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Some features of the life history of the cockscomb prickleback : Anoplaruchus purpurescens Gill Peppar, John Lovell 1965

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SOME FEATURES OF THE LIFE HISTORY OF THE COCKSCOMB PRICKLEBACK, ANOPLARCHUS  PURPURESCENS GILL by JOHN LOVELL PEPPAR B.Sc, University of B r i t i s h Columbia, 1961 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE i n the Department of Zoology We accept t h i s thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA October, 1965 In p r e s e n t i n g t h i s t h e s i s i n p a r t i a l f u l f i l m e n t o f t h e r e q u i r e m e n t s f o r an advanced degree a t t h e U n i v e r s i t y o f B r i t i s h C o l u m b i a , I a g r e e t h a t t h e L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r r e f e r e n c e and s t u d y . I f u r t h e r a g r e e t h a t p e r -m i s s i o n f o r e x t e n s i v e c o p y i n g o f t h i s t h e s i s f o r s c h o l a r l y p u r p o s e s may be g r a n t e d by* t h e Head o f my Department o r by h i s r e p r e s e n t a t i v e s . . I t i s u n d e r s t o o d t h a t c o p y i n g o r p u b l i -c a t i o n o f t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l not be a l l o w e d w i t h o u t my w r i t t e n p e r m i s s i o n . Department o f The U n i v e r s i t y o f B r i t i s h Columbia Vancouver 8, Canada Date ^ 9 l c ^ r £3 • i i ABSTRACT The cockscomb prickleback, Anoplarchus purpures-cens G i l l , family Stichaeidae, ranges from Attu Island and P r i b i l b f Islands, Alaska, to central C a l i f o r n i a . In B r i t i s h Columbia coastal waters i t i s a bottom-dwelling i n t e r t i d a l species, geographically sympatric with A. i n s i g n i s , which appears to prefer deeper water than A. purpurescens. A. purpurescens was collected and studied at an i n t e r t i d a l s i t e at Second Narrows, Burrard Inlet, Vancouver, B r i t i s h Columbia. Morphological v a r i a t i o n within the population studied, was examined by both measurements and meristic counts. Data obtained were used to d i f f e r e n t i a t e the popu-l a t i o n of A. purpurescens used i n the study, from i t s s i b l i n g species A. i n s i g n i s . Food and feeding habits were studied over a wide range i n s i z e , with emphasis on habitat and seasonal d i f f e r -ences shown. Relative importances of various food items re f l e c t e d differences i n a v a i l a b i l i t y of organisms u t i l i z e d as food at various tide l e v e l s . Food intake i s c u r t a i l e d i n adult f i s h approaching and during the breeding season. Marking experiments were designed to examine move-ments, t e r r i t o r i a l i t y and homing behaviour. They showed i i i movements of Anoplarchus to be rather r e s t r i c t e d . F i f t y -eight percent of recaptured marked f i s h showed a homing tendency. Marked f i s h were seldom found more than 50 feet from where o r i g i n a l l y captured. T e r r i t o r i a l i t y was of the home-range type during non-breeding times of the year. With the beginning of pair formation i n advance of spawning, defended t e r r i t o r i a l i t y i s shown. Behaviour associated with courtship, parental care and i n t e r a c t i o n between the sexes subsequent to spawning, i s described. Eggs were successfully hatched and the young are described. Spawning takes place i n the months of January and February. The female Anoplarchus guards and tends i t s eggs. The newly hatched larvae show marked po s i t i v e phototaxis f o r three to f i v e days, suggesting a planktonic existence during t h i s period; they then become negatively phototactic and seek the bottom. Age and growth were examined by the length-frequency method and o t o l i t h analysis. The population was found to be composed of individuals from less than one year of age, to greater than f i v e years of age; representing year classes 1959 to 1963. Females show a s l i g h t l y f a s t e r rate of growth than males and are larger than males at every year of age. The sex r a t i o favours females over almost the iv entire range in length exhibited. The value of exponent n, i n length-weight r e l a t i o n -ship, W = c L n , was found to be 2.98585; log c, -5.31565. The sexes show a si m i l a r trend in values of c o e f f i c i e n t of condition. The c o e f f i c i e n t was at i t s lowest f o r specimens collected during the f i r s t month of the spawning season. Condition was examined on a size , sex and seasonal basis. V TABLE OF CONTENTS Page INTRODUCTION 1 THE STUDY AREA 3 Habitat. . . . . . . . . . . . . . . 5 Animal Associates 7 Predators 9 SYSTEMATICS 11 Capture and Preservation of Specimens. . . . . . . 11 Methods Employed i n the Measurements and Counts 11 Measurements. . . . . . . . . . . . . 11 Counts. . . . . . . . . . . . . . . . 13 C l a s s i f i c a t i o n . . . . . . . . 13 Description of Anoplarchus purpurescens. . . . . . 17 Results of Measurements and Counts . . . . . . . . 19 Measurements 19 Counts. . . . . . . 22 Results of T-Tests on Cockscomb Data. . . . . . 24 Discussion . . 24 FOOD AND FEEDING 26 Introduction . . . . . . . . . . 26 Method 28 Results. . . 29 Composition of Food 29 Correlation of Habitat and Types of Food Eaten. . . . . . . . . . . . . . . . . . . 29 Size of Fish vs Size of Foods Eaten . . . . . . 36 Discussion . . . . . . . . . . . . . 39 MOVEMENTS 46 Introduction . . . . . . . . . 46 Laboratory Study of Tags 49 Description of the Tag and Method of Tagging. . 50 v i Page Results. . . . . . . . . . 52 Discussion . . . . . . . . . . . . . . . . . . 53 Laboratory Study of Anesthetics . . . . . . . . . 54 Laboratory Tests on Propylene Phenoxetol . . . 55 Method 56 Results . . . . . . . . . . 56 Discussion. . . . . . . . . . . . . . . . . 58 Marking Technique . . . . . . . . . 58 Description of Experiments. 60 Results. . . . . . . . . . . . . . . 65 Replacement Studies . 65 Transplant Studies 69 Replacement-Transplant Studies Combined . . 69 Homing vs Non-Homing. ... . . . . . . . . . . 70 Incidence of Non-Homing 70 Time Elapsed to Recapture and Occur-rences of Multiple Recapture. . . . . . . . 71 Extent of Movements . . . . . . . . . . . . 73 T e r r i t o r i a l i t y . . . . . . . 74 Discussion . . . . . . . . . . . . . . . . . . 75 REPRODUCTION 79 Sexual Dimorphism . . . . . . . . . . . . . . . . 79 Gonad Measurements and Fecundity. . . . . . . . . 82 Methods. . . . . . . . . . . . 82 Results . 83 Measurements of Testes. . . . . . . . . . . 83 Measurements of Ovaries . 85 Egg Measurements. . . . . 85 Egg Counts. 85 Discussion . . . . . . 86 Hatching of Eggs and Rearing of Young i n the Laboratory. . . . . . . . . . . . . . . . . . 89 Results. . . . . . . . . . . . . . . . . . . . 90 Female and Egg Mass . 91 Laboratory Spawned Pair of Fish . . . . . . 92 Description of Spawned Out Condition. . . . 93 Description of Young 94 Discussion . . . . . . . . . . . . . . . . . . . 97 Behaviour Associated with Courtship, Parental Care and Between the Sexes Subsequent to Spawning. . . . . . . . . . . . . 101 Courtship 101 Results - Pair Number One . . . . . . . . . 103 Results - Pair Number Two . 104 Parental Care. . . . 108 Observations i n the F i e l d . 109 Observations i n the Laboratory. . . . . . . 109 v i i Page B e h a v i o u r Between Sexes Subsequent t o Spawning. . . . . . . . . . 112 D i s c u s s i o n . . . . . . . . . . . . 113 AGE AND GROWTH I n t r o d u c t i o n . . • • • • • • 119 The Length—Frequency Method . . . . . . . . . . 119 O t o l i t h s 121 R e s u l t s . . . . 122 Length-Frequency A n a l y s i s . . 122 O t o l i t h A n a l y s i s . . . . . . . * . 126 C o r r e l a t i o n o f the Two Methods o f Age D e t e r m i n a t i o n . . . . . 127 Growth Curve. . . . . . . . . . . . . . . . . . 131 Age a t S e x u a l M a t u r i t y . . . . . . . 133 Sex R a t i o . . . . . . . . . . . . . . . . . . . 133 Length-Weight R e l a t i o n s h i p and C o n d i t i o n . . . . . 133 Method. . . . . 133 F a c t o r s f o r C o n v e r s i o n Between S t a n d a r d and T o t a l L e n g t h 135 Length-Weight R e l a t i o n s h i p . . . . . 136 C o e f f i c i e n t o f C o n d i t i o n . 137 D i s c u s s i o n . . . . . . . . . . . . . . 144 SUMMARY 149 LITERATURE CITED 155 v i i i LIST OF TABLES Table Page I. Ranges and means of body measure-ments, shown i n actual measure-ments; 42 males and 42 females. . . . . . . . . 20 I I . Ranges and means of body measure-ments, shown i n frequencies into standard and head lengths; 42 males and 42 females. . 21 I I I . Measurements of cockscomb, shown in actual measurements; 19 immature males, 19 immature females; 13 mature males and 13 mature females 22 IV. Meristic counts; 20 f i s h , 4 cleared and stained . 22 V. Areas of c o l l e c t i o n of samples f o r food analysis, with comments on reference points used and general c h a r a c t e r i s t i c s of areas 30 VI. Total percentage breakdown of 242 stomachs sampled, percentage f r e -quency of occurrence of a l l items found to be food organisms. . 31 VII. C l a s s i f i c a t i o n of organisms found i n food analysis 32 VIII. Total percentage breakdown f o r each of 10 samples examined. Per-centage frequency of occurrence of each food item found i n the stomachs 34 IX. Percentage frequency of occurrence of foods eaten by d i f f e r e n t lengths of f i s h examined. Lengths arranged in standard length groupings of 5 mm each . 37 X. M o r t a l i t i e s of marked and unmarked f i s h during laboratory study of tags. . . . . . . . . . . . . 52 Table ix Page XI. Results of laboratory tests on propylene phenoxeto 1; A. Anesthe-t i z i n g and recovery times f o r three concentrations; B. Anesthetizing and recovery times observed i n 30-minute-duration experiment on three concentrations, with comments on breathing movements observed. . . . . . . 57 XII. Type and dates of marking experi-ments performed; number of f i s h marked and return of marked f i s h . . . . . . 66 XIII. Numbers and percentages ( i n paren-theses) of marked f i s h recaptured, shown fo r the positions of recaptures when found . . . . . . . . . . . 67 XIV. Percentage recapture of marked f i s h f o r each of 14 experiments performed. Mean percentage recapture shown i n A, B, and C, f o r positions of recaptures when found . . . . . . . . 68 XV. Cases of multiple recapture of i n d i -vidual marked f i s h , with dates of each recapture and positions when recaptured . . . . . . . . . . . . 72 XVI. Gonad measurements; 16 testes and 10 ovaries. . 84 XVII. Fecundity; egg counts of 10 ovaries (A) and two egg masses (B). Actual counts indicated by asterisk . 86 XVIII. Modal analysis of each of 13 samples; increases and decreases i n modal siz e between months and between two years of sampling. 125 XIX. Results of age determinations from 34 o t o l i t h s ; 19 males and 15 females. . . . . . 128 XX. Age composition of sexes (19 males, 15 females) by standard lengths, determined from 34 o t o l i t h s . 129 X T a b l e Page XXI. P e r c e n t a g e breadkown by sex o f v a r i o u s l e n g t h s e x h i b i t e d , and p r o g r e s s i v e changes i n sex r a t i o w i t h i n c r e a s i n g s i z e . 134 X X I I . E x c e r p t from d a t a t o i l l u s t r a t e method employed f o r c o m p i l a t i o n of i n f o r m a t i o n on l e n g t h - w e i g h t r e l a t i o n s h i p . 138 X X I I I . L e n g t h - w e i g h t r e l a t i o n s h i p and c o e f f i c i e n t s o f c o n d i t i o n 139 XXIV. D i f f e r e n c e s i n c o e f f i c i e n t s of c o n d i t i o n o f 46 males and 46 f e m a l e s , of s i x samples examined 142 XXV. V a r i a t i o n i n c o e f f i c i e n t s o f con-d i t i o n among i n d i v i d u a l s o f same l e n g t h ; 10 comparisons between i n d i v i d u a l s o f same s e x , 12 com-p a r i s o n s between i n d i v i d u a l s o f o p p o s i t e sex 143 x i LIST OF FIGURES F i g u r e Page 1. The s t u d y a r e a ; photographed from the west s i d e o f the o l d Second Narrows B r i d g e , B u r r a r d I n l e t . A. S e c t i o n r e f e r r e d t o by p i l l a r numbers 1 to 6. B. S e c t i o n r e f e r r e d t o by p i l l a r num-b e r s 7 to 14. 4 2. Counts; f r e q u e n c i e s o f each o f t h e f o u r c o u n t s performed . 23 3. P e r c e n t a g e f r e q u e n c y of o c c u r r e n c e of f o o d items f o r each of 10 samples examined, a r r a n g e d i n a s e a s o n a l sequence 35 4. Frequency o f o c c u r r e n c e o f f o o d s e a t e n by d i f f e r e n t l e n g t h s o f f i s h examined. L e n g t h s a r r a n g e d i n s t a n d a r d l e n g t h g r o u p i n g s o f 5 mm each. . . . . . . . . . . . 38 5. P e r c e n t a g e stomachs w i t h f o o d , o n l y d i g e s t e d m a t t e r and empty. Samples a r r a n g e d i n a s e a s o n a l sequence . . . . . . . 41 6. Diagrammatic r e p r e s e n t a t i o n s of s e c -t i o n s o f the s t u d y a r e a used f o r each m a r k i n g e x p e r i m e n t . Each s e c t i o n (20' x 35' i n the f i e l d ) i s denoted by s o l i d l i n e r e c t a n g e s , w i t h i n w h i c h , o r a d j a c e n t t o , the a r e a s o f o r i g i n a l c a p t u r e a r e shown i n br o k e n l i n e s . Arrows show d i r e c t i o n o f t r a n s p l a n t , l a c k o f arrows denotes a replacement e x p e r i m e n t . Numbers of f i s h marked i n each e x p e r i m e n t i n d i c a t e d 61 7. Graph showing r e l a t i o n s h i p of number of eggs produced t o l e n g t h o f f e m a l e . Unenclosed d o t s r e p r e s e n t d a t a c o l l e c t e d by S c h u l t z and DeLacy (1932) . 88 8. Newly h a t c h e d l a r v a ( l e s s t h a n 24 hours o l d ) ; 7.4 mm t o t a l l e n g t h . A. L a t e r a l v i e w o f l e f t s i d e ; B. V e n t r a l v i e w , f r o m a n t e r i o r t i p o f y o l k s a c , t o t a i l . . . . . . 95 x i i Figure Page 9. Length-frequency d i s t r i b u t i o n s ; t o t a l of 285 specimens, 112 males, 157 females, 16 unsexed. . . . . . 123 10. Length-frequency d i s t r i b u t i o n s of each of 13 i n d i v i d u a l samples; t o t a l of 285 specimens 124 11. Growth curves of male and female Anoplarchus, based on 34 o t o l i t h s ; 19 males, 15 females. Point of age less than one based on 15 measurements. . . . . . 132 12. Length-weight r e l a t i o n s h i p . The curve i s the graph of length-weight equation; dots represent the calculated data . . . . . 140 13. Variations i n the c o e f f i c i e n t of con-d i t i o n of males (broken l i n e ) and females ( s o l i d line) i n r e l a t i o n to season . 145 14. The c o e f f i c i e n t of condition i n r e l a -t i o n to length of f i s h . . . . . . . 145 x i i i ACKNOWLEDGEMENTS The author wishes to express his gratitude to the following persons; - to Dr. N.J. Wilimovsky, f o r his suggestion of the prob-lem and hi s encouragement and supervision throughout the duration of the study. - to Drs. B. McK. Bary,. I.E. Efford and H.D. Fisher, f o r the i r c r i t i c a l reading of the manuscript. - to the Insti t u t e of Fish e r i e s f o r providing f i n a n c i a l assistance to c o l l e c t data and f a c i l i t i e s f o r experi-mentation. - to A.E. Peden, f o r his generosity i n allowing h i s system-a t i c data to be used i n t h i s paper. - to D.A. Peppar, f o r his most appreciated assistance i n the early part of the f i e l d work. - to a l l those persons who gave encouragement and a s s i s t -ance throughout the study. 1 INTRODUCTION Anoplarchus purpurescens, commonly known as the cockscomb blenny or prickleback, i s a common i n t e r t i d a l f i s h of the P a c i f i c Northwest, belonging to the family Stichaeidae (suborder B l e n n i o i d e i ) . The present study deals with various aspects of the l i f e h i story of A. purpurescens. The data were obtained during June, 1963, to the spring of 1965, from a s i t e at Second Narrows, Burrard Inlet, Vancouver, B r i t i s h Columbia. The pricklebacks usually inhabit the cold waters, occurring from the i n t e r t i d a l area to depths of at least 200 fathoms and are bottom dwellers. Clemens and Wilby (1961) report 13 species of pricklebacks occur-r i n g i n the waters off the P a c i f i c Coast of Canada. The suborder Blennioidei i s believed to be di p h y l e t i c , the so-called "eel blennies" (e^g., pholids, stichaeids) are re a d i l y separated from the " t r o p i c a l blennies" (e.g., blenniids, c l i n i d s ) . D e t a i l s of the l i f e h i s t o r i e s of the blennioid f i s h e s , e s p e c i a l l y the "eel blennies", are not well known. Most studies have consid-ered aspects of reproduction, mainly on representatives of the families Blenniidae and Clinidae. Authors such as Guit e l (1893), Pieron (1914), Lebour (1927), Breder (1939, 2 1951) and Qasim (1955), have presented much of the i n f o r -mation known f o r a number of these l a t t e r f a m i l i e s . Studies of the "eel blennies" have been confined mainly to North A t l a n t i c forms. The pholid, Pholis gun-nellu s , has been studied by Ehrenbaum (1904), Gudger (1927) and Qasim (1955), but only aspects of reproduction were examined. D e t a i l s of the l i f e h i s t o r i e s of the P a c i f i c forms are almost unknown. Metz (1912) reported the fi n d i n g of a Xerepes fucorum guarding i t s eggs. Schultz and DeLacy (1932) presented a study of the eggs and nesting habits of Anoplarchus purpurescens. Systematically, A. purpurescens has been well studied, f i r s t l y by Hubbs (1927) and secondly, a reexamin-ation of the species by Peden ( i n press). The reproductive study made by Schultz and DeLacy (1932), however, s t i l l represents a l l that i s known on the biology of the species. 3 THE STUDY AREA The study area was located at Second Narrows, Burrard Inlet, Vancouver, B r i t i s h Columbia; a lati t u d e of 49°18* North and longitude of 123°01' West. The actual s i t e was situated on the north side of the Inlet, beneath and d i r e c t l y adjacent to the old Second Narrows Bridge (Figure 1 ) . The exposed rows of bridge p i l i n g s were used as reference points i n c o l l e c t i o n s made and i n the marking experiments, and reference w i l l be made to bridge p i l l a r numbers. Burrard Inlet i s a large t i d a l body of water, extending some 18 miles (29 Kilometres) eastward from the S t r a i t of Georgia. Six miles (9.6 Km.) east of the i n l e t of English Bay, the Inlet narrows to Lions Gate or F i r s t Narrows, a deep channel about one mile (1.6 Km.) and 1,200 feet (365.7 metres) wide. This channel opens into the main portion of Burrard Inlet, containing Vancouver Harbour. Vancouver Harbour i s about f i v e miles (8.0 Km.) long and one and one-half miles (0.8 Km.) wide. Five miles (8.0 Km.) east of F i r s t Narrows, Burrard Inlet again contracts to form Second Narrows, which i s about one mile (1.6 Km.) long and 2,500 feet (762 m.) wide. From Second Narrows, the Inlet extends eastward some seven miles (11.3 Km.) 4 A. Figure 1. The study area; photographed from the west side of the old Second Narrows bridge, Burrard In l e t . A. Section referred to by p i l l a r num-bers 1 to 6. B. Section referred to by p i l -l a r numbers 7 to 14. 5 The north shore of the Inlet i s deeply scored by r i v e r s , the largest of which are the Capilano, Lynn and Seymour. The Seymour River opens into Burrard Inlet just a few hundred feet from the study area. The shoreline of the north shore consists of t i d a l f l a t s , and f o r the most part, has a very muddy sub-st r a t e . T i d a l v a r i a t i o n at Vancouver (latitude 49°17'N; longitude, 123°07'W) i s as follows (Canadian Hydrographic Service, 1964): higher high water lower low water mean tides 12.4 2.1 large tides 14.2 -1.7 recorded extremes 16.0 -2.6 T i d a l current at Second Narrows i s strong, vary-ing i n v e l o c i t y from 0.5 to 5.5 knots (Canadian Hydro-graphic Service, 1964). Habitat I searched extensively f o r the species i n the Second Narrows area. Animals were only common i n the area beneath and d i r e c t l y adjacent to the old Second Narrows Bridge. Here, unlike areas close by, the muddy substrate 6 is well masked by sand, rocks and gravel, except f o r regions i n the upper-intertidal and lower l o w - i n t e r t i d a l zones. Along the length of the bridge (north-south d i r e c -tion) are aggregations of rocks, highly i r r e g u l a r i n siz e and shape. In the mi d - i n t e r t i d a l zone and upper low-i n t e r t i d a l zone, these rocks tend to be aggregated, and do not press f i r m l y into the substrate, thus providing many spaces f o r habitation by fishes and invertebrates. In contrast, rocks of the upper-intertidal and lower low-i n t e r t i d a l zones tend to be dispersed and also press f i r m l y into the muddy substrate. It i s under the rocks of the mi d - i n t e r t i d a l and upper l o w - i n t e r t i d a l zones where Anoplarchus abounds. The habitat extends approximately 260 feet along the length of the bridge. Collections and studies i n the f i e l d were carried out i n a t o t a l area of approximately 14,300 square feet. The l a t t e r figure includes the area beneath the bridge and the area east and west, d i r e c t l y adjacent to the bridge. The f u l l extent of habitat was only exposed on the very low low tides of less than one foot, but low tides of up to two feet were used f o r m i d - i n t e r t i d a l work. Inhabitation of the i n t e r t i d a l area demands a close association with the substrate of rocks and stones, 7 for shelter during periods of i n t e r t i d a l exposure. With i t s slender and compressed body, Anoplarchus i s able to s l i d e beneath and between rocks where other f i s h and invertebrate species would not fin d access. The shelters not only function as places of refuge during periods of i n t e r t i d a l exposure, but are important sources of bottom-dwelling invertebrates u t i l i z e d as food (for example, polychaete worms and flatworms). During the breeding season, the shelters become the nesting s i t e s . The eggs are l a i d beneath the rocks i n small depressions, eit h e r natural or made by the spawning p a i r , or i n vacant clam s h e l l s . D i s t r i b u t i o n of Anoplarchus over the area studied appears c l o s e l y correlated with the character of the sub-strate (rocks not embedded, substrate beneath damp but not muddy), but also with the d i s t r i b u t i o n of algae. Not only does green algae form a major part of the di e t , but i t i s the habitat of many invertebrates u t i l i z e d as food (for example, amphipods, isopods, mussels and l i t t o r i n e s n a i l s ) . Animal Associates In the i n t e r t i d a l zones inhabited by Anoplarchus, many of the spaces beneath the rocks are used by other f i s h and invertebrate species. The shore crabs, 8 Hemigrapsus nudus and H, oregonensis, are abundant i n the mi d - i n t e r t i d a l zone and occupy the majority of the a v a i l -able spaces beneath the stones i n the uppermost region of th i s zone. Anoplarchus and Hemigrapsus, although some-times being very close together, seldom share the same stone. This l a t t e r s i t u a t i o n was observed by Schultz and DeLacy (1932), who stated that the crabs appeared to occupy the spaces under the more loosely associated stones, while Anoplarchus occurred under the more cl o s e l y f i t t i n g stones. This i s an observation which appears to hold true fo r t h i s area as well. In t h i s zone, the hermit crab, Pagurus, i s common i n the tidepools and occasionally beneath stones. In the lower portion of the m i d - i n t e r t i d a l and lo w - i n t e r t i d a l zones, the larger crabs, Cancer magister and C. productus are located i n tidepools and beneath rocks. In the l o w - i n t e r t i d a l zone, the kelp crab, Puget-t i a , i s commonly seen beneath stones i n the area. M i d - i n t e r t i d a l and deeper, the pholids, Pfaolis  laeta and P. ornata are found, beneath rocks and i n the tidepools. Numbers seen were never great, but observation has shown that Pholis does u t i l i z e the habitat along with Anoplarchus f o r breeding purposes i n January and February. Observed egg masses of Pholis, along with the guarding 9 parents, were located lower down on the beach than those of Anoplarchus, amongst the kelp beds. Tidepool sculpins, Oligocottus maculosus, are common i n the tidepools of the mi d - i n t e r t i d a l zone and occasionally stranded beneath stones i n the area. Other larger c o t t i d s are seldom seen, although the occasional Artedius i s found stranded m i d - i n t e r t i d a l l y or lower. In the l o w - i n t e r t i d a l zone, the c l i n g f i s h , Gobiesox meandricus i s often seen attached to the under-surfaces of stones. Predators Clemens and Wilby (1961) i n t h e i r account of P a c i f i c Coast f i s h e s , mention that many of the larger species feed on small fishes such as sculpins and blennies, but the s p e c i f i c names of those f i s h consumed are not given. Members of the following f a m i l i e s most l i k e l y play a role in the predation of Anoplarchus, at least during some season of the year,, or, some time i n i t s l i f e cycle: Gadidae, Pleuronectidae, Hexagrammidae, Scorpaenidae, Cottidae and possibly Salmonidae. Stomach content analysis was not per-formed on any representatives of these f a m i l i e s , but from the knowledge of t h e i r presence i n the area ( c o l l e c t i o n s made by the Insti t u t e of Fi s h e r i e s ; U.B.C., and the accounts of Clemens and Wilby, 1961) they possibly play some r o l e . 10 A known p r e d a t o r , a t l e a s t i n the F r i d a y Harbour a r e a o f San Juan I s l a n d , Washington, i s the common g a r t e r snake, Thamnophis s i r t a l i s ( B a t t s , 1961). An e x a m i n a t i o n o f the gut c o n t e n t s o f 10 snakes by t h i s a u t h o r showed t h a t f i v e had been f e e d i n g on i n t e r t i d a l f i s h e s . F our stomachs p o s s e s s e d A n o p l a r c h u s ; two snakes w i t h one f i s h p e r stomach and the o t h e r two snakes w i t h t h r e e f i s h p e r stomach. O t h e r f i s h i n the stomachs o f t h e s e snakes were the c l i n g f i s h , G o biesox meandricus and t h e r o c k p r i c k l e -back, X i p h i s t e r a t r o p u r p u r e u s ( = E p i g e i c h t h y s a t r o p u r p u r e u s ) . 11 SYSTEMATXCS Capture and P r e s e r v a t i o n o f Specimens A l l specimens used i n t h i s p a r t o f the work were p a r t o f t h e r e g u l a r c o l l e c t i o n s made from the s t u d y a r e a . E i g h t y - f o u r specimens t a k e n f r o m June, 1963, t o May, 1964, were used f o r the v a r i a t i o n s t u d y . Upon c o l l e c t i o n i n the f i e l d , t h e f i s h were p l a c e d i n t o a 10% s o l u t i o n o f f o r m a l i n and were l a t e r t r a n s f e r r e d t o 40% i s o p r o p y l a l c o h o l . Each c o l l e c t i o n was washed f r e e o f f o r m a l i n w i t h r u n n i n g tap w a t e r f o r s e v e r a l hours b e f o r e t r a n s f e r t o a l c o h o l . The l a t t e r p r o c e d u r e was always completed w i t h i n 24 hours a f t e r c o l l e c t i o n . Thus, no samples were l e f t i n f o r m a l i n f o r more th a n 24 h o u r s , and a s t a n d a r d i z e d p r o c e d u r e was f o l l o w e d f o r a l l samples. Methods Employed i n the Measurements and Counts Measurements Most methods employed i n the measurements f o l -lowed Hubbs and L a g l e r (1958); some however, do not f o l l o w t h e i r g u i d e and are noted as f o l l o w s : 12 S t a n d a r d L e n g t h — d i s t a n c e f r o m a n t e r i o r p o i n t o f head ( t i p o f upper jaw i n t h i s s p e c i e s ) t o p o s t e r i o r p o i n t o f c a u d a l p e d u n c l e ( p o i n t m a r k i n g bases o f c a u -d a l r a y s ) . T o t a l L e n g t h — t h e l a t t e r measurement, p l u s l e n g t h o f c a u d a l f i n . Head Depth-—from p o s t e r i o r end o f cockscomb t o a p o i n t j u s t b e h i n d p o s t e r i o r end o f jaws. Depth of Body at A n u s - - p e r p e n d i c u l a r d i s t a n c e f r o m o u t s i d e edge o f d o r s a l f i n t o a n a l o p e n i n g (measurement t h e r e f o r e i n c l u d e s h e i g h t o f d o r s a l f i n at t h i s p o i n t ) . Cockscomb of Head--three d i m e n s i o n s : l e n g t h , d i s t a n c e a l o n g base o f s t r u c t u r e f rom a n t e r i o r p o i n t t o p o s t e r -i o r p o i n t ; g r e a t e s t w i d t h , w i d t h o f s t r u c t u r e ( o u t e r edge t o o u t e r edge) at a p o i n t j u s t b e h i n d p o s t e r i o r margin o f o r b i t ; g r e a t e s t h e i g h t , p e r -p e n d i c u l a r d i s t a n c e o f s t r u c t u r e near p o s t e r i o r end o f cockscomb ( j u s t p o s t e r i o r t o p o i n t o f w i d t h measurement). I n a l l measurements the s e x e s were t r e a t e d sep-a r a t e l y . As wide a range as p o s s i b l e i n s t a n d a r d l e n g t h o f the body was s e l e c t e d o f each se x . I n the cockscomb a n a l y s i s , the f i s h were s e p a r a t e d as t o immature and mature, as w e l l as t o s e x . 13 Counts Counts were made o f the d o r s a l , a n a l , p e c t o r a l and c a u d a l f i n r a y s and s p i n e s . Methods o f Hubbs and L a g l e r (1958) were employed. I n t h e a n a l f i n , t h e l a s t two r a y s were counted as two r a y s . Counts were made from p r e s e r v e d (a few c l e a r e d and s t a i n e d ) specimens. The s t a n d a r d p r o c e d u r e f o r c l e a r -i n g and s t a i n i n g was employed. The l e n g t h o f the s c a l y a r e a o f the body was a l s o examined, u s i n g c l e a r e d and s t a i n e d specimens f o r t h i s o b s e r v a t i o n . C l a s s i f i c a t i o n A n o p l a r c h u s p u r p u r e s c e n s G i l l , i s a p e r c i f o r m e t e l e o s t b e l o n g i n g t o t h e s u b o r d e r B l e n n i o i d e i , f a m i l y S t i c h a e i d a e (Regan, 1912; Hubbs, 1927; B e r g , 1941; Maku-shok, 1958). Makushok (1958) p l a c e s the f a m i l i e s S t i c h a -e i d a e , P h o l i d a e , A n a r r h i c h a d i d a e and P t i l i c h t h y i d a e w i t h i n the s u p e r - f a m i l y S t i c h a e o i d a e . W i t h i n the f a m i l y S t i c h a e i d a e he p l a c e s e i g h t s u b f a m i l i e s w i t h t h e i r t h i r t y g e nera. The genus A n o p l a r c h u s i s p l a c e d w i t h i n the sub-14 f a m i l y A l e c t r i i n a e a l o n g w i t h the genera A l e c t r i a s and P s e u d a l e c t r i a s . Thus, w i t h i n t h e s u b f a m i l y A l e c t r i i n a e , a c c o r d i n g t o Makushok, t h e r e a re t h r e e genera and f i v e s p e c i e s , A n o p l a r c h u s and P s e u d a l e c t r i a s b e i n g monotypic and A l e c t r i a s h a v i n g t h r e e s p e c i e s . A l e c t r i a s i s c o n s i d e r e d by Makushok t o be the most p r i m i t i v e genus i n the s u b f a m i l y , w i t h A n o p l a r c h u s v e r y c l o s e , b e i n g d i s t i n c t f r om A l e c t r i a s i n h a v i n g the g i l l membranes w i d e l y s e p a r a t e d by the i s t h m u s . P s e u -d a l e c t r i a s has r e t a i n e d the b r o a d l y j o i n e d g i l l membranes which are f r e e f rom t h e i s t h m u s , but has d i v e r g e d f rom A l e c t r i a s i n the f o l l o w i n g f e a t u r e s : l o s s o f s c a l e c o v e r -i n g , absence o f p a l a t i n e t e e t h and r e d u c t i o n i n both p y l o r i c caecae and de r m a l c r e s t . Hubbs ( 1 9 2 7 ), c o n s i d e r i n g A n o p l a r c h u s t o be mono-t y p i c , s u b d i v i d e d t h e genus i n t o t h r e e s u b s p e c i e s , A n o p l a r -chus p u r p u r e s c e n s i n s i g n i s G i l b e r t and Bu r k e , A.p. p u r p u r -escens G i l l and A.p. a r c h o l e p i s Hubbs. C h a r a c t e r s used f o r t h i s s u b s p e c i f i c s e p a r a t i o n were number o f d o r s a l s p i n e s , number o f a n a l r a y s and e x t e n t o f the s c a l y a r e a o f the body. Recent d a t a p r e s e n t e d by Peden ( i n p r e s s ) , demon-s t r a t e s t h a t A n o p l a r c h u s i n s i g n i s G i l b e r t and Burke and A n o p l a r c h u s p u r p u r e s c e n s G i l l , p r e v i o u s l y c o n s i d e r e d 15 s u b s p e c i e s , are v a l i d s i b l i n g s p e c i e s . A. i n s i g n i s d i f -f e r s f r om A. p u r p u r e s c e n s by p o s s e s s i n g h i g h e r m e r i s t i c c o u n t s and h a v i n g a n a r r o w e r w i d t h between the p o i n t s o f attachment o f each g i l l membrane t o the i s t h m u s . The sub-s p e c i e s w h i c h have d i f f e r e n t i a t e d w i t h i n t h e s e two s p e c i e s were not e v a l u a t e d . A. i n s i g n i s ranges from A t t u I s l a n d , A l a s k a , t o Puget Sound and t h e S t r a i t o f Juan de F u c a . , I t shows an i n c r e a s e d number o f d o r s a l s p i n e s and a n a l f i n r a y s o v e r A. p u r p u r e s c e n s . L a t i t u d i n a l v a r i a t i o n i n the number o f s p i n e s a c c o r d i n g t o Peden, i s 57-64; i n the number of a n a l r a y s , 40-46. S c a l e s on t h e p o s t e r i o r p a r t o f t h e body ex t e n d a n t e r i o r l y t o between s e v e n t h and f i f t e e n t h a n a l f i n r a y . A . p urpurescens has a wide range, from A t t u I s l a n d and the P r i b i l o f I s l a n d s , A l a s k a , t o c e n t r a l C a l i f o r n i a . L a t i t u d i n a l v a r i a t i o n i n number of d o r s a l s p i n e s a c c o r d i n g t o Peden, i s 54-59; i n number of a n a l r a y s , i s 36-42. S c a l e s on t h e p o s t e r i o r h a l f o f t h e body e x t e n d a n t e r i o r l y t o between the t w e l f t h a n a l f i n r a y and o r i g i n o f a n a l f i n o r anus. Synonymy: A n o p l a r c h u s p u r p u r e s c e n s . The f o l l o w i n g l i s t o f synonymy f o l l o w s Hubbs (1927), w i t h f u r t h e r a d d i t i o n s w h i c h have o c c u r r e d i n the 16 l i t e r a t u r e s i n c e Hubbs' paper. Ophidium, " S p e c i e s d r i t t e " , K i t t l i t z , D e n k w u r d i g k e i t e n e i n e r R e i s e nach dem r u s s i s c h e n A m e r i k a , e t c , l , 1858, p 225, f i g . 3. An o p l a r c h u s p u r p u r e s c e n s G i l l , P r o c . Acad. Nat. S c i . P h i l a . , 1861, p 262. C e n t r o n o t u s c r i s t a g a l l i G i i n t h e r , C a t . F i s h e s B r i t . Mus., 3, 1861, p 289. An o p l a r c h u s c r i s t a g a l l i G i i n t h e r , i b i d , p 564; J o r d a n , Evermann and C l a r k e , Rept. U.S. Comm. F i s h . , f o r 1928, P t . 2, p 468. A n o p l a r c h u s a l e c t r o l o p h u s J o r d a n and G i l b e r t , P r o c . U.S. Nat. Mus., 3, 1880, p 265; 3, 1880 (18 8 1 ) , p 454 ( i n p a r t ) ; J o r d a n and Jouy, i b i d , 4, 1881, p 4; Jo r d a n and G i l b e r t , i b i d , p 64 ( i n p a r t ) . A n o p l a r c h u s a t r o p u r p u r e u s J o r d a n and G i l b e r t , P r o c . U.S. Nat. Mus., 3, 1880, p 265; Bean, i b i d , 4, 1881, p 245; 4, 1881 (18 8 2 ) , p 468; J o r d a n and G i l b e r t , B u l l . U.S. Nat. Mus., 16, 1883, p 771; Bean, P r o c . U.S. Nat. Miis., 6, 1883 (18 8 4 ) , p 354; J o r d a n , Rept. U.S. Comm. F i s h . , 1885; p 910; T u r n e r , C o n t r . Nat. H i s t . A l a s k a , 1886, p 93; Eigenmann and Eigenmann, Ann. N.Y. Acad., 6, 1892, p 357; J o r d a n and S t a r k s , P r o c . C a l . Acad. S c i . , ( 2 ) : 5 , 1895, p 846; G i l l , P r o c . U.S. Nat. Mus., 18, 1895 17 ( 1 8 9 6 ) , p 150; S t a r k s , P r o c . C a l i f . Acad. S c i . , (2) :6, 1896, p 562; G i l b e r t , R ept. U.S. Comm. F i s h . , 1893 (1896), p 450; J o r d a n and Evermann, B u l l . U.S. Nat. Mus., 47, P t . 3, 1898, p 2422; P t . 4, 1900, f i g . 845; R u t t e r , B u l l . U.S. F i s h . Comm., 1898 (18 9 9 ) , p 192; J o r d a n and G i l b e r t , F u r S e a l s and F u r - S e a l I s l a n d s , 3, 1899, p 483; Osgood, N. Am. Fauna, 21, 1901, p 20; Evermann and G o l d s b o r o u g h , B u l l . U.S. Bur. F i s h . , 26, 1906 (1907), p 338; S t a r k s , Ann. Carn. Mus., 7, 1911, p 212; G i l b e r t and Burk e , B u l l . U.S. Bur. F i s h . , 30, 1910 (1912),,p 88; H a l k e t t , Check f i s h e s Canada, 1913, p 111; M i l e s , P u b l . Puget Sd. B i o l . S t a . 2, 1918, pp 79, 93; K i n c a i d , A n n o t a t e d l i s t Puget Sd. F i s h e s , 1919, pp 41, 42, f i g . 96; Bean and Weed, T r a n s . Roy. Soc. Canada, (3) , 13, 1919 (19 2 0 ) , pp 81, 82. An o p l a r c h u s F r a s e r , Can. F i e l d Nat., 35, 1921, p 48. D e s c r i p t i o n o f A n o p l a r c h u s p u r p u r e s c e n s Body m o d e r a t e l y e l o n g a t e , s l e n d e r , compressed. C a u d a l p e d u n c l e s h o r t , deep and compressed. S k i n w i t h s m a l l embedded s c a l e s , c o n f i n e d t o p o s t e r i o r h a l f o f body and e x t e n d i n g a n t e r i o r l y as f a r as 18 between twelfth anal f i n ray and o r i g i n of anal f i n or anus; portion of body anterior to t h i s naked. L a t e r a l l i n e weakly v i s i b l e ; short. Sensory (seismosensory) canals of head nor-mally developed, opening out with a constant number of pores; nasal 2, i n t e r o r b i t a l 4, p o s t o r b i t a l 7, o c c i p i t a l 4, subor-b i t a l 6, preopercular 6 and mandibular 4 (Makushok, 1958). Head small, moderately pointed i n p r o f i l e ; lacking spines, c i r r i or barbels; with a r o s t r a l - o c c i p i t a l fleshy crest . Mouth terminal, moderately large, oblique. Lower jaw included; l i p s on both upper and lower jaws fleshy. Teeth on jaws, vomer and palatines. Snout pointed i n pro-f i l e , lacking spines or other armature. N o s t r i l s paired, l y i n g just ahead of o r b i t s , one on either side of dermal crest. Eye r e l a t i v e l y small, ovate. Preopercle and opercle unarmed; posterior edge of opercle fleshy. Branchiostegal rays 5. G i l l membranes broadly united, broadly coalesced with isthmus. Fins: dorsal f i n composed of spines only, those in anterior portion weak and f l e x i b l e and acquiring more stoutness and r i g i d i t y towards rear of f i n ; commences above or s l i g h t l y i n advance of pectorals and ends at base of caudal; anal f i n composed e n t i r e l y of soft rays, commences immediately behind anus and ends at base of caudal; caudal f i n with wide base and convex outer border; pectoral f i n s 19 m o d e r a t e l y l a r g e , o u t e r b o r d e r s convex; p e l v i c s l a c k i n g . A n o p l a r c h u s p u r p u r e s c e n s i n the a r e a where c o l -l e c t e d f o r t h i s s t u d y and i n a d j a c e n t w a t e r s , can be d i s -t i n g u i s h e d f r o m a l l o t h e r b l e n n i o i d s by: absence o f p e l v i c f i n s , p r e s e n c e o f the de r m a l c r e s t on the. head, s c a l e s on the p o s t e r i o r p a r t of the body o n l y and the g r e a t w i d t h between the p o i n t s o f attachment o f each g i l l membrane t o th e i s t h m u s . The s p e c i e s was f i r s t r e c o r d e d i n B r i t i s h Colum-b i a n w a t e r s i n 1861 f r o m Vancouver I s l a n d and the mouth o f the F r a s e r R i v e r by A. Gl i n t h e r as C e n t r o n o t u s c r i s t a g a l l i (Clemens and W i l b y , 1961). Specimens were c o l l e c t e d d u r i n g the voyage o f H.M.S. Plumper. The s p e c i e s was c a l l e d t he c r e s t e d b l e n n y i n 1946 by Clemens and W i l b y , t h e American F i s h e r i e s S o c i e t y / American S o c i e t y o f I c h t h y o l o g i s t s and H e r p e t o l o g i s t s recommends s i m p l y "cockscomb" (Clemens and W i l b y , 1961). R e s u l t s o f Measurements and Counts Measurements R e s u l t s o f measurements o f body p a r t s a re shown i n T a b l e s I and I I . F o r t y - t w o males, r a n g i n g i n s t a n d a r d 20 l e n g t h f rom 51.5 t o 123.5 mm, and 42 f e m a l e s r a n g i n g i n s t a n d a r d l e n g t h from 66.2 t o 128.0 mm, were measured. T a b l e I . Ranges and means o f body measurements, shown i n a c t u a l measurements; 42 males and 42 f e m a l e s • Measurement Males Mean Females Range Range Mean Head l e n g t h 8.7- 21.8 15.77 10.7- 20.0 15.50 Head d e p t h 5.6- 15.5 10.62 7.0- 13.8 10.33 Head w i d t h 5.0- 12.8 9.02 6.1- 12.0 8.97 Snout l e n g t h 1.8- 5.7 3.62 2.1- 5.1 3.52 I n t e r o r b i t a l w i d t h 1.2- 2.8 2.00 1.4- 2.5 1.91 O r b i t l e n g t h 1.8- 3.8 2.91 1.9- 3.8 2.94 Depth body a t anus 6.9- 17.5 13.97 9.1- 19.5 14.26 Le n g t h d o r s a l 43.8- 103.0 79.98 56.0-106.8 82.25 L e n g t h a n a l 28.5- 67.1 53.07 38.0- 71.9 54.44 L e n g t h c a u d a l 5.1- 11.3 8.62 5.9-12.1 8.89 Len g t h p e c t o r a l 4.0- 9.9 7.10 4.5- 10.0 7.10 The ranges and means f o r t h e sexes a r e s i m i l a r , and do not show dimorphism. R e s u l t s o f cockscomb measurements a r e shown i n T a b l e I I I . Of immatures, 19 males, r a n g i n g i n s t a n d a r d l e n g t h f rom 51.5 t o 108.0 mm, and 19 f e m a l e s , r a n g i n g i n s t a n d a r d l e n g t h f r o m 65.0 t o 121.0 mm, were used; o f matures 13 males, from 83.0 t o 123.5 mm, and 13 f e m a l e s , from 89.0 t o 128.0 mm, were used. D i f f e r e n c e s between means were t e s t e d s t a t i s t i c a l l y . T a b l e I I . Ranges and means o f body measurements, shown i n f r e q u e n c i e s i n t o s t a n d a r d and head l e n g t h s ; 42 males and 42 f e m a l e s . Males Females Measurement I n t o S.L. I n t o H.L. I n t o S.L. I n t o H.L. Range Mean Range Mean Range Mean Range Mean Head l e n g t h 5.44--7. 12 6. 06 5. 77- -6. 81 6. 30 Head d e p t h 1.33--1.67 1.48 1.33--1. 62 1. 49 Head w i d t h 1.47--1.95 1.74 1.47--2. 13 1. 74 Snout l e n g t h 3.44--5.78 4.43 3.73--5. 72 4. 37 I n t e r o r b i t a l w i d t h 5.84--9.18 7.88 6.68--9. 53 8. 09 O r b i t l e n g t h 4.45--6.25 5.39 4.31--6. 31 5. 28 Depth body a t anus 5.93--8. 10 6. 84 5. 77- -8. 05 6. 86 L e n g t h d o r s a l 1.14--1. 28 1. 19 1. 12- -1. 30 1. 19 L e n g t h a n a l 1.69--1. 90 1. 80 1. 52- -2. 02 1. 79 L e n g t h c a u d a l 1.61--2.04 1.83 1.47--2. 08 1. 74 L e n g t h p e c t o r a l •1.95--2.64 2.23 1.82--2. 58 2. 19 22 Table I I I . Measurements of cockscomb, shown i n actual measurements; 19 immature males and 19 immature females; 13 mature males and 13 mature females. Measurement Immature Mature Range Mean Range Mean Males Length 5.9-12.0 9.05 9.8-15.9 12.6 Greatest height 1.2-2.9 1.96 2.0-4.9 3.28 Greatest width 1.2-2.5 1.91 1.9- 3.6 2.77 Females Length 6.9-13.1 9.26 9.1-12.9 11.6 Greatest height 1.3- 3.0 2.02 2.0- 3.4 2.75 Greatest width 1.4-2.8 1.95 1.8- 2.8 2.22 Counts ii Results of meristic counts are shown i n Table IV; frequencies of counts are shown i n Figure 2. Table IV. Meri s t i c counts; 20 f i s h , 4 cleared and stained. Meristic Range Mean Dorsal f i n spines Anal f i n rays Caudal f i n r a y s - p r i n c i p a l - t o t a l Pectoral f i n rays Length of scaly area - farthest forward - commonest 55-58 38-41 11-13 15-16 9-10 56.75 39.35 12.45 15.75 9.05 3- 4 anal ray 4- 5 anal ray 23 Dorsal spines Anal rays 1 5 r 1 0 - I -•I- l i l L 5 5 5 6 5 7 5 8 3 8 3 9 4 0 4 1 Count Caudal rays Pectoral rays c a i j *5 2 0 r (principal; D CT 11 12 1 3 9 1 0 Figure 2. Counts; frequencies four counts performed. 24 Results of T-Tests on Cockscomb Data A l l tests were carried out on actual measure-ments . (1) Mature males were found to be s i g n i f i c a n t l y d i f f e r e n t , at the 1% l e v e l , i n a l l three dimensions of the cockscomb from immature males. (2) Mature females were found to be s i g n i f i -cantly d i f f e r e n t , at the 1% l e v e l i n length and height, and the 5% l e v e l i n width, from immature females. (3) Mature males were not s i g n i f i c a n t l y d i f f e r -ent (5% level) from mature females i n length of cockscomb, but were s i g n i f i c a n t l y d i f f e r e n t i n height (5%. l e v e l ) , and i n width (1% l e v e l ) . (4) Immature males were not s i g n i f i c a n t l y d i f -ferent (5% level) i n any dimension of cockscomb from immature females. Discussion Available data from c o l l e c t i o n s of Anoplarchus examined by Peden ( i n press), indicates that A. i n s i g n i s prefers deeper water than A. purpurescens. Peden also 25 points out that i n B r i t i s h Columbia A. i n s i g n i s i s d i s t r i b u -ted almost e n t i r e l y i n the subtidal zone. On the basis of information gathered from meristic counts and measurements made i n t h i s study, i t appears that the i n t e r t i d a l population of Anoplarchus used was e n t i r e l y A. purpurescens. 26 FOOD AND FEEDING Introduction The feeding habits or feeding behaviour of fishes are the search f o r and ingesting of food. These should be distinguished from food habits and d i e t , which are materials ha b i t u a l l y or f o r t u i t o u s l y eaten. Food studies based on contents of digestive tr a c t s or of faeces, merely show what an animal w i l l eat. Because of natural fluctuations i n abundance, any one food organism i s not always of constant numerical a v a i l a b i l i t y . Such fluctuations of forage organisms are often c y c l i c and due to factors of t h e i r l i f e h i s t o r i e s or to c l i m a t i c or other environmental conditions. Fishes may be c l a s s i f i e d according to the amount of v a r i a t i o n i n types of food consumed by them: (1) euryphagic — f e e d i n g on a variety of foods; (2) stenOphagic—feeding on a few d i f f e r e n t types of foods; (3) monophagic —-feeding on only a single type of food. The f i s h eats those types of food to which i t i s adapted. The food of f i s h e s may be divided into several categories according to the r e l a t i o n s h i p between fishes and t h e i r food. Nikolsky (1963) divided the food of fishes 27 into three categories, r e a l i z i n g that the c l a s s i f i c a t i o n was to some extent subjective: (1) basic f o o d — t h a t which the f i s h usually consumes, com-p r i s i n g the main part of the gut contents; (2) secondary f o o d — f r e q u e n t l y found, but i n smaller amounts; (3) i n c i d e n t a l f o o d — o n l y r a r e l y enters the gut. A study of feeding habits may be of great help i n reducing l i m i t a t i o n s imposed on food habits by d i f f e r -e n t i a l rates of digestion of various food organisms. Thus, determination of when an animal feeds would f a c i l i t a t e the c o l l e c t i n g of specimens close to or during that time, so that foods i n the stomach would be r e l a t i v e l y whole, undi-gested materials, whether the organisms eaten are s o f t - or hard-bodied. A v a i l a b i l i t y of food organisms was assessed q u a l i t a t i v e l y by observation and c o l l e c t i o n of algae and invertebrates. A quantitative estimate was not attempted. The term " a v a i l a b i l i t y " i s used here as defined by Hess and Swartz (1940), implying those organisms which are capable of being eaten by the f i s h i f i t so desires. Thus, i t i s d i s t i n c t from food "preference", which as Hess and Swartz point out, should only be used to ref e r to a d e f i n i t e exercising of choice by the f i s h . 28 Method Of the f i v e most commonly used methods of analysis of food habits the method of frequency of occur-rence was selected and used i n a l l food analyses presented in t h i s study. This method of analysis characterizes the d i s t r i b u t i o n of a given food species, and together with the va r i e t y of foods, shows the feeding uniformity of a population. It does not, however, indicate the quantita-t i v e value of a given food species. In order to determine frequency of occurrence, the number of i n d i v i d u a l stomachs i n which was found each kind of food item was recorded. Results are expressed as a percentage of the t o t a l number of specimens containing food. Specimens were l e f t intact u n t i l examination of stomachs was to be performed. Standard length, sex and condition of gonads were recorded f o r each specimen. Gonads were l e f t attached to the body cavity f o r future examination. The stomach was cut free from the esophagus and i n t e s t i n e , and s l i t open. The contents were spread on a white-bottomed d i s s e c t i n g tray and sorted using a d i s -secting microscope. Examined contents were then stored i n glass v i a l s . 29 The stomach was the o n l y p o r t i o n o f the gut used f o r r e c o r d i n g f r e q u e n c y o f o c c u r r e n c e o f f o o d m a t e r i a l . I n c a s e s where e i t h e r the stomach was d e v o i d o f f o o d and/or the i n t e s t i n e was s w o l l e n w i t h m a t e r i a l , t h e i n t e s t i n e was examined. Food m a t e r i a l p r e s e n t was not r e c o r d e d as an o c c u r r e n c e o f f o o d . I n f o r m a t i o n g a t h e r e d from such exam-i n a t i o n showed t o what e x t e n t f o o d organisms were broke n down and t h e r e f o r e h e l p e d t o i d e n t i f y m a t e r i a l t h a t was p a r t i a l l y d i g e s t e d i n t h e stomach. R e s u l t s C o m p o s i t i o n o f Food The r e s u l t s a re g i v e n i n T a b l e V I . (See T a b l e V I I f o r more s p e c i f i c i d e n t i f i c a t i o n o f f o o d organisms.) C o r r e l a t i o n o f H a b i t a t and Types o f Food E a t e n E x a m i n a t i o n of d a t a r e v e a l s a c l o s e s i m i l a r i t y i n tho s e organisms e a t e n t o t h o s e organisms p r e s e n t i n zones o f t h e beach where f i s h were c o l l e c t e d . T a b l e V I I I and F i g u r e 3 show the f r e q u e n c y o f o c c u r r e n c e o f each f o o d i t e m f o r each sample o f f i s h examined. The a r e a s o f c o l l e c t i o n o f t h e s e samples and g e n e r a l c h a r a c t e r i s t i c s o f t h e s e a r e a s 30 T a b l e V. Areas o f c o l l e c t i o n o f samples f o r food a n a l y s i s , w i t h comments on r e f e r e n c e p o i n t s used and gen-e r a l c h a r a c t e r i s t i c s o f a r e a s . Date o f Sample R e f e r e n c e B o i n t s , C h a r a c t e r i s t i c s and G e n e r a l Remarks Aug., 1963 Sep., 1963 D e c . , 1 9 6 3 Mar., 1963 May, 1964 A l l c o l l e c t e d between p i l l a r s 8-14, beneath o r v e r y c l o s e l y a d j a c e n t t o the u n d e r s t r u c t u r e of the b r i d g e . Uppermost r e g i o n o f the m i d - i n t e r t i d a l sampled; c h a r a c t e r i z e d by dense growths o f mussels and h e a v i l y - b a r n a c l e d r o c k s . Tangled masses o f nemertean worms among mussels and on p i l i n g s . A few t i d e p o o l s p r e s e n t at t he b a s e s o f outer-most b r i d g e p i l -i n g s . Green a l g a e p r e s e n t . C o l l e c t e d between p i l l a r s 8-12, west s i d e o f b r i d g e . L o w - i n t e r t i d a l ; c h a r a c t e r i z e d by green a l g a e - c o v e r e d r o c k s and k e l p beds. Jun . , 1964 J u l . , 1964 Aug., 1964 Sep., 1964 Oct. , 1964 A l l c o l l e c t e d between p i l l a r s 1-6, beneath and e a s t and west s i d e s of b r i d g e . M i d -i n t e r t i d a l and l o w - i n t e r t i d a l ; c h a r a c t e r -i z e d by mussel beds and h e a v i l y b a r n a c l e d r o c k s i n t h e m i d - i n t e r t i d a l r e g i o n and k e l p beds i n the l o w - i n t e r t i d a l r e g i o n . P r e s e n c e o f p o l y c h a e t e worms v e r y appar-ent . K e l p beds o f gre e n , brown and r e d a l g a e . T i d e p o o l s p r e s e n t , u s u a l l y d r a i n on l o w e s t t i d e s . 3 1 T a b l e V I . P e r c e n t a g e f r e q u e n c y o f o c c u r r e n c e o f a l l items found t o be f o o d o r g a n i s m s . T o t a l p e r c e n t a g e breakdown o f 2 4 2 stomachs sampled. Food Organism No. o f Oc c u r r e n c e s P e r c e n t a g e A l g a e 7 7 3 2 . 3 5 Roundworms 4 2 1 7 . 6 5 Amphipods 3 7 1 5 . 5 5 Flatworms 3 0 1 2 . 6 0 Nemertean Worms 13 5 . 4 6 M u s s e l s 8 3 . 3 6 Shrimp 7 2.94 Isopods 7 2.94 N e r e i d Worms 5 2 . 1 0 L i t t o r i n e S n a i l s 2 0.84 O t h e r 10 4 . 2 0 Stomachs w i t h f o o d p r e s e n t 1 9 5 8 0 . 5 8 - stomachs w i t h i d e n t i f i -a b l e f o o d 1 5 9 8 1 . 5 4 - stomachs o n l y w i t h d i g e s -t e d m a t t e r 3 6 1 8 . 4 6 Stomachs c o m p l e t e l y d e v o i d o f f o o d 4 7 1 9 . 4 2 a r e shown i n T a b l e V. Two more s t r i k i n g s i m i l a r i t i e s w h i c h a r i s e f r o m the d a t a a r e : ( 1 ) h i g h f r e q u e n c y o f o c c u r r e n c e o f numertean worms i n those samples c o l l e c t e d i n t h e upper m i d - i n t e r t i d a l zone; and ( 2 ) t h e i n c r e a s e i n f r e q u e n c y o f o c c u r r e n c e o f p o l y c h a e t e worms ( r e f e r r e d t o as roundworms i n the t a b l e s and f i g u r e s ) and amphipods i n thos e samples c o l l e c t e d i n the m i d - i n t e r t i d a l and lower z o n e s . O b s e r v a t i o n and c o l l e c -t i o n o f organisms i n t h e s e zones has shown the a v a i l a b i l i t y 32 of t h e s e f o o d i t e m s . T a b l e V I I . C l a s s i f i c a t i o n o f organisms found i n the f o o d a n a l y s i s . A f t e r , B o r r a d a i l e and P o t t s (1961); J o h n s t o n and Snook (1927); R i c k e t t s and C a l v i n (1962); S c a g e l (1957). A. A l g a e CHLOROPHYCOPHYTA Order U l o t r i c h a l e s F a m i l y U l v a c e a e - U l v a RHODOPHYCOPHYTA B. I n v e r t e b r a t e s PLATYHELMINTHES C l a s s T u r b e l l a r i a O r d e r P o l y c l a d i d a - N o t o p l a n a NEMERTEA C l a s s E n o p l a O r d e r H o p l o n e m e r t i n i Emplectonema g r a c i l e  Paranemertes ANNELIDA C l a s s P o l y c h a e t a F a m i l y N e r e i d a e - N e r e i s F a m i l y T e r e b e l l i d a e - Thele'pus ARTHROPODA C l a s s C r u s t a c e a S u b c l a s s M a l a c o s t r a c a Order P e r a c a r i d a Suborder Isopoda Cymothoidea- C i r o l a n a Suborder Amphipoda Gammaridea F a m i l y Gammaridae F a m i l y T a l i t r i d a e F a m i l y Amphithoidae O r d e r E u c a r i d a Suborder Decapoda Macrura F a m i l y CragOnidae- Crago 3 3 Table VII (cont'd.) B. Invertebrates MOLLUSCA Class Gastropoda Order Prosobranchiata Suborder Pectinibranchiata Family L i t t o r i n l d a e - L i t t o r i n a Class Lamellibranchiata Order F i l i b r a n c h i a t a Family Mytilidae- Mytjlus edulis 1 Table VIII. Total percentage breakdown for each of 10 samples examined. Percentage frequency of occurrence of each food item found i n the stomachs. % Stomachs Aug. Sep. Dec. Mar. May Jun. J u l , Aug. Sep; Oct. 1963 1963 1963 1964 1964 1964 1964 1964 1964 1964 With food 76.2 82.4 75.0 42.9 94.4 94.7 85.7 82.5 90.9 59.1 -iden. food 57.2 50.0 25.0 35.8 94.4 81.5 78.6 80.0 54.6 59.1 -only d i g . matt. 19.0 32.4 50.0 7.1 0 13.2 7.1 2.5 36.3 0 Empty 23.8 17.6 25.0 57.1 5.6 5.3 14.3 17.5 9.1 40.9 Number sampled 21 34 16 14 18 38 28 40 11 22 Algae 20.0 20.0 0 40.0 37.1 48.8 35.6 26.5 50.0 22.7 Roundworms 6.6 0 20.0 0 31.4 11.6 •27.. 0 14.3 25.0 22.7 Amphipods 6.6 15.0 0 0 20.0 20.9 13.5 22.4 12.5 0 Flatworms 26.6 15.0 20.0 0 0 0 0 22.4 0 50.0 Nemerteans 13.3 30.0 60.0 40.0 0 0 0 0 0 0 Shrimp 0 0 0 0 0 9.3 8.1 0 0 0 Mussels 0 10.0 0 0 0 0 8*1 6.1 0 0 Isopods 6.6 5.0 0 0 2.9 2.3 2.6 4.1 0 0 Nereids 6.6 0 0 0 2.9 2.3 0 0 0 0 L i t t o r i n e s 0 5.0 0 20.0 0 0 0 0 0 0 Other 13.3 0 0 0 5.7 4.7 2.6 4.1 12.5 0 CO 35 60 50 40 30 (3 U § 2 0 <_ <_ U X 10 c IsOh u. S40 c (9 ^ 3 0 o 0_ Aug, 1963 Sept, 1963 Dec, 1963 March, 1964 May, 1964 Month of Sample 20 10 June, 1964 July, 1964 Aug, 1964 Se pt, 1964 I Oct, 1964 Month of Sample 0 Algae H Roundworms ED Amphipods § Flatworms B Nemertean worms DD Shrimp 0 Mussel § Isopods U Nereid worms 0 Littorine snails Figure 3. Percentage frequency of occurrence of food items f o r each of 10 samples examined, arranged i n a seasonal sequence. 36 L i t t o r i n e s n a i l s are confined to the high- and mid - i n t e r t i d a l zones. These s n a i l s were found only i n samples co l l e c t e d i n the upper m i d - i n t e r t i d a l zone. Small shrimp were commonly seen under rocks and algae i n the low-i n t e r t i d a l zone and showed up only i n stomachs of those f i s h that were collected i n the m i d - i n t e r t i d a l zone and lower. The frequency of occurrence of algae i n stomach contents was the greatest of a l l food items, and was almost e n t i r e l y confined to green algae. Traces of red algae were found i n some individuals that were co l l e c t e d at the low-i n t e r t i d a l l e v e l , but occurrences were rare. Brown algae were never found i n stomachs examined. Green algae are available from upper to lower levels of the i n t e r t i d a l , whereas browns and reds are con-fined to lower levels and the s u b - t i d a l . The frequency of occurrence of algae rose with the samples col l e c t e d from the m i d - i n t e r t i d a l and lower but the type of algae remained the same. Size of Fish vs Size of Foods Eaten Table IX and Figure 4 show frequency of occur-rence of various foods eaten by d i f f e r e n t lengths of f i s h Table IX. Percentage frequency of occurrence of foods eaten by d i f f e r e n t lengths of f i s h examined. Lengths arranged i n standard length groupings of 5 mm each. Standard A f e e Round- Amphi- F l a t - Nemer- s h r i m D Mussels Isooods N e r e i d L i t t o r i n e Length & worms pods worms teans y worms s n a i l s 50- 55 100.0 55- 60 60- 65 20.0 40.0 20.0 65- 70 35.7 42.9 7.1 70- 75 40.0 20.0 20.0 20.0 75- 80 33.3 16.7 80- 85 38.5 23.1 15.4 23.1 85- 90 35.0 20.0 15.0 10.0 90- 95 45.8 20.8 20.8 4.2 95- 100 37.5 15.6 25.0 9.4 100- 105 24.4 9.8 ,14.6 9.8 105- 110 29.0 16.1 6.4 22.6 110- 115 34.8 17.4 13.0 21.7 115- 120 28.6 4.8 14.3 14.3 120- 125 50.0 50.0 7.1 7.1 16.7 33.3 10.0 5.0 5.0 4.2 3.1 3.1 3.1 12.2 4.9 2.4 7.3 2.4 6.4 3.2 6.4 3.2 3.2 4.3 4.3 9.5 4.8 4.8 9.5 38 Algae Roundworms Amphipods 8 c 3 .8 o u c CT «S> t-LL 52.5 6 7 5 62.5 97.5 112.5 1275 52.5 6 7 5 82.5 97.5 112.5 127.5 5 2 . 5 6 7 5 8 2 . 5 97.5 112.5 127.5 Flatworms Ncmcrtcans Shrimp I I I I ^ •—•T"^ 52.5 67.5 82.5 97.5 112.5 1275 52.5 6 7 5 8 2 . 5 9 7 5 112.5 127 5 52.5 6 7 5 82.5 9 7 5 112.5 1275 Mussels Isopods Nereids 4P-52.5 67.5 82.5 97.5 112.5 1275 5 2 . 5 67.5 82 .5 9 7 5 T12.5 1275 52.5 6 7 5 8 2 . 5 9 7 5 112.5 127.5 Littorines Other I I 1 1 ( I ) 1 ~ T 5 2 5 67.5 8 2 . 5 9 7 5 112.5 127.5 52 .5 67 .5 8 2 . 5 97.5 112.5 1275 Standard Length (mm) Figure 4. Frequency of occurrence of foods eaten by d i f -ferent lengths of f i s h examined. Lengths arranged i n standard length groupings of 5 mm. each. 3 9 e x a m i n e d . -T h e f o u r m o s t c o m m o n l y f o u n d f o o d o r g a n i s m s i n t h e s t o m a c h s , a l g a e , r o u n d w o r m s , a m p h i p o d s and f l a t w o r m s r e s p e c t i v e l y , w e r e f o u n d t o b e p r e s e n t o v e r t h e g r e a t e s t s i z e r a n g e . N e m e r t e a n w o r m s , s h r i m p , a n d i s o p o d s w e r e c o n f i n e d t o l a r g e r i n d i v i d u a l s . T h e m u s s e l s e a t e n w e r e o f v e r y s m a l l s i z e a n d o c c u r r e d s o m e w h a t s p o r a d i c a l l y o v e r a w i d e r a n g e i n s i z e . N e r e i d w o r m s o c c u r r e d v e r y s p o r a d i c a l -l y o v e r a w i d e s i z e r a n g e , b u t s m a l l e r i n d i v i d u a l s w e r e o n l y c o n s u m i n g p a r t s o f t h e s e w o r m s . L i t t o r i n e s n a i l s w e r e c o n f i n e d t o a l a r g e r s i z e g r o u p . M a t e r i a l s c l a s s i f i e d a s " o t h e r " i n c l u d e d s u c h i t e m s a s s p a w n , c r u s t a c e a n a p p e n -d a g e s a n d b a r n a c l e s h e l l s , a n d w e r e a l m o s t e n t i r e l y c o n -f i n e d t o l a r g e r i n d i v i d u a l s . D i s c u s s i o n S a m p l e s o f f i s h w e r e c o l l e c t e d d u r i n g t h e l o w e s t l o w - t i d e o f t h e d a y . T h i s t i d e o c c u r s f r o m j u s t b e f o r e a n d s h o r t l y a f t e r m i d n i g h t , i n f a l l a n d w i n t e r , t o b e f o r e a n d a f t e r : n o o n , i n s p r i n g a n d s u m m e r . T h e t i d e p r e v i o u s t o t h i s l o w l o w - t i d e t h e r e f o r e , e i t h e r a p p e a r s d u r i n g d a y l i g h t o r d a r k n e s s d e p e n d i n g o n t h e s e a s o n . I t i s d u r i n g t i d e s p r e v i o u s t o t h e l o w l o w - t i d e t h a t A n o p l a r c h u s m u s t f e e d . 40 F i g u r e 5 s h o w s t h e s e a s o n a l c h a n g e s i n p e r c e n -t a g e s o f s t o m a c h s w i t h a n d w i t h o u t f o o d . T h e p e r c e n t a g e o f e m p t y s t o m a c h s a n d s t o m a c h s w i t h o n l y d i g e s t e d m a t t e r , r i s e s i n f a l l a n d w i n t e r s a m p l e s , b u t r e m a i n s a t a l o w l e v e l d u r i n g l a t e s p r i n g a n d summer s a m p l e s . T h e h i g h i n c i d e n c e o f f o o d i n s t o m a c h s o f s p r i n g a n d summer s a m p l e s i n d i c a t e s t h a t t h e f i s h w e r e n o t o n l y f e e d i n g w e l l d u r i n g t h e s e m o n t h s , b u t t h a t t h e y w e r e f e e d i n g d u r i n g t h e h i g h t i d e b e f o r e t h e a r e a w a s u n c o v e r e d b y w a t e r a n d c o l l e c t i o n s m a d e . T h e i n c r e a s e i n p e r c e n t a g e o f e m p t y s t o m a c h s i n f a l l a n d w i n t e r c o u l d b e t a k e n t o i n d i c a t e t h a t f i s h w e r e n o t f e e d i n g t o t h e same e x t e n t a s t h e y w e r e i n s p r i n g a n d s u m m e r . B u t t h e i n c r e a s e i n s t o m a c h s w i t h o n l y d i g e s t e d m a t t e r i n d i c a t e s t h a t f i s h w e r e s t i l l f e e d i n g b u t t h e t i m e o f f e e d i n g h a d s h i f t e d , s o t h a t o r g a n i s m s p r e s e n t i n t h e s t o m a c h s w e r e now d i g e s t e d b e y o n d i d e n t i f i c a t i o n . F e e d i n g w a s t h e r e f o r e t a k i n g p l a c e d u r i n g t h e e a r l y p a r t o f t h e p r e v i o u s h i g h - t i d e o r d u r i n g t h e l o w - t i d e b e f o r e t h e h i g h -t i d e . T h e d a t a t h e n s u g g e s t t h a t A n o p l a r c h u s i s a d a y -l i g h t f e e d e r . T h u s , t h e r e w o u l d t e n d t o be a g r e a t e r p e r -c e n t a g e o f i d e n t i f i a b l e f o o d i t e m s i n t h e s t o m a c h s o f t h o s e f i s h c o l l e c t e d d u r i n g d a y l i g h t h o u r s ( s p r i n g a n d summer s a m p l e s ) , f o r t h e f i s h w o u l d h a v e b e e n a b l e t o f e e d d u r i n g 41 in "D «*- "D © X / K \ / M a o < D (0 c 3 i. 6b O 2 o a E o c o 2 a < o m o m o N m CM 36D+U0DJ0d F i g u r e 5 . P e r c e n t a g e s t o m a c h s w i t h f o o d , o n l y d i g e s t e d m a t t e r a n d e m p t y . S a m p l e s a r r a n g e d i n a s e a s o n a l s e q u e n c e . 42 the previous high-tide. Food i n the stomachs of those f i s h c o l l e c t e d i n f a l l and winter would have been present i n the gut f o r a much greater time and therefore would consist of only digested matter by the time of actual c o l l e c t i o n . Varying rates of digestion and v a r i a t i o n i n actual feeding times, would account f o r those food items which appeared r e l a t i v e l y undigested, i n f a l l and winter samples. Anoplarchus appears to be u t i l i z i n g as food those organisms present and available i n the p a r t i c u l a r habitat at the time of feeding. In samples from August, 1963, to and including, March, 1964, foods found to be present i n the stomachs examined can be c l a s s i f i e d as follows: (1) basic food — nemertean worms, algae and flatwOrms; (2) secondary food — roundworms, amphipods and l i t t o r i n e s n a i l s ; (3) i n c i d e n t a l food — m u s s e l s , isopods and nereid worms. The remaining 1964 samples show a d i s t i n c t change in frequency of occurrence of various food items and thus, in t h e i r r e l a t i v e importance as foods. In these samples, foods found to be present may be c l a s s i f i e d as follows: (1) basic food — algae, roundworms and amphipods; (2) secondary food — flatworms; (3) i n c i d e n t a l f o o d — a l l remaining food organisms could be considered i n thi s category. 43 These changes i n r e l a t i v e importance of various food items show differences i n a v a i l a b i l i t y of organisms u t i l i z e d as food at d i f f e r e n t t i d a l l e v e l s of the i n t e r t i d a l zone. It i s not implied that changes r e f l e c t food prefer-ences, or a d e f i n i t e exercising of choice by the f i s h . Alga was a very frequently occurring food item i n almost every sample studied. Regardless of differences i n habitat, the only alga eaten was green algae (except f o r rare traces of red algae), even i n the l o w - i n t e r t i d a l where a l l three types of algae are re a d i l y a v a i l a b l e . Thus, i n the u t i l i z a t i o n of t h i s food item there appears to be a d e f i n i t e exercising of choice by the f i s h . On the basis of previous categorization of organ-isms u t i l i z e d as food, Anoplarchus can be considered rather stenophagic i n i t s food habits. Food habits of small and large Anoplarchus are not markedly d i f f e r e n t . Those foods c l a s s i f i e d as basic foods were frequently occurring over almost the entire s i z e range studied. Some larger food organisms however, were absent i n stomachs of smaller-sized i n d i v i d u a l s . For the most part, these foods were c l a s s i f i e d as i n c i d e n t a l foods. Nereid worms, largest of organisms u t i l i z e d as food, occur-red rather sporadically over a wide range i n f i s h s i z e . It appears that smaller i n d i v i d u a l s were u t i l i z i n g these worms 44 as food only once they are dead, as small portions only showed up i n the stomachs of these f i s h . Food intake appears to be c u r t a i l e d i n adult Anoplarchus approaching and during the breeding season (Jan-uary and February). Percentages of stomachs devoid of food were highest f o r the samples of December, 1963, March, 1964, and October, 1964; 25.0%, 57.1% and 40.9%, respectively. The May, 1964, sample showed an extremely large increase i n food intake, with only 5.6% of stomachs devoid of food. As w i l l be discussed i n d e t a i l i n a l a t e r section, p a i r i n g up of sexes begins approximately a month before actual spawning takes place. P a i r i n g becomes noticeable i n December, and of course i s extremely prevalent during the period January-March when actual breeding i s taking place. During the time from p a i r i n g to breeding, movements are l i k e l y to be at a minimum and therefore the only food con-sumed would be that which was washed i n close to the breed-ing p a i r . Schultz and DeLacy (1932) expressed the same opin-ion when they examined stomachs of guarding females and found them to be devoid of food. Observation of pairs kept i n aquaria f o r spawning purposes has revealed that the f i s h w i l l feed i f they are provided with food but no deleterious e f f e c t s are observed i f they are not fed. Two such pairs, plus a female and her 45 e g g s , w e r e k e p t f o r a p e r i o d e x c e e d i n g o n e m o n t h w i t h o u t f o o d o f a n y k i n d ; s p a w n i n g a n d h a t c h i n g o c c u r r e d . 4 6 MOVEMENTS I n t r o d u c t i o n T h e m a i n o b j e c t i v e s o f m a r k i n g e x p e r i m e n t s w e r e t o o b t a i n d a t a o n m o v e m e n t s , e x t e n t o f h o m e - r a n g e a n d t h e p o s s i b i l i t y o f a h o m i n g b e h a v i o u r . T h e t e r m " h o m e - r a n g e " i s a d o p t e d h e r e a s i t was d e f i n e d b y D i c e ( 1 9 5 2 ) , n a m e l y " t h e a r e a o v e r w h i c h a n i n d i v i d u a l a n i m a l h a b i t u a l l y t r a v e l s w h i l e e n g a g e d i n h i s u s u a l d a i l y a c t i v i t i e s " . T h e h o m e - r a n g e i n c l u d e s a l l f e e d -i n g s i t e s , b r e e d i n g s i t e s a n d p l a c e s o f r e f u g e h a b i t u a l l y u s e d b y t h e i n d i v i d u a l a n d a l l o t h e r a r e a s r e g u l a r l y t r a v -e r s e d b y h i m . T h i s d e f i n i t i o n h a s no i m p l i c a t i o n o f a n y t e n d e n c y o n t h e p a r t o f t h e i n d i v i d u a l t o e x c l u d e o t h e r s o f t h e s p e c i e s f r o m t h e a r e a . T h e h o m e - r a n g e , o r p a r t o f a h o m e - r a n g e , d e f e n d e d a g a i n s t o t h e r m e m b e r s o f t h e same s p e c i e s , i s t e r m e d a " t e r r i t o r y " ( D i c e , 1 9 5 2 ) . Many o t h e r w o r k e r s i n a n i m a l b e h a v i o u r u s e t h e t e r m t e r r i t o r y t o r e f e r o n l y t o t h i s t y p e o f d e f e n d e d a r e a ( N i c e , 1 9 4 1 ; C o l l i a s , 1 9 5 1 ; C a r p e n t e r , 1 9 5 8 ) . T h e d i s t i n c t i o n b e t w e e n d e f e n d e d a n d h o m e - r a n g e t e r r i t o r y c a n n o t a l w a y s b e m a i n t a i n e d i n p r a c t i c e , s i n c e o u r k n o w l e d g e o f t h e n a t u r a l b e h a v i o u r o f a n i m a l s u n d e r t h e 47 varied natural conditions of t h e i r l i v e s i s often insuf-f i c i e n t to enable us to say to what extent others are excluded from the t e r r i t o r y . Natural populations cannot usually be observed to learn whether or not a f i s h i s defending an area, although marking experiments may i n d i -cate that movement i s r e s t r i c t e d . The features that are defended and even the tendency to defend at a l l , may vary with the season and with the p h y s i o l o g i c a l state of the i n d i v i d u a l (Dice, 1952; Etkin, 1964). Thus, as Etkin (1964) pointed out, using " t e r r i t o r y " as an unqualified term would be useful f o r such cases, and the term could r e a d i l y be changed and q u a l i f i e d as home-range or defended as our knowledge j u s t i f i e s . The idea that some fishes have a home-range has been amply substantiated by several recent workers, notably Newman (1956), Funk (1957), M i l l e r (1957), Williams (1957), Bardach (1958), and Gerking (1959). Most of the work of these authors hinges on the assumption that i f a marked f i s h i s caught two or more times i n a ^restricted area, i t i s l i k e l y that the f i s h has occupied t h i s area f o r a sub-s t a n t i a l period of time. As Gerking (1959) and Williams (1957) pointed out, any quantitative expression about si z e of the home-range and degree of straying from t h i s area, describes 48 techniques of the investigator as much as behaviour of the f i s h . The best that can be hoped is to determine whether or not the movements of a particular species are restricted and to get a general idea about the degree of movement. Homing behaviour of fishes has been quite exten-sively studied, the main emphasis having been on freshwater species. The term "homing" has been used in a variety of -senses. Usually homing implies the return of mature fish to the place of their own genesis for the purpose of spawn-ing, for example, reproductive homing, as defined by Lindsey, et al.- (1959). The typical example of homing of this type is the salmon, but i t has also been shown for the shad (Hollis, 1948; Vladykov, 1950). More general types of homing, not necessarily associated with reproduction, exist in many f i s h . Gerking (1959) in a summary on non-reproductive homing, defined i t as "the choice that a fish makes between returning to a place formerly occupied instead of going to other equally probable places." Williams (1957) implies, in his defin-ition, a periodic return of a certain animal to a certain area that is small compared to the total home-range. Such homing i s known in a variety of intertidal animals. The tidepool fishes show a remarkable homing characteristic. The gill-finned goby, Bathygobius 49 soporator, has been shown by Beebe (1931) and Aronson (1951) to possess such a behaviour. The f i s h leave t h e i r home-pools at high tide and f i n d t h e i r way back at low tide by jumping d i r e c t i o n a l l y across the sand separating the pools. Williams (1957) demonstrated that the wooly sculpin, Clino-cottus analis and the opaleye, G i r e l l a nigricans, come back to the same tidepools on successive low t i d e s . Eastman (1962) showed the same behaviour i n the tidepool sculpin, Oligocottus maculosus. It i s l i k e l y that such a behaviour i s also shown by the surf perch, Amphigonopterus aurora Hubbs, 1921). Among invertebrate i n t e r t i d a l animals, the lim-pets e s p e c i a l l y , have been shown to possess such a behavi-our; Acmaea (Hewatt, 1940), L o t t i a (Wells, 1917) and the European P a t e l l a (Orton, 1914, 1928; Jones, 1948). Some limpets, however, do not home ( V i l l e e and Groody, 1940). Laboratory Study of Tags Before marking i n the study area commenced and during the f i r s t few marking experiments, f i s h were marked and observed i n the laboratory. The f i r s t sample was c o l -lected on June 25, 1963, a second sample on August 8, 1963, and a t h i r d sample on September 13, 1963. 50 The f i r s t sample (14 f i s h ) was d i s t r i b u t e d equally between two aquaria; a s t a i n l e s s s t e e l tank of f i v e gallons capacity and an iron tank of approximately ten g a l -lons capacity. Both aquaria at the s t a r t of the tests were devoid of sand and plants but each had a few rocks. Tagging of the f i r s t sample was not done u n t i l the morning of June 27, 1963. At such time nine f i s h were tagged; four f i s h i n the larger tank and f i v e i n the smal-l e r tank, the remaining f i s h i n each tank were l e f t untag-ged as controls. Description of the Tag and Method of Tagging The tag used consisted of two components, a small (2 by 2 mm) embroidery bead and a short piece of nylon f i s h i n g leader of 3 or 4 pound te s t . The bead was attached to one end of the leader leaving a long t r a i l i n g piece f o r attaching the tag to the f i s h . Tagging of the f i r s t sample of f i s h was perfor-med without the aid of an anesthetic. An anesthetic was used i n a l l the f i e l d work, the use of which w i l l be d e s c r i -bed i n a l a t e r section. Tagging was accomplished i n the following manner. The f i s h was placed on a small cheese-cloth-covered board dampened with sea water. Corners of 51 the c l o t h were folded over the body of the f i s h , leaving only the anterior and mid-portion of the dorsal f i n exposed. In t h i s way wriggling movements were held at a minimum and the area to which the tag was to be applied was i n view. The tag was then threaded on to a f i n e needle and the needle passed through the musclature of the back just under the dorsal f i n . The bead end of the tag was then pulled closer to the body of the f i s h and the leader end sewn through the bead and a knot formed. Superfluous leader material was removed a f t e r the knot was formed. The second sample of f i s h (9 fish) was arranged on August 9, 1963. Five f i s h were tagged and the rest were l e f t as controls. These f i s h were placed i n the larger aquarium. The l a s t sample (8 fish) was placed i n the smal-l e r tank on September 14, 1963. No individ u a l s of t h i s sample were tagged. A l l f i s h were observed d a i l y for any noticeable e f f e c t s as a re s u l t of the tags or any shedding of tags. To simulate more natural conditions, on August 21, 1963, the larger tank was set up with sand and rocks. By equipping the tank i n t h i s manner the f i s h were allowed to burrow under the rocks and present the tags with a 52 condition that they would be more l i k e l y to come across in the f i e l d . Occasional draining of the tank (with the f i s h l e f t i n the tank) increased t h e i r burrowing. Results Table X summarizes the r e s u l t s obtained, showing deaths which occurred and shedding of tags. Table X. M o r t a l i t i e s of marked and unmarked f i s h observed during laboratory study of tags. Date of No. No. Dates of No. No. Marking Marked Controls M o r t a l i t i e s Marked Unmarked June 27, 1963 4 3 July July Aug. 8, 29, 5, 1963 1963 1963 1 1* 1 1 1 June 27, 1963 5 2 Aug. Sep. Sep. 26, 12, 13, 1963 1963 1963 1 1 1 4 Aug. 9, 1963 5 4 Aug. Oct. 26, 17, 1963 1963 3 2 2 2 Sep. 14, 1963 - 8 Sep. Sep. 18, 21, 1963 1963 5 3 * Tag shed. 53 D i s c u s s i o n The swimming o f the f i s h d i d not appear t o be hampered i n any way by the t a g s . A l l f i s h , whether tagged o r n o t , f e d a c t i v e l y , b o t h on the bottom o f t h e tank and mid-water i n t h e t a n k , when g i v e n such f o o d as f r o z e n b r i n e - s h r i m p . Of a l l the f i s h tagged o n l y two d e a t h s c o u l d be d i r e c t l y a t t r i b u t e d t o the w e a r i n g o f t a g s . These two f i s h showed t h e f o l l o w i n g c h a r a c t e r i s t i c s b e f o r e t h e i r subsequent d e a t h : (1) a r e a o f body d i r e c t l y around base of t h e t a g w e l l "worn" and "raw" i n appearance; (2) a d j a c e n t a r e a o f body a p a l e g r a y i s h i n c o l o u r ; ( 3 ) g e n e r a l p a l e n e s s of body c o l o u r ; (4) r e d u c t i o n i n a c t i v i t y d u r i n g f e e d i n g , t o t o t a l absence o f f e e d i n g . Development, o f t h e s e symptoms was c h a r a c t e r i s t i c and once a f i s h r e a c h e d "4" d e a t h r e s u l t e d i n a m a t t e r o f a few d a y s . These symptoms,, h o w e v e r , were th e extreme con-d i t i o n s and were o n l y shown toy two f i s h , one o f which had been w e a r i n g t h e t a g f o r 3 2 days and the o t h e r f o r 3 5 d a y s . A n o t h e r f i s h however, l i v e d f o r 60 days w i t h o u t showing 54 thi s extreme tag damage to the body. The death of t h i s l a t t e r f i s h was d i r e c t l y a t t r i b u t a b l e to an a i r supply f a i l -ure . Sl i g h t "wearing" of the f l e s h around the tag was shown by the other tagged f i s h , but conditions did not develop to the extreme shown by the two f i s h previously described. Deaths, whether the f i s h were tagged or not, were mainly attributable to a i r supply f a i l u r e or some con-tamination of the water supply. One such a i r supply f a i l u r e occurred on August 23, 1963, and was not noticed u n t i l Aug-ust 26, 1963. Five f i s h (3 tagged, 2 untagged) died as a re s u l t of t h i s f a i l u r e . The l a s t sample of f i s h died within a week of c a p t i v i t y . Cause of death was unknown. The tag used was considered useful, even i f i t could only be worn f o r one month i n the f i e l d . Marking experiments planned were to be of short-term and therefore the tag would be r e l i a b l e . Laboratory Study of Anesthetics From the previous lab work on tags and the tagging method i t became obvious that t h i s method would not prove s a t i s f a c t o r y i n the f i e l d unless an anesthetic was used. The anesthetic would help speed up the process of tagging 55 and also help reduce the harmful a f t e r - e f f e c t s of the tag-ging procedure. Tricaine Methanesulfonate (MS222) was the f i r s t t r i e d and used i n the f i e l d . The concentration used was e n t i r e l y a r b i t r a r y and because of t h i s , no f i s h were allowed to remain i n the solut i o n longer than the time i t took them to become anesthetized. Thus the number of f i s h which could be properly handled at any one time was small. At the con-centrations used, the time to anesthetize was on the average about 2 to 3 minutes. About 4 to 5 minutes were required f o r recovery. MS222 was used i n a l l f i e l d experiments conducted i n 1963, but i n 1964, another anesthetic was laboratory tested and used i n a l l subsequent f i e l d experiments. Laboratory Tests on Propylene Phenoxetol Bagenal (1963), showed that propylene phenoxetol was a useful anesthetic f o r periods of up to one hour when at a so l u t i o n of 0.01 to 0.025% and can be used at normal laboratory and aquarium temperatures with f i s h of a wide range i n s i z e . He concluded that f o r anesthetizing f i s h f o r a period of more than three hours, i t must be used with great care since the strength of solution and temperature 56 are c r i t i c a l . On January 10, 1964, a sample of eight f i s h was col l e c t e d and tests were applied on January 11, 1964. Method Three concentrations, 0.05%, 0.025% and 0.1% respectively, were set up with one f i s h per so l u t i o n . The time to anesthetize (ceasing of a l l movement when touched, and r o l l i n g over on back or side) was noted. Each f i s h was then placed i n fresh sea water and the time to recover (maintenance of an upright posture and movement) noted. Three concentrations, 0.05%, 0.025% and 0.033% respectively, were set up with two f i s h i n the f i r s t two concentrations mentioned and one f i s h i n the remaining con-centration. The f i s h were then l e f t i n the solutions f o r a t o t a l of 30 minutes, with the time to anesthetize noted i n each case. Each f i s h of the sample of eight, was used only once. The Temperature varied about 14.0°C. Results Table XI summarizes the data obtained from tests Table XI, Results of laboratory tests on propylene phenoxetol; A. Anesthetizing and recovery times f o r three concentrations; B. Anesthetizing and recovery times observed i n the 30 minute duration experiment on three concentrations, with comments on breathing movements observed. Cone, of Soln. % Time to Anesthetize (min.) Time to Recover (min.) Remarks A. 0.05 0.025 0.10 6 18 4 9 14 20 B. 0.05 5 36 No external signs of breathing (opercular movements ceased); 15 minutes i n fresh sea water before f i r s t signs of breathing observable. 0.025 17 13 Slight opercular movements observable; breathing increased almost instantaneously upon placement into fresh sea water. 0.033 8 21 Opercular movements ceased; breathing increased a f t e r about two minutes i n fresh sea water. cn 58 performed on the anesthetic. Anesthetizing and recovery times are shown with comments on breathing movements obser-ved i n the duration experiment. Discussion After a l l tests were completed, the f i s h were placed i n an aquarium and observed over a two-day period f o r any noticeable harmful e f f e c t s from the anesthetic. No such e f f e c t s were observed. The concentration of 0.033% propylene phenoxetol was chosen to be used i n f i e l d marking experiments. This concentration gave a f a i r l y short (8 min.) anesthetizing time and the recovery time was not as long as the 0.05% soluti o n (21 min. as opposed to 36 min.), a f t e r 30 minutes duration i n the appropriate anesthetic solutions. Consider-ing the use to which the anesthetic i n the f i e l d would be put, the 0.033% concentration appeared to be s u i t a b l e . Marking Technique Preceding each marking experiment, tags were made and set into a perforated sheet of p l a s t i c (3 M x 3"). Three p l a s t i c p a i l s at the time of marking were standard; one f o r c o l l e c t i n g , one f o r anesthetizing and one f o r allowing 59 recovery from anesthetic. C o l l e c t i o n and marking of f i s h was done during the lowest tide of the day. Locating f i s h was simply a matter of turning over stones i n the appropriate habitat. Fish located i n such a manner were captured with gloved hands. A p a i r of nylon-cotton gloves, dampened with sea water, was used f o r handling a l l l i v e f i s h (except those anesthetized). Unnecessary damage to the f i s h due to s l i t h e r i n g through the hands and f a l l i n g to the ground, was kept at a minimum by wearing gloves. Once, captured, f i s h were placed into a p l a s t i c p a i l of fresh sea water. When a number (usually not exceed-ing more than 15 f i s h at any one time) had been collected, they were transferred to the anesthetic sol u t i o n . The actual marking procedure was as follows: The anesthetized f i s h was l a i d out on a dampened cheesecloth-covered p l a s t i c sheet (4" x 8"). A needle which had been previously threaded with the tag, was then passed through the musclature of the back, just under the dorsal f i n , approximately 1 to 1 1/2 inches from the most anterior point of t h i s f i n . At t h i s point, the body of the f i s h i s held somewhat more r i g i d than the posterior half of the body which undulates rapidly during locomotion. The free end of the nylon leader was then c i r c l e d back and sewn through the 60 bead and the tag secured i n place. Free ends were then clipped off with a small p a i r of manicure c l i p p e r s . In most marking experiments, the l e f t pectoral f i n was removed, as a check i n case of any tag shedding. The f i n was clipped at i t s base with the manicure c l i p p e r s . Such f i n clipped f i s h were observed i n aquaria and no d e l i t e r i o u s e f f e c t s were observed. Description of Experiments Sections of the beach where f i s h were marked w i l l be referred to by bridge p i l l a r numbers. The sections used for each experiment are shown diagrammatically i n Figure 6, Experiment 1. July 5, 1963, 18 f i s h marked, between p i l l a r s #9-13, beneath the bridge. Three d i f f e r e n t bead colours were used. Marked f i s h were replaced to where captured. Searching f o r marked f i s h was done over a sub-sequent two-day period. Experiment 2. July 18, 1963, 32 f i s h marked, between p i l l a r s #10-12, beneath the bridge. Three d i f f e r -ent bead colours were used. Marked f i s h were transplanted to tidepools, 15-35 feet away from where captured. Obser-vation f o r marked f i s h was done over a subsequent three-day period. 61a Figure 6. Diagrammatic representations of sections of study area used f o r each marking experiment. Each section (20' x 35' i n the f i e l d ) i s denoted by s o l i d l i n e rectangles, within which, or adjacent to, the areas of o r i g i n a l capture are shown i n broken l i n e s . Arrows show d i r e c t i o n of transplantment, lack of arrows denotes a replace-ment experiment . Number of f i s h marked i n each experiment indicated. 61 Expt.1. Expt.2. Expt.4. r 1 i 12 i i J >• » - — i n i i J !6 i i i I 1 r — i • e i !4| ! J i J Expt.3. Expt.7. !lSr i I 8 i — T V- ! ! -4 h i ! * ~ I 1 1 1 1 L — J l - _ _ J Expt.8. Expt.9. r — - \ 1 1 1 T SB-f 1 1 1 1 L.J Expt.10. IS fr"lY~/! ^ II 7! N" _ I I _ J I . Expt.12. Expt.11. 23 Expt .13. i N 1 3 / l ! N t T ! i 3 • i i ! i i T i [k Expt .14. ^ 1 3 ^ • 1 3 7 : i i i ! j 62 Experiment 3. August 2, 1963, 26 f i s h marked, between p i l l a r s #12-14, beneath the bridge. Two d i f f e r e n t bead colours were used. Marked f i s h were transplanted to tidepools, 15-35 feet away from where captured. Observa-t i o n f o r marked f i s h was done over a subsequent two-day period. Experiment 4. August 17, 1963, 16 f i s h marked, between p i l l a r s #9-10, beneath the bridge. Three d i f f e r -ent bead colours were used. Marked f i s h were replaced, but to tidepools, within 1-3 feet from where o r i g i n a l l y cap-tured. Searching f o r marked f i s h was done over a subse-quent two-day period. Experiment 5. August 27, 1963, three f i s h were colle c t e d during the low ti d e , marked and replaced to where found when the area was just covered by water. Observation was then made from a boat u n t i l the f i s h could no longer be seen. Further observation was made during the next low tide that the area could be searched on foot. Experiment 6. November 7, 1963, 12 f i s h marked, replaced, and the rocks painted. Observation was made during the subsequent high t i d e . Experiment 7. January 11, 1964, 19 f i s h marked, between p i l l a r s #10-11, beneath the bridge. Two d i f f e r e n t 63 bead colours were used. Marked f i s h were transplanted to tidepools, 1-6 feet away from where captured. Observation was made on the following day. Experiment 8. January 12, 1964, 13 f i s h marked, between p i l l a r s #11-14, beneath the bridge. Three d i f f e r -ent bead colours were used. Marked f i s h were transplanted to tidepools, 1-6 feet away from where captured. Observa-ti o n f o r marked f i s h was not done u n t i l January 24, 1964, at which time the enti r e area from p i l l a r s #9-14 was searched. Experiment 9. May 13, 1964, 32 f i s h marked, between p i l l a r s #2-5, adjacent to the bridge, west side. Three d i f f e r e n t bead colours were used. Marked f i s h were transplanted to very small tidepools, just beneath the bridge, a maximum distance of 18 feet but usually 6-10 fee t . Observation was done over a subsequent three-day period. Experiment 10. May 14, 1964, 24 f i s h marked, between p i l l a r s #10-11, beneath the bridge and between p i l l a r s #4-5, adjacent to the bridge, east side. Two d i f -ferent bead colours were used. The former mentioned f i s h were transplanted to a tidepool, a maximum distance of 15 feet. The l a t t e r f i s h were transplanted a maximum of 20 feet to a large draining body of water. Observation was 64 made over a subsequent two-day period. Experiment 11. June 10, 1964, 56 f i s h marked, between p i l l a r s #1-6, adjacent to the bridge, west side. Four d i f f e r e n t bead colours were used. Marked f i s h were transplanted to very small tidepools just beneath the bridge, 6-10 feet away from where captured. Observation was made over a subsequent three-day period. Experiment 12. June 11, 1964, 19 f i s h marked, between p i l l a r s #4-6, adjacent to the bridge, west side. Beads of one colour were used. Marked f i s h were trans-planted as i n l a t t e r experiment. Observation was done over a subsequent two-day period. Experiment 13. July 7, 1964, 32 f i s h marked, between p i l l a r s #4-6, adjacent to the bridge, west side, - and between p i l l a r s #3-4, adjacent to the bridge, east side. Three d i f f e r e n t bead colours were used. Marked f i s h from the west side of the bridge were transplanted as i n the l a t t e r two experiments. Those on the east side of the bridge were transplanted to the draining area used in experiment 10, 8-10 feet from where captured. Obser-vation was done over a subsequent four-day period. Experiment 14. July 8, 1964, 26 f i s h marked, between p i l l a r s #2-4, adjacent to the bridge, west side. 65 Two d i f f e r e n t bead colours were used. Marked f i s h between p i l l a r s #3-4 were transplanted to the small tidepools just beneath the bridge. Those between p i l l a r s #2-3 were trans-planted equally between the tidepools just beneath the bridge and the open water just west of where they were captured, both transplanting distances were of the same magnitude (about 6-10 f e e t ) . Observation was done over a subsequent three-day period. Further observations f o r marked f i s h were made on July 22, 1964, July 24, 1964, August 4, 1964, August 5, 1964, August 7, 1964, September 3, 1964 and September 4, 1964. Table XII outlines the marking experiments per-formed, showing the number of f i s h marked, and the number of returns of marked f i s h i n actual numbers and percentages. Replacement Studies A l l markings of t h i s type were carried out i n 1963, and were confined to sections where only small num-bers of f i s h could be located. Recaptured f i s h were c l a s -s i f i e d as being either, i n the areas of o r i g i n a l capture, or i n some other area. Percentage recapture has been expressed i n two ways: as a percentage of the total.number 66 marked, and as a mean p e r c e n t a g e o f the t o t a l number o f e x p e r i m e n t s performed o f t h i s type ( i . e . , f o u r r eplacement e x p e r i m e n t s , p e r c e n t a g e r e c a p t u r e f o r each, mean p e r c e n t a g e o f f o u r e x p e r i m e n t s ) . T a b l e X I I . Type and d a t e s o f mark i n g e x p e r i m e n t s p e r f o r -med; number o f f i s h marked and r e t u r n o f marked f i s h . E x p t . Date Type o f No. No. P e r c e n -No. Experiment Marked Recap. tage 1 J u l y 5, 1963 repl a c e m e n t 18 3 16.6 2 J u l y 18, 1963 t r a n s p l a n t 32 3 9.4 3 Aug. 2, 1963 t r a n s p l a n t 26 1 3.8 4 Aug. 17, 1963 replacement 16 1 6.3 5 Aug. 27, 1963 repl a c e m e n t 3 1 33.3 6 Nov. 7, 1963 re p l a c e m e n t 12 6 50.0 7 J a n . 11, 1964 t r a n s p l a n t 19 2 10.4 8 J a n . 12, 1964 t r a n s p l a n t 13 0 0 9 May 13, 1964 t r a n s p l a n t 32 5 15.6 10 May 14, 1964 t r a n s p l a n t 24 2 8.3 11 June 10, 1964 t r a n s p l a n t 56 15 26.7 12 June 11, 1964 t r a n s p l a n t 19 4 21.1 13 J u l y 7, 1964 t r a n s p l a n t 32 4 12.5 14 J u l y 8» 1964 t r a n s p l a n t 26 3 11.5 P e r c e n t a g e r e c a p t u r e o f marked f i s h i s shown i n .Table X I I I . The p o s i t i o n s o f the 11 r e c a p t u r e s a r e shown i n t h i s l a t t e r T a b l e and i n T a b l e XIV, where a mean p e r c e n -tage r e c a p t u r e i s shown f o r the f o u r e x p e r i m e n t s performed. The p e r c e n t a g e o f marked f i s h found where o r i g i n a l l y cap-t u r e d was 100%. T a b l e X I I I , Numbers and p e r c e n t a g e s ( i n p a r e n t h e s e s ) o f marked f i s h c a p t u r e d , shown f o r p o s i t i o n s o f r e c a p t u r e s when f o u n d . A l l r e p l a c e m e n t e x p e r i m e n t s were performed i n 1963. 1963 Replacement T r a n s p l a n t E x p e r i m e n t s Experiments 1964 T r a n s p l a n t E x p e r i m e n t s Combined 1963-1964 T r a n s p l a n t E x p e r i m e n t s Combined 1963-1964 R e p l a c e . - T r a n s p l . E x p e r i . Combined No. marked 49 58 221 279 328 No. r e t u r n s 11 ( 22.5) 4( 6.8) 35(15.8) 39(13.9) 50(15.2) No. r e t u r n s found: where marked 11(100.0) 2(50.0) 16(45.7) 18(46.2) 29(58.0) i n t r a n s p l a n t a r e a 1(25.0) 3( 8.6) 4(10.2) 4 ( 8.0) o t h e r area 1(25.0) 16(45.7) 17(43.6) 17(34.0) O) <1 68 T a b l e XIV. P e r c e n t a g e r e c a p t u r e o f marked f i s h f o r each o f 14 m a r k i n g e x p e r i m e n t s p e r f o r m e d . Mean p e r c e n -tage r e c a p t u r e shown i n A, B, and Cj f o r p o s i -t i o n s o f r e c a p t u r e s when f o u n d . P o s i t i o n o f R e c a p t u r e s Where Found Date o f No. Where T r a n s p l . O t h e r M a r k i n g Recap. Marked A r e a A r e a J u l y 5, 1963 3 3(100.0) J u l y 18, 1963 3 2( 66.6) 1(33.3) Aug. 2, 1963 1 1(100.0) Aug. 17, 1963 1 1(100.0) Aug. 27, 1963 1 1(100.0) Nov. 7, 1963 6 6(100.0) J a n . 11, 1964 2 1( 50.0) 1(50.0) J a n . 12, 1964 0 May 13, 1964 5 4 ( 80.0) 1(20.0) May 14, 1964 2 2(100.0) June 10, 1964 15 3 ( 20.0) 2( 13.3) 10(66.6) June 11, 1964 4 2( 50.0) 2(50.0) J u l y 7, 1964 4 2( 50.0) 2(50.0) J u l y 8, 1964 3 2( 66.6) 1( 33.3) A. Replacement e x p e r i m e n t s . Mean p e r c e n t a g e o f marked r e c a p t u r e s f o u n d : where marked -~ 100.0 o t h e r a r e a — — — 0 B. T r a n s p l a n t e x p e r i m e n t s . Mean p e r c e n t a g e o f marked f i s h f ound: where marked -----— 53.69 i n t r a n s p l a n t a r e a — 16.29 o t h e r a r e a - — - — - 29.99 C. R e p l a c e m e n t - T r a n s p l a n t e x p e r i m e n t s combined: Mean p e r c e n t a g e o f marked f i s h f ound: where marked - — 67.17 i n t r a n s p l a n t a r e a — • — • — 11.28 o t h e r a r e a — 20.76 69 T r a n s p l a n t S t u d i e s M a r k i n g s o f t h i s type were c a r r i e d out i n 1963 and 1964. Re c a p t u r e d f i s h were c l a s s i f i e d as b e i n g i n the 1 a r e a s o f o r i g i n a l c a p t u r e , i n t h e a r e a s to whi c h they were t r a n s p l a n t e d , o r i n some o t h e r a r e a . P e r c e n t a g e r e c a p t u r e i s e x p r e s s e d i n t h e same manner as p r e s e n t e d i n the r e p l a c e -ment s t u d i e s . P e r c e n t a g e r e c a p t u r e o f marked f i s h i s shown i n T a b l e X I I I , w i t h t h e e x p e r i m e n t s performed i n 1963 and 1964, shown s e p a r a t e l y and combined. The p o s i t i o n s o f the 39 r e c a p t u r e s a r e shown i n t h i s T a b l e and i n T a b l e XIV, where a mean p e r c e n t a g e r e c a p t u r e i s shown f o r 10 t r a n s p l a n t e x p e r i -ments p e r f o r m e d . The p e r c e n t a g e o f marked f i s h found where o r i g i n a l l y c a p t u r e d was 46.2%; a mean p e r c e n t a g e r e c a p t u r e o f 53.69%. R e p l a c e m e n t - T r a n s p l a n t S t u d i e s Combined The p o s i t i o n s o f the 50 r e c a p t u r e s a r e shown i n T a b l e X I I I , and i n T a b l e XIV, where a mean p e r c e n t a g e r e c a p t u r e i s shown f o r t h e 14 e x p e r i m e n t s p e r f o r m e d . The p e r c e n t a g e o f marked f i s h found where o r i g i n a l l y c a p t u r e d was 58.0%; a mean p e r c e n t a g e r e c a p t u r e o f 67.17%. 70 Homing vs Non-Homing Marked f i s h l o c a t e d i n t h e areas o f t h e i r o r i g i n a l c a p t u r e can be r e f e r r e d t o as "homing" f i s h . A l l o t h e r r e c a p t u r e s o f marked f i s h , whether i n the a r e a s t o which t h e y were t r a n s p l a n t e d , o r l o c a t e d i n o t h e r a r e a s , can be r e f e r r e d t o as "non-homers" o r " s t r a y s " . I n c i d e n c e o f Non-Homing Four f i s h (8.0%) were r e c a p t u r e d i n t h e a r e a s t o which they were t r a n s p l a n t e d ( T a b l e X I I I ) . These f i s h had e i t h e r not moved d u r i n g the subsequent time between o b s e r -v a t i o n s , o r i f they had moved, r e t u r n s were made t o t h e same a r e a s . Seventeen f i s h ( 3 4 . 0 % ) , a mean p e r c e n t a g e o f 20.76%, accounted f o r r e c a p t u r e s l o c a t e d i n o t h e r a r e a s ( T a b l e X I I I ) . T h i s i n c i d e n c e o f s t r a y i n g can be c o n s i d e r e d d i s t i n c t from the l a t t e r , i n t h a t a d e f i n i t e movement away from where o r i g i n a l l y marked and from where t r a n s p l a n t e d , was e x e c u t e d , w i t h no i n d i c a t i o n o f any r e t u r n d u r i n g the time^between o b s e r v a t i o n s . 71 Time Elapsed to Recapture and Occurrences of Multiple Recapture Search f o r marked f i s h was carried out two to three days subsequent to marking (except f o r one replacement study, where i t was made on the high tide immediately f o l -lowing the low). A l l marked f i s h recaptured from replacement experiments were located during t h i s time period. With the replacement experiments there were no instances of multiple recapture of marked f i s h . Recaptures from transplant markings were taken from one to thirty-one days subsequent to marking. Twenty-two f i s h were located at some time i n the three-day observa-tion period d i r e c t l y a f t e r marking, one f i s h was found 15 days afte r marking and the remaining 16 f i s h were located 26 to 31 days subsequent to marking. Four f i s h from the transplant experiments were recaptured f o r a second time. Another f i s h was seen f o r the second time,.but escaped recapture. The dates of each recapture and the positions of these f i s h when recaptured, are shown i n Table XV. In three of f i v e occurrences of multiple recapture, the f i s h were located f o r the second time i n the area occu-pied on t h e i r f i r s t recapture, although only one f i s h was Table XV. Cases of multiple recapture of in d i v i d u a l marked f i s h , with dates of each recapture and positions when recaptured. Date of Date of F i r s t Date of Second Marking Recapture Recapture June 10, 1964 July 9, 1964 July 22, 1964 Recaptured away from where o r i g -i n a l l y marked, but same area each time. July 7, 1964 July 9, 1964 July 10, 1964 Recaptured where o r i g i n a l l y marked each time. July 7, 1964 July 10, 1964 July 11, 1964 Recaptured away from where o r i g -i n a l l y marked, but same area each time. July 8, 1964 July 9, 1964 July 11, 1964 F i r s t recapture where o r i g i n a l l y marked, second i n other area. June 10, 1964 July 10, 1964 Aug. 4, 1964 Was seen only f o r the second time, movements cannot be traced <1 to 73 l o c a t e d i n the s p o t o f i t s o r i g i n a l c a p t u r e , each t i m e . As w i t h a l l r e c a p t u r e s from t r a n s p l a n t e x p e r i m e n t s , r e c a p t u r e d f i s h were r e p l a c e d t o the a r e a s o f o r i g i n a l t r a n s p l a n t . E x t e n t o f Movements R e s u l t s of t r a n s p l a n t e x p e r i m e n t s show t h a t A n o p l a r c h u s i s d e f i n i t e l y c a r r y i n g out movements d u r i n g p e r i o d s o f t i d a l c o v e r a g e . I n d i c a t i o n s a r e , however, t h a t t h e s e movements are r e s t r i c t e d . S t r a y i n g o f f i s h from t h e i r a r e a s o f o r i g i n a l c a p t u r e was o b s e r v e d t o be o f two t y p e s ; f i s h r e m a i n i n g i n t h e i r a r e a s o f t r a n s p l a n t , o r , f i s h s t r a y -i n g t o o t h e r a r e a s . F i s h l o c a t e d i n t h e i r a r e a s o f t r a n s -p l a n t had e i t h e r remained i n the s e a r e a s between o b s e r v a t i o n p e r i o d s , o r had c i r c u l a t e d and r e t u r n e d t o t h e s e a r e a s . The fo r m e r a s s u m p t i o n seems v e r y u n l i k e l y , i n t h a t two o f the f o u r f i s h found i n such a r e a s were l o c a t e d a l o n g time a f t e r t h e t h r e e - d a y o b s e r v a t i o n p e r i o d i m m e d i a t e l y f o l l o w i n g mark-i n g . I t appears t h e r e f o r e t h a t t h e s e f i s h had r e t u r n e d t o the a r e a s a f t e r h a v i n g moved d u r i n g t i d a l c o v e r a g e . The t r a n s p l a n t a r e a s were l o c a t e d a t a range o f 6 t o 35 f e e t from the a r e a s o f o r i g i n a l c a p t u r e . Those f i s h r e c a p t u r e d i n o t h e r a r e a s were l o c a t e d at a d i s t a n c e o f 6 to about 75 f e e t , from the a r e a s o f o r i g -i n a l c a p t u r e . O n l y a v e r y s m a l l number (3 f i s h ) however, 74 were found a t a d i s t a n c e g r e a t e r than 50 f e e t . B o t h i n s t a n c e s o f s t r a y i n g , t h e r e f o r e , i n d i c a t e a r a t h e r l i m i t e d movement. Marked f i s h r e c a p t u r e d from replacement e x p e r i -ments i n d i c a t e a s t r o n g tendency t o remain i n a g i v e n a r e a . R e c a p t u r e s f rom t h e s e e x p e r i m e n t s were a l l found w i t h i n the t h r e e - d a y o b s e r v a t i o n p e r i o d i m m e d i a t e l y f o l l o w i n g t h e i r m a r k i n g . S t r a y i n g was not o b s e r v e d , a l l f i s h were found w i t h i n t h e a r e a s o f o r i g i n a l c a p t u r e . T e r r i t o r i a l i t y O b s e r v a t i o n s o f t h e d i s t r i b u t i o n o f A n o p l a r c h u s made d u r i n g m a r k i n g e x p e r i m e n t s and w h i l e c o l l e c t i n g s p e c i -mens f o r o t h e r p a r t s o f the s t u d y , p o i n t out s e a s o n a l t r e n d s i n d i s t r i b u t i o n o f t h e s p e c i e s o v e r the a r e a . At t i m e s o f the y e a r p r e c e d i n g b r e e d i n g , i t i s common t o f i n d as many as s i x o r more i n d i v i d u a l s beneath the same r o c k . Random sam-p l i n g o f f i s h f o r c o l l e c t i o n s has shown t h a t t h e r e i s no s e x u a l o r i n d i v i d u a l s e g r e g a t i o n a t t h i s t i m e . D u r i n g t h e b r e e d i n g s e a s o n , i n c l u d i n g t h e p e r i o d o f p a i r f o r m a t i o n j u s t , p r i o r t o spawning, s e g r e g a t i o n becomes o b v i o u s . At t h i s t i m e i t i s uncommon t o see more than the one p a i r o f sexes beneath t h e same r o c k , a l t h o u g h v e r y l a r g e 75 rocks may y i e l d as many as two such pairs during the period of p a i r formation just p r i o r to spawning. Later, when egg masses were observed, an i n d i v i d u a l rock yielded only a single egg mass. More than one egg mass was never found beneath a single rock. It appears from these observations, that Anoplar-chus exhibits both home-range and defended t e r r i t o r i a l i t y , the form exhibited depending on the p a r t i c u l a r season involved. At times other than the breeding season, the t e r -r i t o r y would be of the home-range type, as there i s no i n d i -cation on the part of the i n d i v i d u a l to exclude others of the species from the area inhabited. During the breeding season, however, d e f i n i t e segregation i s apparent, with only sexual pairs sharing a small area. It seems l i k e l y that an active defence must be present to maintain t h i s s i t u a t i o n . Discussion The possession of a homing behaviour i s considered a mechanism by which i n t e r t i d a l fishes are safeguarded against being stranded i n some unfavourable s i t u a t i o n . Thus, homing i n fishes such as the tidepool inhabiting sculpins (Williams, 1957; Eastman, 1962), would ensure that the return of the animal to a pool which was permanent; i t s f a i l -ure to do so could r e s u l t i n death. 76 Williams (1957) considered homing and straying with reference to i n d i v i d u a l tidepools, but points out that although the pool i s a convenient unit to deal with, there i s no reason to believe that i t i s the "pool" as a hydro-graphic unit to which the f i s h homes. Considerable l i k e l i -hood of some l o c a l i z a t i o n of individuals within large pools was noted by the author. As Gerking (1959) stressed, the s i t e named "home" i s not as important as the fact that the f i s h e l e c t s to return to a place where he has been p r e v i -ously instead of to some other equally probable place. Recognition of the home area would require accur-ate s p a t i a l f a m i l i a r i t y , with active avoidance of being l e f t i n any, but that p a r t i c u l a r area by the outgoing t i d e s . For Anoplarchus, a species which i s d i r e c t l y associated with the substrate of rocks and vegetation, t h i s would c a l l f o r a great rememberance of the route or routes to the home area and an active seeking of t h i s area on outgoing t i d e s . Results of the marking experiments did not i n d i -cate a strong tendency to home, although both types of experiments did indicate movements of Anoplarchus to be some-what r e s t r i c t e d . The l o c a l i t y sense of animals may take a var i e t y of forms, some of which bear resemblance to the c l a s s i c type of t e r r i t o r i a l i t y (defended area), but others d i f f e r i n that the animal shows no tendency to exclude 77 others of the species from i t s r e s t r i c t e d area of movement. It i s t h i s l a t t e r form which i s termed home range or home-range t e r r i t o r y (Dice, 1952; Gerking, 1959; Etkin, 1964). It appears, based on the r e s u l t s of the marking experiments and observations made of the d i s t r i b u t i o n of the species on a seasonal basis, that Anoplarchus displays both forms of t e r r i t o r i a l i t y . Close association of i n d i v i d -uals preceding spawning,, with apparent lack of any tendency of exclusion, indicates that at t h i s time the indiv i d u a l s are sharing home-range t e r r i t o r i e s . However, during the breeding season, a d e f i n i t e contrast to the l a t t e r s i t u a t i o n i s shown. At t h i s time individuals are highly segregated, both s p a c i a l l y and sexually, into isolated breeding p a i r s . The i s o l a t i o n of such breeding pairs indicates that each pa i r may be maintaining a small breeding t e r r i t o r y . A t e r r i t o r y of the defended type i s maintained by aggressive actions of the t e r r i t o r y holder. Behaviour of two breeding pairs was observed under laboratory conditions. The behaviour observed w i l l be discussed i n d e t a i l i n a l a t e r section on reproduction, but i t should be pointed out here that aggression was displayed between members of a breeding pair, both before and a f t e r spawning. Before spawn-ing, aggression was shown by the male only. Following spawning however, the female warded off any advances of the 78 male towards that part of the tank where she was providing parental care to the eggs. Such a behaviour displayed i n the f i e l d would serve to ward off the sexual partner and any other intruders which may happen to infring e upon the nesting s i t e , and thus maintain the established breeding t e r r i t o r y . Guarding females were observed i n the laboratory to provide such care to eggs up to the time that they hatched. Thus, the male l i k e l y defends the breeding t e r r i -tory u n t i l spawning occurs. After spawning, the female takes over and defends the t e r r i t o r y u n t i l the eggs have hatched. 79 REPRODUCTION Sexual Dimorphism Early descriptions of the colouration of A, purpurescens although sometimes quite extensive, f a i l e d to point out the sexually dimorphic nature of the colouration. G i l l (1861) i n hi s o r i g i n a l d e s c r i p t i o n of the species, stated the colour to be a dark purple. Jordan and Evermann (1898) gave a f a i r l y detailed and extensive account of the colouration and pointed out v a r i a t i o n i n colouration between in d i v i d u a l s . However, they did not indicate the sexually dimorphic nature of the colouration. In t h e i r d e s c r i p t i o n of the eggs and nesting habits, Schultz and DeLacy (1932) mentioned that the sexes of Anoplarchus were coloured d i f f e r -ently, but t h e i r account of the differences was extremely b r i e f . Clemens and Wilby (1961) attempted to show the dimorphic nature of the colouration between the sexes and the extent of v a r i a t i o n between in d i v i d u a l s , but as with Jordan and Evermann,their account was not extensive. In the desc r i p t i o n to follow, colour w i l l be des-cribed only as i t emphasizes dimorphism between the sexes. The d e s c r i p t i o n i s based on both l i v i n g and preserved speci-mens . 80 The general body colouration of the male i s usually darker and less interrupted i n i t s d i s t r i b u t i o n on the body, than that of the female. Colour i s highly v a r i -able; l i g h t gray to dark gray, with olivaceous overtones; brown to dark brown, with or without reddish overtones; purple to almost black. Regardless of colour shown, colour-ation of the male i s uniformly d i s t r i b u t e d and lacks the usually highly mottled quality of the female. Colour of the female i s not as variable as the male; l i g h t to dark gray, with olivaceous overtones, marked with brownish gray r e t i c u -l a t i o n s ; dark brown, less r e t i c u l a t e d , lacking reddish overtones. The b e l l y i s pale. Speckling of the body i s more subtle i n the male due to i t s usual darker colouration. Cockscomb and under-surface of the head are pale, rather yellowish and devoid of any speckling or mottling i n the male. The female, i n d i r e c t contrast, shows a highly spec-kled and mottled head, on both the l a t e r a l and undersurfaces, e s p e c i a l l y marked on the jaws and throat. This l a t t e r feature of colouration enables one to quickly i d e n t i f y the sex of preserved specimens. With the approach of the breeding season, the most s t r i k i n g features of the sexually dimorphic nature of the colouration become obvious. The male at t h i s time develops a bright orange to red colouration on i t s f i n s . The anal and pectoral f i n s become bright orange, somewhat more 81 s t r i k i n g on the anal f i n . The caudal and dorsal f i n s show a reddish colouration, usually more pronounced on the cau-dal f i n . The non-breeding male shows a s i m i l a r colouration but more subtle and usually confined to the anal and pec-t o r a l f i n s . Fins of the female are highly speckled and lack bright colouration. The dorsal f i n shows d i s t i n c t grayish to white blotches, which extend v e n t r a l l y on the dorsal part of the back. Along with the differences i n colouration between the sexes at t h i s time of the year, i n both sexes the cocks-comb enlarges with sexual maturity. This structure of the male becomes larger than that of the female i n both height and width. The females average a larger s i z e than the males. In a l l but two of the 13 samples col l e c t e d f o r length f r e -quency, the mean length of females was greater than that f o r males. Among 269 f i s h c o l l e c t e d , the longest female was 128.0 mm i n standard length, and the longest male, 123.5 mm. The mean length of females (157 f i s h ) was 97.05 mm, and of males, 95.01 mm (112 f i s h ) . 82 Gonad Measurements and Fecundity Methods Sexually mature f i s h , distinguished on the basis of granular ovaries and enlarged, opaque testes, were s e l -ected from the September, 1963, December, 1963 and January, 1964, samples. The spawning egg masses examined were obtain-ed on February 9, 1964, and February 12, 1964. The masses were fixed i n 10% formalin and then placed into 40% isopro-py l alcohol. Gonad measurements were made with a pa i r of draftman's di v i d e r s and a small metric scale. A l l measure-ments were recorded to the nearest one-tenth of a m i l l i -meter. Sixteen males were examined, over a range i n stand-ard length of 83.0 to 123.5 mm. The testes were measured fo r length of both l e f t and ri g h t lobes and greatest o v e r a l l width. Ten females were examined, over a range i n standard length of 89.0 to 128.0. The ovaries were measured f o r length and greatest width. The measured gonads were then placed into numbered v i a l s , Actual egg counts were performed on one ovary and one egg mass. A l l remaining counts were estimated using the Von Bayer method (Von Bayer, 1908). 83 To obtain the actual counts, the ovary or egg mass was broken with a p a i r of d i s s e c t i n g pins and the eggs separated. T a l l y i n g of count was done with a small hand recorder. Breaking up of egg masses frequently caused eggs and t h e i r corresponding yolks to become separated. In such cases, yolks only were counted. Average diameters of eggs from the ovaries and masses were obtained by the Von Bayer method. Consultation of the table giving number per quart f o r eggs of various diameters was then done and t h i s figure converted to number per cubic centimeter by d i v i d i n g number per quart by 946.4. Volumetric displacement of the ovary or egg mass was then determined ( i n water) and the figure obtained multiplied by the number of eggs per cc, to obtain the estimate of number of eggs i n the ovary or egg mass. Results Measurements of Testes The lengths f o r each lobe and greatest width of the entire structure, are shown fo r each of the 16 males studied, i n Table XVI. In each i n d i v i d u a l examined the l e f t lobe of the t e s t i s was s l i g h t l y longer than the r i g h t . The l e f t lobe ranged i n length from 8.4 to 17.8 (mean 12.6) i \ 84 mm and the r i g h t l o b e from 7.1 t o 17.0 (mean 11.3) mm. T h i s d i f f e r e n c e i s due p r i m a r i l y t o the a n a t o m i c a l a r r a n g e -ment o f organs i n t h e c o e l o m i c c a v i t y . I n A n o p l a r c h u s the u r i n a r y b l a d d e r l i e s t o the r i g h t s i d e i n t h e body c a v i t y . The l e f t l o b e o f t h e t e s t i s e x t e n d s p o s t e r i o r and a d j a c e n t t o t h i s s t r u c t u r e , whereas, t h e r i g h t lobe, i s i n h i b i t e d f r om d o i n g s o , and ends a t the t i p o f the b l a d d e r . T a b l e XVI. Gonad measurements; 16 t e s t e s and 10 o v a r i e s . T e s t e s O v a r i e s L e n g t h S t a n d a r d L e f t R i g h t G r e a t e s t S t a n d a r d G r e a t e s t L e n g t h Lobe Lobe Width L e n g t h L e n S t h width 83.0 8.9 8.4 3.3 90.0 13.5 11.2 ,5.4 91.8 10.0 8.0 4.5 94.8 14.5 13.8 6.8 95.0 16.8 15.5 5.4 95.5 11.7 10.7 5.2 98.3 8.4 7.1 3.8 101.0 10.0 8.6 5.1 102.0 9.8 9.0 5.0 104.0 15.5 12.8 6.9 105.0 13.1 11.5 6.9 105.2 11.1 10.6 5.7 109.9 17.8 17.0 6.5 115.0 12.5 10.9 5.0 118.0 13.9 12.2 6.1 123.5 13.6 12.2 4.7 Mean 12.6 11.3 5.4 89.0 21.0 8.9 91.5 18.4 7.1 92.0 20.7 8.9 106.5 20.1 9.5 109.0 23.0 10.3 109.2 23.6 10.8 111.5 25.1 11.9 117.0 27.0 12.3 120.0 28.0 11.4 128.0 25.8 10.1 Mean 23.3 10.1 85 Measurements of Ovaries The lengths and greatest width of the structure, are shown f o r each of 10 females studied, i n Table XVI. The ovary i s a p a r t i a l l y bilobed structure; differences i n lengths of the two lobes did not appear to warrant d i f f e r -e n t i a t i o n i n the measurements. Ovaries ranged i n length from 18.4 to 28.0 (mean 23.3) mm and i n greatest width from 7.1 to 12.3 (mean 10.1) mm. Egg Measurements The average egg diameters obtained f o r each of the 10 ovaries and two egg masses studied, are shown i n Table XVII. U n f e r t i l i z e d eggs from the ovaries ranged i n average diameter from 0.787 to 0.965 (mean 0.864) mm and f e r t i l i z e d eggs from the masses from 1.270 to 1.448 (mean 1.372) mm. Egg Counts Actual and estimated numbers of eggs f o r the 10 ovaries and two egg masses studied, are shown i n Table XVII. Counts from ovaries ranged from 2,001 to 3,183 (mean 2,738) eggs and from the masses, 2,288 to 3,082 (mean 2,685) 86 eggs. Table XVII. Fecundity; egg counts of 10 ovaries (A.) and two egg masses (B.). Actual counts indicated by aste r i s k . Standard Ave. No. of Length Dia. Eggs A. 89.0 0.889 2,001 91.5 0.838 2,387 92.0 0.864 2,183 106.5 0.787 2,880 109.0 0.838 2,984 111.5 0.889 3,002 114.0 0.838 3,183 115.0 0.889 3,093* 117.0 0.914 3,065 120.0 0.965 2,606 B. 119.5 1.448 3,082* — — 1.270 2,288 Discussion In t h i s species, i t i s clea r that the entire egg-production of each female i s being concentrated into a single spawning act. This i s evident from the close s i m i l a r i t y of counts of u n f e r t i l i z e d ovarian eggs and f e r t i l i z e d eggs of the l a i d masses, and also from observations of spawning r e s u l t s i n the laboratory and subsequent gonadal examination of the f i s h involved. Spawned f i s h (male and female) were 87 found to be completely "spent", (gross examination only was performed) and could not therefore have spawned again f o r a year. This condition appears common for northern f i s h species (Qasim, 1955). The mean number of eggs obtained f o r the ovaries was 2,738; and f o r the egg masses, 2,685. Schultz and DeLacy (1932) reported a mean number of 1,613 eggs, f o r s i x unspawned females, with a mean standard length of 88.0 mm, as compared with a mean standard length of 106.6 mm, i n the present study. Gudger (1927) reported only 686 eggs i n a mass l a i d by Pholis gunnellus. The number of eggs plotted against length of female, (Figure 7) produces a straight l i n e r e l a t i o n s h i p , with number of eggs increasing as s i z e of female increases. Data were used from the study of Schultz and DeLacy as well as the present study, f o r construction of the graph. The mean average diameter of u n f e r t i l i z e d eggs was 0.864 mm, and f o r f e r t i l i z e d eggs, 1.372 mm. Schultz and DeLacy reported a range i n diameter of u n f e r t i l i z e d eggs of 0.693 to 1.112 mm, a mean was not shown but i t was stated that i t i s about the same s i z e as the diameter of the yolk material of f e r t i l i z e d eggs; a range i n the l a t t e r of 0.819 to 1.070 mm, was shown. They reported an average diameter of f e r t i l i z e d eggs of 1.441 mm. Discrepancies 88 4 0 0 0 r 3000 CD UJ 5 2 0 0 0 L. JO E D z 1 0 0 0 70 8 0 9 0 100 110 120 Standard Length of Female (mm) Figure 7. Graph showing relationship of number of eggs produced to length of female. Unenclosed dots represent data gathered by Schultz and DeLacy (1932) . 130 89 between the two studies are probably related to differences i n s i z e ranges examined and sample s i z e s . The f e r t i l i z e d eggs of Anoplarchus are about the same s i z e as those of Blennius pholis, which are 1.4 mm, as reported by Lebour (1927), but smaller than the eggs of Pholis gunnellus, which average about 2.0 mm i n diameter (Gudger, 1927). Size of the egg does not appear to corre-late with size of f i s h or number of eggs produced. Within the small sample of males studied, there did not appear to be any c o r r e l a t i o n between s i z e of f i s h and length or width of the testes. This lack of c o r r e l a -t i o n may be due to the small sample size and/or to d i f f e r -ent degrees of "ripeness". The length and width of the ovary however, does appear to correlate with s i z e of the female. Hatching of Eggs and Rearing of Young i n the Laboratory In 1964 one attempt was made to hatch eggs of Anoplarchus. Four egg masses were coll e c t e d from the study area on February 12, 1964, and placed into a well aerated, but non-circulating, tank of sea water. Temperature was controlled only by i n s u l a t i n g the entire set-up with sheets of glass-wool., The tank was kept i n darkness. 90 D u r i n g the b r e e d i n g season o f 1965, f u r t h e r a t t e m p t s were made t o h a t c h eggs and a l s o o b s e r v e spawning b e h a v i o u r . On J a n u a r y 16, 1965, a g u a r d i n g female and h e r egg mass was p l a c e d i n t o a t h r e e - g a l l o n aquarium. The aquarium was equipped w i t h a c o o l i n g tube ( g l a s s t u b i n g w i t h r u n n i n g c o l d t a p w a t e r ) , a e r a t i o n , s and-covered bottom and a few s m a l l s t o n e s t o p r o v i d e a s h e l t e r . The tank was kep t c o v e r e d , a l l o w i n g o n l y a minimum of d a y l i g h t t o e n t e r . Over the d u r a t i o n o f the o b s e r v a t i o n p e r i o d , w a t e r temper-a t u r e f l u c t u a t e d between 10° t o 12°C. As w e l l as the f e m a l e and h e r egg mass, two p a i r s o f f i s h were a l s o c o l l e c t e d on the same day. The purpose of t h e i r c o l l e c t i o n was f o r a t t e m p t s t o o b s e r v e , spawning b e h a v i o u r . R e s u l t s The 1964 attempt t o h a t c h eggs was u n s u c c e s s f u l . The egg masses wh i c h were at v a r y i n g s t a g e s o f development when c o l l e c t e d (one mass was "eyed", the o t h e r s n o t ) , s u r -v i v e d f o r 11 da y s . One mass reached a s t a g e v e r y c l o s e t o h a t c h i n g , as w i t n e s s e d by o b s e r v a t i o n o f movements w i t h i n the eggs, but h a t c h i n g d i d not o c c u r . F l u c t u a t i n g temper-a t u r e may have been a f a c t o r i n t h e subsequent d e a t h o f t h e eggs. 91 Attempts made in 1965, under controlled condi-tions, were successful. Two hatchings were observed; one from a spawning of one of the pairs of f i s h i n the lab and the other from the egg mass and female c o l l e c t e d . Female and Egg Mass Continual parental care was provided by the female, from the time of placement into the aquarium, to the time of hatching. The observed behaviour of the female w i l l be described i n a l a t e r section. The egg mass at time of c o l l e c t i o n was "uneyed", and reached the eyed stage by the fourth day of incubation. On the twelfth day of incubation, the mass possessed a golden sheen, due to the eyes of the embryos within. The f i r s t hatched f i s h was observed on January 29, 1965, the thirteenth day of incubation. Further hatching was pro-longed. Only about 12 f i s h were observed u n t i l the morning of February 3, 1965, when the aquarium was clouded with hatched larvae. The female at t h i s time was observed to be out from the shelter of stones and was removed and preser-ved f o r subsequent examination. Young were removed from the aquarium at 24-hour in t e r v a l s , over the 18-day period from f i r s t hatching to 92 death of the remaining young. Laboratory Spawned Pair of Fish The p a i r of f i s h was placed into a f i v e gallon aquarium on January 20, 1965, equipped i n the same manner as the tank previously described. Instead of stones f o r a shelter, an abalone s h e l l was placed in one corner of the tank. Water temperature was cooler, and over the duration of observations fluctuated between 7° to 9°C. The f i s h spawned, unfortunately without personal observation of the spawning act, on February 23, 1965. The male was removed from the aquarium. Observations made of t h e i r behaviour p r i o r to spawning w i l l be described i n a l a t e r section. The egg mass was located beneath the s h e l l and could be observed only by l i f t i n g the s h e l l and disturbing the guarding female. Actual observations of the eggs over the incubation period were therefore kept infrequent. The female provided continual parental care. The f i r s t hatched larvae were observed on March 11, 1965, the sixteenth day of incubation. Hatching did not appear to be prolonged as shown by the previous obser-vations. When the f i r s t larva was observed, the tank was clouded with young. The female was removed at t h i s time 93 and p r e s e r v e d . Hatched f i s h were not sampled, as the p r e v i o u s h a t c h i n g p r o v i d e d a number s u f f i c i e n t t o d e s c r i b e t h e young. The young s u r v i v e d t o March 19, 1965, e i g h t days a f t e r h a t c h i n g . On t h i s day, t h e a i r s u p p l y t o the tank was o b s e r v e d to be o f f and t h e r e f o r e may have been one o f the causes c o n t r i b u t i n g t o the d e a t h o f the young. D e s c r i p t i o n o f t h e Spawned Out C o n d i t i o n The two f e m a l e s had s t a n d a r d l e n g t h s o f 118.0 mm ( f e m a l e and egg mass) and 113.3 mm ( f e m a l e o f p a i r #1). The l a t t e r f emale spawned w i t h a male o f 133.8 mm s t a n d a r d l e n g t h . A l l f i s h were found t o be c o m p l e t e l y " s p e n t " , e x c e p t f o r a v e r y s m a l l number of ova r e m a i n i n g i n t h e o v a r i e s . The b e l l i e s o f b o t h sexes were shrunken and f l a t -t e n e d , and "worn" o f the o u t e r l a y e r o f pigmented s k i n i n the f e m a l e s . The s i d e s o f t h e b o d i e s o f f e m a l e s , a d j a c e n t t o the b e l l i e s , were w r i n k l e d , q u i t e n o t i c e a b l y b e f o r e t h e i r p r e s e r v a t i o n . The gonopore r e g i o n was p i n k i s h , r a t h e r "raw" i n appearance. 94 D e s c r i p t i o n o f Young The newly hatched l a r v a i s t r a n s l u c e n t and v a r i e s from 7.4 t o 7.6 mm i n t o t a l l e n g t h . At s e v e r a l p o s i t i o n s on t h e body, the l a r v a i s pigmented w i t h melanophores, con-s t a n t i n t h e i r d i s p o s i t i o n . An o v a l y o l k sac i s p r e s e n t , w i t h an o i l d r o p l e t a n t e r i o r i n p o s i t i o n ( F i g u r e 8 ) . The head i s f r e e o f p i g m e n t a t i o n . A p a i r o f l a r g e chromatophores are p r e s e n t on t h e a n t e r o - l a t e r a l s u r -f a c e o f t h e y o l k s a c , i m m e d i a t e l y i n f r o n t of the o i l d r o p l e t . D o r s a l t o t h e s a c , t h e r e i s one (may be absent) l a r g e chromatophore on t h e gut t u b e . A l o n g the v e n t r a l mid-l i n e o f the body, r u n n i n g f rom the a n t e r i o r end o f the y o l k sac t o a p o i n t j u s t ahead of the a n a l p a p i l l a , t h e r e i s a row o f 10 to 14 l a r g e chromatophores. Two t o t h r e e l a r g e w i d e l y - s p a c e d chromatophores a r e p r e s e n t v e n t r a l l y on the gut tube, from t h e y o l k sac to the a n a l p a p i l l a . Two l a r g e chromatophores a r e s i t u a t e d on t h e l a t e r a l s u r f a c e s o f the a n a l p a p i l l a , and a p a i r o f e q u a l s i z e median t o the l a t -t e r , a t the base of t h e p a p i l l a . From a p o i n t j u s t p o s t e r -i o r t o the a n a l p a p i l l a , a row o f 36 t o 47 s m a l l chromato-phores r u n s a l o n g the v e n t r a l m i d - l i n e o f the body t o a p o i n t near the c a u d a l p e d u n c l e . On t h e c a u d a l p e d u n c l e , s e p a r a t e d f rom the l a t t e r , f o u r t o s i x chromatophores a r e p r e s e n t . 95 Figure 8. Newly hatched larva (less than 24 hours o l d ) ; 7.4 mm t o t a l length. A. L a t e r a l view of l e f t side; B. Ventral view, from anterior t i p of yolk sac, to t a i l . 96 The f i n - f o l d i s c o n t i n u o u s , b e g i n n i n g d o r s a l l y f r om a p o i n t m a r k i n g the v e r t i c a l a n t e r i o r e x t e n t o f the y o l k s a c , and v e n t r a l l y from the p o s t e r i o r end o f the y o l k s a c . The f o l d widens o v e r the t r u n k , and narrows o v e r the c a u d a l p e d u n c l e , both d o r s a l l y and v e n t r a l l y . Body c o l o u r o f p r e s e r v e d specimens i s a p a l e cream. S u r f a c e o f the eyes a r e a s i l v e r y b l a c k . F o r t y - e i g h t hours a f t e r h a t c h i n g , the l a r v a e a v e r -age 7.85 mm i n t o t a l l e n g t h . The y o l k sac w i t h e n c l o s e d o i l d r o p l e t i s s t i l l p r e s e n t , a l t h o u g h reduced c o n s i d e r a b l y i n s i z e . No changes were o b s e r v e d i n d i s p o s i t i o n o r number o f chromatophores. By n i n e t y - s i x h o u r s , the l a r v a e average 8.05 mm i n t o t a l l e n g t h . The y o l k sac a t t h i s s t a g e i s v e r y s m a l l . The chromatophores a l o n g the v e n t r a l m i d - l i n e o f the body, from the a n t e r i o r end of t h e y o l k sac t o a p o i n t j u s t ahead o f t h e anus, had e n l a r g e d and become r a t h e r d i f f u s e ; a count o f 17 was o b t a i n e d f o r one i n d i v i d u a l , an i n c r e a s e o f t h r e e o v e r the h i g h e s t count o b t a i n e d f o r a t w e n t y - f o u r hour l a r v a . O t h e r chromatophore c o u n t s and d i s p o s i t i o n were the same as t h e l a t t e r s t a g e d e s c r i b e d . By f i v e days a f t e r h a t c h i n g t h e l a r v a e averaged 8.18 mm i n t o t a l l e n g t h and t h e y o l k sac i s c o m p l e t e l y 97 absorbed. Twelve days a f t e r h a t c h i n g the young average 8.50 mm i n - t o t a l l e n g t h . Chromatophore d i s p o s i t i o n i s s t i l l s i m i l a r t o the l a t t e r s t a g e d e s c r i b e d , but c o u n t s o f t h o s e on the v e n t r a l m i d - l i n e of the body c o u l d not be made because o f t h e i r e n l a r g e d and d i f f u s e n a t u r e . O t h e r chrom-atophore c o u n t s were the same as b e f o r e . One f i s h o f t h i r t e e n days o f age showed o n l y one d i f f e r e n c e t o t h e l a t t e r s t a g e d e s c r i b e d ; an i n c r e a s e by one, i n the chromatophore count of t h o s e p r e s e n t v e n t r a l l y a l o n g the gut t u b e , between y o l k sac and anus. F u r t h e r changes o b s e r v e d i n c l u d e d enlargement o f the mouth and jaws, o p e r c u l a r bones and b r a n c h i o s t e g a l s , and the p e c t o r a l f i n s . The f i n - f o l d o f the o l d e s t i n d i v i d u a l was s t i l l c o n t i n u o u s . D i s c u s s i o n H a t c h i n g t i m e s o f 13 and 16 d a y s , were o b t a i n e d f o r t e m p e r a t u r e s of 10° t o 12°C and 7° t o 9°C, r e s p e c t i v e l y . I n the f i r s t h a t c h i n g ( f e m a l e and egg mass) the time o f appearance o f the f i r s t h a t c h e d l a r v a t o complete h a t c h i n g , was p r o l o n g e d . The f i r s t l a r v a was seen on t h e t h i r t e e n t h day o f i n c u b a t i o n but h a t c h i n g was not complete u n t i l the 98 e i g h t e e n t h day. The second h a t c h i n g ( l a b o r a t o r y spawned f i s h ) d i d not show t h i s p r o l o n g a t i o n i n the h a t c h i n g of young; however; t h e a c t u a l time o f spawning was not c r i t i c -a l l y d e t e r m i n e d , and the e s t i m a t e of s i x t e e n days t o h a t c h -i n g may be somewhat low. The t e m p e r a t u r e of t h i s s e t - u p was l o w e r t h a n the p r e v i o u s d e v e l o p m e n t a l t e m p e r a t u r e . P h o l i s  g u n n e l l u s was r e p o r t e d t o have a p e r i o d o f i n c u b a t i o n o f 42 t o 70 d a y s , a t about 6°C (Ehrenbaum, 1904). The l e n g t h o f h a t c h e d l a r v a o f A n o p l a r c h u s was 7.4 t o 7.6 mm which i s s m a l l e r t h a n t h a t r e p o r t e d f o r P h o l i s  g u n n e l l u s by Ehrenbaum (1904), who s t a t e d the l a r v a t o be about 9 mm a t h a t c h i n g . I t i s however, much l a r g e r t h a n the c l i n i d , P a r a c l i n u s marmoratus, w h i c h i s o n l y 4.1 mm a t h a t c h i n g ( B r e d e r , 1939). The young of A n o p l a r c h u s show marked p o s i t i v e p h o t o a x i s . A l i g h t p l a c e d a t one end o f t h e aquarium brought the l a r v a e t o the s u r f a c e at t h i s end. They were a l s o o b s e r v e d t o g a t h e r a t the s u r f a c e a t the back o f t h e tank where d a y l i g h t f r o m a window be h i n d e n t e r e d t h e aquar-ium. T h i s p o s i t i v e p h o t o t a c t i c r e s p o n s e appeared t o l e s s e n a f t e r t h r e e days o f age, and by f i v e d a y s , the young appeared n e g a t i v e l y p h o t o t a c t i c , and would remain c l o s e t o o r on the bottom. On c l e a n i n g o f t h e aquarium subsequent t o t h e d e a t h of the young, t h e l a r g e s t p e r c e n t a g e o f dead f i s h were 99 l o c a t e d beneath the s h e l t e r s p r o v i d e d . T h i s tends t o con-f i r m t h a t the young were n e g a t i v e l y p h o t o t a c t i c a t t h i s s t a g e . T h i s l i g h t r e a c t i o n may have c o n s i d e r a b l e s i g n i f i -cance t o the h a b i t s of the young i n the f i r s t few days o f l i f e . P o s s e s s i o n o f a p o s i t i v e p h o t o t a c t i c r e s p o n s e would b r i n g the h a t c h e d f i s h out from beneath t h e r o c k s and i n t o the c u r r e n t s when l o c o m o t o r y powers were not d e v e l o p e d t o any g r e a t d e g r e e , t h u s becoming an element o f the p l a n k t o n . Once the p o s i t i v e r e s p o n s e t o l i g h t was l o s t , t h e l a r v a e would seek t h e bottom and d i s p e r s a l would depend more on t h e i n d i v i d u a l once l o c o m o t i o n was d e v e l o p e d to a h i g h e r degree, r a t h e r t h a n on c u r r e n t s . Because o f f a i l u r e t o keep l a r v a e a l i v e f o r any l e n g t h o f t i m e i n the l a b o r a t o r y , and t h e l a c k o f knowledge o f t h e i r movements d u r i n g the f i r s t few months o f t h e i r l i f e i n t h e f i e l d , n o t h i n g f u r t h e r i s known about t h e development and metamorphosis o f l a r v a l f i s h . The young o f the 1964 b r e e d i n g s e a s o n were f i r s t f ound i n the i n t e r t i d a l a r e a on June 11, 1964. At t h i s s t a g e they measured between 29.0 and 30.8 mm s t a n d a r d l e n g t h . A l l appeared a d u l t - l i k e i n form, w i t h s m a l l cockscombs and were f u l l y pigmented. S t i l l s m a l l e r f i s h o f 25.5 and 27.0 mm, were c o l l e c t e d i n J u l y and August o f 1964. 100 The f i r s t young located (June sample) were found beneath rocks i n the kelp beds of the l o w - i n t e r t i d a l zone. In l a t e r samples the young were located over habitat i n both the mid- and l o w - i n t e r t i d a l zones. The l a t t e r data suggests that there had been a movement of young from the sub-tidal into the i n t e r t i d a l zone, with a progressive movement up the beach. This movement to the i n t e r t i d a l area appears to take place when the young are 25 to 30 mm i n length. Breder (1941) stated the larvae of Paraclinus  marmoratus to be p o s i t i v e l y phototactic at hatching, but remained so f o r a maximum period of only one day, then sought the bottom. He concluded that they had a planktonic existence f o r a very short time. In contrast, Lebour (1927), reported that the young of Blennius pholis and B. gattoru-gine, remain i n the plankton up to 18 mm and 20 mm or more in length, respectively. Young of B. pholis of 25 mm were found i n small rock pools near high water mark. Evidence as witness by the phototactic responses shown by Anoplarchus young, suggests that they are plank-tonic elements f o r at least three to f i v e days but then are negatively phototactic and seek the bottom. Thus, they are s i m i l a r to the young of Paraclinus marmoratus, but maintain the p o s i t i v e phototactic response f o r a longer period of 101 time. It would appear that migration of young to the i n t e r t i d a l area i s accomplished by locomotion and not by a more passive planktonic migration. Behaviour Associated with Courtship, Parental Care and Between the Sexes Subsequent to Spawning Courtship As previously stated, two pairs of f i s h were brought into the laboratory i n an attempt to observe court-ship and spawning. The f i r s t of such pairs (Pair #1) was placed into an aquarium on January 20, 1965. The f i s h were not fed. Daily observations were made as to the a c t i v i t y of the f i s h . An abalone s h e l l was placed at one end of the tank, to provide a shelter, as well as a possible s i t e for the spawning. An observation i n the f i e l d of an egg mass being guarded i n a large butter clam valve indicated that Anoplar-chus l i k e c e r t a i n other blennioid f i s h e s , does u t i l i z e such structures as spawning s i t e s . Schultz and DeLacy (1932) reported the f i n d i n g of an egg mass of Anoplarchus i n a clam s h e l l , while Gudger (1927) and Ehrenbaum (1904) reported the f i n d i n g of eggs of Pholis gunnellus i n oyster s h e l l s . The p a i r upon introduction into the aquarium 102 i m m e d i a t e l y s i t u a t e d beneath the s h e l l and d u r i n g t h e major-i t y o f o b s e r v a t i o n s , were found i n such a p o s i t i o n . The f i s h were noted t o be v e r y i n a c t i v e u n t i l J a n u a r y 30, 1965, when b o t h sexes appeared t o be r a t h e r r e s t l e s s . The p a i r was ob s e r v e d w i t h the a i d o f d a y l i g h t f r om an above window, f o r two h o u r s . O b s e r v a t i o n was made th r o u g h a s m a l l window c u t i n t o the c a r d b o a r d s u r r o u n d i n g the t a n k , a t a d i s t a n c e of about two t o t h r e e f e e t . P a i r number two was s e t up on March 1, 1965. The female was b u l g i n g w i t h eggs and f i r s t o b s e r v a t i o n s appeared t o i n d i c a t e a r e a d i n e s s t o spawn. The sexes were k e p t sep-a r a t e d between o b s e r v a t i o n s by a c l e a r p a r t i t i o n o f g l a s s , i n an attempt t o e l i m i n a t e u n w i t n e s s e d spawning. Upon f i r s t i n t r o d u c t i o n t o the ta n k , t h e f i s h were f e d l i v e earthworms. Up u n t i l t h i s t i me t h e f i s h had o n l y been f e d s p a r s i n g l y , on f r o z e n a d u l t b r i n e - s h r i m p . Earthworms were a c c e p t e d r e a d i l y and none was r e g u r g i t a t e d . The tank was s e t up f i r s t l y w i t h o u t a s h e l t e r , but l a t e r an abalone s h e l l was added. A l l o b s e r v a t i o n s were made i n the e v e n i n g s , and t h e r e f o r e , a lamp had t o be used f o r i l l u m i n a t i o n . A c a r d -board box w i t h s l i t s t h r o u g h w h i c h one c o u l d o b s e r v e was p l a c e d o v e r t h e aquarium and the lamp s e t a t one end, w i t h 103 the l i g h t d i r e c t e d t o the s i d e . Such an arrangement a l l o w e d o n l y enough l i g h t t o e n t e r to o b s e r v e the movements of the f i s h , not q u i t e s u f f i c i e n t however, t o d e s c r i b e i n d e t a i l c o l o u r changes. The f i s h were a l l o w e d 10 t o 15 minutes t o a c c l i m a t i z e t o t h e l i g h t b e f o r e o b s e r v a t i o n s were commenced. O b s e r v a t i o n s were made o v e r 15 p e r i o d s , each p e r i -od v a r y i n g i n l e n g t h from 1 1/2 t o 3 1/2 (mean 2 1/2) h o u r s . I f t h e f i s h showed any i n d i c a t i o n s o f awareness of the o b s e r v e r , o b s e r v a t i o n s were c u r t a i l e d f o r a few m i n u t e s . W i t h each p e r i o d , o b s e r v a t i o n s ceased when the f i s h were no l o n g e r a c t i v e . R e s u l t s - P a i r Number One T h i s p a i r was o b s e r v e d f o r o n l y two h o u r s . As w i t h P a i r #2, the f e m a l e was t h e most a c t i v e s e x . Her move-ments were s t r i c t l y c o n f i n e d t o the a r e a a d j a c e n t to the s h e l l , r e p e a t e d l y swimming beneath and o v e r the s h e l l , f r e -q u e n t l y p e r c h i n g on t o p . The male i n c o n t r a s t , was r a t h e r q u i e t , r e m a i n i n g near and u s u a l l y p a r t l y under, th e s h e l l . O c c a s i o n a l b u r s t s i n t o a c t i v e swimming at t h e back o f the tank however, d i d o c c u r . One approach o f t h e f e m a l e towards th e male, w h i l e 104 the l a t t e r f i s h was beneath the s h e l l , was met with two strong bitings to the head. The female did not swim away but moved i n closer to the male. Observations of Pair #2 showed that such agonistic behaviour by the male usually e l i c i t e d fanning by the female. No such fanning behaviour was observed by P a i r #1. Several other approaches were made by the female to the side of the male. No further displays of agonistic behaviour were observed and the male did not appear to respond to the closeness of the female i n any c h a r a c t e r i s t i c manner. His usual reaction was to remain motionless or slowly move away. Similar to P a i r #2, movements of either sex were clo s e l y watched by the opposite sex. Subsequent to January 30, 1965, frequent checks were made as to the a c t i v i t y of the f i s h but no observations indicated a readiness to spawn. On February 23, 1965, an egg mass was found beneath the s h e l l , with the female c o i l e d and fanning. The spawning was unwitnessed. Results - Pair Number Two The f i r s t observation period (2 hours) was on 105 March 2 , 1965. The f i s h were very active and the d i f f e r e n t behavioural patterns observed were found, when subsequent observations were made, to represent almost the whole spec-trum of behavioural patternsobserved over the entire 15 observation periods. The behaviour observed w i l l be d e s c r i -bed i n d e t a i l and can be considered t y p i c a l of a l l behaviour observed. Data gathered from other observations w i l l be used to supplement and quantify the f i r s t period observa-tio n s . Upon removal of the p a r t i t i o n , the sexes almost immediately came together. The approach by each sex was about equal and e l i c i t e d an erection of dorsal and anal f i n s of both f i s h . The p a i r was together f o r only a few seconds. Investigative behaviour was then shown, with both f i s h moving about parts of the tank where t h e i r movements were previously in h i b i t e d due to the p a r t i t i o n . Movements of e i t h e r f i s h were keenly followed by the other, both v i s u -a l l y and p h y s i c a l l y . Frequent approaches of one sex to the other were observed and f o r the most part these advances were made by the female. In approaching the male, a c h a r a c t e r i s t i c pose was assumed; body held somewhat U-shaped, undulated slowly, bringing about a somewhat sideways movement towards the male and ending with the l a t e r a l surface against the length 1 0 6 of the male's body. With these movements dorsal and anal f i n s were f u l l y erected and caudal f i n expanded. Coloura-t i o n was very l i g h t . Such advances by the female usually e l i c i t e d a strong aggressive attack by the male; a b i t i n g to her head or pectoral area. These b i t i n g s were sometimes repeated several times. In one observation period of two hours dur-ation, the male was observed to bite the female a t o t a l of 10 times, i n quick successions of two or three bi t i n g s at a time. With other approaches by the female the male showed no aggressive behaviour and moved away from the l a t t e r . Agonistic behaviour towards the female, i n the form of quick darting bites, preceded sometimes by what appeared to be a threatening posture (turning of the head and s l i g h t puffing of the opercles), i n only a few instances caused the female to swim away from the male. Her usual response was to fan ( i d e n t i c a l to the fanning c h a r a c t e r i s t i c i n parental care of the eggs), and edging closer to the male, sometimes against the body of the male, with the fan-ning undulating against his side. The male's usual response to such a behaviour was a movement away, but sometimes he remained motionless against the female. Movement of the sexes away from each other was accompanied by a lowering of the dorsal and anal f i n s , and a 107 somewhat lightening of colouration of the male, although at times t h i s l a t t e r was not obvious because of the pool i l l u m -i nation of the tank. The colouration of the female was always a l i g h t gray, i n contrast to the dark brown coloura-ti o n of the male. The anal and pectoral f i n s of the male were a bright orange and the caudal f i n reddish. The dorsal f i n was s l i g h t l y tinged with red as well . One c h a r a c t e r i s t i c behaviour shown by the male was. that of digging depressions i n the sand. In almost every observation period t h i s behaviour was expressed. The body was turned s l i g h t l y on i t s side, head held r i g i d against the end of the tank, and the posterior portion of the body v i b r a -ted very ra p i d l y , sending out a spray of sand behind the f i s h . The process was repeated two or three times i n succes-sion, with the end res u l t a depression was formed i n the sand large enough to accompany one f i s h i n a c o i l e d p o s i t i o n . The male did not occupy these depressions f o r long periods of time, nor did he show much aggression towards the female i f she approached such holes. Approaches by the female, when the hole was being occupied by the male sometimes e l i c i t e d what appeared to be a threat response ( r a i s i n g of the f i n s and gaping) but i n other instances showed a complete lack of response. On March 30, 1965, an egg mass was found beneath 108 the s h e l l , w i t h the female c o i l e d and f a n n i n g . She had r e l e a s e d h e r eggs sometime d u r i n g the n i g h t o r e a r l y morn-i n g . The l a s t o b s e r v a t i o n p e r i o d was on the e v e n i n g o f March 29, 1965, a t which time the f i s h appeared r a t h e r i n a c t i v e . P a r e n t a l Care I n c o n t r a s t t o many o t h e r b l e n n i o i d f i s h e s , i t i s t h e female which s t a y s w i t h the eggs and p r o v i d e s p a r e n t a l c a r e . W i t h each egg mass l o c a t e d , S c h u l t z and DeLacy (1932) found a f e m a l e t o be a s s o c i a t e d w i t h i t . I n one case they found a male nearby, but i t escaped c a p t u r e . R e s u l t s borne o u t by t h i s s t u d y were s i m i l a r ; a l l l o c a t e d egg masses were b e i n g guarded by a f e m a l e ; no males were l o c a t e d near the s i t e o f t h e eggs and f e m a l e . B r e d e r (1939) summarized d a t a known f o r a number of s p e c i e s and r e p o r t e d t h e males o f the f o l l o w i n g s p e c i e s t o guard the eggs; B l e n n i u s g a t t o r u g i n e , B. montagui, B. p h o l i s , B. o c e l l a r i s , B. sphynx and C l i n u s a r g e n t a t u s . B o t h p a r e n t s may guard t h e n e s t of P h o l i s g u n n e l l u s . I n the p r e v i o u s l y named s p e c i e s , t h e eggs are a d h e s i v e . No f a n n i n g o f the eggs has been d e s c r i b e d ( B r e d e r , 1939). 109 O b s e r v a t i o n s i n the F i e l d L i f t i n g o f a r o c k c o v e r i n g a n e s t i n g s i t e d i d not appear t o d i s t u r b the f i s h f r om i t s g u a r d i n g p o s i t i o n . The female was found t o c o i l about t h e egg mass i n a U-shape, somewhat more t i g h t l y under t h e s e c o n d i t i o n s t h a n when co v e r e d by w a t e r . No movements were made by the female u n t i l t he egg mass was l i f t e d f r o m i t s r e s t i n g p o s i t i o n . T h i s i s a fo r m o f b e h a v i o u r q u i t e d i s t i n c t f r o m the u s u a l f l e e i n g o b s e r v e d when such d i s t u r b a n c e s o c c u r a t o t h e r t i m e s of t h e y e a r . O b s e r v a t i o n s i n t h e L a b o r a t o r y A f e m a l e w i t h h e r egg mass was p l a c e d i n t o an aquarium on J a n u a r y 16, 1965 ( h a t c h i n g o f eggs p r e v i o u s l y d e s c r i b e d ) and h e r subsequent b e h a v i o u r o b s e r v e d . The tan k was s e t up w i t h a s m a l l s h e l t e r o f s t o n e s a r r a n g e d i n such a way t h a t o b s e r v a t i o n s c o u l d be made from the f r o n t of t h e ta n k . The eggs were p l a c e d under the s h e l t e r and the n t h e fem a l e i n t r o d u c e d . I n i t i a l o b s e r v a t i o n s were c u r t a i l e d f o r a day t o a l l o w t h e fe m a l e t o become accustomed to h e r s u r -r o u n d i n g s . The tank was kep t i n darkness between o b s e r v a -t i o n s . 110 Two days a f t e r f i r s t i n t r o d u c t i o n t o t h e tank s i g n s o f p a r e n t a l c a r e were o b s e r v e d . The egg mass was ob s e r v e d t o be out from the s h e l t e r of s t o n e s , i n one c o r -ner o f t h e t a n k , h a v i n g e i t h e r f l o a t e d out o r been moved out by t h e f e m a l e . The femal e was c o i l e d about the eggs and f a n n i n g r h y t h m i c a l l y w i t h t h e p o s t e r i o r h a l f o f h e r body. A l a t e r o b s e r v a t i o n t h e same day r e v e a l e d b oth mass and f i s h t o be under t h e s h e l t e r of s t o n e s , the f e m a l e c o i l e d and f a n n i n g . F o r the d u r a t i o n o f r e m a i n i n g p e r i o d s of o b s e r v a t i o n up t o h a t c h i n g , t h e female remained i n t h i s p o s i t i o n and was never ob s e r v e d out from the s h e l t e r . The d o r s a l and a n a l f i n s a r e e r e c t e d and the c a u d a l f i n expanded, d u r i n g f a n n i n g a c t i v i t y . I n e f f e c t , t h i s i n c r e a s e s the s u r f a c e a r e a of t h e body i n c o n t a c t w i t h t h e w a t e r . The p r e s e n c e o f l i g h t (a lamp was used d u r i n g t h e o b s e r v a t i o n p e r i o d s ) appeared t o i n c r e a s e t h e r a t e o f f a n -n i n g , and i f l e f t f o r more than two o r t h r e e m i n u t e s , caused the f e m a l e t o become q u i t e r e s t l e s s beneath t h e s h e l -t e r . F a n n i n g o f t h e eggs was m a i n t a i n e d up u n t i l t he time o f complete h a t c h i n g . As was p r e v i o u s l y d e s c r i b e d , the f i r s t h a t c h e d f i s h was o b s e r v e d on J a n u a r y 29, 1965, but the mass was not c o m p l e t e l y h a t c h e d u n t i l F e b r u a r y 3, I l l 1965. Over t h i s extended period of hatching, the fanning behaviour was maintained. On February 3, 1965, the female was observed to be out from the shelter and was removed. At the time of removal, she showed much aggression towards the intrusion by the net used to capture her, darting and b i t i n g at the mouth of the net. Similar observations were made of the second female, whose spawned eggs were located beneath an abalone s h e l l placed i n one corner of the aquarium. The c o i l e d p o s i t i o n with fanning was maintained f o r the duration of the incubation period. At the time of her removal (day of hatching of the eggs, March 11, 1965), the same aggression was shown to the in t r u s i o n by the net as that shown by the previous female. One example of parental care being provided to an u n f e r t i l i z e d egg mass was observed. The eggs were released on March 30, 1965. The male at the time of the egg laying was partitioned away from the female. The eggs were guarded with fanning f o r s i x days. By A p r i l 9, 1965, c o i l i n g about the eggs was seldom seen and the female began to eat the eggs. By A p r i l 14, 1965, a l l eggs had been devoured. 112 Behaviour Between Sexes Subsequent to Spawning Observations were made of the behaviour of P a i r #2 subsequent to the female's release of eggs. At the time of egg releasing the sexes were partitioned by a cl e a r sheet of glass. The p a r t i t i t i o n was placed diagonally i n the aquarium. The egg mass was located beneath the s h e l l at one end of the tank. The male could see and get close to the female (within 2 to 3") but was kept from actual contact with the female by the glass p a r t i t i o n . The p a r t i t i o n was removed and the male allowed to move f r e e l y over the ent i r e tank. At the end of 14 minutes of observation the male had not moved to the end of the tank where the female and eggs were located. The female during t h i s time remained with the eggs, fanning, except f o r short movements around the s h e l l with a return to the eggs each time. These l a t t e r movements indicated an awareness of the male's presence. Her head was turned i n the d i r e c t i o n of the male, appearing to watch the male but returning shortly to the eggs. The male did not come close to the nesting s i t e u n t i l 27 minutes of observation had been completed. At which time the female turned and darted out from under the s h e l l , b i t i n g the male to the head, and then to the t a i l as 113 the male moved away. The male did not return to the region of the nesting s i t e f o r the remainder of the observation period. The p a r t i t i o n was replaced. Further observations were made two days l a t e r ( A p r i l 1, 1965), with the p a r t i t i o n i n place. Approaches by the male were met with the same expression of agonistic behaviour as before. The female darted to the p a r t i t i o n , snapping against the glass. The l a s t observations were made on A p r i l 5, 1965; the same behaviour was displayed. Discussion The female Anoplarchus purpurescens guardsand tends i t s eggs. Such care i s characterized by a c o i l i n g about the eggs i n a U-shape, dorsal and anal f i n s erect, caudal expanded, and fanning. Schultz and DeLacy (1932) stated that the female on guard was party c o i l e d around the eggs, but made no mention of any fanning behaviour. The male plays no role i n parental care, and i s repulsed from the region of the nesting s i t e by vigourous expressions of agonistic behaviour by the female. The l a t -ter consists of darting out from the nest, with eventual b i t i n g . 114 Care i s provided continually, from just a f t e r spawning, to the time of complete hatching of the eggs. Courtship display was observed, although the actual spawning act was unwitnessed. The term courtship i s adopted here as i t was defined by Morris (1958), as "the heterosexual communications systems leading up to the con-summatpry sexual act of f e r t i l i z a t i o n , " In some species t h i s behaviour consists of one continuous complex, the courtship of the stickleback as an example. In other species, however, there are two quite d i s t i n c t behaviour complexes— pair-formation and pre-copulatory display—which are separ-ated i n time. A. purpurescens shows a courtship behaviour of the l a t t e r type. It i s assumed that the courtship display observed was pre-copulatory display, as the f i s h were brought into the laboratory a f t e r they had paired up i n the f i e l d . The i n i t i a l pair-forming courtship was not observed and there-fore cannot be distinguished from the subsequent pre-copulatory display. The behaviour observed appeared to be rather l i m i t e d . Baerends and Baerends-von-Roon (1950) showed i n t h e i r study of c i c h l i d f i s h e s , that i n species where both partners take part i n parental duties, there i s a greater d i s t i n c t i o n between pair-formation and pre-copulatory 115 ceremonies, than i n those species in which only one sex performs parental duties. Anoplarchus may show s i m i l a r courtship, and because of not observing the i n i t i a l p a i r -formation display, the remaining pre-copulatory display appeared that much more li m i t e d . Courtship display observed showed the male to be very aggressive to approaches by the female. This type of courtship i n which the animal tends to attack i t s mate, i s commonly shown by f i s h e s , the stickleback's courtship being the most widely used example. Pugnacity i s however, a feature well known f o r gobies and most t e r r i t o r i a l f ishes (Tavolga, 1954). Morris (1954), observed t h i s type of court ship.ibroa c o t t i d , Cottus gobio. The male of t h i s species i s extremely aggressive to the female, with courtship often beginning by the male swallowing the female'shhead : i n one huge b i t e . The female i f ripe with eggs, responds sexually to the assault of the male and enters the: nest, or allows herself to be carried i n by him. The female Anoplarchus showed a somewhat s i m i l a r response to the aggressive actions of the male. There was no tendency on her part to f l e e the male when attacked, but rather, she would move closer to him, c h a r a c t e r i s t i c a l l y with a fanning movement of her body. The moving i n close to the male appeared to be a d i r e c t response to the male's 116 a g g r e s s i v e a t t a c k . G u i t e l (1893) o b s e r v e d a somewhat s i m i -l a r b e h a v i o u r f o r t h e female C l i n u s a r g e n t a t u s , who a t t r a c t s males by means o f body q u i v e r s . The male A n o p l a r c h u s d i d not show any f u r t h e r a g g r e s s i v e b e h a v i o u r once the f e m a l e had p o s i t i o n e d v e r y c l o s e , r e m a i n i n g r a t h e r m o t i o n l e s s o r u s u a l l y moving s l o w l y away. There appeared t o be a d e f i n i t e l a c k o f s e x u a l r e s -ponse t o t h e f e m a l e . M o r r i s (1954) p o i n t e d out t h a t even i n t h e v e r y a g g r e s s i v e c o u r t s h i p of C. g o b i o , the male does not o n l y a t t a c k the f e m a l e , but a l s o t h r e a t e n s h e r , t h u s r e v e a l i n g that h i s f l e e i n g d r i v e i s a l s o a c t i v a t e d , i f o n l y s l i g h t l y ( t h r e a t b e i n g the r e s u l t of a c o n f l i c t between a t t a c k i n g and f l e e i n g t e n d e n c i e s ) . The male A n o p l a r c h u s a l s o showed what appeared t o be a t h r e a t p o s t u r e b e f o r e a c t u a l a g g r e s s i v e a t t a c k ; t u r n i n g towards the f e m a l e , g a p i n g , e r e c t i n g the f i n s and sometimes e x p a n d i n g t h e o p e r c l e s . The l a t t e r b e h a v i o u r has been shown f o r b l e n n i o i d f i s h e s ; B l e n n i u s  sphynx ( G u i t e l , 1893), B. o c e l l a r i s ( P i e r o n , 1914) and P a r a c l i n u s marmoratus ( B r e d e r , 1939, 1941). B o t h T i n b e r g e n (1952) and M o r r i s (1956) have sug-g e s t e d r e a s o n s why t h e r e i s so much a g g r e s s i o n shown i n c o u r t s h i p . T i n b e r g e n f e e l s t h a t because the males have t o f i g h t one a n o t h e r , the f e m a l e s even when d i m o r p h i c , cannot 117 help stimulating the male's aggression s l i g h t l y . Morris modified t h i s view somewhat, by s t a t i n g that t e r r i t o r i a l males cannot help treating intruding females as objects to be attacked as well as courted, because such males respond aggressively primarily to intrusion onto t h e i r t e r r i t o r i e s and only secondarily to the nature of the intruding object. He suggests therefore, that the basis of pair-bond i s removal of the attacking and f l e e i n g tendencies, which per-mit the now unsiippressed sexual tendency to keep male and female together. Thus, i t i s possible to perform i n i t i a l (pair-forming) courtship displays and then, having formed a pair-bond, wait f o r some time before attempting to achieve f e r t i l i z a t i o n . The successful spawning which took place by Pair #1, indicates that aggressive tendencies were overcome by sexual tendencies, allowing f e r t i l i z e d spawning to take place. With Pair #2, however, a successful spawning did not occur and the male appeared to remain aggressive towards the female f o r the duration of observation periods performed, r e s u l t i n g i n subsequent egg release by the female. Subsequent to spawning, during parental care duties, a d i s t i n c t change i n behaviour i s shown by the female towards the male. She i s now very aggressive to the male's approaches to the nesting s i t e , b i t i n g and chasing him away. A few such attacks appeared s u f f i c i e n t i n keeping 118 the male away from the nesting s i t e . 119 AGE AND GROWTH Introduction , The Length-Frequency Method Most fishes have a more or less sharply defined breeding season, i . e . , the young are born during a d e f i n i t e season.of the year. If a large c o l l e c t i o n of a species i s made over a short period of time (on one day, f o r example), the individuals of the sample should group themselves around c e r t a i n modal s i z e s . Thus several peaks appear i n the length-frequency curve of a multiple age population of f i s h , each peak (mode) t h e o r e t i c a l l y representing an age group or year c l a s s . The distinctness of each of these modes would depend on the extent of overlap of length-frequency d i s t r i b u t i o n s of consecutive age groups. The overlap would of course depend on length of the breeding season, on r a p i d i t y aad uniformity of growth within the species, and on length of the period of sampling. It i s sometimes possible to follow the growth of f i s h during one or several years by using length-frequency d i s t r i b u t i o n s . Length d i s t r i b u t i o n s are plotted from monthly samples and curves arranged one above the other on a d i a g r a m . With growth of the f i s h , i t s length d i s t r i b u t i o n 120 c u r v e f o r each age group i s d i s p l a c e d to the r i g h t . V a l i d i t y o f t h e l e n g t h - f r e q u e n c y method f o r a pproximate age d e t e r m i n a t i o n has ample s u b s t a n t i a t i o n , and has been a p p l i e d w i d e l y i n age and growth s t u d i e s . However, t h r e e b a s i c p r e r e q u i s i t e s a r e n e c e s s a r y f o r good r e s u l t s , and the f o l l o w i n g c h a r a c t e r i s t i c s o f the sample s h o u l d be c l o s e l y met: (1) composed o f a l a r g e number o f i n d i v i d u a l s ; (2) c o l l e c t e d i n a r e s t r i c t e d p e r i o d o f time ( i n a s i n g l e day, p r e f e r a b l y ) ; (3) good r e p r e s e n t a t i o n of a l l the s i z e - and age-groups i n t h e p o p u l a t i o n . These l a t t e r optimum c o n d i t i o n s o f the method were p r e s e n t i n two r e s p e c t s , i n the a n a l y s i s t o f o l l o w . A l t h o u g h sample s i z e s were not g r e a t , each c o l l e c t i o n was made on a r e s t r i c t e d p e r i o d of t i m e , u s u a l l y on one day (some were c o l -l e c t e d on two days, but were not s e p a r a t e d i n time by more t h a n one week), and each c o l l e c t i o n r e v e a l e d a good range i n l e n g t h s . T h i r t e e n samples were u t i l i z e d , p r o v i d i n g 285 s p e c i -mens. The f i r s t sample was c o l l e c t e d i n June, 1963 and the l a s t i n O c t o b e r , 1964. L e n g t h - f r e q u e n c y c u r v e s were p l o t t e d f o r each sample, and f o r the t o t a l c o l l e c t i o n and t h e s e x e s . Modal a n a l y s i s was performed on the s e a s o n a l s e r i e s of 13 121 samples. The v a r i o u s l e n g t h s e x h i b i t e d were grouped i n t o s t a n d a r d l e n g t h g r o u p i n g s o f f i v e m i l l i m e t r e s each. . O t o l i t h s A s t u d y o f o t o l i t h s was performed t o s u b s t a n t i -a t e the l e n g t h - f r e q u e n c y method of age and growth d e t e r m i n -a t i o n . A t o t a l o f 70 o t o l i t h s were examined, u t i l i z i n g t h e samples from June, 1963 t o June, 1964. The o t o l i t h s o f A n o p l a r c h u s a r e v e r y s m a l l (0.8 t o 2.2 mm), and f a i r l y t r a n s p a r e n t . No p r o c e s s i n g o f the o t o l i t h was r e q u i r e d , e x cept f o r a drop o f 100% g l y c e r i n , used as a c l a r i f y i n g agent. Opaque w h i t e and c o m p l e t e l y c l e a r o t o l i t h s were d i s c a r d e d . The o t o l i t h s were s t u d i e d w i t h r e f l e c t e d l i g h t , a g a i n s t a b l a c k background, which produces an a l t e r n a t i o n o f wide w h i t e zones and narrow d a r k ones. The a n n u a l r i n g was t a k e n as the b o r d e r between t h e i n t e r n a l narrow zone and the e x t e r n a l broad one. A s m a l l r i n g w hich was f r e q u e n t l y found around t h e c e n t r a l c o r e of t h e o t o l i t h , was d i s r e g a r d e d and not counted as the f i r s t a nnual r i n g . The o t o l i t h c o u l d o n l y be read w i t h any c e r t a i n t y i n a p p r o x i m a t e l y o n e - h a l f of t h e 70 o t o l i t h s examined. T h i r t y - f o u r age d e t e r m i n a t i o n s were made from the o t o l i t h s . 122 R e s u l t s L e n gth-Frequency A n a l y s i s The l e n g t h - f r e q u e n c y d i s t r i b u t i o n of the e n t i r e c o l l e c t i o n o f 285 f i s h i s shown i n F i g u r e 9, a l o n g w i t h t h o s e o f the males and f e m a l e s . The p l o t o f the t o t a l c o l -l e c t i o n , i n d i c a t e s a p o s s i b i l i t y o f f o u r modes, which i s f u r t h e r s u b s t a n t i a t e d by t h e r e s u l t s o f the modal a n a l y s i s of each of 13 samples examined. F i g u r e 10 shows the l e n g t h -f r e q u e n c y d i s t r i b u t i o n s o f each of 13 samples, and the modal a n a l y s i s i s p r e s e n t e d i n T a b l e X V I I I . Such a n a l y s i s r e v e a l e d a t o t a l o f f o u r modes, whi c h c o u l d p o s s i b l y r e p r e -s e n t age groups 0, I , I I and I I I . The a s s u m p t i o n t h a t the f i r s t s i z e group r e p r e s e n t s young o f the y e a r i s not w i t h o u t f o u n d a t i o n . As i s g e n e r a l l y known, most f i s h e s grow more r a p i d l y (in l e n g t h ) i n e a r l y l i f e t h a n l a t e r . T h i s r a p i d e a r l y growth would t e n d t o make young s i z e groups more c l e a r l y d i s t i n g u i s h a b l e i n a f r e q u e n c y d i s t r i b u t i o n t h a n the o l d e r ones. F u r t h e r m o r e , t h e spawning season o f A n o p l a r c h u s i s i n J a n u a r y and F e b r u a r y . The f i r s t v e r y s m a l l f i s h (25 t o 30 mm) were l o c a t e d i n June (1964), but were s t i l l b e i n g l o c a t e d in September and O c t o b e r , a t a s i z e o f 30 t o 50 mm. T h e i r s i z e p r e c l u d e s the p o s s i b i l i t y o f t h e i r h a v i n g been spawned i n t h e p r e v i o u s y e a r . Figure 9 Length-frequency d i s t r i b u t i o n s ; t o t a l of 285 specimens, 112 males, 157 females, 16 unsexed. 124 Figure 10. 2- June, 1963 n=6 4 - Aug. 1963 n = 21 12 10 8 6 4 2 > 2 o C 4 cr 2 o 6 Sept, 1963 n=34 July, 1964 Aug, 1964 n = 46 Sept, 1964 n = 15 ^ Oct, 1964 n=25 I I I I i i i I I i i i i i i i i i i i i i 22.5 37.5 52.5 67.5 82.5 975 112.5 127.5 Standard Length (mm) Length-frequency d i s t r i b u t i o n s of each of the 13 i n d i v i d u a l samples; t o t a l of 285 specimens. T a b l e X V I I I . Modal a n a l y s i s o f each o f 13 samples; i n c r e a s e s and d e c r e a s e s i n modal s i z e between months and between the two y e a r s o f s a m p l i n g . Date o f Sample Age Groups 1 I I I I I J u n . Aug. Sep. Nov. Dec. J a n . Mar. May, J u n . J u l . Aug. Sep. 1963 1963 1963 1963 1963 1964 1964 1964 1964 1964 1964 1964 O c t . , 1964 30.0 27.5 45.0 42.5 60.0 72.5 82.5 92.5 92.5 82.5 87.5 67.5 65.0 67.5 72.5 90.0 97.5 92.5 102.5 102.5 102.5 107.5 95.0 97.5 97.5 92.5 102.5 105.0 112.5 117,5 120,0 117.5 117.5 122,5 115.0 117.5 117.5 117.5 126 Age group 0 i s l a c k i n g f rom a l l samples c o l l e c t e d i n 1963, and does not show up u n t i l t he June,, 1964 sample. The apparent l a c k o f young o f the y e a r i n t h e samples o f 1963 i s p r o b a b l y a t t r i b u t a b l e t o s e l e c t i v i t y i n s a m p l i n g . These samples were a l l c o l l e c t e d i n the uppermost mid-i n t e r t i d a l and m i d - i n t e r t i d a l i n a r a t h e r r e s t r i c t e d a r e a i n comparison t o the t o t a l s t u d y a r e a (between p i l l a r s #8-14, beneath the b r i d g e ) . Once s a m p l i n g was f u r t h e r e d t o o t h e r a r e a s , s t a r t i n g w i t h t h e May, 1964 c o l l e c t i o n , s m a l l f i s h o f 50, 60 and 70 mm were o b t a i n e d . These f i s h were o f a s i z e too l a r g e t o be c o n s i d e r e d 1964 young o f t h e y e a r (as e v i -denced by l a t e r 1964 samples) and must have been young o f the 1963 b r e e d i n g s e a s o n , now age group I f i s h . The s m a l l -e s t f i s h o b t a i n e d i n 1963 was a f i s h o f 68.5 mm, i n June; an age group I f i s h , h a t c h e d i n 1962. O t o l i t h A n a l y s i s Ages were d e t e r m i n e d from 34 o t o l i t h s , f r om a range i n s t a n d a r d l e n g t h o f 51.5 t o 123.5 mm, and o v e r a time p e r i o d o f August, 1963 t o June, 1964. The r e s u l t s a re p r e s e n t e d i n T a b l e s XIX and XX. F i v e age groups were found t o be p r e s e n t i n t h i s s i z e range; I , I I , I I I , IV and V, w i t h t h e number o f a n n u l a r r i n g s v a r y i n g from one t o f i v e . Thus, ages from 1+ t o 5+ y e a r s , r e p r e s e n t i n g y e a r c l a s s e s from 1959 t o 1963, were 127 p r e s e n t . The assumption made i n the l e n g t h - f r e q u e n c y a n a l -y s i s , t h a t f i s h o f 50, 60 and 70 mm, t a k e n i n t h e May, 1964 sample, do not r e p r e s e n t young of the y e a r , i s f u r t h e r sub-s t a n t i a t e d by the o t o l i t h r e a d i n g s . F i s h from 50 t o 80 mm were found, t o p o s s e s s one a n n u l a r r i n g and t h e r e f o r e must have been hat c h e d i n t h e p r e v i o u s y e a r ( i . e . , 1963 y e a r c l a s s ) . F i s h from 80 t o 90 mm, may have one o r two a n n u l a r r i n g s , t h u s , may be 1+ o r 2+ y e a r s of age. From 90 t o 100 mm, f i s h a re u s u a l l y 2+ y e a r s o f age, but: may be 3+. F i s h f r o m 100 t o 120 mm, are p r e d o m i n a n t l y 3+ y e a r s of age, but may be 4+. S i z e s o f 120 mm and g r e a t e r , may be 4+ o r 5+ y e a r s o f age. C o r r e l a t i o n o f t h e Two Methods of Age D e t e r m i n a t i o n L e n g t h - f r e q u e n c y a n a l y s i s i n d i c a t e d t h e p o s s i b i l -i t y o f f o u r age g r o u p s , from young of the y e a r , age group 0, t o age group I I I . I t i s apparent however, as e v i d e n c e d by by the o t o l i t h r e a d i n g s , t h a t t h e s i n g l e modes a r e not r e p r e s e n t i n g f i s h o f a s i n g l e age. The l a t t e r i s p r o b a b l y a t t r i b u t a b l e t o the s m a l l sample s i z e s , and i s not c o r r e c t e d by t h e p l o t o f the t o t a l c o l l e c t i o n ( F i g u r e 9) because of wide s e p a r a t i o n i n time o f the c o l l e c t i o n o f each o f t h e 13 i n d i v i d u a l s amples. There i s c l e a r - c u t d i s t i n c t i o n o f young 128 T a b l e XIX. R e s u l t s o f age d e t e r m i n a t i o n s from 34 o t o l i t h s ; 19 males, and 15 f e m a l e s . Males Females Date o f T No. T No. Sample L e n g t h R i n g g L e n g t h R i n g s Aug., 1963 85.0 1 84.0 1 101.0 3 92.1 2 104.5 3 106.0 3 108.0 3 115.0 3 Sep., 1963 94.8 2 98.8 2 109.9 3 101.5 3 114.0 3 118.2 3 D e c , 1963 118.0 4 J a n . , 1964 83.0 2 92.0 2 98.3 3 114.0 3 123.5 4 120.0 5 Mar., 1964 86.0 2 86.2 2 May, 1964 51.5 1 66.2 1 53.9 1 67.2 1 56.9 1 61.1 1 62.0 1 119.2 3 J u n . , 1964 84.0 1 61.2 1 75.2 1 T a b l e XX. Age c o m p o s i t i o n o f sexes (19 males, 15 f e m a l e s ) by s t a n d a r d l e n g t h s , determined from 34 o t o l i t h s . 1 2 3 4 5 Sta n d a r d Males Females Males Females Males Females Males Females Males Females L e n g t h 50- 60 3 60- 70 2 3 70- 80 1 8 0 - 9 0 2 1 3 90-100 1 3 1 100-110 4 2 110-120 1 4 1 120-130 1 1 Sums 7 5 4 3 6 6 2 0 0 1 Mean Length 64.91 70.76 87.50 94.30 106.82 111.45 120.75 120.00 130 o f the y e a r (age group 0) but w i t h i n c r e a s i n g f i s h s i z e , the modes tend t o o v e r l a p more and more, and thus do not r e p r e - , s e n t s i n g l e ages. The o t o l i t h a n a l y s i s c o v e r e d a range i n s i z e w h i c h the l e n g t h - f r e q u e n c y method i n d i c a t e d t o r e p r e s e n t age group I t o I I I . O t o l i t h r e a d i n g s however, showed f i v e age groups, from-1 t o V, t o be p r e s e n t o v e r t h i s range i n s i z e , r e p r e -s e n t i n g ages from 1+ t o 5+. Thus, on t h e b a s i s o f o t o l i t h a n a l y s i s , when the young o f the y e a r a r e i n c l u d e d , the p o p u l a t i o n o f A n o p l a r c h u s s t u d i e d i s composed o f i n d i v i d u a l s f r o m l e s s t h a n One y e a r o f age, t o g r e a t e r t h a n f i v e y e a r s o f age, r e p r e s e n t i n g y e a r c l a s s e s 1959 t o 1963. I f o n l y t h e l e n g t h - f r e q u e n c y method o f a n a l y s i s had been employed, i t would have been i m p o s s i b l e t o show the age d i f f e r e n c e s between i n d i v i d u a l s w i t h i n s m a l l s i z e r a n g e s . Many-under- and o v e r - e s t i m a t e s would have been made. How-e v e r , the method does have v a l u e i n a p p r o x i m a t i n g the age groups p r e s e n t and i n t h i s r e s p e c t i n d i c a t e d f i s h o f age 1 t o age 4 t o be p r e s e n t . But t o s e p a r a t e i n d i v i d u a l s o f d i f -f e r e n t ages from a s i n g l e mode, a method such as o t o l i t h a n a l y s i s would have t o be employed. 131 Growth Curve T a b l e XX shows mean l e n g t h s o f the s e x e s , f o r each o f the determined ages. These mean l e n g t h s were used t o p l o t growth c u r v e s f o r the s e x e s , as p r e s e n t e d i n F i g u r e 11. The p o i n t f o r young of the y e a r (age< 1) i s based on 15 specimens, c o l l e c t e d between June, 1964 and O c t o b e r , 1964, o f a range i n l e n g t h from 25.5 to 47.3 (mean 36.5) mm. Rate o f growth of the f e m a l e s f r o m < l t o 1+ y e a r s of age i s c o n s t a n t , whereas t h a t o f the males d e c r e a s e s . Thus, a t 1+ y e a r s o f age, the f e m a l e s . t e n d t o be s l i g h t l y l a r g e r t h a n the males. From 1+ to 4+ y e a r s o f age, both c u r v e s show a s i m i l a r d e c r e a s e i n r a t e o f growth, w i t h f e m a l e s r e m a i n i n g a t a l a r g e r s i z e t h a n males at each age. Females o f 4+ y e a r s o f age were l a c k i n g f rom the sample o f o t o l i t h s s t u d i e d , thus the pronounced d e c r e a s e i n growth r a t e i n d i c a t e d f rom 3+ t o 5+ y e a r s o f age i s p r o b a b l y o v e r -emphasized as i t i s shown. A l t h o u g h the c u r v e s a re o n l y based on 34 o t o l i t h r e a d i n g s , t h e complete range i n s t a n d a r d l e n g t h ; e x h i b i t e d was examined and age d i f f e r e n c e s between s m a l l ranges i n s i z e were a n a l y z e d . 132 140r Age (years) Figure 11. Growth curves of male and female Anoplarchus, based on 34 o t o l i t h s ; 19 males, 15 females. Point of age< 1 based on 15 measurements. 133 Age a t S e x u a l M a t u r i t y The s exes show t h e i r f i r s t s i g n s o f m a t u r i t y a t a l e n g t h o f 80 t o 90 mm. At t h i s t i m e , t h e t e s t e s a r e e n l a r -g i n g and t h e o v a r i e s becoming g r a n u l a r . Spawning t a k e s p l a c e f o r t h e f i r s t t i me a t an age o f 2+ o r 3+ y e a r s . Sex R a t i o Of 269 f i s h i d e n t i f i e d t o s e x , males accounted f o r 112 i n d i v i d u a l s (41.64%) and f e m a l e s , 157 i n d i v i d u a l s ( 5 8 . 3 6 % ) . A s e x r a t i o i n f a v o u r o f f e m a l e s o f 1.4 : 1. T h i s n u m e r i c a l s u p e r i o r i t y o f f e m a l e s i s c o n s t a n t o v e r almost th e e n t i r e range i n s t a n d a r d l e n g t h . P r o g r e s s i v e changes i n s e x r a t i o are d e t a i l e d i n T a b l e XXI , a l o n g w i t h t h e p e r c e n t a g e breakdown by s e x o f t h e v a r i o u s l e n g t h group-i n g s . Length-Weight R e l a t i o n s h i p and C o n d i t i o n Method P r e s e r v e d specimens f r o m the 1963 samples o f August, September, and December and t h e 1964 samples o f J a n u a r y , May and J u l y , were used. W e i g h i n g was;performed 134 T a b l e X X I . P e r c e n t a g e breakdown by s e x o f the l e n g t h s e x h i b i t e d , and p r o g r e s s i v e sex r a t i o w i t h i n c r e a s i n g s i z e . v a r i o u s changes i n S t a n d a r d L e n g t h (mm) . No. Males % o f T o t a l % Sex No. Females % o f . , T o t a l % Sex 50.0- 55.0 2 1.79 100.0 0 55.0- 60.0 0 0 60.0- 65.0 3 2.68 60.0 2 1.27 40.0 65.0- 70.0 6 5.36 50.0 6 3.82 50.0 70.0- 75.0 2 1.79 25.0 6 3.82 75.0 75.0- 80.0 2 1.79 25.0 6 3.82 75.0 80.0- 85.0 4 3.57 33.3 8 5.10 66.7 85.0- 90.0 13 1 1 . 6 1 56.5 10 6.37 43.5 90.0- 95.0 11 9.82 35.5 20 12.74 64.5 95.0- 100.0 18 16.07 47.4 20 12.74 52.6 100.0- 105.0 17 15.18 40.5 25 15.92 59.5 105.0- 110.0 15 13.39 45.5 18 11.46 54.5 110.0- 115.0 8 7.14 36.4 14 8.92 63.6 115.0- 120.0 9 8.04 34.6 17 10.83 65.4 120.0- 125.0 2 1.79 40.0 3 1.91 60.0 125.0- 130.0 0 1 0.64 100.0 112 100.00 157 IQOSipO 135 i n A ugust, 1964. A t o t a l o f 92 f i s h were examined (46 males and 46 f e m a l e s ) . B e f o r e w e i g h i n g , a l l c o n t e n t s o f t h e body c a v i -t i e s were removed. Each f i s h was p l a c e d on paper t o w e l l i n g , a l l o w e d to d r y o f e x c e s s m o i s t u r e , and t h e n b e f o r e each was weighed, wrapped and squeezed l i g h t l y . W e i ghing was done on a M e t t l e r p r e c i s i o n b a l a n c e , t o t h e n e a r e s t 0.01 gram. F a c t o r s f o r C o n v e r s i o n Between S t a n d a r d and T o t a l L e n g t h The method p r e s e n t e d by Beckman (1945) was employed. I n t h e o r i g i n a l c o m p i l a t i o n o f s t a n d a r d and t o t a l l e n g t h s , each m i l l i m e t r e o f t o t a l l e n g t h was made a column h e a d i n g and each s t a n d a r d l e n g t h was r e c o r d e d i n t h e a p p r o p r i a t e column. Average s t a n d a r d l e n g t h s and t o t a l l e n g t h s were computed f o r 5 - m i l l i m e t r e i n t e r v a l s o f t o t a l l e n g t h . The r a t i o o f s t a n d -a r d t o t o t a l l e n g t h was d e t e r m i n e d f o r each o f t h e s e i n t e r -v a l s by d i v i d i n g the average s t a n d a r d l e n g t h by t h e average t o t a l l e n g t h . F o r example, two f i s h i n the 85 t o 90 mm group o f t o t a l l e n g t h , averaged 86.6 mm i n t o t a l l e n g t h and 79.5 mm i n s t a n d a r d l e n g t h . The v a l u e 79.5 / 86.6 = 0 . 9 1 5 i s the r a t i o f o r t h a t g r o u p . A r e v e r s e o f the l a t t e r , 86.6 / 79.5 = 1.089 g i v e s t h e r a t i o f o r c o n v e r s i o n o f s t a n d a r d l e n g t h t o t o t a l l e n g t h f o r t h a t group. 136 The o b t a i n e d c o n v e r s i o n f a c t o r s f o r t o t a l l e n g t h t o s t a n d a r d l e n g t h ranged from 0.912 t o 0.918, w i t h no t r e n d i n i n c r e a s i n g o r d e c r e a s i n g w i t h i n c r e a s i n g l e n g t h . The average r a t i o o f 0.9145 was computed and r e p r e s e n t s t h e f a c -t o r f o r c o n v e r s i o n o f t o t a l l e n g t h t o s t a n d a r d l e n g t h . . The o b t a i n e d c o n v e r s i o n f a c t o r s f o r s t a n d a r d l e n g t h t o t o t a l l e n g t h ranged from 1.088 t o 1.094, the a v e r -age r a t i o o f 1.0907 r e p r e s e n t s the f a c t o r f o r c o n v e r s i o n Of s t a n d a r d l e n g t h t o t o t a l l e n g t h . Length-Weight R e l a t i o n s h i p The e q u a t i o n used i n c o m p i l a t i o n of t h e l e n g t h -w e i g h t r e l a t i o n s h i p was t h a t o f t h e g e n e r a l p a r a b o l a , W = c L n , where W = weight i n grams; L = t o t a l ' l e n g t h i n m i l l i m e t r e s , and c and n a r e c o n s t a n t s . T h i s l a t t e r equa-t i o n , a c c o r d i n g t o H i l e ( 1 9 3 6 ) , g e n e r a l l y g i v e s a b e t t e r r e s u l t : i n •. the e x p r e s s i o n , o f t h e l e n g t h - w e i g h t r e l a t i o n s h i p t h a n does the c u b i c p a r a b o l a , W = c L^, where W = weight i n grams, L = t o t a l length-, i n m i l l i m e t r e s , and c i s a c o n s t a n t . The e q u a t i o n , W = c L n , e x p r e s s e d i n l o g a r i t h m i c f o rm becomes a s t r a i g h t l i n e : l o g W.= l o g c + n l o g L.- The v a l u e s o f l o g c and n were d e t e r m i n e d from the f o l l o w i n g e q u a t i o n s : 137 l l o g W • l ( ' l o g L ) 2 . - f l o g L • i_(log L»* log W) l O g C = : N • £(log L ) 2 - ( £ log L ) 2 and log W - (N • log c) n = £ log L The method employed i n compiling the data was that presented by Beckman (1945), and i s i l l u s t r a t e d i n Table XXII. This Table i s an excerpt from the o r i g i n a l tabulation. The obtained values f o r log c and n are as f o l -lows : log c = - 5.31565 and n = 2.98585 By su b s t i t u t i n g these values i n the logarithmic form of the equation, W = c L n , the calculated weights were determined. Table XXIII contains the length-weight data, including standard and t o t a l lengths i n millimetres, emprical weight i n grams and the calculated weights i n grams. The length-weight r e l a t i o n s h i p i s presented graphically i n Figure 1 2 . C o e f f i c i e n t of Condition The c o e f f i c i e n t of condition, K, was determined by the use of the following formula, K = 100,000 W / L ? , where T a b l e X X I I . E x c e r p t from d a t a t o i l l u s t r a t e t h e method employed f o r c o m p i l a t i o n of i n f o r m a t i o n on l e n g t h - w e i g h t r e l a t i o n s h i p T o t a l L e n g t h Standard Length No. of F i s h Ave. Weight l o g L l o g W l l o g L X l o g W ( l o g L) C a l c . l o g W C a l c . Weight 73.2 67.1 10 1.62 1.82672 0.12710 0.23218 3.33691 0.13866 1.37 105.3 96.5 11 5.59 2.02243 0.74741 1.51158 4.09022 0.72302 5.29 126.5 116.0 6 9.57 2.10209 0.98091 2.06196 4.41878 0.96088 9.14 t-1 CO 00 139 Table XXIII. The length-weight relationship and c o e f f i -cients of condition. Empirical Calculated Weight N Weight K (Gms) (Gms) 3 61.5 67.1 1.34 1.37 0.564 10 67.1 73.2 1.62 1.78 0.536 4 72.7 79.3 2.06 2.26 0.529 3 77.3 84.3 2.68 2.72 0.576 3 81.8 89.2 3.28 3.22 0.598 6 86.9 94.8 4.09 3.86 0.619 10 92.2 100.6 5.28 4.61 0.671 11 96.5 105.3 5.59 5.29 0.622 11 102.6 111.9 7.00 6.17 0.644 13 107.5 117.3 7.95 7.29 0.638 6 112.3 122.5 8.67 8.30 0.611 6 116.0 126.5 9.57 9.14 0.524 3 121.5 132.5 10.18 10.50 0.566 1 128.0 139.6 9.04 12.26 0.431 vr_ _.f Standard Total F?*sh ^ n g ^ h L ? n * f h (mm) (mm) 140 Total Length (mm) 60 70 80 90 100 110 120 130 140 — i 1 1 1 1 1 1 1 1 60 70 80 90 100 110 120 130 Standard Length (mm) Figure 12. Length-weight r e l a t i o n s h i p . The curve i s the graph of the length-weight equation; dots repre sent the calculated data. 141 W = weight i n grams and L = l e n g t h ( s t a n d a r d ) i n m i l l i m e t r e s . C o e f f i c i e n t s o f c o n d i t i o n c a l c u l a t e d f rom the cube r e l a t i o n -s h i p d e s c r i b e r e l a t i v e h e a v i n e s s i n d e p e n d e n t l y o f the gen-e r a l l e n g t h - w e i g h t r e l a t i o n s h i p , and a r e more s a t i s f a c t o r y measures o f c o n d i t i o n t h a n the q u a n t i t y c, i n t h e e q u a t i o n , W = c L n , where the v a l u e o f n i s d e t e r m i n e d e m p i r i c a l l y ( H i l e , 1936). The ranges and means o f c o e f f i c i e n t s o f c o n d i t i o n o b t a i n e d f o r each o f s i x samples examined are shown i n T a b l e XXIV, where the x v a l u e s , f o r the sexes a r e p r e s e n t e d s e p a r -a t e l y . The average v a l u e s f o r t h e grouped l e n g t h s a r e p r e s -ented i n T a b l e X X I I I , and g r a p h i c a l l y i n F i g u r e 14. Comparisons between i n d i v i d u a l s o f t h e same l e n g t h , sex and time o f c o l l e c t i o n , r e v e a l a c l o s e s i m i l a r i t y i n v a l u e s o f t h e c o e f f i c i e n t . Once comparisons a r e made between d i f f e r e n t l e n g t h s , o p p o s i t e sex and c o l l e c t i o n s s e p a r a t e d i n t i m e , d i f f e r e n c e s in the c o e f f i c i e n t s e n l a r g e . Some examples o f . v a r i a t i o n between i n d i v i d u a l s o f the same, l e n g t h are shown i n T a b l e XXV. D i f f e r e n c e s in the c o e f f i c i e n t o f c o n d i t i o n between •the sexes were not c o n s i s t e n t , • a l t h o u g h t h e males were found, t o have a s l i g h t l y higher, mean v a l u e i n - a l l - b u t - o n e sample ( T a b l e X X I V ) . The mean v a l u e s f o r both sexes show a s e a s o n a l t r e n d i n i n c r e a s i n g i n v a l u e from summer and f a l l months, T a b l e XXIV. D i f f e r e n c e s i n c o e f f i c i e n t s o f c o n d i t i o n o f 46 males and 46 f e m a l e s , of s i x samples examined. Males Females Date of •Av. C o e f f i c i e n t Av. C o e f f i c i e n t Sample No.' Length Range Mean No. L e n g t h Range Mean Aug., 1963 9 95.6 0.477-0.732 0.633 11 100.9 0.497-0.804 0.624 Sep., 1963 5 95.0 0.530-0.779 0.695 6 101.1 0.500-0.857 0.674 Dec.., 1963 8 101.1 0.460-0.825 0.694 8 106.1 0.645-0.755 0.707 J a n . , 1964 3 112.3 0.472-0.547 0.499 4 113.5 0.431-0.566 0.489 May, 1964 9 64.3 0.483-0.598 0.535 9 71.2 0.492-0.551 0.524 J u l . , 1964 12 93.8 0.492-0.674 0.582 8 97.0 0.480-0.609 0.565 T a b l e XXV. V a r i a t i o n i n c o e f f i c i e n t s o f c o n d i t i o n among i n d i v i d u a l s o f t h e same l e n g t h ; 10 comparisons between i n d i v i d u a l s o f the same s e x , 12 com-p a r i s o n s between i n d i v i d u a l s o f o p p o s i t e s e x . Same Sex O p p o s i t e Sex Date of L e n g th C o e f f i c i e n t L e n g t h C o e f f i c i e n t Sample Range Range Aug., 1963 108.9 0.565=0.645 92.2 0.732-0.804 96.9 0.570-0.701 108.9 0.565-0.588 Sep. , 1963 94.9 0.740-0.779 95.0 0.702-^0.740 103.8 0.689-0.857 Dec., 1963 105.1 0.676-0.742 89.5 0.752-0.755 109.1 0.715=0.718 91.7 0.460-0.729 J a n . , 1964 114.5 0.479-0.493 May, 1964 61.3 0.562-0.563 66.3 0.493-0.522 61.8 0.562-0.567 67.1 0.538-0.551 J u l . , 1964 86.6 0.576-0.607 68.0 0.480-0.595 105.8 0.569-0.605 103.7 0.551-0.674 109.5 0.598-0.609 112.0 0.548-0.579 144 r e a c h i n g a peak i n December, d e c r e a s i n g i n Ja n u a r y ( t h e f i r s t month o f the spawning p e r i o d ) , and t h e n r i s i n g a g a i n i n t he s p r i n g and summer ( F i g u r e 1 3 ) . The d e c r e a s e shown i n t h e J a n u a r y sample tends t o s u b s t a n t i a t e the p r e v i o u s c o n c l u s i o n s made c o n c e r n i n g f e e d i n g h a b i t s ( d i s c u s s i o n o f fo o d h a b i t s ) o f the s p e c i e s w i t h t h e approach and d u r i n g t h e b r e e d i n g s e a s o n . D i s c u s s i o n The l e n g t h - f r e q u e n c y method of age a n a l y s i s was a p p l i e d and i t s v a l i d i t y f o r t h e samples s t u d i e d q u a l i f i e d by o t o l i t h a n a l y s i s . On t h e whole, l e n g t h i s a poor i n d e x o f age. The amount o f o v e r l a p between c o n s e c u t i v e age groups i s too g r e a t and i n most ca s e s a f i s h o f a g i v e n l e n g t h might have any o f s e v e r a l ages. The l a r g e s t c o n t r i b -u t o r t o t h i s l a t t e r s i t u a t i o n was l i k e l y t h e s m a l l sample s i z e s . O t o l i t h a n a l y s i s was used t o supplement the l e n g t h - f r e q u e n c y method. The number o f age d e t e r m i n a t i o n s made by t h i s method was not g r e a t , a t o t a l o f 70 o t o l i t h s were s t u d i e d , w i t h each o t o l i t h examined a number o f t i m e s . E i t h e r a c o n s i s t e n t assignment o f age was made, o r the o t o l i t h s were marked as u n r e a d a b l e . T h i r t y - f o u r age d e t e r -m i n a t i o n s were made. I f o n l y the l e n g t h - f r e q u e n c y method .8 145 § 7 "O 8 .a £ .5r "0 o U •4r A u g , 1 9 6 3 Scp t ,1963 DG C , 1 9 6 3 Jan,1964 May,1964 July, 1964 M o n t h o f S a m p l e Figure 13. Variations i n c o e f f i c i e n t of condition of males (broken line) and females ( s o l i d line) i n r e l a -t i o n to the season. c o •5 «6 c o o o .5h +-> c o <*— o o u 5 0 6 0 Figure 14. 7 0 8 0 9 0 1 0 0 110 120 130 S t a n d a r d L e n g t h ( m m ) C o e f f i c i e n t of condition i n r e l a t i o n to length of f i s h . 146 o f a n a l y s i s had been employed age d i f f e r e n c e s between i n d i -v i d u a l s w i t h i n s m a l l s i z e ranges would have been i m p o s s i b l e t o show. The mean l e n g t h s o f t h o s e i n d i v i d u a l s used f o r t h e o t o l i t h r e a d i n g s were used t o p l o t growth c u r v e s f o r t h e s e x e s . The c u r v e s can be c o n s i d e r e d r a t h e r p r e l i m i n a r y , due t o t h e i r b a s i s on o n l y 34 f i s h , but t h e age d e t e r m i n a -t i o n s were made from a good range i n l e n g t h and d i d p o i n t out d i f f e r e n c e s i n age between i n d i v i d u a l s o f a p p r o x i m a t e l y t h e same l e n g t h . S u f f i c i e n t v a r i a t i o n i n i n d i v i d u a l growth showed t h a t t h e l a r g e s t i n d i v i d u a l need not n e c e s s a r i l y be t h e o l d e s t . I t i s a l s o t r u e f u r t h e r , t h a t t h e l a r g e s t f i s h w i t h r e s p e c t t o l e n g t h i s not always the h e a v i e s t * ( T a b l e s XIX and X X I I I ) . The g e n e r a l e q u a t i o n , W = c L n , was used t o d e s -c r i b e t h e l e n g t h - w e i g h t r e l a t i o n s h i p . The v a l u e s o f c and n i n t h i s e q u a t i o n were d e t e r m i n e d e m p i r i c a l l y . T h i s e q u a t i o n , i n c o n t r a s t t o t h e well-known cube law, W = c L^, has been shown t o be a much more s a t i s f a c t o r y method of d e s c r i b i n g t h e l e n g t h - w e i g h t r e l a t i o n s h i p i n f i s h e s ( H i l e , 1936). As H i l e p o i n t e d o u t , the use o f t h e cube law i n t h i s l a t t e r c a p a c i t y has met w i t h i n d i f f e r e n t s u c c e s s , due t o the f a i l u r e t o d e s c r i b e a c c u r a t e l y t h e r e l a t i o n s h i p s o f l e n g t h t o w e i g ht i n many forms o f f i s h e s . W i t h s c a r c e l y an 147 e x c e p t i o n , t h e w e i g h t a t a g i v e n l e n g t h i s g r e a t e r t h a n t h e c a l c u l a t e d w e i g h t from t h e law, so t h a t i f the s p e c i f i c g r a v i t y o f the f i s h e s remains c o n s t a n t t h e y must i n c r e a s e somewhat more i n o t h e r d i m e n s i o n s t h a n i n l e n g t h . The e q u a t i o n , W = c L^, was however, used i n t h e d e t e r m i n a t i o n s o f the c o e f f i c i e n t s o f c o n d i t i o n . C o e f f i c -i e n t s based on e m p i r i c a l exponents f a i l t o r e f l e c t d i f f e r -ences i n form o r r e l a t i v e h e a v i n e s s , w h i l e t h o s e based on the cube r e l a t i o n s h i p o f f e r a d i r e c t measure of r e l a t i v e h e a v i n e s s independent o f the g e n e r a l l e n g t h - w e i g h t r e l a t i o n -s h i p s and comparable as measures o f r e l a t i v e h e a v i n e s s between f i s h o f any l e n g t h ( H i l e , 1936). The f o l l o w i n g r e a s o n i n g i s put f o r t h by H i l e . Inasmuch as r e l a t i v e h e a v i -ness i s shown t o be dependent on f a t n e s s ( c o n d i t i o n ) , changes w i t h l e n g t h i n r e l a t i v e h e a v i n e s s must be c o n s i d e r e d a l s o t o r e p r e s e n t changes o f c o n d i t i o n . I n v i e w o f t h i s f a c t i t does not appear v a l i d t o measure c o n d i t i o n i n terms o f a q u a n t i t y t h a t tends t o be c o n s t a n t f o r f i s h o f a l l l e n g t h s r e g a r d l e s s o f a c t u a l changes t h a t may o c c u r i n r e l a t i v e h e a v i n e s s o f form w i t h change i n l e n g t h . S i n c e t h e q u a n i t y c i n the e q u a t i o n , W = c L n , tends toward t h i s c o n s t a n c y and f a i l s t o measure r e l a t i v e h e a v i n e s s , i t must f a i l a l s o t o measure the d i f f e r e n c e s o f f a t n e s s ( c o n d i t i o n ) upon wh i c h d i f f e r e n c e s i n r e l a t i v e h e a v i n e s s depend. '. 148 The c o e f f i c i e n t o f c o n d i t i o n was used c h i e f l y as a measure o f the s t a t e o f n o u r i s h m e n t . The f l u c t u a t i o n s o b t a i n e d were due t o some changes i n c o m p o s i t i o n o f t h e body t i s s u e s , presumably an i n c r e a s e o r d e c r e a s e i n t h e f a t c o n t e n t , and not t o t h e amount of f o o d i n t h e a l i m e n t a r y t r a c t o r t o the growth o f t h e f s e x o r g a n s , as t h e d i g e s t i v e t r a c t and organs and t h e gonads were removed b e f o r e w e i g h t s were t a k e n . Used i n t h i s way, t h e c o e f f i c i e n t has v a l u e i n s u p p l e m e n t i n g the f o o d h a b i t s a n a l y s i s . 149 SUMMARY (1) S y s t e m a t i c s A n o p l a r c h u s p u r p u r e s c e n s i s a p e r c i f o r m e t e l e o s t b e l o n g i n g t o the s u b o r d e r B l e n n i o i d e i , f a m i l y S t i c h a e i d a e . The genus A n o p l a r c h u s i s p l a c e d w i t h i n t h e s u b f a m i l y ALec-t r i i n a e , a l o n g w i t h t h e genera A l e c t r i a s and P s e u d a l e c t r i a s . A. p u r p u r e s c e n s and i t s s i b l i n g s p e c i e s , A. i n s i g n i s , a r e g e o g r a p h i c a l l y s y m p a t r i c , but i n B r i t i s h Columbian c o a s t a l w a t e r s , t h e l a t t e r p r e f e r s deeper w a t e r t h a n A. p u r p u r e s c e n s . The range o f A. p u r p u r e s c e n s i s wide, f r o m A t t u I s l a n d and t h e P r i b i l o f I s l a n d s , A l a s k a , t o cen-t r a l C a l i f o r n i a . I n t h e a r e a where c o l l e c t e d and a d j a c e n t w a t e r s , A. p u r p u r e s c e n s can be d i s t i n g u i s h e d f r om a l l o t h e r b l e n -n i o i d s by; absence o f p e l v i c f i n s , p r e s e n c e o f t h e der m a l c r e s t on the head, s c a l e s on t h e p o s t e r i o r h a l f o f t h e body o n l y and t h e g r e a t w i d t h between the p o i n t s o f attachment of each g i l l membrane to t h e i s t h m u s . I t may be d i s t i n g -u i s h e d f r o m A. i n s i g n i s by i t s l o w e r m e r i s t i c c o u n t s and the g r e a t e r w i d t h between the p o i n t s o f attachment o f each g i l l membrane t o t h e i s t h m u s . The s p e c i e s was f i r s t r e c o r d e d i n B r i t i s h 150 Columbian w a t e r s i n 1861 from Vancouver I s l a n d and t h e mouth o f t h e F r a s e r R i v e r by A. G i i n t h e r , as C e n t r e not us  c r i s t a g a l l i . (2) H a b i t a t A n o p l a r c h u s i s a b o t t o m - d w e l l i n g form, i n h a b i t i n g the i n t e r t i d a l zone, from upper m i d - i n t e r t i d a l t o low-i n t e r t i d a l . I t s o c c u r r e n c e i n the uppermost p a r t o f t h e m i d - i n t e r t i d a l , p l a c e s i t h i g h e r up i n t h e i n t e r t i d a l zone t h a n any o t h e r " e e l b l e n n y " i n t h e a r e a . A n o p l a r c h u s i s v e r y c l o s e l y a s s o c i a t e d w i t h the s u b s t r a t e o f r o c k s and a l g a e , o c c u p y i n g a v a i l a b l e spaces beneath t h e r o c k s f o r day to day a c t i v i t i e s as s h e l t e r and f e e d i n g and s p e c i a l r e q u i r e m e n t s as spawning. A n i m a l a s s o c i a t e s i n c l u d e a number o f bottom-d w e l l i n g c r u s t a c e a n s and f i s h e s . (3) Movements M a r k i n g e x p e r i m e n t s d i d not i n d i c a t e a s t r o n g tendency t o home but t h e y d i d show movements o f A n o p l a r c h u s t o be r a t h e r r e s t r i c t e d . Of the t o t a l number o f f i s h r e c a p t u r e d , 58.0% showed a homing tendency. The s t r a y i n g was o b s e r v e d t o be o f two t y p e s ; e i t h e r t o the t r a n s p l a n t a r e a o r , t o some o t h e r a r e a . S t r a y i n g o f the l a t t e r t y pe 151 was shown by 34.0% o f t h o s e f i s h r e c a p t u r e d . Marked f i s h however, were r a r e l y found more than.50 f e e t f rom where o r i g i n a l l y c a p t u r e d , A n o p l a r c h u s d i s p l a y s b o t h home-range and defended t e r r i t o r i a l i t y . I n d i v i d u a l s s h a r e a home-range t e r r i t o r y d u r i n g n o n - b r e e d i n g t i m e s o f t h e y e a r , but d u r i n g t h e b r e e d i n g s e a s o n , i n d i v i d u a l s a r e . h i g h l y s e g r e g a t e d , s p a c i -a l l y and s e x u a l l y , i n t o i s o l a t e d b r e e d i n g p a i r s . At t h i s t i m e defended t e r r i t o r i a l i t y i s e x h i b i t e d . The s m a l l b r e e d i n g t e r r i t o r y appears t o be main-t a i n e d by a g g r e s s i o n on t h e p a r t o f the male b e f o r e spawn-i n g , and by the f e m a l e once spawning i s c o m p l e t e . The l a t t e r d e f e n c e i s m a i n t a i n e d t h r o u g h the p e r i o d o f egg i n c u b a t i o n , t o the time o f h a t c h i n g . (4) Food and F e e d i n g A n o p l a r c h u s appears t o be a d a y l i g h t f e e d e r , u t i l i z i n g as f o o d t h o s e organisms p r e s e n t and a v a i l a b l e i n t h e p a r t i c u l a r h a b i t a t b e i n g r e s i d e d i n a t the :.time o f f e e d -i n g . R e l a t i v e i m p o r t a n c e s o f the v a r i o u s f o o d i t e m s r e f l e c -t e d d i f f e r e n c e s i n a v a i l a b i l i t y o f the o r g a n i s m s u t i l i z e d as f o o d i n d i f f e r e n t t i d a l l e v e l s o f the i n t e r t i d a l zone. I n u t i l i z a t i o n o f one f o o d i t e m , a l g a e , t h e r e appeared t o be a d e f i n i t e p r e f e r e n c e f o r green a l g a e . 152 A n o p l a r c h u s can be c o n s i d e r e d f a i r l y s t e n o p h a g i c i n i t s f o o d h a b i t s . B a s i c f o o d s i n c l u d e ; a l g a e , p o l y c h a e t e worms ( e x c l u d i n g n e r e i d worms), nemertean worms, amphipods and f l a t w o r m s . M u s s e l s , l i t t o r i n e s n a i l s , i s o p o d s and n e r e i d worms compose the r e m a i n i n g p o r t i o n o f t h e d i e t . The b a s i c f o o d s a r e u t i l i z e d by A n o p l a r c h u s o f a l l s i z e s . Food i n t a k e i s c u r t a i l e d i n a d u l t f i s h a p p r o a c h i n g and d u r i n g t h e b r e e d i n g s e a s o n . (5) R e p r o d u c t i o n Sexes a r e d i m o r p h i c ; c o l o u r a t i o n and markings o f the body a t a l l t i m e s o f t h e y e a r , cockscomb when s e x u a l l y mature and s i z e . F e c u n d i t y i s h i g h (2,001 t o 3,183 e g g s ) . The e n t i r e egg p r o d u c t i o n o f each f e m a l e i s c o n c e n t r a t e d i n t o a s i n g l e spawning a c t . Number o f eggs i n c r e a s e s w i t h i n c r e a s i n g f e m a l e s i z e . Average d i a m e t e r o f the eggs i s 0.864 mm and 1.372 mm, f o r u n f e r t i l i z e d and f e r t i l i z e d eggs, r e s p e c t i v e l y . The sexes form p a i r - b o n d s b e f o r e a c t u a l spawning. P r e - c o p u l a t o r y d i s p l a y shows the male t o p l a y a v e r y a g g r e s s i v e r o l e , w i t h t h r e a t d i s p l a y and a c t u a l b i t i n g . Spawning t a k e s p l a c e i n t h e months o f J a n u a r y and F e b r u a r y . 153 The f e m a l e A. p u r p u r e s c e n s guards and tends i t s eggs; c h a r a c t e r i z e d by c o i l i n g about t h e eggs and f a n n i n g . D u r i n g p a r e n t a l c a r e o f eggs the male i s r e p u l s e d f rom the n e s t i n g s i t e . Care i s p r o v i d e d c o n t i n u a l l y t o complete h a t c h i n g o f the eggs. H a t c h i n g t i m e s o f 13 and 16 days were o b t a i n e d f o r t e m p e r a t u r e s o f 10 t o 12 eC and 7 t o 9°C, r e s p e c t i v e l y . The newly h a t c h e d l a r v a i s 7.4 t o 7.6 mm i n t o t a l l e n g t h , f a i r l y t r a n s p a r e n t and pigmented a t s e v e r a l p o s i -t i o n s on t h e body w i t h melanophores. An o v a l y o l k sac w i t h s i n g l e o i l d r o p l e t i s p r e s e n t f o r f i v e d a y s . The l a r v a e show marked p o s i t i v e p h o t o a x i s f o r t h r e e t o f i v e d a y s , t h e n are n e g a t i v e l y p h o t o t a o t i c . (6) Age and Growth On the b a s i s o f l e n g t h - f r e q u e n c y and o t o l i t h a n a l y s e s , t h e p o p u l a t i o n o f A n o p l a r c h u s a t t h e time o f the p r e s e n t s t u d y was composed o f i n d i v i d u a l s from l e s s t h a n one y e a r o f age, t o g r e a t e r t h a n f i v e y e a r s o f age; age group 0, t o age g r o u p : V , r e p r e s e n t i n g y e a r c l a s s e s 1959 t o 1963. Females show a s l i g h t l y f a s t e r r a t e o f growth' t h a n males and are l a r g e r t h a n males a t a l l ages. 154 Spawning f o r the f i r s t time occurs at an age of 2+ or 3+ years. A sex r a t i o i n favour of females of 1.4 : 1 was shown f o r 269 f i s h . The sex r a t i o remains i n favour of females over almost the entire range i n length exhibited. The value of the exponent n, i n the length-weight equation, W = c L n , was found to be 2.98585; log c -5.31565. 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