnacles. Continous s t i r r i n g of the culture was provided by a magnetic s t i r r e r . fin equal number of empty barnacle s h e l l s were used in the control cf the feeding experiment. The numbers of diatom were counted usinq a haemacytometer and the t o t a l consumption of the barnacle group was found by the difference in the t o t a l number of diatcms before and after the experiment. The energy value of Skeletpnema costaturn consumed was then calculated based on (1) the number and dry weight conversion factor and the chemical component analysis of Skeletonema costatum given by Parsons et a l (1961) and (2) assigning a c a l o r i f i c value of 5.€5, 4.10 and 9.50 c a l mg for protein, carbohydrate and f a t , respectively (Crisp, 1971). For a e r i a l exposure, the barnacle groups were put in an incubator with a known, constant temperature (ranging from 10 to 20°C). The duration of a e r i a l exposure, the a i r temperature, 27 the feeding temperature, the feeding time and feeding concentration varied among the three barnacle groups. The 0X consumption of each barnacle group was then calculated from the series of equations derived from the e a r l i e r a e r i a l and aguatic respiration experiments (see 4.3); taking into account the duration of exposure and submergence and the respective a i r / water temperature that each p a r t i c u l a r group was subjected to. The t o t a l energy loss i n a e r i a l and aguatic respiration ,for each barnacle group was then calculated. fit the end of the experiment, the barnacles were k i l l e d and the t o t a l energy content stored i n the body and ovarian tissues was determined. The production cf body tissue and cvarian tissue f o r each group during the experimental period was then found by the difference in the t o t a l energy content of the respective item before and aft e r the 36 day period. The production of s h e l l was assumed to be zero i n constructing the laboratory budgets since s h e l l growth of B. glandula i s minimal a f t e r 7 months cf adult l i f e (see re s u l t s and discussion). 3.7 Population dynamics and energy flow of the natural barnacle Emulation A one year sampling program (Hay, 1976 to Apr i l , 1977) 28 was carried out i n order tc provide an estimate of the energy flow of the natural barnacle population. Because of the high degree of heterogeneity of the i n t e r t i d a l environment , i t was necessary to confine the sampling to a r e l a t i v e l y homogeneous, small area. A granite rocky shore about 100 m tc the east of the P a c i f i c Environment I n s t i t u t e {Fig. 2) was choosen for this study, due to i t s a c e s s i b i l i t y . A l l the sampling was carried out on a narrow horizontal s t r i p {approximately 6 by 0.6 m, located from 2.6 to 3.3 m above chart datum and which was determined to be the 'optimal* v e r t i c a l range of qlandula ) marked on t h i s shore. At the beginning of the sampling program, a matrix of 300 (6 x 50) spots was painted on the s t r i p of the shore. Eight of them were chosen monthly at random (by finding their co-ordinates in a random number tab l e ) . A 0.01 m2 quadrate was then placed on the chosen spot and the barnacles sampled. Further sampling from the same spot was subsguently avoided. The body tissue, egg and s h e l l of each i n d i v i d u a l barnacle in the samples were dissected and dried as described above. They were then weighed to the nearest 0.1 mg with a Perkin Elmer AD 2 autoblance. During the spat f a l l s i n flay and September 1976, the number of spats in eight 2 cm2 guadrats were counted to provided an estimate on the t o t a l number of spats s e t t l i n g i n an area of 0.01 m2. The mean dry weight of an 29 i n d i v i d u a l spat was estimated by weighing the body tissue of 50 barnacles . then pooled for every month. A l l the population parameters i n this study, unless s p e c i f i e d , are expressed i n terms of an area of 0.08 m2. The cumulative percentages of each 0.1 mg group were then plotted on a prob a b i l i t y paper to d i f f e r e n t i a t e different size classes (Harding, 1949). The modal value of the mean weight of each class was then found at the intercept of the line at 50 % and the standard deviation given by the distance between 50 and 16 (or 86) %. The production and mortality (in mg dry weight) of each class were then calculated using the following equations (Crisp, 1971) :-The data obtained from the eight 0.01 m2 guadrats were t P = 2 N A w 0 t 0 where P = Production M = Mortality w = Average mean weight over period 30 N = Number of individuals in each class N .= Average value cf N over period = 1/2 ( N + N } t t +A t The production and mortality in dry weight were then converted i n t o c a l o r i e s by using the c a l o r i f i c values and percentages of ash-free weight obtained from the corresponding category of barnacle tissue i n the same month in 1975 -1976 . The energetic cost represented by aguatic and a e r i a l r e s p i r a t i o n of each class was calculated, using the same method as already described in 3.4; taking into account : (1) the number of individuals i n each class and (2) the hour of submergence and emergence at 3 m t i d a l l e v e l for each month , calculated from a l o c a l tide table and (3) the 'long term* mean ai r / water temperatures f o r each month (Environment Canada / Atmospheric Environment, 1973; Institute of Oceanography, the University of B r i t i s h Columbia, 1S73). (see Appendix 1 and 2). The energy l o s t i n molting and faecal production was calculated, using the molting and faecal production data obtained i n 1975-1976 . Since the molting frequency and faecal production data i n Fig. 16 and 17 were derived for the cohort s e t t l e d i n May, 1975; the data may not be s t r i c t l y applicable to the > 1 year settlements and the September 1976 settlement. However, such an adoption of data was assumed to have introduced a negligible error since molting and faecal production were 31 found to be r e l a t i v e l y unimportant in the energy budget of B. glandula (see res u l t s and discussion). An annual energy budget was then constructed for each size class of the natural barnacle population frcm May, 1976 to A p r i l , 1977. As before, consumption was found by the summation method. The energy flow of the barnacle population was then egual to (C - F) . 3.8 Istimaticn of error in volyed in using long term mean temperatures in cal c u l a t i n g respiration energetic cost long term mean air and water temperatures were used in the present ca l c u l a t i o n of R.AEE. and R.AQ. i n the annual energy budget eguations as well as in estimating the resp i r a t i o n energetic cost of the natural population (see 3.4 S 3.7). Since the barnacles are normally subjected to large fluctuations of ai r and water temperatures i n the i n t e r t i d a l environment, the use of mean temperatures (especially the 'long term* mean values used in the present study) may have poss i b i l y introduced a large error i n estimating R.AER. and R.AQ. The following experiment was therefore designed to estimate such an error involved. The changes in a i r / water temperatures frcm 1st July, 1976 to 30th July, 1976 were continuously recorded by a thermograph (Ryan Model F) secured on the p i l i n g of P a c i f i c 32 Environment I n s t i t u t e at 3 ra t i d a l l e v e l . The t o t a l time i n hours subjected to a pa r t i c u l a r a i r / water temperature for the whole month was then determined. The energetic cost i n aguatic and a e r i a l r e s p i r a t i o n of two hypothetical i n d i v i d u a l barnacles : A (body tissue = 1.28 mg, ovarian tissue = 0.45 mg, egg masses = 0.06 mg) and B (body tissue = 4.01 mg, ovarian tissue - 3.87 mg, egg masses = 1.05 mg), subjected to <1) the recorded fluctuating a i r and water temperatures and (2) the Mcng term' mean a i r and water temperatures (Environment Canada / Atmospheric Environment, 1973; Ins t i t u t e of Oceanography, the University of B r i t i s h Columbia, 1973) during the period of 1st to 30th July, 1976, were calculated separately and compared. 33 (4) RESULTS C a l o r i f i c values a.njj energy; staijdlpa crop The seasonal changes in the c a l o r i f i c value of ovarian tissue, egg masses, naupliar embryos and body tissue are shown in Fig. 3. In general, the highest c a l o r i f i c value was found i n ovarian tissue, followed i n decreasing order by : egg masses > naupliar embryos > body tissue. The seasonal variations i n the c a l o r i f i c value of a single category of barnacle tissue were generally small. The result of the chemical analysis and c a l o r i f i c value of barnacle s h e l l s are given i n Table 1. The c a l o r i f i c value of the barnacle s h e l l was very low { 0.134 ± 0.03 c a l rag dry wt. -*) when compared to the tissues. The seasonal changes i n the energy standing crop of the body tissue, egg, s h e l l and the t o t a l energy standing crop (the summation of the energy standing crop of body tissue , egg and shell) of i n d i v i d u a l barnacles from various aggregation patterns are shown in Fig. 4 to 7. The energy standing crop of body tissue, egg and t o t a l 34 energy standing crop for a l l the aggregation patterns showed a rapid increase in the f i r s t four months after the settlement (June to September) and reached a peak i n September or October. These values decreased to a lower l e v e l i n the winter mcnths (November to March). A marked increase was observed in A p r i l , followed immediately by a sudden decrease in May. In comparing the energy standing crop of body tissue, egg, s h e l l and the t o t a l energy standing crop between individuals from d i f f e r e n t aggregation patterns, these values showed a general trend of decrease with an increasing degree of aggregation ; following the order : • Isolated * > * Pair * > *9» > '81' > * Crowded ' . These values for the various aggregation patterns were compared for each month with a Duccan's new multiple range test ( L i , 1964). The r e s u l t of t h i s s t a t i s t i c a l analysis showed that the majority of these comparisons are s i g n i f i c a n t (p=0.05). Without exception, the * Isolated • in d i v i d u a l s always had a s i g n i f i c a n t l y higher value in their energy standing crop of body tiss u e , egg, as well as t o t a l energy standing crop when compared with that cf t i e ' Crowded * individuals . The egg : body tissue r a t i o and s h e l l : tissue r a t i o (here tissue i s defined as the t o t a l of body tissue and egg), when expressed i n terms of energy , served as a good in d i c a t i o n of the r e l a t i v e proportion of energy that in d i v i d u a l barnacles 35 had channeled to the building up of s h e l l , body tissue and egg. These two r a t i o s were calculated for each i n d i v i d u a l barnacle from the various aggregation patterns for every month and are shown in Fig. 8 and 9, repectively. The monthly egg : body tissue r a t i o s and s h e l l : tissue r a t i o s for individuals from the various aggregation patterns were then compared with a Duncan 1s new multiple range test fp=0.05) ( L i , 1961). Except i n August, the egg : body tissue r a t i o showed a clear trend of decrease following the order : * Isolated • > » Pair » > »9» > »81» > ' Crowded * . A large number of comparisons among the mean values of t h i s r a t i o are s i g n i f i c a n t ; and without exception, the * Isolated * individuals always had a s i g n i f i c a n t l y higher egg : body tissue r a t i o when compared with the * Crowded ' individuals . For a l l the months investigated, the s h e l l : tissue r a t i o showed an apparent trend of decrease following the order : * Crowded » > • 81 * > • 9 • > » Paired 1 > » Isolated • . Sim i l a r l y , a Duncan's new multiple range test ( L i , 1964) showed that the majority of the comparisons among the means were s i g n i f i c a n t (p=0.05). When considering only the ' Isolated * and * Crowded * indivi d u a l s ,the former always had a s i g n i f i c a n t l y lower s h e l l : tissue r a t i o when compared with the l a t t e r . ^•2 Production 36 The seasonal changes i n the mean P.BT., P.E. and P.S. of individ u a l s from the various aggregation patterns are calculated and shown in Fig. 10 to 12. Shell production showed a progressive increase from June , reached a peak i n September, and then became neg l i g i b l e after December. Positive values of P.BT. were generally found in June, July, August, September, February, March and A p r i l ; and negative P.BT. were found in the other months of the year. P.BT. showed a marked positive and negative peak i n Ap r i l and May, respectively. Positive values of P.E. were found from June to September; with a marked peak in September. A negative peak of P.E. was observed in November. Although the values of P.E. were e r r a t i c during the winter months {December to March), negative values of P.E. were generally observed. Positive values of P.E. were found for a l l the aggregation patterns i n A p r i l . Negative P.E. was, again, found f o r • 9 * , * 81 • , and • Crowded * in d i v i d u a l s i n May while individuals from * is o l a t e d * and * Pair * s t i l l had a positive P.E. at t h i s time. 4.3 Sespiraticn Covariance analysis showed no s i g n i f i c a n t difference (p=0.05) between the a e r i a l and aguatic 0 2 consumption rate of the ovarian t i s s u e . S i m i l a r i l y , no s i g n i f i c a n t difference 37 (p=0.05) was found between the a e r i a l and aguatic 0 2 consumption rate cf the egg masses. The data frcm the aguatic and a e r i a l 0 2 consumption rate were therefore pooled for these two items. 0 2 consumption rate as a function of temperature for ovarian tissue, egg masses and naupliar embryos i s shown i n Fig. 13. Regression l i n e s were f i t t e d by the least sguares method for each set of data. The relationships between the 0 2 consumption rate and temperature for the various tissues are given by the following equations :-log 0 2 consumption rate of ovarian tissue = 0.017 temperature °C - , 1.274 (r=0.810; n=32) log 0 2 consumption rate of egg masses = 0.03 2 temperature °C 0.663 {r=0.985; n=32) log 0 2 consumption rate of naupliar embryos = 0.025 temperature °C - 0.663 (r=0.941; n=24) Serial and aguatic 0 2 consumption rates of the body tissue are plotted against the body weight at 5° , 10° , 15° and 20° C and shown i n Fig. 14 and 15, respectively. The l i n e s are f i t t e d by the least sguare method. In order to predict the a e r i a l and aguatic 0 2 consumption rate of an i n d i v i d u a l barnacle with any given body weight and air/water temperature, the following multiple l i n e a r regression equations were generated frcm the data :-38 log a e r i a l 0 2 consumption rate of body tissue = (-0.075 ± 0.027) + (0.034 ± 0.001 temperature °C ) - (0.168 ± 0.023 log mg body weight) (r=0.892; n=79) log aquatic 0 2 consumption rate of body tissue = (0.309 ± 0.022) + (0.021 + 0.001 temperature °C ) - ( 0.334 ± 0.016 log mg body weight) (r=0.9l4; n=60) ***** f a e c a l production and molting rate The monthly molting freguency and faecal production rate for an i n d i v i d u a l barnacle are shown i n Fig. 16 and 17, respectively. Both the molting frequency and faecal production values varied p o s i t i v e l y with the body tissue and egg production . High values cf molting frequency and faecal production were found i n august , September, March and A p r i l , when the plankton was presumably abundant in the waters . Values were low and negli g i b l e i n December and January when the temperature was low and the food was scarce. 4.5 Annual energy, budgets for i n d i v i d u a l barnacles frem the various aggregation patterns Five separate annual energy budgets were constructed for the individuals from each of the aggregation patterns and are shown i n Table 2. The percentage of energy chanelled to each budget item i s shown in brackets by assuming that consumption 39 eguals 100 %. An exceptionally high assimilation e f f i c i e n c y hut low gross production e f f i c i e n c y and net production e f f i c i e n c y were found from the annual budgets. (91.8 to 92.7, 22.8 to 26.1 and 24.7 to 28.6 %, r e s p e c t i v e l y ) . When comparing the proportion of energy used i n the various budget items, i t i s apparent that B. qlandula channeled a very large proportion (64.8 to 67.7%) of energy intake into r e s p i r a t i o n . Egg production was the second most important (12.3 to 15.3%), followed in order of decreasing importance by : production cf s h e l l (6.1 to 7.21) > production of body tissue (3.9 to 4.6S) > molting (1.1 to 2.3%). The degree of aggregation in the various patterns was ranked a r b i t a r i l y (1,2,3,4,5 for ' Isolated 1 , • Pair * , * 9 ' , 1 81 * and 1 Crowded * , r e s p e c t i v e l y ) . A simple c o r r e l a t i o n test showed that the annual values of C, A, P, P.BT., P.E. and P.S. of an i n d i v i d u a l barnacle showed a very good and s i g n i f i c a n t (p=0.05) negative c o r r e l a t i o n (r=-0.934, -0.934, -0.94 7, -0.811, -0.929 and -0.995 , respectively) with the degree of aggregation .The ' Isolated • individuals have a much higher value of C, A, P, P.BT., P.E. and P.S. (2.0 to 2.5 times higher) when compared to that of the * crowded 1 individuals . 4.6 laboratory energy budgets The laboratory energy budgets are presented i n Table 3. The consumption values derived frcm both the summation method 40 and actual feeding experiments are also shown. The consumption values determined by summation were 7.41 higher and 16.2 and 15.6% lower than those determined by actual feeding experiments for the three barnacle groups. Gross and net production e f f i c i e n c i e s were not calculated from the laboratory energy budgets since production gave negative values i n a l l three barnacle groups. The assimilation e f f i c i e n c i e s derived from the three independent barnacle groups were s i m i l a r l y high (99.3 ± 0.1%) and agreed with that derived from the annual energy budgets. 4»7 Population dynamics and energy flow of the natural barnacle EP.£.22a t i on Three size groups were readily distinguishable ca the shore at the beginning of the sampling in May 1976. From a small to a large body weight, they were designated as ' > 1 year settlement (A) * , * > 1 year settlement (B) ' and 1 > 1 year settlement <C) ' respectively. Subsequently, barnacles s e t t l e d at a high density on the study shore i n May and September 1976. These two additional settlements were designated as ' May, 1976 settlement * and * September, 1976 settlement * correspondingly. The various size groups of B A glandula d i f f e r e n t i a t e d for each month are shown in F i g . 18. 41 Five separate annual energy budgets were constructed for each size group and shown in Table 4. When comparing the energy flow and production of the various size groups, i t was obvious that the • flay, 1976 settlement ' has contributed to the majority of the energy flow and production of the whole population (64.4 and 53.71, r e s p e c t i v e l y ) . The data of the 1 September, 1976 settlement * are based on less than one year's measurement and not d i r e c t l y comparable to the other size groups, although t h e i r energy flow and production values are only second to that of the * May, 1976 settlement » . Following in the order of decreasing values of energy flew and production are : ' > 1 year settlement (A) • > ' > 1 year settlement (E) * > • > 1 year settlement (C) * . An annual energy budget expressed i n terms of Kcal m - 2 year-* was calculated for the natural barnacle population (by summing up the values of the respective items frcm the various size group and converted into an area of 1 m2) and i s shown i n Table 5. The annual energy flow and production of the whole population were estimated to be 6666.6 and 2897.1 Kcal m - 2 year -* , respectively. The energy l o s t to mortality for each s i z e group and for the whole population were calculated and are shown i n Table 6. The t o t a l energy l o s t to mortality was 201.5 Kcal 0.08m-2 year -* , which was egualivant to 2518.8 m - 2 y e a r - 1 . Some 6 0.8 % 42 of the t o t a l energy l o s t to mortality was in form of barnacle s h e l l s . 4.8 Estimation of error involved i n using long term mean temperatures i n cal c u l a t i n g r e s p i r a t i o n energetic cost The t o t a l energetic cost of respiration (aguatic and aerial) calculated f o r the two hypothetical barnacle i n d i v i d u a l s (a and B ) based on (1) the recorded fluctuating a i r and water temperatures and (2) the Mong term* mean values of the a i r and water temperatures during the period of 1st to 30th July , 1976, are shown i n Table 7. The energetic cost of resp i r a t i o n derived from (1) and (2) differed from each other by 11.1 and 25.7 % . 43 Fig. 1 The various experimental aggregation patterns on the s e t t l i n g plates (a) • Isolated * (b) ' Pair « (c) • 9 ' (d) • 81 ' <e) * Crowded * Cd) Fig. 2. The experimental reeky shore near the P a c i f i c Environment I n s t i t u t e , Pest Vancouver, B r i t i s h Columbia (area of s t r i p approximately eguals to 0.6 by 6 m) 4 8 4 9 Fig. 3 . Seasonal changes in the c a l o r i f i c values of various tissues of BJJ . , glandula , fron* June, 1 9 7 5 to Hay, 1 9 7 6 O V A R I A N T I S S U E 7.0 n 51 Fig. 4. The monthly mean energy standing crop c f body tissue o f i n d i v i d u a l barnacles ( B A glandula ) f ront the various aggregation patterns I I ISOLATED lfr/Vfo.-«l PAIR liillllllllllllllliliiiiiiiiil 9 tsaeasaa 81 53 F i g . , 5. The monthly mean energy standing crop of eggs of i n d i v i d u a l barnacles ( glandula ) from the various aggregation patterns E N E R G Y S T A N D I N G C R O P O F E G G ( C A L I N D I V I D U A L 1 ) 55 Fig. 6. The monthly mean energy standing crop of s h e l l s of i n d i v i d u a l barnacles ( B x qlandula ) from the various aggregation patterns 56 57 Fig. 7. The monthly mean t o t a l energy standing crop of i n d i v i d u a l barnacles ( qlandula ) from the various aggregation patterns I I ISOLATED K-y'.'-J PAIR IllIMIMlllll 9 BgaaBSSa 81 oo 59 Fig. 8. Difference i n energy p a r t i t i o n between individuals ( B. glandula ) from the various aggregation patterns. Egg : Eody tissue r a t i o I I ISOLATED I-:'-'--.--.--* PAIR IBiM 81 S E P OCT NOV DEC JAN FEB MAR APR MAY 7 5 7 6 76 o 61 Fig. 9. Difference i n energy p a r t i t i o n between individuals ,{ B A qlandula ) from the various aggregation patterns. Shell : Tissue r a t i o < Z> CO to X CO Z < I I ISOLATE D PAIR tfllllllliilllllllllllilll 9 H 81 C R O W D E D 63 Fig. 10. The monthly mean production of body tissue of i n d i v i d u a l barnacles ( B.. glandula ) from the various aggregation patterns 65 Fig . 11. The monthly mean production of egg of i n d i v i d u a l barnacles ( B A glandula ) from the various aggregation patterns 6 7 Fig. 12. The monthly mean production of s h e l l of i n d i v i d u a l barnacles { B A qlandula ) from the various aggregation patterns 40 I ISOLATED K-xW-:yy-:.1 PAIR teaaaaa 81 SEP OCT NOV DEC CD 69 Fig. , 13. 0 2 consumption rate of egg masses, ovarian tissue and naupliar embryos of j r . glandula under different temperatures. •EGG MASSES TEMPERATURE ( °C ) 71 Fig. 14. A e r i a l 0 2 consumption rate of body tissue of B. glandula in r e l a t i o n to body weight and temperature O C O N S U M P T I O N ( ju l mg" ' h"1 ) 73 Fig . 15. aguatic 0 2 consumption rate of body tissue of B qlandula in r e l a t i o n to body weight and temperature O CONSUMPTION (^Img''^') 75 Fig.,16. Molting frequency cf B A gjanduja over the period June, 1975 to May, 1976.,500 in d i v i d u a l s were observed for each date Mean no. of molt / i n d i v i d u a l / month r — ^ 1 i r — i — i 1 2 n 01 s i + cn 77 Fig. 17. Faecal production rate of JBX glandula over the period June, 1975 to May, 1976. 500 individuals were observed for each date Faecal production rate ( mg faeces / mg body tissue / month ) 79 Fig. 18. The s i z e frequency d i s t r i b u t i o n {in mg dry wt.) of the barnacles sampled frcm an area of 0.08 m2 from the experimental shore during the period May, 1976 to A p r i l , 1977, Each class i s represented by separated l i n e s on the probability paper and the number of each class i s given with the l i n e . The modal value of each class can be found at the 50 % intercept • ' May, 1976 settlement • A ' September, 1976 settlement * • •••> 1 year settlement (A) ' o » > 1 year settlement (B) 1 • ' > 1 year settlement (C) 1 PERCENTAGE ? ? ? ? ! ? ? ? ? j ? ? ? ? ? ? ? ? ? ? f 9 o ^ o f o o o o f 08 81 Table 1. Proximate analysis and c a l o r i f i c value of barnacle { J i glandula ) s h e l l ] % of protein % of c h i t i n • % of CaC0 3 c a l o r i e s / mg of s h e l l 1.82 ± 0.09 0.76 ± 0.06 97.42 ± 0.04 0.134 ± 0.03 00 83 Table 2. Annual energy budgets for individual barnacles { glandula ) from the various aggregation patterns ; over the period Hay, 1975 to A p r i l , 1976. Data i n c a l o r i e s and the percentages of energy chanelied to each items are shown i n brackets by assuming that consumption eguals 100 % . (For meanings of abbreviations, see Terminology in text) C P. BT. P.E. P. S. R.AER. R.AQ. M F A P R P/A P/R ISOLATED 1402 8 55 .0 215.0 100.6 215 .7 693.3 15.8 107.5 1295.4 370.6 909.0 0.286 0.408 (100 0) (3 .9) (15.3) (7 • 2) (15 .4) (49.4) (1.1) (7.7) (92 3) (26.4) (64.8) PAIR 1304 4 50 . 1 180.1 91 .0 204 8 655.1 15.8 107.6 1196.8 321.2 859.9 0.268 0.374 (100 0) (3 .8) (13.8) (7 .0) (15 7) (50.2) (1.2) (8.3) (91 8) (24.6) (65.9) 1264 7 51 .0 164.7 75 .6 202 0 654.1 15.8 101.4 1163.3 291.4 856.2 0.250 0.340 9 (100 0) (4 .0) (13.0) (6 0) (16 0) (51.7) (1.3) (8.0) (92 0) (23.0) (67.7) 81 1061 1 6 49 3 160.6 64 7 158 3 535.3 15.8 77.5 984 1 274.6 693.6 0.279 0.396 (100 0) (4 .6) (15.1) (6 1) (14 9) (50.4) (1.5) (7.3) (92 7) (25.9) (65.4) 699. 5 27 .5 86.0 46 2 101. 6 370.3 15.8 52.2 647. 3 159.6 471.9 0.247 0.338 CROWDED (100. 0) (3 (12.3) .9) (6 6) (14. 5) (52.9) (2.3) (7.5) (92. 5) (22.8) (67.5) 85 Table 3. Laboratory energy budgets for three groups of B_. qlandula (n=40) , with independent measurement for each budget item (in c a l o r i e s ) . * C s = Consumption, derived from summation methcd, * C F = Consumption, derived from feeding experiments. (For meanings of other abbreviations, see Terminology i n text) P BT RAER M F * * V A:C g A:C F r— x 100 % . L F 1 26.7 -356.7 527.2 2392.0 44.4 22.3 2655.9 2473.0 0.992 0.991 + 7.4 % 2 -33.4 -571.6 421.7 2183.9 50.3 18.7 2069.6 2470.0 0.991 0.992 - 16.2 X 3 119.2 -222.3 831.8 2011.8 53.2 21.2 2814.9 3336.0 0.992 • 0.994 - 15.6 % 00 87 fable 4. Annual energy budgets for each in d i v i d u a l size class of barnacles on the experimental shore over the period May, 1976 to A p r i l , 1977. Data expressed in Kcal 0.08m-2 y e a r - 1 . Consumption was found by the summation method. The percentages of energy channelled to each item were shown in brackets by assuming that consumption eguals 100 % C P.BT. P.E. P.S. R.AER. R.AQ. M F A P R P/A P/R September i g 7 6 125.5 3.5 19.5 72.8 4.2 19.4 4.4 1.7 123.8 95.8 23.6 0.774 0.406 settlement (100.0) (2.8) (15.6) (58.0) (3.3) (15.5) (3.5) (1.4) (98.6) (76.3) (18.8) "^1976 • 352.7 18.3 54.9 51.2 36.7 155.5 26.6 9.3 343.2 124.4 192.2 0.362 0.647 settlement (100.0) (5.2) (15.6) (14.5) (10.4) . (44.1) (7.6) (2.6) (97.4) (35.3) (54.5) > 1 year 49.5 2.5 6.8 0 6.3 29.9 1.9 2.2 47.4 9.3 36.2 0.196 0.257 settlement (A) (100.0) (5.0) (13.7) (0) (12.7) (60.3) (3.9) (4.4) (95.6) (18.7) (73.0) > 1 year 12.4 0.3 1.4 0 1.7 8.1 0.3 0.6 11.8 1.8 9.8 0.148 0.179 settlement (B) (100.0) (2.5) (11.5) (0) (13.6) (65.1) (2.0) (5.1) (94.9) (14.1) (78.7) > 1 year 7.5 -0.1 0.6 0 1.1 5.2 0.3 0.4 7.1 0.5 6.3 0.069 0.078 settlement (C) (100.0) * (0) (7.5) ' (0) (14.8) (68.8) (4.2) (5.6) (94.4) (6.6) (83.7) 00 00 89 Table 5. The annual energy budget for the barnacle population on the experimental shore over the period May, 1976 to A p r i l , 1977. Data expressed i n Kcal m - a y e a r - 1 . Consumption was found by the summation method. The r e l a t i v e percentages of energy chanelied to each budget item were shown in brackets , by assuming that consumption eguals 100 % c P.BT. P.E. P.S. R.AER. R.AQ. M F A P R P/A P/R 6844.9 306.9 1040.2 1549.1 625.1 2725.4 • 419.0 178.3 6666.6 2897.1 3350.5 0.435 0.865 (100.0) (4.5) (15.2) (22.6) (9.1) (39.8) (6.1) (2.6) (97.4) (42.3) (48.9) V O 91 Table 6. The annual mortality loss for each in d i v i d u a l barnacle size class on the experimental shore over the period May, 1976 to A p r i l , 1977 .Data expressed i n Kcal 0.08 m~2 year-* . Body tissue Egg Shell Total September 1 9 7 6 settlement 3.3 11.7 72.7 87.6 ^ 1 9 7 6 settlement 22.9 26.7 23.8 73.4 >• 1 year settlement (A) 4.6 5.8 21.2 31.5 >^ 1 year settlement (B) 1.2 0.9 3.0 5.1 "y* 1 year settlement (C) 1.0 1.0 2.0 4.0 Total 32.9 46.0 122.6 201.5 9 3 Table 7. Comparison between the use of actual fluctuating a i r / water temperatures and *lcng tern* mean a i r / water temperatures in c a l c u l a t i n g r e s p i r a t i o n energetic cost for two hypothetical barnacle individuals 1 (1) Total r e s p i r a t i o n energetic cost (in cals) calculated from the 'long term' mean a i r and water temperatures for July (2) Total r e s p i r a t i o n energetic cost ( i n cals) calculated from f l u c t u a t i n g a i r and water temperatures as recorded by the thermograph for July,1976 ( l ) - ( 2 ) X 100 % (2) * Individual # A 14.4 16.2 -11.1 % * Individual // B 36.2 r 48.7 -25.7 % Individual // A : Body tissue = 1.28 mg, Ovarian tissue = 0.45 mg, Egg masses = 0.06 mg Individual // B : Body tissue = 4.01 mg, Ovarian tissue = 3.87 mg, Egg masses = 1.05 mg 95 (5) DISCUSSION The production of i n d i v i d u a l barnacles generally showed a positive correlation with the abundance of the plarkton in the waters. For i n d i v i d u a l s from a l l the aggregation patterns, high production values for both s h e l l , body tissue and egg were generally found i n august and September. At t h i s time the abundance of phytoplankton was high {Stockner and C l i f f , 1976) and the adults were at t h e i r juvenile stage. Shell production became neglibible after December (7 months after the barnacle settlement). Negative production of body tissue and egg was generally observed in November to January, when phytoplankton abundance was the lowest for the year (Stocker and C l i f f , 1976). High production of body tissue and egg was observed in A p r i l , during the spring bloom of the phytoplankton. Onexplicably, negative production of tissue was, however, found in Hay when the plankton abundance was presumably s t i l l high. Large seasonal fluctuations of body and ovarian tissue were observed in the present study for B A glandula . This suggests that the barnacles consumed a large amount of energy i n summer when food was abundant, and stored t h i s i n the form of body and ovarian tissues. They could then maintain themselves by remobilizing t h e i r energy when food became scarce i n the winter months. Ovarian tissue , which has a high c a l o r i f i c value and 96 biomass, appears to be a more important energy reserve than body tissue . Barnes and Barnes (1S67) found that the ovarian development was d i r e c t l y related to the abundance of food . Barnes and Archituv {1976) found that the ovarian tissue and body tissue in B.. ^langicjgs were net separated compartments and,can be mobilized during starvation. It has also been shown that the mussel Mjtilus edulis can store up glycogen reserves i n summer and r e - u t i l i z e them during winter times or starvation (Thompson and Bayne, 1972; Gabbott and Eayne, 1S73) . S i m i l a r l y , Holland and Spencer (1973) have demenstrated the loss of organic matter and the r e - u t i l i z a t i o n of the l i p i d reserves i n the starved oyster { Ostrea edulis ) spats. Such an adaptation i n energy storage may enable these animals to cope with the large seasonal fluctuation of food a v a i l a b i l i t y i n their environment. In an o v e r a l l analysis of the annual energy budgets, i t i s obvious that respiration was the most important budget item . A large proportion (64.8 to 67.7%) of energy intake of this animal was l o s t in r e s p i r a t i o n and became unavailable to the next trophic l e v e l . The next important budget item was egg production (12.3 to 15.31); followed in the order of decreasing importance by : s h e l l production (6.1 to 7.2%) > body tissue production (3.9 to 4.6%) > molting (1.1 to 2.3%). The energy channelled to egg production was some 54% 97 of the t o t a l energy channelled to production, which was some 3 times more than that to body tissue production and about 2 times that of s h e l l production. The energy committed to reproduction would be even higher than that to egg production, since semen forms a s i g n i f i c a n t portion of body tissue In the barnacles (Barnes and Barnes, 1969). (The production of semen was incorporated into the production of body tissue in the present study). Such a large energy commitment to reproduction i s comparable to that of the oyster Crassqstrea v i r g i n i c a (Dame, 1976) and C, gigas (Bernard, 1973) (48% and 32.51 of t o t a l energy to production, r e s p e c t i v e l y ) . Grahame (1977) suggested that r-selected species would have a higher reproductive e f f o r t than k-selected species. Per those r-selected animals such as barnacles and oysters, which must suffer a very high mortality during t h e i r planktonic stages and their early settlement, i t may be advantageous for them to channel a larger proportion of th e i r energy intake into reproduction i n order to ensure the propagation of the species. Moreover, the adult barnacles may suffer from a large mortality from catastrophies or intensive predation (Dayton, 1971) i n the f i r s t year after they have s e t t l e d . The large energy coumitment to reproduction shortly a f t e r t h e i r settlement (August and September) i n B•_ glandula may well be of s e l e c t i v e advantage to such an opportunistic type of l i f e cycle. ft very high assimilation e f f i c i e n c y (91.8 to 92.7% 98 from the annual energy budgets and 99.1 to 99.4% from the laboratory energy budgets) was obtained i n the present study for S A glandula , and results agree with the only other data on barnacles ( Balanus iraproyisus , Kuznetsova, 1973) .Based on the re s u l t of her feeding experiments, Kuznetsova (1973) found that the assimilation e f f i c i e n c y of B. improyisus was very high (94%) when fed with barnacle n a u p l i i , and was lower when fed cn the algae Asterione12a and Cladophora (86 and 66%, re s p e c t i v e l y ) . The high value of assimilation e f f i c i e n c y found for B^ glandula in the present study i s , however, not uncommon for carnivores. For example, the crustacean aacrocyclgps albidus feeding cn Paramecium has been reported to have a maximum assimilation e f f i c i e n c y of 97% (Klekowski and Shushkina, 1966; in Lawton, 1970). The carnivorous f i s h Megalop cyprincides has an assimilation e f f i c i e n c y of 92% when feeding on the prawn Metapenaeus monpceros (Pandian, 1967). Forster and Gabbott (1971) found that the assimilation e f f i c i e n c y of the prawns ( Palaemon serratus and Pandalus platycergs ) may be as high as 95.1 % when fed with a protein concentrated diet. Dagg (1S76) also reported a high assimilation e f f i c i e n c y (90.4 %) for the amphipod Calliojaius laeyiusculus feeding on the copepod Calanus sp. . However, most of the herbivores and other animals feeding on phyto-plankton and zoo-plankton usually have much lower assimilation e f f i c e n c i e s when compared with B A glandula . For example, the oyster Crassostrea v i r g i n i c a , which i s s i m i l a r to glandula in some aspects (e. g. both are s e s s i l e , f i l t e r 99 feeding macro-invertebrates l i v i n g i n the i n t e r t i d a l zone) have a much lower (42%) assimilation e f f i c i e n c y (Dame, 1976). The assimilation e f f i c i e n c y f o r the marine ccpepod Calamus hyerboreus and the fresh water copepod Diaptomus g r a c i l i s , both feeding on phytoplankton, had an assimilation e f f i c i e n c y of 60 and 78% respectively (Conover, 1966; Kibby, 1971). The marine crustacean Euphausia p a c i f i c a feeding on Artemia larvae had an assimilation e f f i c i e n c y of 84% (Lasker, 1966). Overestimation of assimilation e f f i c i e n c y would r e s u l t i f faecal production was underestimated. However, B. qlandula produces large, sheathed faecal p e l l e t s , which do not break up even when the water i s agitated. Their faecal p e l l e t s can therefore be recovered e a s i l y . Leaching or b a c t e r i a l degradation of faecal p e l l e t s (Johannes and Satomi, 1967) before c o l l e c t i o n may, however, have introduced an error i n the present faecal production estimations, since the fa e c a l p e l l e t s were not collected immediately after t h e i r production. Welch (1968) compiled data from the l i t e r a t u r e and found that the gross and the net production e f f i c i e n c y of aguatic poikilotherms are usually 15 to 35% and 20 to 90%, respectively. He further suggested that net production e f f i c i e n c y i s inversely proportional to assimilation e f f i c i e n c y . Although a high net production e f f i c i e n c y and gross production e f f i c i e n c y would be expected for the barnacles since they are 100 s e s s i l e and a r e l a t i v e l y small amount of energy would be required in their normal a c t i v i t i e s , low net production e f f i c i e n c y (24.7 to 28.6%) and gross production e f f i c i e n c y (22.8 to 26.4%) were found for glandula i n this study. The r e s u l t s thus give seme support to the suggestion of Welch (1968). The P/R r a t i o of j*. glandula derived from the present in d i v i d u a l energy budgets (0.34 to 0.41) resembles that for those 'short l i v e d poikilotherms' summarized by McNeill and Lawton (1970). McNeill and Lawtcn (1970) have alsc derived equations to predict P from R, and vice versa. In, applying the present values cf P and R for B A glandula to t h e i r equations; the calculated values of P and R were about 4.0% lower and 1.8% hiqher than the actual values, respectively. The present findings therefore support the findings of McNeill and Lawtcn on 'short lived poikilotherms•. Laboratory energy budget studies indicated that the consumption values derived from the summation method clos e l y agreed with those from the feeding experiments. The result appears to suggest that the present energy budget eguaticr i s reasonably accurate and ether sources of energy loss (e. g. DOB) may presumably be n e g l i g i b l e i n B.. qlandula . Moreover, the assimilation e f f i c i e n c i e s derived from the laboratory energy budgets were s i m i l a r l y high (99.1 to 99.4%) and agreed with that (91.8 to 92.7%) from the annual energy budgets. 101 Because of the large negative values of productier of body tissue and production of egg , consumption values derived from the summation method showed negative values for * Pair ' , • 81 • and *" Crowded * i n November (see appendix 3). Respiration alone cannot account for the large reabsorption of tissue . toss of energy to other sources unaccounted i n the budget eguation (e. g. DOM) may be possible but not l i k e l y , since the energy budgets balanced c l o s e l y i n the laboratory. Such a large discrepancy i n the balance of the equation i n November i s d i f f i c u l t to explain. when comparing the energy standing crop of body tissue, egg, s h e l l as well as the t o t a l energy standing crop of in d i v i d u a l barnacles frcm the various aggregation patterns, these values showed a clear decreasing trend as the degree of aggregation increases, following the order : ' Isolated * > * Paired • > ' 9 ' > * 81 • > * Crowded • . Without exception, the ' Isolated » indiv i d u a l s always have a s i g n i f i c a n t l y higher value in a l l these items when compared with the ' Crowded • individuals . This suggests that aggregation s i g n i f i c a n t l y reduces the t o t a l energy standing crop, egg, s h e l l and body tissue of an i n d i v i d u a l barnacle. In comparing the annual energy budgets derived for i n d i v i d u a l barnacles from the various aggregation patterns, i t 102 was, found that the consumption, production, production of tody tissue, egg, and s h e l l decreased with an increased degree of aggregation . The annual values of consumption, production , production of egg, body tissue and s h e l l for the ' Isolated 1 i n d i v i d u a l s was some 2.0 to 2.5 times higher than that of the * Crowded * ind i v i d u a l s , The present findings therefore, have demonstrated that crowding had a s i g n i f i c a n t , negative e f f e c t upon the consumption and production of ind i v i d u a l barnacles. The reduction in i n d i v i d u a l consumption with an increase i n the degree of crowding may po s s i b i l y r e f l e c t an increase in the in t e n s i t y of food competition among the barnacles . Amongst a l l the budget items, production of egg appears to be most sensitive to crowding. The annual production of egg of the * Isolated » individ u a l s was some 2.5 times higher than that of the * Crowded » individuals . However, the annual production of egg of the 1 Crowded * population i n a unit area of 0.61 mz was some twice that of the * Isolated ' population; because the number of individ u a l s i n the former was some 5 times larger than in the l a t t e r . For animals such as barnacles, which must suffer from a high mortality during their planktonic stages as well as during settlement, i t would appear to be far more important for the animals to maximize the absolute output of egg per unit area rather than to maximize the reproductive output per i n d i v i d u a l , in order to ensure that the population can be sustained. 103 fin increase in the degree of aggregation i s always associated with a decrease i n egg : body tissue r a t i o of the individu a l s ; i n d i c a t i n g that the uncrowded i n d i v i d u a l barnacles would channel a larger proportion of their energy into egg output, when compared with the crowded individuals . I t would appear that the • Isolated * ind i v i d u a l s , acguiring an adequate amount of energy from consumption, could afford to channel a larger amount of energy into egg output and hence produce more progeny. On the other hand, the crowded individuals , acguiring only a l i m i t i n g amount of energy , apparently conserve a larger proportion of t h e i r energy in building up body tissue and ether uses such as res p i r a t i o n . Crisp (1964) found that i n t r a s p e c i f i c competition r e s u l t i n g from high population density l i m i t s the body size and fecundity of the barnacle Balanus balanoides . Si m i l a r l y , the fecundity of the water f l e a Daphnia p u l i c a r i a (Frank, 1952); the great t i t Parus major (Klinjver, 1951) and the pend s n a i l Lymnaea elodes (Eisenberg, 1966) was found to be reduced when the population density was high. It appears that the barnacles may adjust t h e i r fecundity i n response to a high population density or a low supply of food . Comparison among the mean s h e l l : tissue r a t i o of individuals frcm the various aggregation patterns suggested yet another difference i n energy p a r t i t i o n i n the crowded and uncrowded barnacle indiv i d u a l s .The mean s h e l l : tissue r a t i o s 10 4 of the crowded in d i v i d u a l s (e. g. • Crowded ' and • 81 ' ) was always s i g n i f i c a n t l y higher than that of the uncrowded (e. q. • Isolated » ) ind i v i d u a l s ; i n d i c a t i n g that the crowded individuals may channel a larger proportion of th e i r energy i n s h e l l formation. Barnes and Powell (1950) have described the 'hummock formation* i n the crowded population of Balanus balanoides . They found that the crowded in d i v i d u a l s in the middle of a hummock have a t a l l e r s h e l l than less crowded ones on the periphery. Hummock formation was also observed i n the present study on the » Crowded * population of J A glandula . I t appears that i t would be an advantage for the crowded individuals to use a larger proportion of their energy i n building up a t a l l e r s h e l l so as to increase the general feeding area of the whole population and hence reduce the intensity of i n t r a s p e c i f i c competition for food within the crowded s e s s i l e population. Drastic seasonal changes in population structure was observed in the present population of B*. glandula . This was mainly due to the high recruitment rate of the larvae, the high mortality as well as the rapid growth rate of the young barnacles ( i . e. the • May, 1976 settlement • and « September, 1976 settlement * ). The increase i n energy standing crop (production) of a population should be egual to mortality i f the population i s stable (Hughes, 1971a; Dame, 1976). The annual mortality and production of the present Bj. glandula population 105 were both very high and sim i l a r (2 522.8 and 2896.5 Kcal m~2 y e a r - 1 respectively), suggesting that the barnacle population under current investigation was guite stable with a high turn over rate. In the present study, the energy flow and production of the natural barnacle population was estimated as 6666.6 and 2897.1 Kcal m ~ 2 yr - 1 ,respectively . In his i n t e r t i d a l association - energy flow model, Perkins (1975) estimated that the energy flow and net production (in which the production of s h e l l has not been considered) of a barnacle population i n Monterey Bay, C a l i f o r n i a { B_. glandula and Chthamalus sp., in which J3A glandula was the great majority) were 10410 and 557.6 Kcal m~2 year-* , respectively. The value of energy flow derived from his study i s therefore considerably higher than my estimation. The current estimation for production i s , on the other hand, much higher than that derived from the study of Perkins. Moreover, the low production e f f i c i e n c y (9.2 %) calculated from the study of Perkins i s not comparable to that found i n the present study. Such discrepancies may possibly due to (1) a difference i n the l o c a l i t y of study and (2) production of s h e l l was not taken into account i n Perkin's study. The energy flow of 6666.6 Kcal m - 2 y e a r - 1 derived for the barnacle population i n the present study, when compared vith that of other eguilibrium animal populations, i s among the 106 highest and only lower than that of the oyster Crassgstrea 2i£flinica (9788 Kcal m - 2 y e a r - 1 , according to Dame, 1S76). Barnacles normally feed on phyto- and zoo-plankton. The present high value of energy flow indicated there must be a large source of planktonic food f o r the barnacles and that the barnacles are s i g n i f i c a n t consumers of the near shore plankton. The high values of energy flow, production and egg production strongly suggest the functional importance of the barnacles in both the l i t t o r a l and near-shore plankton systems, and the production of 2897.1 Kcal m~2 y e a r - 1 found f o r the barnacles i n the present study i s also very high i n comparison with other studies. Dame (1976) found that 30% cf the standing crop energy of the oyster Crassostrea v i r g i n i c a was stored in s h e l l . In t h i s study, some 53.5% of production of the barnacle population was in form of s h e l l . This portion cf energy was therefore not available to those predators which only prey on the soft part of the barnacles (e. g. the gastropod Thais sp.,and the nemertean Emplectonema sp.). It i s also doubtful that barnacle s h e l l could be u t i l i z e d by other predators which prey cn whole barnacles (e. g. the s t a r f i s h Pisaster ochraceus ). This portion of energy probably passes to the decomposers. A s i g n i f i c a n t portion of energy was, however, elaborated in forms of egg and body tissue (1040.2 and 306.9 Kcal m~2 y e a r - 1 respectively) and became available to a variety of barnacle predators. The high value of egg production also indicated that 107 barnacles channelled a large amount of energy t c production of pelagic naupliar larvae during t h e i r spawning seasons (in A p r i l and August). In the case of the natural barnacle population, a very large amount of energy (2518.8 Kcal m ~ z yr - * ) was l o s t i n mortality , mostly i n form of barnacle s h e l l . Intensive crowding was observed on the experimental shore, and may have accounted for the high mortality found in the present barnacle population, esp e c i a l l y for new settlements. This form of i n t r a - s p e c i f i c competition has been observed and documented fo r barnacles by several other workers (Connell, 1960; 1961; Meadow, 1969; Branscomb, 1976). D r i f t i n g logs are common at the present study s i t e , and log battering was implicated by Dayton (1971) as a s i g n i f i c a n t source of barnacle mortality at Puget Sound. Small patches of destroyed barnacles were freguently observed at the present study s i t e , i n d i c a t i n g that log battering may have contributed to mortality. Predators, especially the s t a r f i s h Pisaste'r ochraceus and the nemertean Bmplectonema g r a c i l e (Kozloff, 1973), have been observed near the present experimental population, and may also be responsible for a portion of mortality. The gastropod Thais sp., which was considered as an important predator of B A , glandula (Connell, 1970) was, however, not found near the present study area. In coraparision with the older age groups, the younger 108 age groups usually contribute to the greater part of the t o t a l population production (Crisp, 1971). For example, S t r e i t (1S76) found that the production by the smallest size class was a very considerable part of the whole population i n the gastropod Ancjlus f l u v i a t i l i s , and the young specimens shewed considerably higher production e f f i c i e n c y than the adults. a s i m i l i a r r e s u l t was obtained i n the present study of the barnacle population : the energy flow, production, as well as the production e f f i c i e n c i e s (net and gross) decrease following the order : * May, 1976 settlement » > * September, 1976 settlement • > « > 1 year settlement (a) ' > ' > 1 year settlement (B) » > * > 1 year settlement (C) * . This suggests that the contribution to the energy flow and production of the barnacle population would decrease i n the older age groups, p o s s i b i l y due to both the reduction i n t h e i r numbers and f i l t e r i n g rate as well as their production potentials. Most of the assimilated energy i n the cider age groups was used i n respir a t i o n . Connell (1970) found that Thais eliminated nearly a l l the E x glandula at the lower t i d a l l e v e l within the f i r s t year after t h e i r settlement, and B± glandula replaced themselves and supplied Thais- by regular recruitments. I t appeared that the e f f e c t of predation by Thais could therefore maintain the barnacles i n a productive state. The majority of the energy flow and production of the population during the study period was contributed by the 1 May, 109 1976 settlement * (343.2 and 124.4 Kcal m~2 year- 1, which accounted f o r 64.4 and 53.7 % of the t o t a l energy flow and production of the whole population, respectively). The annual egg production of the • May, 1976 settlement * comprised 66.0 % of the t o t a l egg production of the whole population . The * May, 1976 settlement * was, therefore, also the most important egg producing group i n the population. The energy flow and production data of the * September, 1976 settlement * can not be d i r e c t l y comparable to the other size groups since the data were derived from measurement of l e s s than a year. However, the energy flow and production of the * September, 1976 settlement ' were the second highest i n the whole population and would therefore also be important i n contributing to the energy flow and production of the whole population. The energetic cost of resp i r a t i o n calculated for the two hypothetical i n d i v i d u a l barnacles (section 4.8) based on (1) the »long term 1 mean air/water temperatures and |2) the actual recorded temperatures d i f f e r e d to each other by 11.1 and 25.7 55, respectively. I t appears, therefore, that the use of long term mean air/water temperatures did not incur a large error in respi r a t i o n calculations i n the present study. 1 1 0 (6) LITERATURE CIT ED Andrews, R., D. C. Coleman, J. E. E l l i s and J . S. Singh, 1974. Energy flow rela t i o n s h i p in a short grass p r a i r i e ecosystem. Proc. 1st I n t l . Congr. Ecol. : Structure, functioning and management of ecosystems. The Hague, The Netherlands. September 8 -14, 1974. 22-28. Barnes, H. and Y. Achituv, 1976. The u t i l i z a t i o n of various biochemical e n t i t i e s i n gcnadally mature Balanus balanoides (L.) under starvation and feeding in the absence of copulation. J. Expt. Bar. B i o l . 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The 'long term' mean a i r and water temperatures used in the present study (Data from Environment Canada / atmospheric Envircment and I n s t i t u t e of Oceanography, the University of B r i t i s h Columbia) Mean A i r Mean water temperature ( C) temperature ( January 2.8 6.5 Feburary 4.7 6.1 March 5.9 6.8 A p r i l 8.6 9.2 May 12.0 13.8 June 14.7 15.4 July 17.0 17.9 August 16.8 17.6 September 14.5 14.4 October 10.4 13.2 November 6.3 8.6 December 4.1 7.6 125 appendix 3 . The monthly energy budget for i n d i v i d u a l barnacles < J L L ) from the various aggregation patterns {a) ' i s o l a t e d » <b) * Pair 1 (c) « 9 ' • (e) • 81 * (f) 1 Crowded * Isolated C P.BT. P.E. P.S. R.AER. R.AQ. M F A P R J u n e 7 5 23.4 4.8 8.9 4.1 3.3 . 2.3 0 0 23.4 17.7 5.6 Jul y 94.0 14.5 16.4 11.4 10.9 35.1 2.9 2.9 91.1 42.2 46.0 August 240.6 28.4 72.9 22.2 23.9 78.3 3.2 11.7 228.8 123.4 102.2 September 383.3 29.9 180.8 31.1 29.2 88.3 2.4 21.6 361.7 241.8 117.5 October 130.0 -8.6 29.5 19.6 14.1 73.8 1.4 0.4 129.7 40.5 87.8 November 29.3 0 -53.5 12.3 13.6 53.9 . 1.2 1.9 27.4 -41.3 67.4 December^,. •35.7 -18.2 -22.5 0 14.4 62.0 0 0 35.7 -40.7 76.4 J a n u a r y ^ 27.8 0 -41.0 0 16.6 51.5 0 0.6 27.2 -41.0 68.2 Feburary 69.3 0 -9.9 0 15.5 56.1 0.5 7.2 62.2 -9.9 71.6 March 76.1 0 -5.6 0 16.7 48.3 2.0 14.7 61.4 -5.6 65.0 A p r i l 209.7 41.7 15.4 0 32.7 81.9 2.2 35.8 173.9 57.1 114.5 ^ 7 6 83.7 -37.4 23.7 0 24.9 61.9 0 10.6 73.1 -13.7 86.8 Pair — . — C P .BT. P.E. P.S. R.AER. R.AQ.. M F A P R June^,. 21.6 4.5 7.6 4.1 3.2 2.2 0 0 21.6 16.2 5.4 Ju l y 89.6 14.7 12.9 9.5 11.1 35.7 2.9 2.9 86.7 37.1 46.8 August 22A.0 27.3 64.1 21.5 22.4 74.0 3.2 11.5 212.5 112.9 96.4 September 311.2 27.9 131.3 21.9 26.5 80.5 2.4 26.8 290.5 181.0 107.0 October 118.2 -5.7 17.7 20.6 13.4 70.6 1.4 0.4 117.8 32.5 84.0 November -3.5 -11.4 -51.2 0 11.0 45.3 1.2 1.6 -5.1 -62.6 56.3 December^ 42.9 -8.8 -19.0 4.6 12.2 53.9 0 0 i2.9 -23.2 66. 1 January-, / b 61.5 -7.8 1.7 8.9 14.0 44.2 0 0.4 61.0 2.8 58.2 Feburary 63.5 2.4 -12.3 0 14.4 52.4 0.5 6.1 57.4 -9.9 66.8 March 65.7 3.9 -13.9 0 15.5 45.2 2.0 13.2 52.6 -10.0 60.7 A p r i l 247.7 58.4 29.0 0 33.6 83.4 2.2 41.0 206.7 87.4 117.0 May ? 6 62.0 -55.3 12.2 0 27.6 67.7 0 9.8 52.3 -43.0 95.3 C P.BT. P.E. P.S. R.AER. R.AQ. M F . A P R J u n e 7 5 19.7 4.1 7.1 3.4 2.9 " 2.1 0 0 19.7 14.7 5.0 July 82.7 13.9 9.0 10.2 10.3 33.6 2.9 2.7 79.9 33.1 43.9 August 269.0 32.6 86.7 14.6 28.0 91.5 3.2 12.5 256.5 133.9 119.5 Septenber 260.8 14.0 109.9 19.0 23.8 73.7 2.4 18.0 242.8 142.9 97.5 October 113.7 0 16.8 15.3 12.6 67.3 1.4 0.4 113.3 32.1 79.9 November 38.2 -6.7 -2S.1 13.2 11.1 45.8 1.2 1.6 36.6 -21.6 56.9 Dece-ber,, / 5 39.3 -6.0 -25.1 0 13.1 57.3 0 0 39.3 -31.1 70.4 January-, / 0 46.9 -5.5 -10.8 0 15.0 47.7 0 0.5 46.4 -16.3 62.6 Feburary 58.0 -9,9 0 0 13.3 48.9 0.5 5.1 52.9 -9.9 62.2 March 94.4 20.3 -7.5 0 16.3 47.4 2.0 16.0 78.4 12.8 63.7 A p r i l 229.8 32.0 54.1 0 30.3 76.4 2.2 34.7 195.1 86.2 106.7 May ? 6 12.4 -37.9 -47.4 0 25.2 62.6 0 ,10.0 2.5 -85.4 87.8 81 C P.E. P.S. R.AER. R. AQ. M A P R 15.0 3.4 4.6 2.8 2.4 1.8 0 0 15.0 ' 10.8 4.2 July 68.3 9.2 8.9 9.5 8.2 27.8 2.9 1.9 66.4 27.5 36.0 August 145.4 18.3 34.1 10.9 15.9 55.5 3.2 7.6 137.8 63.2 71.4 September 182.7 13.1 59.8 16.2 18.6 60.4 2.4 12.2 170.5 89.1 78.9 October 197.6 14.4 83.5 22.1 11.9 64.1 1.4 0.3 197.3 120.0 76.0 November -28.9 -6.9 -74.4 0 9.5 40.3 1.2 1.4 -30.3 -81.3 49.7 December^,. 69.0 -5.4 16.4 3.3 9.8 45.0 0 0 69.0 14.3 54.8 January.,. / b 23.0 -6.4 -22.4 0 11.9 39.4 0 0.4 22.5 -28.8 51.3 Feburary 72.2 5.3 0 0 12.6 47.4 0.5 6.3 65.9 5.3 60.1 March 75.2 0 4.2 0 14.2 42.1 2.0 12.8 62.4 4.2 56.2 A p r i l 196.7 19.1 73.3 0 21.1 56.1 2.2 24.9 171.8 92.4 77.2 May ? 6 45.4 -14.7 -27.4 0 22.2 55.6 0 9.6 35.8 -42.0 77.3 Crowded -C P.BT. P.E. P.S. R.AER. R.AQ. M F A P R June.,. 14.1 2.8 3.3 4.3 2. 1 1.6 0 0 14.1 10.5 3.7 July 27.9 3.4 0 1.8 3.9 15.2 2.9 0.9 27.0 5.1 19.0 August 162.9 22.1 47.7 19.0 14.1 49.9 3.2 7.0 156.0 88.7 64.0 Sep tember 147.3 6.4 50. 1 16.2 14.3 48.3 2.4 9.7 137.7 72.6 62.6 October 46.0 0 -16.4 5.0 8.3 47.6 1.4 0.2 45.8 -11.5 55.9 November -40.4 -14.9 -51.5 0 4.0 20.3 1.2 0.6 -40.9 -66.4 24.3 December^ 78.7 8.1 32.9 0 6.4 31.4 0 0 78.7 41.0 37.8 January., /o 15.3 -4.5 -12.1 0 6.8 .24.9 0 0.3 15.1 ' -16.6 31.7 Feburary 59.6 6.7 8.3 0 7.8 32.1 0.5 4.2 55.4 15.0 39.9 ' March 28.1 -4.6 -8.9 0 7.5 25.1 2.0 7.1 21.1 -13.5 32.6 A p r i l 141.7 18.6 51.9 0 13.4 38.5 2.2' 17.1 124.6 70.5 51.9 May 7 f i 18.2 -16.4 -19.3 0 13.0 35.5 0 5.3 12.8 -35.7 48.5 

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