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The systematics of the prickly sculpin, cottus asper : an investigation of genetic and non-genetic variation.. 1965

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The University of B r i t i s h Columbia FACULTY OF GRADUATE STUDIES PROGRAMME OF THE FINAL ORAL EXAMINATION FOR THE DEGREE OF DOCTOR OF PHILOSOPHY of RICHARD JOSEPH KREJSA B.Sc, Michigan State College, 1954 M.A., University of C a l i f o r n i a , Los Angeles, 1958 TUESDAY, JUNE 8th,1965, AT 4:00 P.M. IN ROOM 3332, BIOLOGICAL SCIENCES BUILDING COMMITTEE IN CHARGE Chairman: 1= McT. Cowan D< H. Chi t t y H. D. Fisher W. S. Hoar C. C. Lindsey G„ E. Rouse N. J . Wilimovsky P. A. Dehnel. External Examiner: Dr. F e r r i s Neave P a c i f i c B i o l o g i c a l Station Nanaimo, B „ G. ' THE SYSTEMATICS OF THE PRICKLY SCULPI.N, COTTUS ASPER, AN INVESTIGATION OF GENETIC AND ""NON-GENETIC VARIATION WITHIN A POLYTYPIC SPECIES, ABSTRACT The p r i c k l y sculpin., Cottus jaspjar, i s a polytypic species c h a r a c t e r i s t i c a l l y represented by very p r i c k l y , fresh-water spawning "inland" forms., and less p r i c k l y , brackish-water spawning " c o a s t a l " forms. Its wide- spread geographic d i s t r i b u t i o n , pronounced phenotypic v a r i a b i l i t y , and the e f f e c t s of the contrasting environ- ments i n which i t occurs are the subject of t h i s investigation,. A complete nom.encl.at.ural h i s t o r y and synonymy of the species for the period 1836-1936 i s presented. Morphological evidence i s interpreted i n the l i g h t of f i e l d and laboratory studies of the migration and reproductive behaviour. Differences found i n d i s t r i b u t i o n and i n t e n s i t y of body p r i c k l e s (modified scales), geographical d i s t r i b u - tion," and migratory behaviour, support, the contention that the polytypy of "coastal." and "inland" forms of Cottus asper has a genetic basis, Other morphological evidence i s equivocal i n support of t h i s i n t e r p r e t a t i o n . Differences i n f i n ray counts, e.g, pectoral rays, between " c o a s t a l " and "inland" forms are correlated p o s i t i v e l y with the presence or absence of s a l i n i t y i n the environment. However, it. i s not known whether such differences are the r e s u l t of i n d i v i d u a l modifications induced by the l o c a l environment, or due to long term genetic f i x a t i o n . Evidence from studies of c l o s e l y - r e l a t e d species supports the i n t e r p r e t a t i o n that Gj__ti_s asper i s poly- typic species which has become, and i s i n the process of becoming, modified into several c h a r a c t e r i s t i c geno- types. Evolution within t h i s "asper species group" probably occurred i n three stages, each correlated with past geologic h i s t o r y . During the f i r s t stage, perhaps i n the Pliocene or Late Miocene, " c o a s t a l " and "inland" forms were derived from an ancestral marine c o t t i d . The second stage, during the Pleistocene, was characterized by fragmenta- t i o n of the gene pool of the "inland" form into a series of geographically i s o l a t e d populations which have since evolved into valid'--species. The t h i r d stage, i n Recent Time since the retreat of the C o r d i l l e r a n g l a c i e r s , has occurred p r i m a r i l y within the northern representatives of the " c o a s t a l " form.. At. least two derivatives can now be distinguished within the " c o a s t a l " form, GRADUATE STUDIES F i e l d of Study; Zoology Functional Morphology of Marine Zoogeography T e r r e s t r i a l Zoogeography Fishes C c C„ Lindsey J„ C, Briggs M, D. V. Udvardy Related Studies: Paleontology Philosophical Problems V. J. Okulitch B. Savery PUBLICATIONS Krejsa, R.J. 1960. The eastern, t r o p i c a l P a c i f i c fishes of the genus Blenniolus, including a new i s l a n d endemic, Copeia, 1960, No. 4, 322-336. M c A l l i s t e r , D.E. and Krejsa, R.J. 1961. Placement of the prow-fishes, Zaproridae, i n the superfamily Stichaeoidae. Nat. H i s t . Pap., Nat. Mus. Canada, I I , 1-4. Krejsa, R.J C 1964, Reproductive behavior and Sexual d i dimorphism i n the manacled sculpin, Synchirus g i l l i Bean. Copeia, 1964, No, 2, 448-450. THE SYSTEMATICS OF THE PRICKLY S C U L P I N , COTTUS A S P E R ; AN INVESTIGATION OF GENE T I C AND NON-GENETIC VARIATION WITHIN A POLYTYPIC SPECIES b y RICHARD JOSEPH KREJSA B . S o , M i c h i g a n S t a t e C o l l e g e , 195U M . A . , U n i v e r s i t y o f C a l i f o r n i a , L o s A n g e l e s , 1958 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY i n the Depar tment o f ZOOLOGY We accept this thesis as conforming t o the r e q u i r e d s t a n d a r d THE UNIVERSITY OF BRITISH COLUMBIA J U N E , 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 of the requirements f o r an advanced degree at the U n i v e r s i t y of • B r i t i s h Columbia, I agree that the 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 reference and study* I f u r t h e r agree that per- m i s s i o n f o r extensive copying of t h i s t h e s i s f o r s c h o l a r l y purposes may be granted by the Head of my Department or by h i s r e p r e s e n t a t i v e s . I t i s understood t h a t , c o p y i n g or p u b l i - c a t i o n of t h i s t h e s i s f o r f i n a n c i a l gain s h a l l not be allowed without my w r i t t e n permission* Department of. o o The U n i v e r s i t y of B r i t i s h Columbia, Vancouver 8 ? Canada Date 8 0 une I ̂  4 S i i ABSTRACT The prickly sculpin, Cottus asper, is a polytypic species characteristi- cally represented by very prickly, fresh-water spawning "inland" forms, and less prickly, brackish-water spawning "coastal" forms. Its widespread geo- graphic distribution, pronounced phenotypic v a r i a b i l i t y , and the effects of the contrasting environments in which i t occurs are the subject of this i n - vestigation. A complete nomenclatural history and synonymy of the species * for the period 1836-1936 is presented. Morphological evidence is interpreted in the light of f i e l d and laboratory studies of the migration and reproduc- tive behavior. Differences found in distribution and intensity of body prickles (modified scales), geographical distribution, and migratory behavior, support the contention that the polytypy of "coastal" and "inland" forms of Cottus asper has a genetic basis. Other morphological evidence i s equivocal in support of this interpretation. Differences in f i n ray counts, e.g. , pectoral rays, between "coastal" and "inland" forms are correlated positively with the presence or absence of salin i t y in the environment. However, i t is not known whether such differences are the result of individual modifications induced by the local environment, or due to long term genetic fixation. Evidence from studies of closely-related species supports the inter- pretation that Cottus asper i s a polytypic species which has become, and is in the process of becoming, modified into^several characteristic genotypes. Evolution within this '?asper species group",probably occurred in three stages, each correlated with past geologic history. During the f i r s t stage, perhaps in the Pliocene or Late Miocene, "coastal" and "inland" forms were derived from an ancestral marine cottid. The second stage, during the Pleistocene, was characterized by fragmentation of the gene pool of the "inland" form into a series of geographically isolated populations which have since evolved into valid species. The third i i i stage, in Recent Time since the retreat of the Cordilleran glaciers, has occurred primarily within the northern representatives of the "coastal" form0 At least two derivatives can now be distinguished within the "coastal" form. TABLE OF CONTENTS ABS TRACT o f l o e e o o o o o o o o e o o o o o o o o o o TABLE OF CON TEN TS 0 0 0 0 0 0 4 0 0 0 0 0 0 0 0 0 0 0 0 LXST OF TABLES 0 * 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 LXST OF FX GURES o o o o o e o o o o o o o o o o o o o o ACKN OWLEDGEMENTS o e o e o e o o o o o o o o o o o o o e INTRODUCTION o . • 0 o 0 o o 0 a e 0 0 0 0 0 0 0 0 0 0 e o . Literature Re view o e e o o o o o o o o o o o o o o MORPHOLOGICAL ANALYSIS OF PRICKLY SCULPIN POPULATIONS . Materials and Methods . o o a o o o o o o o o o o o Museum Col l e c t i o n s o»»»*t»*»»»**».» Material Examined , . . o o . o o o « « « <> « <, Counts and Measurements « . . » . . » o . « > « C r i t i q u e of the Methods pf Determining P r i c k l i n g D i s t r i b u t i o n and Intensity . . « o o o o <> « « o S t a t i s t i c a l Methods o o . o o . o o o o o o o o O O O O O V O O O O O O O O O O O O O O O O Nomenclatural History . o . o o o « o o o o o o D i s t r i b u t i o n O o o o « o o o o o o o o » o o o o o Ran ge 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 9 0 0 0 Results; Analysis of Geographic V a r i a t i o n « , « «- o P r i c k l i n g D i s t r i b u t i o n , 0 . 0 0 0 0 0 0 0 0 0 0 P r i c k l i n g Intensity o o o o o o o o o o o o o o Pectoral Fin Asymmetry . . . . 0 « 0 « « 0 0 . 0 Pectoral Fin Ray Count 8 0 > . • . 0 0 Spiny Dorsal Fin 0 0 0 0 0 0 0 0 . 0 0 0 0 0 0 0 Soft Dorsal Fin o o o o o o - . o o o e o o o o o Combined Dorsal Fin Elements . . « ' . . . . . . « V Alldl PlTl o o c o e o o o o o o o o A A O O O o o o o o o e o e 3S PsXVIC Pin o a o o o « « o f l o e o o * o o e o o < j o o ( » 9 o o 0̂ Latitudinal Variation in Ranges of Mean Fin Ray Counts , . . . 40 Lateral Line O 0 o e o 6 e . . o o o o o o o o . . . 0 . 0 0 0 43 Discussion of Meristic Variation Within C. asper < , . „ . . . 0 . 43 Summary of Morphological Analysis . . . . . . . . . « <> . . . . . 49 STUDIES ON THE LIFE HISTORY OF THE PRICKLY SCULPIN . . . . . . . . . . . 51 Present Knowledge of Spawning Migration in Cottus asper . . . . . 51 Spawning Period and Temperature Relationships . . . . . . . . . . 52 Materials and Methods . c r , * = ( c ? • « • « • . . • • 55 Sampling Localities and Study Streams t t t » » 55 Results of Field Studies . . . . o o . . . . . . . . . . . . . . 58 Salinity Tolerance in Prickly Sculpin Eggs e . . . . . . . . 62 DlSCUSSXOn o . e o o o e o . o o o ' o . e t i G ? 3 s . f l e . 0 o 0 64 Summary of Life History Studies . . . . . . . . . « . « . . . . . 65 DISCUSSION o o ~ . . o o o o e o o o 6 6 6 6 . 0 0 0 0 0 0 0 0 0 0 0 0 0 65 Morphological Variation Within„Cottus asper . . . . . . 66 The Phylogeny of Cottus asper and Closely-Related Species . . . . 70 Stage l o o o . e o o e e e o o o o o o o e o o o o o o o o . 72 State l i e 0 0 6 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 . 0 0 6 9 0 . 0 74 Stage' IIIo o o . 0 0 0 0 0 0 0 0 0 . 9 6 0 0 4 . 0 0 0 . a e 79 LITERATURE CITED 6 o o o o . . o . . o 6 o « . . . . 0 . . e . « « > 0 83 APPENDIX 0 0 0 0 0 0 0 0 0 0 9 0 0 6 0 0 0 0 0 . 0 0 0 0 0 0 6 0 0 . 9 90 TahleS 0 0 0 0 0 0 6 6 6 0 0 0 . 0 0 0 . 0 0 0 0 6 6 6 6 0 0 0 0 90 1. List of Collections, Localities, and Other Pertinent Information O 6 0 O 6 . « « 6 . . 6 O « « . 6 O 0 90 2. Weighted Mean Percentage Prickling and Resorption for Individual Populations of Cottus asper Used in Compiling Table I o . 6 0 6 . e o o o o . o . . 6 . 6 0 9 98 3 , Non-Weighted Mean Percentage Virtual Prickling in Each of 1 4 Body Sections for A l l Populations (Specimens ^ 6 0 mm S 0 L < i ) o o o o o o o o o 8 o a o o o » » 1 0 0 4 „ Mean Relative Prickling Intensity Index Values for A l l Populations of Cottus asper , « . . . . 1 0 2 50 Pectoral Fin Asyfflme-try in Cottus asper by Populations o o o o o o o o o o o o a o a o o o o o a o o 1 0 4 60 Summary of Meristic Counts (Means and Ranges) for A l l Populations of Cottus asper o o o o o o o a o o o a e o o 1 0 7 7 o Monthly Mean Temperature in °C» for Selected Coastal and Inland Localities, e o o o o s o o a o a t o o a s s a 1 0 9 v i i LIST OF TABLES Table Title I Mean Percentage Prickling and Resorption Among Aggregate Populations of Cottus asper i . . . . . . . . . . . 27 II Mean Relative Prickling Intensity for Populations of Cottus asper Aggregated by Locality and Prickling Affinity. Skin Section " j " . . . . . . 30 III Incidence of Occurrence, Direction, and Amount of Pectoral Fin Asymmetry Within and Among Aggregated Populations of Cottus asper . . . . . . . . . . 32 IV Lateral Line Pored-Scale Counts by Size Groups for Selected Representative Populations of Cottus asper . . . . 44 V Summary of Variable Characteristics for Selected Phenotypes of Cottus asper . . . . . . . . . . . . . . . 0 . 67 VI Summary of Meristic Data for Cottus asper and Other Species in ithe "asper species group" ° ° . 0 0 9 . 6 0 0 . 77 v i i i LIST OF FIGURES Figure Title 1. "Skin Map" Used for Recording Distribution and Intensity of Prickling in Cottus asper 0 . . . » • 7 2. Areas of Prickling and Areas of Resorptions A. in Coastal Cottus asper . . . . . . . . . . 9 Bo in Inland Cottus asper o o o o o o .. • 9 3. Distributional Range of Cottus asper . . . . . . . . . . . . 25 4. Virtual, Resorbed, and Apparent Prickling in CottUS aSper o o ' o . f l . o o . o . . . . . . . . . . . . . . 26 5. Mean Pectoral Fin Ray Count by Latitude . . . . , . . . . * 34 6 0 Mean Spiny Dorsal Fin Ray Count by Latitude . . 36 7. Mean Soft Dorsal Fin Ray Count by Latitude . , . . . . . . . 37 8. Hubbs-Hubbs Plot of Soft Dorsal Fin Ray Counts for Inland Population Samples Pooled into Aggregates of Two Degrees Latitude . . . . . o . . . . . . . . . . . . . 38 9. Mean Combined Dorsal Fin Ray Count by Latitude . . . . . . . 39 10. Mean Anal Fin Ray Count by Latitude . . . . . . . . . . . . 41 11. Approximate Ranges of Mean Fin Ray Counts Plotted Against Latitude for Inland, Coastal, Coastal Lake, and Coastal Derivative Populations . . . . . . . . 42 12. Monthly Mean Temperatures in °C. Arranged by Latitude for Coastal and Inland Localities Encompassing the Distributional Range of Cottus asper . . . . . . . . . . . . 54§ 13. Collection Localities for Spawning Populations of Cottus asper Used in Life History Studies . . . . . . . . . 56 14. Collection Localities Within L i t t l e Campbell R. . . . . . . 57 15. Collection Sites, Tidal Influence, and Stream Gradients in the L i t t l e Campbell R. . . . . . . . . . . . . 59 Figure Title 16„ Monthly Distribution of Young-of-the-Year Cottus asper Within the L i t t l e Campbell R„ Taken from Pooled Bi = Weekly Samples o o o o . o o o o o e o o o o o , , . . . . . 61 ,17. Distribution of Yearling, Sub-Adult, and Adult Cottus asper Within the L i t t l e Campbell R. . . . . . . . . . 63 18. Known Ranges of Various Phenotypes of Cottus asper Plotted Against the Distributional Ranges of Closely- Related Species in the "asper species group" . . . . . . . . 71 19. Phylogeny of Cottus asper. Stage I. Evolution in the Pliocene (? ) o o o O O O O O O O 0 0 0 0 0 0 e o a o o a 73 20. Phylogeny of Cottus asper. Stage II, Evolution of "asper species group" During the Pleistocene 78 21. Phylogeny of Cottus asper. Stage III. Evolution of Coastal Form Following Retreat of Cordilleran Glaciers During Recent Time . . , , , e s '. , . , . . , , , , « . . . 80 X ACKNOWLEDGEMENTS I wish to express my indebtedness to the many persons and organizations who have given me encouragement and assistance during the period of this i n - vestigation. The British Columbia Fish and Game Branch provided financial support during the summer of 1960 and also the much appreciated assistance of Dr. G. F. Hartman and Mr. C„ A„ G i l l in f i e l d studies. The National Research Council of Canada provided studentships in 1961 and 1962. The Vancouver Public Aquarium provided research space for l i f e history studies and I wish to thank the Curator and staff for a l l their help and kindnesses. Almost every graduate student enrolled in the Institute of Fisheries volunteered assistance in f i e l d collections at one time or another during the period 1960-1963. I offer a collective thanks to a l l 0 Dr. C. Co Lindsey's assistance in supervision of the entire investi- gation cannot be measured in words. His door was always open to me and I took advantage of that fact on innumerable occasions. Drs. N. J. Wilimovsky provided encouragement and many helpful sug- gestions and criticisms throughout the investigation, and above a l l , friendship. Dr. R. H, Rosenblatt provided much inspiration during the period of his ( . . . ... v i s i t in Van'couver0 Dr. Carl E. Bond shared his wealth of knowledge about the members of the genus Cottus. Committee members Drs 0 W. S, Hoar, E, C. Black, P. A, Dehnel, and D. Wo Chitty read the manuscript and offered helpful suggestions and c r i t i - cisms throughout the study. Mrso Mary Williams typed the f i n a l copy and her perserverance i s greatly appreciated. xi Finally, I wish to offer my sincere gratitude for the help and en- couragement of my wife Julie, who, while patiently tolerating a l l the indignities encompassed within the title "wife of graduate student", found time to act as literary critic, typist, and friendo I INTRODUCTION The role of the environment versus the role of heredity in determining the phenotypic variations found among and within natural populations has long been a problem of central interest to students of evolution. Because fishes apparently undergo modification by their environment, they have proved to be useful in the study of phenotypic variation. For example, several authors have shown that non-genetic morphological or func*- tional variation in fishes can be caused by maternal care and diet prior to egg deposition, and to the effects of environment during f e r t i l i z a t i o n and development, and after hatching. In spite of such documentation many systematic studies ta c i t l y assume that differences discovered between i n * dividuals, or within and/or among series of specimens, are genetic. In some species, breeding experiments have shown that, under controlled conditions, the integrity of the parental phenotype i s maintained in the F^ generation. Such results have been interpreted as being genetically deter- mined. Except for a few notable examples, there have been no studies on fishes which have attempted to correlate the morphological and physiological evidence and to relate both to the l i f e history of the species concerned. Such an approach i s attempted in. this investigation. The prickly sculpin ranges geographically over about 3000 miles of Pacific North Temperate coastline and inland as far as 300 miles. Structural modifications correlated with latitudinal changes in temperature 0.& sal i n i t y should, i f present, be revealed by morphological analysis. Like the stic k l e - back Gasterosteus aculeatus, and the darter Etheostoma (Boleosoma) nigrum, the prickly sculpin exists in two modes of morphological variability? one, a fresh-water spawner, has extensive squamation on certain regions of the body; the other, a brackish-water spawner, has l i t t l e or none. Since prickly sculpin eggs are spawned naturally in environments which are known to vary in at least one major factor, i.e,, s a l i n i t y , and since, under contrasting environmental circumstances, i t shows so many similarities with the stickle- back, the prickly sculpin is an excellent subject for variational studies. Prior to the present study, regional systematic treatments of this widespread species resulted in a proliferation of generic and specific taxa, a l l referable to Cottus asper. A complete nomenclatural history and synonymy of the species for the period 1836-1936 is presented. Morphological evidence based.on examination of 1604 specimens from 190 collections throughout the range of the species is correlated with that from field and laboratory studies of the migratory and reproductive behavior. The thesis is presented in two sections corresponding to the morpho- logical and li f e history approaches followed in this investigation. A - general discussion considers the morphological and ecological variation with- in the species and attempts to reconstruct the phylogeny of Cottus asper and closely-related species. Literature Review Hubbs (1943) has listed many reasons why fishes are particularly well- suited for determination of genetic vs. environmental responses. He has also summarized (1926) much of the early work especially in regard to the struc- tural consequences of varying developmental rates. Numerous workers have found that different morphological or meristic series in fishes are "plastic," i.e., sensitive to environmental influence before egg deposition, during fertilization, during development, and after hatching. Taning (1944) ascribed at least one instance of mass vertebral de- formities and low egg survival to an accident befalling the female trout : before the eggs were removed from her. Hubbs and Stavenhagen (1958) have shown that low survival of different egg lots was caused by carotenoid deficiency in the maternal diet.' Kinne (1962) reviewed some of the recent literature on structural and functional properties of fish egg membranes and suggested that the spawning 3 medium affects the physical properties of the immediate environment of the embryo and that these induced properties "tend to persist in later different salinities." On the basis of his own studies, of Cyprinodon macularius, he interprets such conditioning by the spawning medium, leading to irreversible functional or structural adjustment in the embryo, as a special case of imprinting, or "irreversible non-genetic adaptation." Battle (1929); Mottley (1934); Taning (1944); Gabriel (1944); Heuts (1949); Lindsey (1954);, and Seymoilr (1959), have demonstrated the effects of environmental influences, especially temperature, during different develop- mental stages prior to hatching. Strawn (1961) found that the diet of newly-hatched larvae can affect the morpbGi&gy'* Lindsey (1954) found that certain meristic characters are thermolabile up to 20 days after hatching. Lagler and Bailey (1947) studied the inheritance of differential mor- phological characters in two subspecies of Etheostoma (Boleosoma) nigrum, a darter. The main differences between the subspecies lie in the extent of the squamation on the nape, cheek, and breast, one being typically naked or with few scales, and the other being well, i f not fully, scaled. Lagler and Bailey found that the.F generation of each subspecies, when reared and hatched under uniform environmental conditions in the laboratory, maintained the integrity of the parental phenotype. They concluded that these char- acters were genetically fixed. «.•••....•• After a vast and careful examination of some 9000 individuals, Bertin (1925) concluded that in populations of the stickleback,. Gasterosteus aculeatus, throughout Europe, most of the variable characters, especially the lateral body plates, are direct individual adaptations to environmental influences, especially temperature. He described four forms based primarily on body plate numbers. Heuts (1947a), however, showed that, at least in Belgian sticklebacks, 4 only two main modes of variability exist, each correlated with definite ecological characteristics, each with differences in breeding habits and distribution. He also showed (1947b) that certain physiological characters are correlated with morphological ones and he attempted to demonstrate that natural selection favors different complexes of genes in the different ecological niches in which the species occurs. He considers the two types of sticklebacks to be valid genetic races. Robins and Miller (1957) have called attention to the fact that because variation is so marked and haphazard in the sculpins of the genus Cottus, interpretation of the species limits within the group is frequently difficult. They consider Cottus to be one of the=most perplexing groups of North American freshwater fishes. In considering a character such as prickles on the body, they state that so much variation occurs between populations that its geo- graphic consistency in one or two forms is difficult to interpret. They also consider that a character such as "armature, like body prickles, may greatly assist in the identification of a species at any one locality, but is often unreliable for relating or distinguishing allopatric forms." MORPHOLOGICAL ANALYSIS OF PRICKLY SCULPIN POPULATIONS Materials and Methods Museum Collections. - Specimens examined include those in the following institutional collections (abbreviations listed are used in the text)! AB Bureau of Commercial Fisheries, Biological Laboratory, Auke Bay, Alaska BC University of British Columbia, Vancouver, Canada CAS California Academy of Sciences, San Francisco NMC National Museum of Canada, Ottawa, Ontario OSC Oregon State University, Corvallis SNHM Natural History Museum, Stanford University, Palo Alto, California (SU) 5 UMMZ University of Michigan Museum of Zoology, Ann Arbor USNM United States National Museum, Washington, D, C, UW University of Washington, Seattle Additional non-catalogued British Columbia material used in this study was made available through the personal field collecting of the following individuals. Abbreviations used are those which preface field collection numbers s K- R. J, Krejsas Little Campbell R. , and miscellaneous others A- G. F. Hartman, D. W. Wilkie, and C. A. Gills South Alouette R. V- " " " " " 11 «•• " " " . Vedder R, SW- " " » R. J. Krejsa, " " " " s Sweltzer Cr, S- J. S, Nelson, and P. J. McCarts Peace R. drainage. Material Examined, - A total of 1604 specimens from 190 collections was utilized in this study. Data for 158 specimens, from 16 collections, was culled from the published accounts of Snyder (1905, 1908b, and 1913), and Rutter (1908). Counts and measurements for 63 specimens, from 6 collections, were taken by Dr. D. E. McAllister of the National Museum of Canada, The remaining 1383 specimens were examined by the author. A l i s t of collections B together with pertinent data, is presented in Appendix Table 1, Counts and Measurements. - Measurements were taken to within 0.1 mm using break-arm dividers with one flat point and one needle point, in accordance with the methods of Hubbs and Lagler (1958). Counts for median fins and pelvic fins follow the methods outlined by Robins and Miller (1957). Other counts were taken as follows. Pectoral Fin Rayss Counts from both the right and left side were summed by population and then divided by the total number of fins. Pectoral fin asymmetry in Cottus asper was investigated to ascertain the incidence of occurrence, the direction, and the amount of asymmetry in this species, and 6 to determine whether individual populations, population aggregates, or geo- graphical regions could be characterized by these criteria. Distribution of Prickles on the Bodyi The areal distribution of body prickles is defined as the area of skin surface covered, or formerly covered, by prickles. It was recorded as follows. A superficial median incision was made through the dorsum and venter of a single adult specimen of Cottus asper 80 mm in standard length (S.L.). A transverse incision was made ventrolater- al ly from the median supratemporal pore to a point medial and posterior to the pelvic fin base. The skin was then peeled off the left side of the body as far posterior as the hypural fold where i t was excised. It was then placed flat on a sheet of paper and a tracing made around the perimeter. The resulting blank outline tracing was then subdivided into 14 lettered sections based on morphological landmarks. The subdivided put- line of the original skin hereinafter referred to as the "skin map," was stenciled onto sample work sheets on which a l l counts and measurements were recorded (Fig. IB), To estimate the areal distribution of prickles, a specimen is examined under the binocular dissecting microscope and the perimeter of the prickly area on the left side is outlined by scratching the mucous layer with a sharp needle point. This "apparent" outline is copied onto the work sheet skin map,with colored ink. The mucus is then scraped from the entire left side with the aid of a syringe-needle jet of compressed air, exposing those partially-resorbed prickles which do not project through the mucous layer. As will be discussed later, some prickles in Cottus asper are completely resorbed during maturation. Therefore, further treatment is necessary to expose the extent of the prickling in its entirety, i.e., the "virtual" prickling. Vigorous rubbing or massaging of the skin with the air needle will expose the areas formerly covered by prickles. This technique pre- sumably causes the breakdown of the sub-epidermal chromatophores. The 7 A L A T E R A L A S P E C T • D O R S A L A S P E C T , L E F T SIDE Figure lo — "Skin Map" Used f o r Recording D i s t r i b u t i o n and Intensity of P r i c k l i n g i n Cottus aspero 8 melanin granules thus released become aggregated in the empty dermal prickle pockets and the areas of resorption become visible. Thus resurrected, the original or " v i r t u a l " areal distribution of prickling can be recorded-in ink of another color over the "apparent" prickling distribution already recorded on the skin map (Fig. 2). Since each of the 11 lettered sections of the skin map represents a fraction of the t o t a l body surface on the l e f t side of the animal, a polar planimeter was used to determine the total area occupied by each section. Sectional areas were then converted into a set of fractional constants, each : representing a percentage of the total area of the skin map. These constants were used in calculations of the absolute prickling in each section. For s t a t i s t i c a l treatment, the total areal distribution of prickles was tabulated as followss A copy of the skin map was permanently printed onto a piece of transparent plastic. On this overlay, each of the lettered sections was further partitioned into subdivisions of 5, 10, and 25%. The plastic overlay was matched with the completed skin map on the work sheet and the percentage area covered by prickles in each numbered section was visually estimated to within 5%. The estimates for each section were then transcribed and summed for a l l specimens in the sample and a mean percentage "apparent" and mean percentage " v i r t u a l " prickling was determined. Multiplication of each of these sectional mean percentages by the appropriate fractional con- stant (previously determined) resulted in a product which i s an estimate of the mean percentage absolute prickling ("virtual" and/or "apparent") for each section. The sum of a l l 14 sectional products i s an estimate of the mean per^ centage areal prickling ("virtual'' or "apparent") for the population sample. Subtraction of the population mean percentage "apparent" areal prickling from the population mean percentage "virtual" areal prickling results in a remainder which represents the population mean percentage absolute /respcption AREA OF RESORPTION VIRTUAL PRICKLING Figure 2 «—Areas of Prickling and Areas of Resorptions A. in Coastal Cottus asper; B, in Inland Cottus asper. 10 for any given sample. The mean percentage relative resorption in any given population sample is obtained by the formulas Mean % relative resorption = 100% - inn/Mean V"Apparent"\ \Mean % "Virtual" / An examination of the condition of the gonads in more than 1000 speci- mens indicated that Cottus asper usually becomes reproductively mature at a size of 60 mm S e l i o On this basis 9 treatment of a l l prickling data was divided into groups of specimens smaller than 9 or equal to and larger than, 60 mm. Prickling Intensity; The size of prickles in Cottus asper varies directly with the' size of the animal. However, the intensity, i . e , , the number per unit area, does not appear to be a direct function of size, It varies among individuals of a l l sizes and even on different sections of the same individual, Because of the impracticality of recording absolute prickle numbers for large numbers of specimens, a relative index of range 0 through 5 was established. In this index zero indicated no prickling, and five i n - dicated the most intense prickling. When a single prickle was found in an otherwise naked area, i t was given an arbitrary value of 0.5, For s t a t i s t i c a l purppses, a single isolate was considered to have an areal distribution which was proportional to the lettered body section of the skin map in which i t was found, i.e., 4% i n sections 'a' and 'b'; 10% in sections 'd', 'e 1, 'h 1, 'k', and 'n'; and 5.5% in sections '1' and 'm'. If more than 4 prickles were found in an isolated patch, an index value of "1" was assigned and the areal distribution was estimated to the nearest unit percentage. To determine the nature of differential mean intensity index values, sample counts of prickles were taken from sq, cm, sections of skin adjudged to have index values of one and five. In rounded numbers, an index value of 11 "1" is equivalent to 400 prickles per sq. cm,, whereas a value of "5" i s equivalent to an estimated 2500 prickles per sq. cm. Mean values were inter- polated from these counts. Lateral Lines Previous workers (Robins and Miller, 1957) have con- sidered the lateral line as incomplete i f the number of pored-scales did not reach the hypural fold. The number of completely pored late r a l line scales varies with size and among populations. Close examination of sculpins re- veals that although a specimen may have an "incomplete" late r a l line ( i . e . , pored-scale count), unenclosed neuromast hillocks are present and readily v i s i b l e . Thus the most reliable measure of serial repetition along the later a l l i n e , for a l l size categories other than young-of-the-year, is a count of the complete pored-scales plus the pa r t i a l l y , or completely un- enclosed neuromast hillocks which appear sequentially along the trunk and out onto the caudal f i n . In practice, this count was made only to the hypural fold and the mean number of pored-scales plus hillocks was considered to represent a "complete" l a t e r a l line count. "Complete" counts were made; for less than one half of the specimens examined in this study and, in most small specimens, the count was l i s t e d only as "incomplete." The number of collections i s few in which a l l size ranges from juvenile to adult are re>- presented. The number in which "complete" lateral line counts have been taken i s even fewer. Critique of the Methods of Determining Prickling Distribution and Intensity. - The method of determining distribution proved unreliable on extremely old and/or badly preserved specimens. In some the skin was so desiccated that massage was impossible without damage to the specimen. In others, a l l the pigment had been bleached after years of preservation and no amount of massaging would make the empty prickle pockets appear. In such specimens only the "apparent" prickling distribution was determined and recorded. Estimates of prickling distribution on badly preserved or 12 desiccated specimens were not used in compiling population means. Tiny prickles in peripheral areas of immature specimens, and partially resorbed prickles on heavily pigmented mature specimens were difficult to see, even with the aid of the microscope. Therefore the estimates of "apparent" prickling both in mature and immature specimens are less than the "virtual" prickling in immature specimens. The accuracy of the visual estimate in peripheral areas was checked against alizarin-stained specimens, and was found to be within -10% on specimens > 60t>,mm, and within ±5% in specimens < 60 mm. While 60 mm appears to be a valid estimate of the average size at which most Cottus asper attain reproductive maturity, not. a l l spawn at that size. Some populations, e.g., from Queen Charlotte Is., are known to have females which spawn at a size of only 42 mm_ S.L., while in other populations, a few females do not spawn until a size of 70 mm is attained. The discrepancy in Table I between the mean percentage "virtual" prickling and the slightly lower (from 2-7%) "apparent" prickling in fishes < 60 mm is attributed to.; this cause. Another discrepancy in the data reported in Table I is that mean dif- ferences of from 4 to 10% occur between the "virtual" prickling percentages in specimens < 60 mm, and those >60 mm from the same population. These differences are attributed to the inability to "resurrect" a l l the prickle pockets completely. Statistical Methods. - Most of the raw data has been converted to means or percentages. Obvious differences were not tested. When a comparison of means was thought necessary, the two most divergent values were examined fir s t . If no significant differences were revealed, the less divergent o means were not tested. The single-classification test, or the Student t-test, was used at the 5% critical level to determine acceptance or rejec- tion of an hypothesis of equal means. Graphical presentation of certain 13 meristic characters is according to the format of Hubbs and Hubbs (1953), Synonymy It has been almost 130 years since Cottus asper was first described by Sir John Richardson, Early revisionary work by Girard in 1851 and 1852 was incomplete because of lack of specimens. Recent regional works (Robins and Miller, 1957; McAllister, 1957; McAllister and Lindsey, 1959; and Bond, 1963) have included the species as part of a geographical area or river drainage system, but there has never been a comprehensive treatment of the species throughout its entire range. In the period 1836-1936, 38 reports of Cottus asper were cited in the literature. In the same period, 32 additional cita-? tions occurred which were either misidentifications or synonyms properly- referable to Cottus asper. In none of the systematic treatments published since the original description in 1836 has there been a synonymy containing more than 7 citations. McAllister (1957) listed 15 citations in his un- > published M.A, thesis. The present synonymy, consisting of 70 citations published during the period 1836 to 19536, is thought to be complete. Since 1936, the species has been cited in numerous fishery journals and publications. Therefore, only the major systematic, or otherwise noteworthy, citations occurring in the past 29 years have been recorded. Cottus asper Richardson, 1836 Cottus asper. - Richardson, 1836s295, pi, 95, Fig. 1 (original description and figure; Columbia R,; collected by Dr, Gairdner (probably near Fort Vancouver, Washington Territory)), Storer, 18468.260, and 1846bs8 (northwestern coast of N. America). Girard, 1850s409, and 1851asl89 (discusses propriety of present nomenclature). Eigenmann, 1895&118 (abundant in Fraser system from tidewater to 1900 feet; Mission, Sicamous, Kamloops, and Griffin L., British Columbia; and Umatilla, Oregon). Seale, 1896s854 (Lake Washington), Gilbert and 14 Evermann, 1895:201 (description; comparison to Sacramento R. form; Walla Walla R. at Wallula, and Lake Washington, Washington). Gilbert, 1896:418 (description; stream entering Departure Bay, Vancouver Island). Jordan and Evermann, 1896:439 (synonymy; streams of the Cascade Range, from Vancouver Island to Oregon). Gilbert, 1898:1 (Columbia R.; notes absence in Klamath R.). Jordan and Evermann, 1898:1944 (description; synonymy; Walla Walla; Departure Bay; about Port Townsend; streams of the Cascade Range, from Vancouver Island to Oregon). Evermann and Meek, 1898:83 (Lake Washington). Meek, 1899:231 (Lake Southerland, Olympic Pen., Wash- ington). Snyder, 1905:337 (description; affinities; habitat pre- ference; San Franciscito, Madera, San Antonio, Guadalupe, Coyote, and Alameda Creeks, a l l flowing into San Francisco Bay). Evermann and Goldsborough, 1907a:306 (characters; prickling descriptions; Deep Bay, Naha R., and Steelhead Cr,, Loring, Alaska; Hunter Bay, Yes Bay, and McDonald L., Alaska). Evermann and Goldsborough, 1907b:110 (Fraser R, at Mission, Shuswap, L. at Sicamous, Thompson R. at Kamloops, and Griffin : L . R u t t e r , 1908:145 (Cottopsis parvus first placed in synonymy with asper; synonymy, in part, except Uranidea semiscaber (sic) centropleura Eigenmann and Eigeniiann; summary of 9 localities in Sacramento R, system). Snyder, 1908a:269 (Russian R., California)* Snyder, 1908b:184 (characters; prickling description; summary of Al localities: from Lake Washington, Columbia and Sacramento R., and river basins in between). Nichols, 1909:172 (head of Chilkoot L., Alaska), Evermann and Latimer, 1910:138 (4 localities in Marin Co., and 2 localities in San Francisco Bay, California; 12 localities in Olympic Pen., Washington). Snyder, 1913:72 (characters; Pajaro R., California). Snyder, 1916:381 (Papermill and Walker Creeks, California), Kermode, 1917:20 15 (Hanceville, B r i t i s h Columbia (Chilcotin R« drainage)),, Jordan, 1919;249 (designates Cottus asper Richardson, as the orthotype of Cottopsis Girard), Bean and Weed, 1920g76 (mouth of Fraser R,). Hubbs, 1921s7 (re-identification of San Luis Cr,9 California, speci= mens misidentified by Jordan as C, gulosus (1895sl41); range ex= tension to Ventura R,, California),, Crawford, 1927sl77 (streams flowing into Puget Sound). Schultz, 1929s48 (listing only), Schultz, 1930;14 (most streams and lakes of western Washington), Jordan, Evermann, and Clark, 1930g383 (synonymy; streams of the Cas- cade Range, southeastern Alaska to Oregon; south to Sacramento R,), Kermode, 1931.19 (Cowichan L,„ Vancouver Island), Evermann and Clark, 1931s56 (summary of 32 recorded localities in California), Schultz and DeLacys 1936as 128 (synonymy; coastal streams from Alaska to Ventura Co,, California; freshwater and brackish water; review of most records from Puget Sound to Oregon, and addition of 26 more localities), Schultz and DeLacy8 1936bs213 (additional synonymy; 3 new localities), Schultz, 1936sl79 (keys to species of Cottus; coastal streams from Alaska to Ventura Co 0 8 California; freshwater and brackish), Dymond_ 1936.71 (description; throughout souths western British Columbia8 including southern Vancouver Island; 16 localities listed), Sumner, 1942'gl=>25 (common in tidewater areas along the Oregon coast), Hubbs and Wallis s 1948s141 (identification of "Cottus sp." recorded by D i l l , 1946s54), Bailey and Dimick_ 1949sl4 (comparison with Cottus hubbsi), Shapovalov and D i l l , 1950s387 (listing only), Wilimovsky, 1954s285 (southeast Alaska to California). Robeckg et al, 1954sB=65 (Columbia R,8 above Trinidad, Washington; cited as "prickly sculpins Cottus sp,", these may include Cottus rhotheus in part,) Lindsey, 1956s777 (Pacific Slope of No America from Alaska to California; British Columbia mainland from 16 Columbia, Fraser, and Skeena systems, Stikine R, headwaters, and Peace R. (Summit L., Heart' L,, Angusmac Cr,, and McLeod L.)), Robins and Miller, 1957:229 (Cottopsis parvus again removed from synonymy of C. gulosus ), Lindsey, 1957:657 (British Columbia: Columbia R.; Fraser R.; Skeena R,; Coast drainages south of Skeena; Nass R.; Stikine R.; Peace R,). Wilimovsky, 1958:62 (key to species in Alaska; southeast Alaska to California). Shapovalov, D i l l , and Cordone, 1959 ( l i s t i n g only), Carl, Clemens, and Lindsey, 1959:158 (description; Pacific drainages from Chilkoot L,, Alaska to Ventura R, , California, In British Columbia: lakes and rivers of the Columbia, Fraser, Dean, Skeena, Nass, and Stikine systems; coastal rivers of the mainland and Vancouver Island, and Queen Charlotte Islands; headwaters of Peace R, system from Summit L, to McLeod L,). McAllister and Lindsey, 1959:70 (description; synonymy; intra- specific variation; l o c a l i t i e s as in Carl, Clemens, and Lindsey,- 1959), McAllister, 1960:42 (collection in salt water, Pt, Atkinson, British Columbia). Bond, 1961:36 (key to species in Oregon; prickling variation; Columbia R, drainage). Bond, 1963:79 (synonymy l i f e history observations; oxygen, temperature, and salin i t y toler- ance of adults; f i s h associates; habitat preference; l i s t s 35 new collection l o c a l i t i e s in Coastal Oregon, 30 l o c a l i t i e s from Columbia R. drainage in Oregon, and also some lakes in the southwest corner of Rainer National Park, Washington), Bailey and Bond, 1963:19 (recognition of several species groups within Cottus in western N. America; characters and l i s t of species in the "asper species group"). Centridermichthys asper. - Richardson, 1844:76 (River Oregon = Columbia R.), Gunther, 1860:170 (description; synonymy; fresh waters of the Oregon and Washington Territories). Lord, 1866a:130 [ l i f e history . 17 spawning behavior, in part; Puget Sound; "streams flowing through the Sumass and Chilukweyuk prairies" (Sumas R0 and Chilliwack R, (?)), British Columbia; in part, a l l streams east and west of the Cascades], Cottopsis asper. - Girard, 1851bs303 (introduction of Cottopsis gen, nov.; synonymy; limited to River Oregon (= Columbia R,)). Girard, 1851csl85 (not seen). Girard, 1852s61 (definition of Cottopsis gen, nov., based on Richardson's description of Cottus asper; synonymy; Columbia R.). Girard, 1859s51 (description; synonymy; based on 8 specimens from Astoria and Fort Dalles, Oregon, and Fort Steilacoom, Puget Sound, Washington Territory). Suckley (in Cooper and Suckley, 1859) 1959:351 (description; synonymy; small fresh water streams emptying into Puget Sound; Ft. Steilacoom; and Columbia R, 200 miles above mouth), Suckley, 1860.351 (description; synonymy; small fresh- water streams emptying into Puget Sound; Ft, Steilacoom; and Columbia R, 200 miles above mouth), Jordan and Jouy, 1882s5 (Puget Sound; Columbia R,; Mare Island and Sacramento R,, California), Uranidea aspera. = Jordan and Gilbert, 1883s694 (description; synonymy; streams west of the Sierra Nevada and Cascade Mountains), Jordan,- 1885sll0 (sub-genus Cottopsis and a l i s t of species therein), Trachidermus richardsoni, - He eke 1,, 1840 ;162 (synonymy; Columbia R.), (not Cottus richardsoni of Agassiz B 1850). Cottopsis parvus. ••<• Girard, 1856bsl44 (original description; Presidio (on San Francisco Bay), California). Girard, 1857sll (description; Presidio, and Monterey, California), Girard, 1859s54 (description; synonymy; Monterey, Presidio, Fort Reading, and Petaluma, California), Cooper, 1868s492.(listing only), Jordan1877§5 (as the young of Cottopsis asper). 18 Centridermichthys parvus. - Gunther, 1860:170 (descriptionj synonymy; fresh waters of California). Lord, 1866b:352 ( l i s t i n g ; "frequenting the same l o c a l i t i e s as..." Centridermichthys asper3. Uranidea aspera var. parvus. - Jordan and Gilbert, 1883:694 (Sacramento R. forms), Cottus gulosus parvus. - Jordan and Evermann, 1898:1945, and Jordan, Evermann, and Clark, 1930:383.(Cottopsis parvus: Monterey, Presidio, Fort Reading, and Petaluma, California). Uranidea gulosa, in part. - Jordan and Gilbert, 1883 :695 (misidentifications: a l l specimens from "Vancouver's Island" and probably those from "about Port Townsend," cf. Jordan and Evermann, 1898:1944), Kermode, 1909:87 ( l i s t i n g only; probable misidentification: Shawnigan L., Vancouver Island). Cottus gulosus, in part. - Jordan, 1895:141 (misidentification: San Luis Cr., near Avila, California, cf. Hubbs, 1921:7). Jordan and Evermann,- 1898:1945 (misidentification: a l l specimens from San Franciscito Cr., Santa Clara Co., California)." Jordan, Evermann, and Clark"*. 1930':383 (probable misidentifications: specimens from Loring and Boca de Quadra, Alaska). Evermann and Clark, 1931:57 (misidentifications: Presidio, Monterey, Fort Reading, Petaluma, and San Luis Cr,, California), Evermann and Clark, 1931:12, 13 (misidentifications: Monterey, Presidio, Fort Reading, and Petaluma), Bean and Weed, 1920:76 (questionable identification: 4 specimens from Victoria, Vancouver Island, British,Columbia). Wilimovsky, 1954:285 (doubts validity of southeast Alaska record), Centridermichthys gulosus. - Lord, 1866b:352 ( l i s t i n g ; "frequenting the same l o c a l i t i e s as..." Centridermichthys asper). Cottus sp. - D i l l , 1946:54 (San Joaquin R, near Friant, California: identi- fication as asper by Hubbs and Wallis, 1948:141). 19 Nomenclatural History The specific name asper i s currently well founded in the genus Cottus t to which i t was originally designated by Richardson in 1836, But, as shown in the preceding synonymy, the binomen was extremely unstable for the f i r s t 100 years after.its introduction,, After an i n i t i a l period of uncertainty regarding i t s a f f i n i t y to marine or to freshwater Cottoids, three main no- menclatural d i f f i c u l t i e s are encountered? the often-repeated misidentification as Cottus gulosus (Girard)j the failure to recognize that Cottopsis parvus a n d Cottus asper are conspecific; and the failure to recognize the specific relationship of asper to other species in the genus Cottus, The almost immediate placement of asper into Trachidermus by Heckel (1840), and then into Centridermichthys by Richardson (1844), reflects the early opinion that asper was more closely a l l i e d to the marine Cottoids, Girard (1851b; 1852) recognized i t s a f f i n i t i e s with the freshwater genus Cottus, but distinguished i t from that genus by erecting the genus Cottopsis, based on the presence of palatine teeth and the "skin beset with prickles, Instead.of being smooth and scaieless 0" Lacking any specimens, Girard de- fined Cottopsis on the basis of Richardson's original description but, on p 0 63, he misquoted Richardson especially in regard to the absence of scales,, Girard's lack of specimens proved unfortunate since soon thereafter (1856b) he named and described Cottopsis" parvus e from the Presidio (in San Francisco), California, comparing i t not w i t h C asper, but with Cottopsis^ gulosus Girard, also newly described (1856a)B from the San Joaquin R,, California, In his later report (1859)9 Girard had 8 specimens of C,. asper in his pos- session, from the Columbia R, and Puget Sound, Obviously he again failed to . recognize the conspecificity of asper and parvus and he followed Richardson's original description rather than compare them with specimens of parvus, which he seems to have reserved for comparison with gulosus, Jordan (1877) referred^, in passing, to Cottopsis parvus as the young of 20 C. asper, Jordan and Jouy (1882) however, listed specimens of Cottopsis asper from Mare Island and Sacramento R,, California, and from Puget Sound and the Columbia R, Lass than a year later, Jordan and Gilbert (1883) placed asper into the genus Uranidea DeKay, subgenus Cottopsis, based on the presence of palatine teeth, and the g i l l membranes being broadly united to the isthmus. In the same report, they refer to the Sacramento R, form of •>, U, aspera as "var, parvus, smaller in size, paler in color and with the inter- orbital space concave, narrower than eye," Eigenmann (1895) used the valid name to describe specimens from the Fraser and Columbia R,, as also did Gilbert and Evermann (18.95), who sug- gested that the nominal species was separable "at least subspecifically from the Sacramento River form," Seale (1896), and Gilbert (1896), used the valid name for northern specimens. But obviously, Jordan (1895), and Jordan and Evermann (1896), s t i l l thought in terms of a distinct Californian^species (gulosus), and a distinct northern species (asper). Jordan misidentified a specimen of asper from San Luis Cr.?near Avila, California, as gulosus. Jordan and Evermann listed the range of the nominal species from Vancouver Island to Oregon, and of gulosus, from California Coast Range streams and inland in the San Joaquin R. Gilbert (1896) referred to Cottus asper of the Columbia and Cottus gulosus_ of the Sacramento as "two species so extremely similar that i t is difficult to distinguish them," Jordan and Evermann (1898) repeated the suggestion of Gilbert and Evermann that the nominal species is separable, at least subspecifically, from the Sacramento R, form "Cottus gulosus." That Jordan* and Evermann perceived neither the conspecific relationship °f parvus to asp_er9 nor the limits of the valid species Cottus gulosus, be- comes more fully evident on the next page (p, 1945) of the 1898 report. Their description of Cottus gulosus (Girard) is taken from misidentified specimens of C. asper collected in San Franciscito Cr,, Santa Clara Co., California. These were large specimens "3 to 7 inches in length" and, most significantly, the count for anal rays i s given as "A, 16 to 18." Both of these characters separate asper from gulosus. Furthermore, they include Cottopsis parvus Girard, from Monterey, the Presidio, Fort Reading, and Petaluma, California, in the synonymy of gulosus. Snyder (1905) collected and correctly identified Cottus asper from the same loc a l i t y , San Franciscito Cr. He was probably the only one of his time to understand and explain the true relationships of asper, parvus, and gulosus., On p. 337, he states? Recent authors have identified the common Sacramento form which represents the Cottus asper of the Columbia River with the Cottopsis gulosus of Girard. They have sometimes considered the Sacramento form as identical with C. asper and have placed the name gulosus in the synonymy of the latter. At- other times they have considered the species as a slightly differentiated form worthy of recognition in nomenclature, and have used the name gulosus to designate i t . The former view con- cerning the species i s probably correct. The association of the name gulosus with i t , however, i s without warrant. The latter belongs to a species easily distinguished from C. asper B differing notably in having a much shorter anal f i n . There are usually fewer dorsal spines and rays B a more limited distribution of prickles, and an almost uniform absence of palatime teeth. In C. asger the dorsal has 8 to 10 spines and 19 to 22 articulated rays, Fhe anal 16 to 18 rays, while in C, gulosus the dorsal has 7 to 9 spines, 17 to 18 rays, the anal 12 —to 14 rays, 1 Snyder then continues with a note on habitat preference: In i t s distribution C. asper appears to be largely confined to the lower courses of the streams", being especially abundant near tide water, while C. gulosus is found further up, where the water i s clear and the current rapid Rutter (1908) correctly placed Cottopsis parvus into the synonymy of Cottus asper, presumably recognizing that the two were conspecific. However, he incorrectly synonymized Uranidea semiscabra centropleura Eigenmann and ; Eigenmann, which i s properly referable to Cottus gulosus. Snyder was the f i r s t to consider a series of specimens of the nominal species throughout i t s entire range, as then known, and, in the same issue of the Bull. Bur. Fisheries in which Rutter had correctly synonymized parvus, he noted the extreme variation of prickling investment. While recognizing 22 the variation between streams, he also noted that the prickling variation i s common among individuals from the same stream. Although e x p l i c i t l y aware of Snyder's comments on asper and gulosus, Evermann and Goldsborough (1907&)identified 16 specimens of gulosus from, Loring and Boca de Quadra, Alaska, The r e l i a b i l i t y of these identifications is questionable. Kermode's l i s t i n g (1909) of Uranidea gulosa from Shawnigan L,, Vancouver Island, i s probably a misidentification of £, asper. Snyder (1913j 1916) again recorded the occurrence of C, asper and gulosus in differing habitats of the same stream, Hubbs (1921) recognized Jordan's earlier misidentification of gulosus from San Luis Cr,, California. He also commented on the var i a b i l i t y of prickling in £. asper from several streams. It would seem that with the accession of Snyder's insight into the problem, the valid name was destined for s t a b i l i t y . However, Jordan, Evermann, and Clark (1930) repeated the earlier error of Jordan and Evermann (1898) by including Cottopsis parvus as a synonym of Cottus gulosus. In a similar manner, they also incorporated the error of Evermann and Goldsborough (1907a):,, previously cited, by including the misidentif ied specimens of asper from Loring and Boca de Quadra, Alaska in the l i s t of records for gulosus,- In the same work, Jordan and Evermann extended the range of asper (cited in 1898 as "streams of the Cascade Range, from Vancouver Island to Oregon") by appending the phrase "south to Sacramento River," Evermann and Clark (1931) also retained Cottopsis parvus in the synonymy of C. gulosus and perpetuated Jordan's misidentification of the San Luis Cr. gulosus, which Hubbs had correctly re-identified as asper ten years before (1921), > Schultz and DeLacy's catalogue (1936) included a comprehensive l i s t i n g of Washington and Oregon l o c a l i t i e s for C, asper. However, some of the lo c a l i t i e s are doubtful since they frequently misidentified C, asper as gulosus and/or perplexus. They also incorrectly maintained the presence of 23 C_. gulosus in Alaska. Robins and Miller (1957) presumably overlooked the earlier citation of Rutter (1908) and removed Cottopsis parvus from the synonymy of gulosus, placing i t in synonymy with asper, supposedly for the f i r s t time. McAllister and Lindsey (1959) f i r s t suggested the probable existence of "coastal" and "non-coastal" populations of Cottus ajper on the basis of mor- phological and, perhaps, behavioral differences. Bond (1961) hinted at the possibility of polytypy in Cottus asper when he stated in his key that the body i s "well covered with prickles, especially in inland waters and in young individuals from coastal waters. Bond (1963) gives the most comprehensive treatment yet recorded for Cottus asper and 12 other species in the genus. His- study 8 however, is more concerned with < interspecific rather than intraspecific relationships within the genus, <.*•., Bailey and.Bond (1963) have indicated their concern for the supraspecifie relationships within the genus Cottus by their recognition of several species groups, one of which i s the "asper species group." Distribution Range. - Pacific Slope Drainage of North America? Coastal streams from Seward, Alaska, to Ventura R.. California; lakes and streams of the Queen ; Charlotte Islands and Vancouver Island; and a l l major Pacific drainages from the headwaters of the Stikine R. in British Columbia, to the Kern R._ San Joaquin R. drainage. Californian- The-following are exceptions? Fraser R. .in the area of the Rocky Mountain Trench, east of Prince George, British Columbia (area not yet collected); Kootenay Lake drainage of the Columbia R, in British Columbia; Upper Snake R. of the Columbia R, drainage in Washington and Oregon; Middle Fork of the Willamette R. in Oregon, above Oakridge; Klamath R, Basin in Oregon; and Sacramento R, drainages above Lake Shasta, California, Arctic Slope Drainage of North America? Headwaters of the Peace R. in British Columbia?' from Summit L. to McLeod L. 9 Crooked R. drainage; from Tacheeda L., 24 Parsnip R. drainage; from Tchentlo L.* Nation R, drainage. The present distributional range of Cottus asper i s illustrated in Fig. 3. Results; Analysis of Geographic Variation Prickling Distribution. - As previously defined (p. 7 ) » the "v i r t u a l " prickling represents the sum of two component percentages, i . e . , the "appa- rent" prickling and the resorbed prickling. Fig. 4 illustrates graphically that mature specimens from populations of C_, asper from coastal l o c a l i t i e s are considerably less prickled, both apparently and virtually, than inland l o c a l i t i e s . However, most inland populations from the Peace, Nass, Stikine, and parts of the Skeena R. drainage systems are coastal in their prickling a f f i n i t i e s . Hereafter, these populations w i l l be referred to as "coastal derivative." Likewise, certain populations from lakes on Vancouver Id, and the Queen Charlotte Islands (Q0C,I.) are also coastal in their prickling a f f i n i t i e s . Fig, 4 also indicates that the population from Carquinez Strait, at the mouth of the Sacramento R. in San Pable Bay, California, has a mean "virtual" prickling percentage-greater than that of any coastal popu- lation, and lower than a l l but one of the inland populations. For comparative purposes, the Carquinez Strait population w i l l hereafter be referred to as the "intergrade" population. Table I summarizes the mean percentage areal prickling and resorption for a l l populations grouped into aggregates according to l o c a l i t y and prickling a f f i n i t y . If the data for specimens under 60 mm S,L, are considered separately, there i s a significant difference in mean percentage prickling, both "apparent" and "v i r t u a l , " between the coastal and inland populations. Ex- pressed in terms of a ratio between coastal-inland populations, a ratio of "apparent" prickling 1 s 1.8, and "vi r t u a l " prickling 1 ? 1,5, is obtained. The aggregate means for the "coastal derivative" populations are essentially Figure 3.—Distributional Range of Cottus asper. 26 Figure 4„—Virtual, Resorbed, and Apparent Prickling in Cottus asper. TABLE I. Mean Percentage Prickling and Resorption Among Aggregate Populations of Cottus asper. Aggregate Population Specimens <60 mm S.L. Specimens ^60 mm S,L, % Resorption Designation Virtual Apparent Virtual Apparent Relative Absolute 1 Coastal it 43.5 (9) 36,0 (12) 35.0 (16) 17.8 (19) 48.9 17.2 2 Coastal Derivative 44.8 (6) 41,8 ( 6) 34,7 ( 6) 24.2 ( 6) 32.0 10,5 3 Intergrade 60.1 (1) 57,1 ( 1) 52,9 ( 1) 35,6 ( 1) 32.8 17.3 Inland1* 67,2 (16) 65.3 (18) 63.2 (18) 53.2 (19) 16.1 10.0 Brackets enclose numbers of population means in determination of aggregate mean percentages. 1 Populations 1-31, listed in Appendix Table 1 2 Populations 32-43, listed in Appendix Table 1 3 Population 73, listed in Appendix Table 1 4 Populations 44-72, listed in Appendix Table 1 28 the same as those for the coastal, and they d i f f e r significantly from the aggregate means for the inland populations. The mean for' the intergrade population l i e s closer to the inland aggregate. When specimens larger than 60 mm S.L. are considered, the same magnitude of difference i s noted between the coastal and inland "virtual" percentages, i.e. , a ratio of 1 s 1.8, but the "apparent" prickling percentages are even more divergent, having a ratio of 1 s 3, Since the "vi r t u a l " ratio remains constant, the divergent "apparent" ratios indicate a differential percentage resorption between mature specimens from coastal and inland populations. This differential i s further suggested by the inverse relationship which exists between the mean percentage "apparent" prickling and the mean per- centage relative resorption. It indicates that in coastal populations a greater percentage of the " v i r t u a l " prickling area i s being resorbed than in inland populations. The r e a l i t y of this differential resorption i s shown in the figures for mean percentage absolute resorption which indicate that, in coastal forms, resorption occurs over almost twice as much area as in inland forms. The differential can be expressed in terms of a coastal-inland r a t i o , i«e©9 Xe 7 t Xe Prickling and resorption percentages on which Table I is based are given for a l l populations in Appendix Table 2. Population analysis of the non-weighted mean percentage "v i r t u a l " prickling in each of the 14 body sections, of specimens ^ 60 mm S.L., i n - dicates that in coastal and "coastal derivative"populations, body sections 'a', 'k', '1', and 'n' are completely naked except for the rare occurrence of an isolated prickle. Similarly, in coastal and "coastal derivative" popu- lations, only small portions of sections 'b', 'd', 'e ?, 'h', and 'm' are prickled. When compared to the mean percentage "v i r t u a l " prickling of inland populations, a l l nine of these sections are significantly different. Further- more, in the remaining five sections Cc', ' f ' , 'g», ' i ' , and ' j ' ) , the mean 29 percentage "virtual" prickling i s always higher in the inland populations. These results corroborate the suggestion, made for the aggregate population means, that the areal distribution of prickling in coastal and inland popu- lations of C. asper i s different. Data for this analysis are presented in Appendix Table 3. Prickling Intensity. - In general, the intensity of prickling, i . e . , the number of prickles per unit area, i s greater in inland f i s h than in coastal f i s h . In coastal populations, where prickles are almost never pre- sent in sections 'a', 'k', '1', and 'n', no comparison of intensity can be made. Rarely, however, when an isolated prickle or group of prickles occurs in one of these sections, the intensity is always less than that of the corresponding section in an inland.population. The same holds true for sections 'b', 'd*, 'e', 'h', and.''m'.f The comparison of intensity must therefore be made in one of the sections in which prickles are always present on both inland and coastal specimens,- Section ' j ' was chosen as representa- tive. Table II shows that the mean relative intensity is greater in the inland populations than in coastal or "coastal derivatives," Interpolation of mean intensity values given in Table II into approximate number of prickles per sq, cm. (cf. Methods) yields counts of ca. 1325 prickles per sq. cm. in coastal forms (2.94), and ca. 1820 per sq, cm, in inland forms/fo.se). Expressed in terms of differences between coastal and inland intensities, a . ratio of 1 ; 1.4 is obtained. Comparison to the ratio obtained for area: prickling resorption, I.e.", 1,7 ; 1^ indicates that even though inland f i s h have more prickles per unit area, more resorption (on a prickle for prickle basis) occurs in coastal f i s h . Mean relative intensity values for individual populations of Cottus asper are given in Appendix Table 4, Table II also shows that in mean intensity, of prickling, the "coastal derivative" populations aggregate is near the coastal mean whereas the "intergrade" population i s closest to the inland mean. If the mean intensity TABLE II. Mean Relative Prickling Intensity for Populations of Cottus asper Aggregated by,Locality and Prickling Affinity. Skin Section ' j * . Aggregate Specimens <60 mm S.L, Specimens ̂ 60 mm S.L. Population Sample Intensity Sample Intensity Designation Size Mean (Range) Size Mean (Range) 1 ft ft Coastal 136 (13) 2.94 (1-5) 235 (18) 2,64 (1-5) 2 Coastal Derivative 103 (11) 2.74 (1-4) 43 ( 7) 2.95 (1-5) 3 Intergrade 8 ( 1 ) 3,75 (3-5) 20 ( 1) 3.45 (2-4) Inland **. 238 (21) 3.86 (2-5) 265 (19) 4.20 (3-5) * Brackets enclose numbers of populations included in aggregate sample. 1 Populations 1-31, listed in Appendix Table 1 2 Populations 32-43, listed in Appendix Table 1 3 Population 73 only, listed in Appendix Table 1 4 Populations 44-72, listed in Appendix Table 1 31 values of certain coastal lake forms, i.e., Home L., Buttle L., Skidegate L., and Chilkoot L., are considered (Appendix Table 4), they are found to be slightly lower than the coastal means. These data corroborate the differ- ences already shown to exist in mean areal distribution of prickling between coastal and inland populations. • Pectoral Fin Asymmetry, - Of 59 populations investigated for pectoral fin asymmetry, 56 had some specimens with asymmetrical pectoral fin element counts. Of 1163 paired counts, from 59 populations, 23.6% (275 pairs) were asymmetrical (Table III). This percentage incidence of occurrence is similar to the percentages reported for other cottids by Hubbs and Hubbs (1945)., Of the 56 populations of Cottus asper exhibiting asymmetry, 39 (69.6%) were sinistral, 11 (19.6%) were dextral, and 6 (10,7%) had equal numbers of right and left asymmetrical fins.- This apparent direction of asymmetry within populations is not statistically significant since the sample sizes of asymmetrical fins within each population were very small. However, if-, the asymmetrical fins are grouped according to locality and supposed de- r rivation (based on prickling), or pooled for a l l populations (Table III, columns 5 6 6), the overall direction of asymmetry in Cottus asper is signi- 2 ficantly sinistral ( X = 21.56, cC--= .05, 1 d.f., for the pooled sample). While Cottus asper is significantly sinistral in the overall direction of its asymmetry, the amount of sinistrality within populations varies from 0 to 100%. When the populations are grouped according to locality and supposed derivation (Table 111^ col. 5), the amount of sinistrality for 24 coastal populations is 66.7%; for 12 coastal derivative populations, 69.2%; and for 23 inland populations, 60.8%. These differences among aggregate populations are not significant. Additional analysis of data on a north- south basis revealed no significant differences in direction or amount of asymmetry. Total pectoral fin counts on which Table III is based are given for a l l TABLE III. Incidence of Occurrence, Direction, and Amount of Pectoral Fin Asymmetry Within and Among Aggregated Populations of Cottus asper. Locality No. Pairs Symmetrical Asymmetrical Sinistral Dextral ft Coastal 460 ftft 367 (79,8) 93 (20.2) 62 (66.7) 31 (33.3) Coastal Derivative 152 - 113 (74.4) 39 (25.6) 27 (69.2) 12 (30.8) Inland 551 408 (74.0) 143 (26.0) 87 (60.8) 56 (39.2) Pooled Total 1163 888 (76.4) 275 (23.6) 176 (64,0) 99 (36.0) Includes three coastal island lake populations aft Bra eke t s en close aggregat e mean percent age s 33 populations in Appendix Table 5. It has been shown for several species of fish that the lowermost pectoral ray is shorter on the side which has the higher count (Hubbs and Hubbs, 1945} Hubbs, 1963). Cottus asper appears to follow this rule. Of 69 specimens, from 17 populations having asymmetrical pectoral fin counts, 66 had a shorter lowermost ray on the side with the higher count, and only 3 had the lowermost rays equal on each side. While Cottus. asper demonstrates a significant sinistrality in the .j . direction of its pectoral fin asymmetry, the amount of this sinistrality within and among populations cannot be used to separate individual popu- lations or geographically aggregated groups of populations. Pectoral Fin Ray Count. - The mean number of pectoral fin rays in- creases progressively from south to north in the inland populations (Figs, 5 & 11). Coastal populations show a similar increase northward only to about 42-44° N. Latitude where the mean pectoral fin ray counts diverge from the inland counts and decrease toward-the north. Mean counts for the "coastal derivative" population are found to be equal to or less than inland popu- lation means, but always higher than those of the coastal means, at the same latitudes. The means of certain coastal island lake and coastal main- land lake populations (Buttle, Home, Skidegate.and Owikeno L., cf. Appendix Table 1) are almost a f u l l count higher than the means of nearby stream populations from which they probably were derived. Data from Carquinez Strait "intergrade" population and two additional populations from the San Francisco-San Pablo Bay area, thought to be intergrades, (cf, Appendix Table 1, population numbers 74 6 76) were compared. At the low latitudes in which they occur, their mean pectoral ray counts were indis- tinguishable from either coastal-or inland means. Spiny Dorsal Fin. - The average spine number is clustered about nine for a l l coastal populations except:the southernmost California populations 34 60 - O 58 56 "o 54 52 - 50 I _ l_48 cr Q46 z o 44 42 40 38 36 34 I I I I I I I I I I I I o o o o ® 4 J , ^ A ® A o o e o o iA A A AAA e O AA K A A G O O 0G O o A A A O A 4 o o o I I I J L. _L O COASTAL ® COASTAL DERIVATIVE @ COASTAL LAKE A INLAND A INTERGRADE 16 17 P E C T O R A L RAYS Figure 5„—Mean Pectoral Fin Ray Count by Latitude, 35 which are slightly lower (Fig, 5) 0 The San Joaquin-Sacramento. R« inland populations have a mean count of eight or slightly higher.. This wide diver- gence tends to accentuate a slight, but discernible, geographic cline for the inland populations as a whole. The mean counts are slig h t l y higher than nine in the high latitudes, slightly lower in the mid-latitudes, and much lower in the low latitudes (cf. Fig. 11), A comparison of the ranges (Appendix Table 6) indicates high counts of nine to eleven in the north, and low counts of seven to nine in the south of the distributional range of the species. : The intermediate positions of the three "intergrade" populations within the San Francisco-San Pablo Bay area should be noted. The means of the "coastal de- rivative 1' populations tend to be closer to those of the inland populations. Softs Dorsal Fin. - The mean soft-ray number varies considerably with latitude. Counts are slig h t l y higher than 20 in the high latitudes, closer to 21 in the mid-latitudes, and between 19 and 20 in the low latitudes. As can be seen in Fig. 7, the inland Sacramento and, especially, the San Joaquin R. populations have considerably lower mean counts than their coastal counter- parts at the same latitudes. If the soft-ray counts for a l l inland popu- lations are summed into aggregates of 2-degrees latitude and plotted according to the Hubbs-Hubbs format (Fig. 8), the V-shaped geographic cline within the means and the ranges becomes immediately apparent (cf. Fig. 11), Combined Dorsal Fin Elements, - Combining the spiny and soft dorsal elements tends to minimize the variation in means at the middle of the range, while accentuating the divergence between coastal and inland populations in the highland, especially, the low latitudes (Fig. 9), The divergence between the San Joaquin-Sacramento populations and their coastal counterparts at the same latitude i s especially enhanced (cf. Fig. 11), The intermediate posi- tion of the intergrade population means and the position of the "coastal de- rivative" means w i l l be discussed later. Anal Fin. - The mean number of anal rays varies around 17 for a l l 36 60 58 - 56 54 - 52 - 50 - H I 4 8 cr O 46 z o 4 4 4 2 4 0 3 8 36 3 4 0 O O O O ^ A O A A M (X) o o o o O o A O o o O o O COASTAL ® COASTAL DERIVATIVE O COASTAL LAKE A INLAND A INTERGRADE 8 10 DORSAL SPINES Figure 6,—Mean Spiny Dorsal Fin Ray Count by Latitude, 60 - —i i i 1 O 1 0 58 • O 56 O A 5 4 #>A A - 5 2 • (8) G e A A - 50 x A . ^ e J Z 4 8 cr Q 4 6 z o 4 4 - <* A 4) & A A 0 0 42 A o - 4 0 - A - A 03 i 38 - A A 4 0 ° - G C O A S T A L A A A A ,o ® C O A S T A L DERIVATIVE 36 A £ C O A S T A L L A K E A INLAND 34 1 I ^ ~ - A INTERGRADE 1 1 1 I -1 19 20 21 2 2 S O F T D O R S A L R A Y S Figure 7.—Mean Soft Dorsal Fin Ray Count by Latitude. 38 Standarcy—^ fMean A Deviation^ .• 57h 5 5 53f 51 49 X cr O 45h 43 41 39 37 35 - 8 ,-2 Standard Errors of the Mean h - - 7 / Range- ' ' ' 19 2 0 21 S O F T D O R S A L R A Y S 22 Figure 8,—Hubbs-Hubbs Plot of Soft Dorsal Fin Ray Counts for Inland Population; Samples Pooled into Aggregates of.Two Degrees Latitude, 60 58 56 54 52 50 H 4 8 o r 0 4 6 Z o 44 42 40 38 36 34 T r 1 r o o o o e <8> e o A O O • O O o O G ? A A A O O o o A O O O _i L J L O COASTAL ® COASTAL DERIVATIVE © COASTAL LAKE A INLAND A INTERGRADE 27 28 29 30 C O M B I N E D D O R S A L E L E M E N T S Figure 9,—Mean Combined Dorsal Fin Ray Count by Latitude, 40 populations (Fig. 10). There is a tendency for the mean to be slightly lower in the northern and southern ends of the distributional range, whereas in the mid-latitudes the means are clustered about 17. This pattern tends loosely to follow the V-shaped curve observed in the number of soft dorsal rays (cf. Fig. 11). Pelvic Fin. - The pelvic fin almost invariable consists of 1 spiny plus 4 segmented rays. Among more than 1000 specimens examined for this charac- ter, there were only 20 deviants from-the usual left-right count of 4-4, , In one specimen both fins, and another specimen, the left fin was missing. In neither specimen was there any apparent injury. Of the remaining 18, 7 deviants (4 damaged) were on the left side, and 11 (4 damaged) on the right. Thus even in the most stable meristic character, rare deviations occur, A summary of meristic counts for a l l populations is given in Appendix Table 6. , s Latitudinal Variation in Ranges of Mean Fin Ray Counts. - Figure 11 is derived from each of the Figures: -5, 6, 7, 9, and 10. When the means of the inland and the coastal populations are enclosed by sets of lines approxi- mating the range of means, a family of curves representing a l l the meristic data for a l l latitudes is generated. Several trends are apparent in Figure lis 1) The curves for inland and coastal population means, though over- lapping in some areas, are widely divergent in others; 2) All curves, except for the spiny dorsal and the inland pectoral means, demonstrate a characteristic V-shaped pattern with an inflection occurring at about 42-44° N. Latitude; 3) For pectoral fins, the greatest amount of divergence between coastal and inland means exists in the high latitudes. The opposite is true for the dorsal fin elements; and 4) In areas where coastal and inland curves are divergent, the means 41 60 58- 56- 54- 52 50 48 h- * 4 6 O o 4 4 | 42 40 38 36 34 -i 1 r -i 1 1 r _J , r T 1 1 r o o o <2> o 8 o o 8 e 0 ^ ° A A O G O O A 6 o cP A o o <8> o A _ l . I J _ O COASTAL ® COASTAL DERIVATIVE 0 COASTAL LAKE A INLAND A INTERGRADE 16 17 A N A L F I N R A Y S 18 Figure 10,—-Mean Anal Fin Ray Count by Latitude, 15 16 17 Pectoral 8 10 Spiny Dorsal 18 20 22 Soft Dorsal 26 28 30 Combined Dorsal 16 17 Anal 18 Figure 11.—Approximate Ranges of Mean Fin Ray Counts Plotted Against Latitude for Inland ( Coastal ( ) B and Coastal Lake and Coastal Derivative CciiiiBiiiiii) Populations. Curves fitted by eye from Figures 5, 6g 7, 9, and 10. 43 for coastal lake and "coastal derivative" populations always approximate the inland rather than the coastal curve. The interpretation of these curves and the similarity of their charac- teristic V-shape to curves generated from laboratory rearing experiments under controlled temperatures will be considered in the discussion on meristic variation within Cottus asper (below). Lateral Line. - The number of lateral line pored-scales varies directly with size up to about 60 mm S.L., the average size at which most Cottus asper first become reproductively mature (Table IV), In most specimens larger than 60 mm, vi the pored-scale count approaches, or equals, the "com- plete" lateral line count. One obvious exception is in a spawning popu- lation collected from the estuary of a small stream in Juskatla Inlet, Queen Charlotte Islands, in which many of the small specimens 40-49 mnu S.L. were in spawning condition. Their pored-scale counts approached orr, equalled the mean "complete" lateral line count (Table IV). Another ex- ception to the generalization is that afforded by specimens from the coastal lake populations found on the British Columbia or Alaska mainland, or on the offshore islands. Most mature specimens are characterized by their low pored-scale counts (Table IV). Of the coastal lake populations, small specimens were available only from Skidegate L. A l l of these were im- mature and a l l had low pored-scale counts, but the "complete" counts of this, and other coastal lake populations, were equivalent to the mean counts for other coastal and inland populations. Tentative analysis of mean "complete" lateral line counts indicates a tendency toward higher counts in the north. Paucity of counts throughout the entire range of Cottus asper precludes any further comparisons at this; time. Discussion of.Meristic Variation Within Cottus asper Lindsey (1961; 1962) has reviewed literature which demonstrates that TABLE IV. Lateral Line Pored-Scale Counts by Size Groups for Selected Representative Populations of C. asper Population Size 1 7 1 8 1 9 2 0 2 1 2 2 2 3 2 4 2 5 2 6 _ 2 7 2 8 ^ 2 9 3 0 3 1 3 2 _ 33 3 4 3 5 3 6 _ 11 3 8 3 9 4 0 4 1 _ 4 2 _ 4 3 _ Saltery Cr. A-laska ^ 6 0 eno •oes oo «o o o o o ==> — <• — o oo — o = o _ = 1 3 9 3 — . » ~ * **"complete" X = 4 0 . 0 6 Cultus L o B.C. oo o — oo oo o « o o =,= oo — - •=» 1 1 3 3 2 2 ^ — > • "complete" 5 0 - 6 0 = = oo oo oo = o == — = - — - = == — - = • » = — — — — — , X = 4 0 . 2 7 4 0 = 4 9 oo = = oo oo oo = = — — — ==. OO ==• 2 OQ 1 == =•= — otto 3 0 = 3 9 oo = = -o = o = - — 1 — 1 2 1 1 — ==. — 2 0 = 2 9 ~ " " 1 ^ = = 1 — == = = ="•= =«= •=•= <==. — ~ —= — Graham Id. g QCI 5 * 6 0 oo oo oo <=c oo oo « = ~ = == ==. OO OS 1 1 4 1 3 3 1 Juskatla Inlet 5 0 = 5 9 oo _ = oo — - = — — — = - — — == 3 2 3 4 J=«= CO "complete " 4 0 = 4 9 = - o o - - • - * =•= — — 1 M 1 2 3 1 9 5 1 5 = . X = 3 9 . 3 3 3 0 = 3 9 1 1 2 2 IB<B •<=> - - . 1 = - 3 2 ~ ~ — 2 0 = 2 9 = = == == = = — * ~ = — — —= —— 1 o o «-.<= - — < = „ = ~ == •="• — Moresby Id., QCI > 6 0 oo oo o o oo o o ™ - — — 2 1 2 1 - = 3 == 2 4 2 3 « = - == Skidgate L . 5 0 = 5 9 = =, oo oo oo oo — — —=• == - = 1 „„ 1 2 - = == 1 == —= =•= "complete'' 4 0 = 4 9 oo oo o — o o — o oo — — 1 OO 1 1 —= 1 — — =•"= == =— == = == === == == ==• == X = 3 9 . 7 Q 3 0 = 3 9 — — 1 —\ Buttle L . Van. Id. > 6 0 ~ = = ~ 1 — 1 1 1 — 1 — >=•= — o c a , —«= == •== "complete" 4 0 - 4 9 CB-3 •><-> COB o v a - e-o BO == = — DCS X = 3 9 . 7 1 - - Horne L . , Van. Id. 2 "complete" > 6 0 oo . _ — ~ — 1 1 2 2 " 2 - = — - = = = = ., 5T= 3 9 o 4 5 f * "complete" counts are mean counts of latera l line pored-scales plus unenclosed neuromast hill o c k s . ** Saltery Cr. represents a typical coastal population i Cultus L. represents a typical inland population; Juskatla Inlet represents a coastal island stream population; Skidegate,Buttie, and Horne L, represent coastal lake populations 45 in fishes reared experimentally over a wide range of temperatures, a V- shaped, or inverted V-shaped, curve i s typical for meristic series, es- pecially vertebrae. Generally, an intermediate temperature produces the minimum count, but sometimes, a maximum count. Lindsey (1962) states that the inflection in the curve might not be discovered i f a narrow range of temperature were used and that demonstration of a V-shaped, rather than a simple direct or inverse, relationship between meristic counts and tempera- tures sometimes requires rearing at extremes close to the upper or lower level of temperature tolerance of the species. Such V-shaped curves have seldom been shown to exist in natural f i s h populations. That they exist in Cottus asper, a latitudinally wide-ranging species, might be interpreted to indicate that, at the ends of i t s d i s t r i - butional range, the prickly sculpin's eggs are subjected to developmental temperatures close to the upper and lower levels of tolerance. However,.we have elsewhere indicated (p. 53) that the usual spawning range for Cottus ..' asper i s from 8 to 13° C, It i s assumed that in a l l populations spawning i s initiated within this 5° range of temperature which proceeds in somewhat of a thermal "wave" from south to north in both inland and coastal l o c a l i - ties (Fig, 5). This i s not to say that they a l l spawn at the same tempera- ture within that range. Furthermore, because the monthly rate of increase of ambient temperature is greater in the north (Fig, 12, and Appendix Table 7), the duration of exposure to any given temperature within the 5° tempera- ture range may be shorter. Eggs subjected to these conditions would pre- sumably have a faster development than eggs which developed under relatively more thermostable conditions such as occur in the south. Low meristic counts are generally (although not invariably) associated with faster rates of development. Based on the information given above, northern eggs should produce offspring with low meristic counts. But when plotted against l a t i - tude, the mean pectoral f i n ray counts of inland Cottus asper populations 46 produce a linear curve which is typically clinal, i.e., with more rays in the high latitudes, fewer in the low latitudes. The coastal population means demonstrate a V-shaped curve with low counts in the high and low latitudes, and higher counts in the mid latitudes. Such results are enig- matic. Whether a temperature range of only 5° C. is enough to e l i c i t a V- shaped response has not been experimentally confirmed for Cottus asper, but Lindsey (1962) obtained such a curve in the mean counts of pectoral fin rays in the stickleback, Gasterosteus aculeatus, over a developmental range of 15 to 20° C. It is apparent from Fig. 11 that V-shaped curves are characteristic for the median soft fin ray counts of Cottus asper whereas for the paired pectoral fins, only the coastal populations show a V-shaped curve. The in- land means appear to be the only exception, at least in the high latitudes. The existence of such a divergence between coastal and inland means indicates that temperature alone is not the determining factor. Interpretation of the divergent results between coastal and inland means is facilitated by an analysis of mean pectoral fin ray counts for coastal lake and coastal derivative populations. In the former, the mean counts do not f a l l , as would be expected, on the curve for coastal popula- tions but they are directly in line with, and indistinguishable from, the higher means of inland populations. In the derivative populations, the means are found to be equal to, or slightly less than, the inland means but always greater than the coastal means at the same latitude (Fig, 5), This further suggests that temperature alone cannot be the determining factor. There is only one universal factor which, at any given latitude, varies between coastal populations and a l l others. This factor is salinity and/or the lack of access to i t in the spawning population. At least two of the four coastal lake populations are restricted to spawning in fresh water 47 because of man-made or natural barriers, i.e., Horne L. and Buttle L. on Vancouver Island. Even though the mouth of Skidegate L. on Moresby Island, QCI, is only about 10 miles from the sea, the population of Cottus asper from which the meristic data were sampled, is known to spawn in fresh water inlet streams. The spawning locality of the Owikeno L., B. C, population is unknown. That of the Chilkoot L., Alaska, population is also unknown but the fin ray counts are typically inland whereas the prickling pattern is definitely coastal in its affinity. It may therefore be supposed that spawning occurs in fresh water. It may also be presumed that a l l the coastal derivative forms now spawn in fresh water. Therefore, in the high latitudes, increased-mean pectoral fin ray counts in coastal lake or "coastal derivative" populations, and in inland populations, is correlated with the lack of salinity in the environment. Any discussion.of the nature of the differences between coastal and a l l other populations is a matter of conjecture with the evidence now at hand. Two opposing hypothesis can be postulated: 1) The observed differences reflect environmentally induced changes of the individual phenotypes within each population; or 2) The differences have become genetically fixed in the populations over a long period of time. Consideration of these alternatives rests on an interpretation of the relationship of the coastal lake and'coastal derivative" populations to the main coastal population. In the areal distribution and intensity of their prickling, the coastal lake and "coastal derivative" forms show very close affinity with the coastal populations. Furthermore, because of the glacial history of the coastal islands on which they are found, the near-naked forms from Buttle L., Horne L., and Skidegate L,, can be assumed to have been de- rived from coastal stream forms which colonized the island after the retreat H8 of the Cordilleran ice sheet which covered the area during the late Wisconsin Glaciation. The probability that they were colonized by inland forms is remote since the nearest inland forms would presumably have been found in unglaciated areas of the lower Columbia R. basin (cf. Fig, 20), In similar manner, the occurrence of the "coastal derivative" popula- tions only in parts of northern British Columbia relatively recently ex- posed by the retreating glaciers argues that these populations are probably derived from coastal ancestors which followed the retreat of the glaciers northward and then inland. It can be assumed that they are more recent than their coastal lake counterparts,.and perhaps even more recent than those in- land forms which followed the retreat of the ice in the Fraser R. drainage (cf. final discussion, p. 7 9 ) ; . As was previously stated, the increased mean pectoral fin ray counts in coastal lake and "coastal derivative" populations is correlated with a lack of salinity. One can explain the phenotypic changes in pectoral fin ray counts by postulating an environmental inhibition or alteration of the metabolic pathway usually traversed by the gene which controls the pheno- typic expression. The lack of salinity in the environment might produce such an alteration. But, since not a l l of the "coastal derivative" means are equal to those of the inland populations, i t can also be argued that,., because of a selective pressure caused by the absence of salinity in the environment, the coastal lake populations have become, and the "coastal derivative" populations are becoming, genotypically modified toward the means characteristic of the inland forms of Cottus asper. Such modification might be envisioned in terms of a mutation in one of the steps in the meta- bolic pathway of the controlling gene. Whatever the cause of the differences, the evidence at hand is not now sufficient to determine its nature. A definitive experiment regarding the nature of the differences would consist of taking the eggs of coastal lake 49 or'coastal derivative" forms and rearing them in brackish water using fresh water reared eggs as a control. If the mean fin ray counts of the F^ gener- ation were significantly lower than the control means, the result could be interpreted as being due to.environmental induction rather than to long term genetic fixation. In either event, the mechanisms involved in the - actual modification of the phenotype and/or genotype would s t i l l remain a matter for more refined genetic experiment. Summary of Morphological Analysis 1. Cottus asper is found in a l l major Pacific Slope drainages from Seward, Alaska, to Ventura, California. On the Artie Slope drainage its distribution is limited to the Upper MacKenzie R. drainage only in the headwaters of the Peace R. in British Columbia. 2. Some of the prickles found in the skin of Cottus asper are re- sorbed during reproductive maturation which occurs at about 60 mm S.L. Because of this, a l l population means were calculated for groups of speci- mens less than or greater than 60 mm S.L. 3. Estimates of the percentage of total body surface covered with prickles are calculated from the formula: % "virtual" prickling = % "apparent" prickling + % area of . ; - resorbed prickling. 4. Coastal populations of Cottus asper are, "apparently" and "virtually," less prickled than inland populations. Differences in "appar- ent" prickling between coastal and inland populations are more pronounced in specimens ^ 60 mm S.L., i.e., a coastal-inland ratio of 1 : 3 exists. 5. Inland populations of £. asper from the headwaters of the Peace, Ness, Stikine, and Skeena R. systems in northern British Columbia, are coastal in their prickling affinities and are referred to as "coastal dê r rivative" populations. 6. A population of C. asper from the mouth of the Sacramento R, in 50 California i s intermediate in i t s prickling a f f i n i t i e s and i t i s designated as an "intergrade" population. 7. Asymmetrical pectoral f i n element counts were found in 56 of 59 populations of £. asper. Of 1163 paired counts, from 59 populations, 23.6% .were asymmetrical. When the asymmetrical fins from a l l populations were pooled, the overall direction of asymmetry was significantly sinistral(64% s i n i s t r a l , 36% dextral). 8. Differences in amount of s i n i s t r a l i t y within and among aggregate- populations are not significant. No c l i n a l trends are evident. 9. Mean pectoral f i n ray counts for a l l populations are similar in the low and mid-latitudes.of the distributional range, but are widely divergent between coastal and inland populations in the high latitudes. 10. Means of "coastal derivative"populations are equal to or less than those of inland population means but always higher than those of coastal means at the same latitude, 11. Means of coastal lake forms are a f u l l count higher than the means of coastal stream populations from which they were probably derived. They are indistinguishable.from inland means at the same latitude. 12. When plotted against latitude, mean f i n ray counts produce curves which are different for inland and coastal populations. 13. For pectoral f i n s , the, greatest amount of divergence between i n * land and coastal means exists in the high latitudes. The opposite i s true for the dorsal f i n elements, 14. Except.for the spiny dorsal and the inland pectoral means, a l l meristic curves demonstrate a characteristic V-shaped pattern with the in- flection occurring at about 42 to 44°~N, Latitude, 15. The pelvic f i n almost invariably consists of 1 spiny plus 4 seg- mented rays, 16. Counts of the later a l line pored-scales are extremely variable 51 with size up to about 60 mm S.L, In Cottus asper, a "complete" lateral line count cannot be considered reliable unless a count i s made of the number of pored-scales plus the number of unenclosed neuromast hillocks, STUDIES ON THE LIFE HISTORY OF THE PRICKLY SCULPIN Present Knowledge of Spawning Migration in Cottus asper The existence of a seaward spawning migration of prickly sculpins in coastal streams has been suggested, or implied, for at least 30 years (Taft, 1934; Pritchard, 1936} Sumner,;1953; Shapovalov and Taft, 1954; Hunter, 1959; and McAllister and Lindsey, 1959). Although some of these authors have observed the presence of reproductively mature Cottus asper in the intertidal areas of coastal streams, none has demonstrated that intertidal spawning actually occurs. Populations of Cottus asper occurring in lakes and streams far enough inland to preclude the poss i b i l i t y of an annual seaward spawning migration are presumed to spawn in fresh water0 The approximate or exact spawning sites of some of these populations have now been determined from the pre- sence of larvae (Nicola L., B, C.), and egg clusters or gravid females (Pothole L. 8 near Merritt, B.C.). Other l o c a l i t i e s are close enough to the sea to imply the existence of a short seaward migration on the part of the Cottus asper populations l i v i n g therein, but access to the sea i s prevented by natural or manmade barriers, e.g.. Buttle L„ and Horne L. on Vancouver Island s B.C. Spawning of prickly sculpins in these areas is necessarily restricted to fresh water. S t i l l other'localities, frequented by migratory salmonids, are close enough to the sea to imply a seaward migration on the part of Cottus asper, but i t does not occur. For example, the spawning sites of the following, prickly sculpin populations in the lower Fraser Valley in British Columbia 52 are known from capture of gravid f i s h and/or egg masses: South Alduette R.; Kenworthy Cr. and Chilqua Slough (both are inlet streams to Hatzic L.); Squakum L.; and Cultus L. In addition, spawning fish have been captured in inlet streams of S k i d e g a t e i n the Queen Charlotte Islands, along with migratory juvenile salmonids. The outlet of Skidegate;L. i s only about 13 miles from the sea. A newly-hatched larva of Cottus asper (?) has been taken in a plankton net in the Second Narrows region of Owikeno L. (about 30 miles from the sea), on the coast of central British Columbia. It i s presumed that the parents spawned in fresh water. Spawning Period and Temperature Relationships. - Egg clusters were collected from several l o c a l i t i e s in-the lower Eraser valley and in the L i t t l e Campbell R. Gonads were examined in over 1100 of the preserved speci- mens from a l l latitudes. These-data indicate that egg deposition begins in the south of the distribution range (low latitudes) in February, and pro- gresses northward u n t i l late July. Males usually attain f u l l reproductive maturity about a month before, and remain in spawning condition for almost a month after, the period of oviposition in females. Gravid females have been found over a <+~week period in Squakum L., and 6 weeks in the L i t t l e Campbell R„ Ripe males have been taken over an 8 to 12 week period, re- spectively, in these same l o c a l i t i e s . The earliest date on which a ripe male, in nuptial dress and with flow- ing sperm, was collected i s February 6th, in San Francisco Bay. The earliest collection of gravid females was in Waddell Cr., California, on February 2<+th, In the north end of the range, gravid females,were taken as late as June 20th in Petersen Cr., near Juneau, Alaska, and on July 22nd in streams entering Juskatla Inlet, Queen Charlotte Islands. Gravid females have also been collected from Middle R., near Takla L., on June 28th, and from Meziadin L., B.C., on July 25th. Field records and personal observation indicate that natural spawning 53 temperature range from 8 to 13° C, Egg rearing experiments at various temperatures resulted in complete mortality in 18° C., and less than 50% survival at 15° C, The annual mean range of temperatures experienced by inland populations of Cottus asper is twice as great as that experienced by coastal populations at the same latitude (Appendix Table 7), The monthly mean temperature range between northern and southern localities is from 8.8 to 19,5° C. on the coast, and from 9.6 to 29.6° C. inland (Appendix Table 7). A priori, one might expect that inland populations would experience a greater range of developmental temperatures than do coastal populations. Apparently, how- ever, they do not. When monthly mean temperatures, representing inland and coastal localities encompassing the distributional range of Cottus asper« are plotted against latitude, the mean temperature differences between in- land and coastal localities during the spawning period are almost negligible (Fig, 12). In fact, the common temperature range of 8 to 13° C. (shaded bar, Fig, 12) can be followed as a thermal "wave" progressing from the south in February, to the north in June, Approximate spawning times, determined from examinations of gonadal condition in over a thousand specimens in a l l latitudes, are in general agreement with this south-north progression a l - though the latitudinal range over-which spawning occurs in March, April,-and May is remarkably consistent (Fig. 12). Theoretically, inland forms have a shorter period of exposure to spawn- ing temperature of 8 to 13° C,, thanido coastal forms (Fig, 12), This has been born out by field data from the;two most frequently collected spawning sites, i,e,, Squakum L,, and Little Campbell R. According to Fig. 12, the inception of spawning in inland streams should lag behind that of coastal streams at similar latitudes. This is because upstream or inland areas remain; colder for a longer period than coastal areas. This is apparently true in the lower Fraser valley. For example, LEGEND; Coastal (• •); Inland (o-—-o); Empirical Spawning Temperature Range (8-l3°C .)E Latitudinal Range of Probable Spawning in: Coastal 1\ \ \ \ \ l ; Inland K / / / \ \ and Both IX Y Y Y M Forms. State of Maturation: VII = Near Spawning, Vlll= Spawning Imminent, Eggs and Sperm Flowing; IX= Spent. Figure 12.-Monthly/Mean Temperatures in °C„ Arranged by Latitude for Coastal and Inland Localities Encompassing the Distributional Range of Cottus asper. Values obtained from Appendix Table 7. Solid or f i l l e d v e r t i c a l bars 8 l e f t side of graph, indicate actual ranges of latitude over which specimens have been found in given state of maturity. 55 the following localities are all within 15' of 49° N. Latitude, and gravid females and/or eggs have been collected from them during the following dates; March 7th to May 10th in Little Campbell R. (122° 46' W. Long.); March 25th to April 15th in S. Alauette R. (122° 35' W. Long.); and April 30th to May 27th in Squakum L. (122° 00' W. Long.). Some of the implications of these temperature and spawning data have al- ready been discussed*(p. 45), Materials and Methods > s To document the supposed occurrence and to determine the success of intertidal spawning of prickly sculpins in: the coastal streams, the Little Campbell R. was chosen as a study stream. In1960-1961, a series of 18 permanent collecting sites (Fig. 14) was sampled at bi-weekly intervals for a period of one year, and at monthly intervals for an additional six months. The lower reaches of the river, stations C-l to C-3, were also sampled • several times in late Winter and early Spring of 1962 and 1963, to obtain live specimens for experimental studies, A 3 mm mesh woven nylon seine, 3 meters wide x 2 meters deep, was mounted on collapsible telescoping aluminum poles and used for all field collections. Salinities were measured with three salinity hydrometers, densities; 1.000 to 1.011; 1.010 to 1,021; and 1,020 to 1.031 sp, gr, of NaCl. Sampling Localities and Study Streams The primary study area was the Little Campbell R, (Campbell Cr.), which is approximately 15 miles long and empties into Semiahmoo Bay between White Rock, British Columbia, and Blaine, Washington (inset of Fig. 13), The stream drainage consists of an area of approximately 28 square miles. Collection-sites are shown in Fig, 14. Station 0-1 is located on a sand-mud flat outside the main river channel. Station C-0 is located at the mouth, in the main channel. Station C-l is located about 50 yds, inside the Figure 13.—Collection Localities for Spawning Populations of Cottus asper Used in Life History Studies. Figure 14.—Collection Localities Within L i t t l e Campbell R. 58 mouth and Station C-1A about 75 yds. inside the mouth. Station C-2 i s locat- ed 1/4 mile, and C-3 i s 1/3 mile, from the mouth. Fig. 15 shows collection sites in relation to salinity gradients and elevation. Stations C-0 through C-9 are a l l subject to t i d a l fluctuations in depth, whereas stations C-0 through C-3 are t i d a l l y inundated with mixo- haline waters. Barnacles (Balanus sp.) are found at stations C-0 through C-3, and permanent beds of the oyster, Crassostrea gigas, are located be- tween Stations C-l and C-1A, and at C-2, Typical fish associates in areas C-0 through C-3 are Leptocottus armatus and Platichthys stellatus; Oligo- cottus maculosus and Clinocottus acuticeps are commonly found up to Station C-2, Additional live specimens for use in the experimental studies were collected from the following l o c a l i t i e s (Fig, 13)s Nile Cr, and Big Qualicum R,, Vancouver Island; South Ald;uette R,; Kenworthy and Edwards Cr, (Hatzic L o ) ; Squakum L, (Lake Erroch); and Sweltzer Cr, (outlet of Cultus L.) Results of Field Studies The prickly sculpin i s distributed primarily in the lower four miles of the L i t t l e Campbell R, (Fig, 217), From late February to early March, the prickly sculpin undertakes a migration downstream to the estuary. The only area in the lower four miles of stream in which a suitable spawning sub- strate (large cobbles, f l a t rocks) occurs i s a stretch about 100 yards long lying one quarter mile upstream from the mouth (Station C-2, Figs, 14 S 15), The males, which come into spawning condition earlier in the season than-the females, select nesting sites under large cobbles or flat rocks in areas of the stream bed with current velocities equal to or less than one cubic foot/ second (at low tide). It i s apparently important that the substrate surface be relatively rough in texture since the adhesive eggs adhere only tempo- r a r i l y to a smooth surface. Old automobile exhaust pipes, or muffler tubes, A L T I T U D E I N F E E T cn O cn O cn 6S 60 are preferred nesting sites when available in the environment (as they are in the Little Campbell R.), Females aggregate upstream (about Station C-3) above the main spawning area and then move individually onto the spawning beds where they are courted by the males both outside.and inside the nest. After the male selects a female to occupy his nest, further courtship and pre-spawning behavior occurs within the nest. The adhesive eggs are laid in a jelly- enclosed cluster on the ceiling of the spawning chamber. Ovariotomy of preserved gravid females from throughout the range of C. asper yielded counts of 336 mature ova in a 4.9,5 mm S.L. female to 5,652 mature ova in a female 119.5 mm S.L. The largest female examined was 192 mm S.L,, but she was spent. A conservative estimate would be about 10,000 eggs for this female. Egg numbers, in masses collected in the field, varied from 700 to 4000. However, one male may court and successfully mate with as many as 10 different females. As many as 10 egg masses, in varying stages of develop- ment from newly-fertilized to near-hatching, have been found in the nest of a single brooding male. An estimated 25,000-30,000 eggs were present in this one nest. After spawning, the spent females leave, or are chased from, the nests and they again aggregate above the spawning areas and begin feeding. The males remain in the nests, fanning and protecting the eggs, and do not eat until hatching is completed. Much of the courtship and pre-spawning be- havior, as well as most of the paternal brooding behavior, has been docu- mented but will not be reported here. Fig. 16 illustrates numbers and distribution of Cottus asper young-of- the-year, 12-25 mm S.L., taken in a total of 700 seine hauls. In late Spring and throughout the Summer, the young-of-the-year are found in great numbers around and below the spawning site. The concentration is greatest around Station C-1A where there is a bed of fine, pea-size gravel adjacent T CO 23 III1 £ 1 9 h - C O | I O 9 o LU O 4 O * 3 - J 2 —J IA LU |, Op Q, a, 0-1 O o o o o I- o x o o o o o o o o o o o X o o o o o o o o o X o o o o o o o o o o o X o o o o o o o o o X o o o o o o o o o o X X X o o X X o o X X X X X x X o o o X o o X X o o o o o X o X o o o - 4 9 8 s 7 — 6 f" 5 CO 32 CO m I > O £ CO ° 2 CD L E G E N D ; X= NO collection made; 0=Collectlon, but no fish; #=1-6 fish; •=7-18; W=l9-36; W = 37-60, W = 6 l - 9 0 l \ \ \ \ \ M Intertidal Zone; EMnnsnninSpawning Zone; ^ ^ ^ B l I S p a w n i n g Season. Figure 16„—Monthly Distribution of Young-of-the-Year Cottus asper Within the L i t t l e Campbell Re Taken from Pooled Bi-Weekly Samples, 62 to a large oyster bed. In mid-Summer, there is a definite upstream migration of the young-of- the year. In both 1960 and 1961, the increasing abundance of young-of-the- year at Stations C-4 and C-5 was.correlated with the decreasing abundance of specimens in the estuarine areas of the river (Fig. 16). Fig. 17 illustrates the disjunct distribution of yearlings, sub-adult, and adult prickly sculpins in the Little Campbell R. Especially in Spring, 1961, an increased number of Cottus asper were present in the lower reaches of the river, around the spawning site. Over the 18-month sampling period, no Cottus asper were captured in stations upstream of C-22, With three exceptions, none was taken in the fast flowing, high gradient area of the stream below C-20 and above C-ll (Fig. 15), This area is densely populated with the coastrange sculpin, Cottus aleuticus. Newly metamorphosed young-of-the-year are found in abundance around and especially below the main spawning beds. Laboratory studies on the behavior of Cottus asper larvae, done in extension of the.experimental salinity tolerance experiments, indicate that, at 12° C,, the larvae 5 to 7 mm in total length begin swimming immediately upon hatching. They remain pelagic, as lightly-pigmented, transparent larvae, for a period of 30 to 35 days before metamorphosing and settling to the bottom. Salinity Tolerance in Prickly Sculpin Eggs Attempts to rear arti f i c i a l l y fertilized eggs beyond the critical yolk- sac resorption stage were unsuccessful. Although a few sculpins were reared to a point where meristic series and prickling patterns were fixed, sample sizes were too small to determine whether the salinity of the fertilization and incubation medium was correlated with fin ray numbers, or the intensity and distribution of prickles. Alternate criteria, based on the percentage hatching success of "in- land" and "coastal" eggs in brackish and fresh water, indicated a phenomenon CO 23- O Z r : 2 r p CO , 2 11 O 9 o LU < X 0 X 0 X 0 0 0 X X X X X X X X _ 9 • • • • • 0 0 0 0 0 X X • • X 8 7 0 0 0 0 0 0 0 0 0 0 X 0 X 0 0 x • 1 6 0 0 0 0 0 0 0 0 0 • 0 0 X 0 0 5 • 0 0 0 0 0 • • 0 0 • X X X 0 •- 4 • • X • X • 0 X 0 X X X X 0 # • _ 3 J J A S 0 N D J F M A M J J A S O I 960 1961 m CO m 1 > O w <Q CO O S CD LEGEND: X-NO collection made; OCollection, but no fish; •=l-6 fish; •=7-18; W=l9-36, W = 37-60; W = 6 h 9 0 1 \ \ \ \ \ \i Intertidal Zone; EinnnnsnnniSpawning Zone; Mli j i l i i ff l Spawning Season. Figure 17.—Distribution of Yearling, Sub-Adult, and Adult Cottus asper within the Lit t l e Campbell R, 64 of non-genetic adaptation on the part of the "inland" eggs. The low hatching success of "inland" eggs which had been f e r t i l i z e d and incubated in brackish water could be significantly increased by f i r s t f e r t i l i z i n g them in brackish water. It was also found that acclimation of "inland" parents to brackish water at least three days prior to f e r t i l i z a t i o n significantly increased the percentage hatching success of their eggs over those eggs from parents which had been maintained in fresh water prior to f e r t i l i z a t i o n . Data for these egg salin i t y tolerance experiments is on f i l e in the Institute of Fisheries, Department of Zoology, University of British Columbia. Discussion This study confirms the existence of a downstream spawning migration of the prickly sculpins in the L i t t l e Campbell R. It not only documents the occurrence of intertidal spawning in Cottus asper, but indicates that this spawning is successful. Figure 16 shows a lag of about two months in the appearance of young- of=the~year C, asper, 12 to 25 mm S.L., after the f i r s t recorded spawning.in March. Water temperatures in March are normally from 8 to 10° C„ Eggs spawned early in March probably have an incubation period several days longer than the 15=16 day period found to be typical at 12° C, in the labo- ratory. The lag shown in Figure 16 i s probably due to an incubation period of three weeks followed by a pelagic larval stage of 4 to 5 weeks. Plankton tows taken during late April in the shallow back-eddies of the stream have captured a few larvae 9 to 10 mm S.L, (not recorded in Fig, 16), An upstream migration of adults precedes that of the young=of-the-year in late Summer, This is probably related to the food habits of the two groups and also to the fact that a later return of the young-of-the-year coincides with lower water levels in the stream, when reduced velocity f a c i l i t a t e s easier access upstream. 65 Summary of Life History Studies 1. The study stream, the Little Campbell R., is a small coastal stream, the lower one half mile of which is subject to tidal inundation of mixohaline waters. 2. In late Winter and early Spring, the adults and juvenile prickly sculpins migrate downstream to the estuarine region of the Little Campbell R., the only region in the lower four miles in which suitable spawning sub- strate is available« 3. Males set up nesting sites under large cobbles and rocks and courtship occurs both outside and within the nest* 4. Spawning occurs from March throughout early May. 5. Newly-hatched larvae begin swimming immediately and remain pelagic for a period of 30 to 35 days before metamorphosing and settling to the bottom. 6. In May, metamorphosed young-of-the-year (approximately 12 mm S.L.) begin appearing only in those collections taken in the estuarine portion of the river. They occur in great abundance until September when the numbers decrease in the estuary and increase in upstream, non-estuarine waters. 7. During the non-migratory phase of its life history, the Cottus asper population in the Little Campbell R. is distributed disjunctly in the low gradient, low velocity, portions of the stream. DISCUSSION The purpose of this investigation was to determine which characters in Cottus asper were genetically determined and which were due to non- genetic modification. Properly answered, these questions could help the systematist to a better understanding of the selective forces that produce variations within a species. This knowledge would, in turn, facilitate a 66 more stable classification of taxa. Morphological Variation Within Cottus asper The interpretation of the morphological variation will be considered in the light of experimental and l i f e history data obtained in this investi- gation. Table V summarizes several of the variable characteristics for aggregate populations of Cottus asper. Coastal vs. Inland Populations. - The data presented in Tables I and V8 and in Figure 4, reveal consistent differences in prickling patterns which coincide exactly with the geographic locality of collections. Coastal fish have less area of the body covered by prickles (ratio 1 t 1.8), and less prickles per unit area (ratio 1 : 1,4) than inland fish. The existence and maintenance of such a remarkable discontinuity of characters between closely related but geographically distinct populations, distributed over a wide range of latitudes, must be interpreted as being genetically deter- mined. Furthermore, these populations, which can be separated on the basis of prickling patterns and geographic distribution, can also be separated by other characters. For example, consider the migratory behavior. Weakly-prickled Cottus asper living in coastal streams having open access to' the sea undertake a downstream migration to the estuarine regions where they spawn. Densely-prickled Cottus asper living in distant inland ;, streams, where access to the sea is almost impossible, undertake only local migratory movements. But densely-prickled forms living in some inland lakes and streams relatively close to the sea, where access to the sea is open and relatively easy, do not migrate seaward. They also undertake only local movements and spawn within the fresh,water system in which they are found. These primary differences in behavior patterns correlated with distinct prickling patterns further corroborate the contention that coastal and in- land forms of Cottus asper are genetically distinct. The evidence based on meristic counts is equivocal. When plotted TABLE V» Summary of Variable Characteristics for-Selected Phenotypes of Cottus asper,, Phenotype Descriptions COASTAL Northern COASTAL DERIVATIVE Northern COASTAL LAKE Southern iNTERGRADE INLAND Distribut ion s Streams of Pacific Coast from Alaska to California River drainages north of Fraser R, drainage in British Columbia Island and mainland lakes close to sea in British Columbia and Alaska Mouth of Sac= ramento R» in San Pablo and San Francisco Bays A l l major inland drainage sys= terns on Pacific Slope except drainages north of Fraser River., Prickling Virtual Areas Intensity! % Resorptions Low Low High Low Low Lower Lowest Lowest Lower "-.Higher Higher High Highest Highest Lowest Meristics Pectoral Ray Numbers @ High Latitude Combined Dorsals at Low Latitudes Low Highest Higher Highest Higher Highest Low TABLE V (cont'd). Summary of Variable Characteristics for Selected Phenotypes of Cottus asper. Phenotype Description: COASTAL Northern COASTAL DERIVATIVE Northern COASTAL LAKE Southern INTERGRADE INLAND Migratory Behavior: Seaward Migration ? ? ? Local Migra- tory movement Natural Spawning Environment: Brackish Presumably Fresh Presumably Fresh Presumably Brackish Fresh Egg Survival in other than natural media: Excellent; as in natural medium ? ? ? Poor; signifi- cantly less ex- cept i f parents or eggs pre- acclimated 69 against latitude, the means for inland and coastal populations form V-shaped curves which, though overlapping in certain areas, are widely divergent in others (Fig. 11). For example, the pectoral rays of inland and coastal populations are similar in the low latitudes but at a point about 42-44° N, Lat., the coastal means begin to decrease, whereas the inland means con- tinue to increase, in fin ray numbers. Such divergence in mean counts be- tween coastal and inland populations can not be explained on the basis of temperature alone. Salinity in the spawning environment is the only factor which, at any given latitude, can be shown to vary directly between inland and coastal populations. The nature of the differences between coastal forms and a l l the popula- tions probably derived from them, or between coastal and inland forms, is a matter of conjecture. Phenotypic modifications now shown to exist can, on the basis of present evidence, be interpreted either as being individual changes induced by the local environment, or, as having become genetically fixed in the populations over long periods of time. Theoretical arguments for each interpretation have been presented ( p, 48 )» but the final deter- mination of the underlying nature of existing differences in fin ray counts awaits the test of successful rearing, under controlled conditions, to a stage where fin ray numbers are fixed. Derivative and Intergrade Populations, - Certain sub-groups of prickly sculpins have been found which are morphologically distinctive but which have close affinity to either the coastal or inland forms. An analysis of these populations corroborates the contention that, at least in prickling pattern and migratory behavior, coastal and inland forms of Cottus asper are genetically discrete. Even though the coastal lake and "coastal derivative" populations have prickling patterns similar or identical to coastal forms, the absolute per- centage resorption is more like that of the inland forms (Tables I and V), 70 This might be expected since presumably they no longer spawn in brackish water and the osmotic stresses experienced while spawning in fresh water must be different. The prickling pattern of the southern "intergrade" population (Carquinez Str.,Calif.) is similar to that of the densely prickled inland population forms from which it may have been derived. The amount of prickling resorp- tion, however, approaches that of the coastal percentage. If we assume, as we have above, that the percentage resorption is greater in brackish water this would indicate that they spawn in estuarine conditions. In their mean fin ray counts, other populations from the San Pablo-San Francisco Bay area are intermediate between the coastal and inland populations. With the in- formation now available, i t would be difficult to determine whether these are true inland derivatives which have colonized the streams of the Bay region or whether they are intergrades between inland and coastal forms. :, The Phylogeny of Cottus asper and Closely-Related Species The prickly sculpin, Cottus asper, is a polytypic species. Figure 18 shows the known range of various phenotypes of Cottus asper, determined in this study, plotted against the distributional ranges of closely-related species which comprise the "asper species group" proposed by Bailey and Bond (1963). From a taxonomic viewpoint several interpretations of this polytypy can be made: 1) Cottus asper is a phenotypically variable species within which can be recognized 4 or 5 races; or 2) Cottus asper represents a species complex comprised of at least two sibling species, each with one or more derivative subspecies (or races i f the phenotypic differences can be shown to be en- vironmentally induced). 3) Cottus asper represents a polytypic species complex in which a spectrum of evolutionary grades, from incipient subspecies through gure 18c— Known Ranges of Various Phenotypes of Cottus asper Plotted Against the Distributional Ranges Closely-Related Species in the "asper species group". 72 f u l l species, can be traced. Evidence presented in this study favors the l a t t e r interpretation. The genus Cottus i s a fresh water derivative of the predominantly marine family Cottidae. The genus Cottus is most closely related to the Asiatic marine genus Trachidermus, represented on the Pacific Coast of N, America by Leptocottus, a closely-allied genus. Although their interpreta- tions of relationships within the genus Cottus were not always accurate, I concur with the earlier opinion of Jordan and Evermann (1898, p. 1942) that, of the species of Cottus occupying the Pacific Slope, Cottus asper appears to be more closely a l l i e d to Trachidermus. It has been concluded that differences in prickling patterns and migratory behavior between the geographically separated coastal and inland populations of Cottus asper are genetic (p.66 ). The discontinuity in such conservative characteristics indicates that the isolation of the inland populations from the coastal gene pool has been of long-standing, perhaps since the Pliocene or late Miocene. Evidence to support this speculations i s based on the geologic history of the area now occupied by Cottus asper and closely-related species in the "asper,!species group" discussed below. Stage I. - It must be assumed that at sometime in the distant past, i.e., perhaps Pliocene or e a r l i e r , the ancestral form of Cottus asper was derived from a marine ancestor similar to Trachidermus. After this i n i t i a l separation, some of the ancestral Cottus asper penetrated inland and, through geological agencies and/or biological processes, became isolated from the coastal gene pool. Such an isolation i s represented schematically as Stage I in the evolution of Cottus asper (Fig. 19). The dashed arrows indicate probable routes of invasion of coastal types inland via either the Columbia R. or the Sacramento R. system. During the Pleistocene, access to the north was blocked by the Cordilleran ice sheet, lobes of which advanced and re- treated several times. The limits ^hown in Fig, 18 indicates the maximum 73 Figure 19,~Phylogeny of Cottus asper. Stage I, Evolution in the Pliocene (?)„ Similar pattern of background indicates continuous range, 7 4 southward extent of ice cover during the late Pleistocene; earlier ad- vances were somewhat further southward, Zoogeographically, the disjunct populations of Cottus asper and "asper species group" represented in Fig, 18 are members of one of three great Pacific Slope fresh water ichthyofaunae, i.e,, the Columbia; the Klamath; and the Sacramento, It i s here proposed that at some time or times before the basins B especially the Klamath, became completely isolated from one another by alternate periods of floods and desiccation, lava flows, and mountain building, a continuous exchange of fauna was possible. The geo- logical evidence of contemporaneous connections between the (Pleistocene) pluvial lake basins is wanting, but the ichthyological evidence c a l l s for such a connection some time in the past (Hubbs and Miller, 1948), For ex- ample, the cyprinid genus Ptychocheilus i s absent from the Klamath system but closely related species are present in the Columbia system to the north, and in the Sacramento system to the south. The sucker, Catostomus snyderi, and local representatives of the dace Rhinichthys occurring in the Klamath basin may be related to the forms in the Columbia and the Sacramento (Hubbs and Miller, 1948), Stage II. - The fluctuating regimen of the pluvial lakes in the Great Basin has been correlated approximately with the advances and retreats of the Cordilleran glaciers ( F l i n t , 1957; Hubbs and Miller, 1948). It i s here suggested that during this period of extensive flooding and desiccation, the gene pool of the previously continuous inland population of Cottus asper was divided up into a series of geographically isolated populations, some in the Klamath basin, some in the upper Sacramento, and some in the Deschutes R. drainage of the Columbia. To-the south of Klamath, Cottus gulosus evolved as a fast water, upstream derivative of inland Cottus asper and spread southward and out to the Pacific Coast, Among other differences, the fecundity of gulosus is intermediate between the high egg complement of 75 asper and the low complements of non-related species of Cottus (Bond, 1963). Perhaps one of the most intensive selective factors impinging on the incipient gulosus was that favoring a large egg which would produce a non-pelagic larva in the fast water environment (Cottus asper has pelagic larvae). Cottus pitensis, a disjunct r e l i c t form from the upper Sacramento R. above Lake Shasta, and from the upper Pit R. and Goose L, drainage, is thought to be derived from gulosus (Bailey and Bond, 1963j Bond, 1963). To the north of Klamath, Cottus perplexus was probably derived from inland asper as the northern ecotypic counterpart of gulosus, in so far as i t also prefers upstream areas and has an intermediate egg complement. By an unknown connection, perhaps through the outlet of Pluvial Fort Rock L. which contains fo s s i l s of possible faunal relationship with the Klamath Lakes, Lake Lahontan, Lake Bonneville, and the Columbia River (Hubbs and Miller, 1948), perplexus worked i t s way into the Deschutes R, and down to the Columbia R. where i t spread both upstream and downstream (Fig, 20), Cottus marginatus is probably a r e l i c t form of a formerly more widespread distribution of perplexus (Bond, 1963). Ecologically variable, and closely related to perplexus, is Cottus klamathensis, a disjunct r e l i c t found in Klamath L,, Oregon and Pit R, California (Gilbert, 1898} Bailey and Dimick, 1949} Robins and Miller, 1957; Bailey and Bond, 1963} and Bond, 1963). A Pleistocene connection between the Klamath, Goose L,, and Pit R, basins has been suggested by Hubbs and Miller (1948), and Robins and Miller (1957). The fecundity of klamathensis i s as high as in asper, and other characters suggest a close relationship with, and an early derivation from, the ancestral asper or perplexus. For example, Gilbert (1898, p, 11) commented on the resorption of body prickles occurring in klamathensis; "These, [the prickles of the young] become grad- ually absorbed with age, adults being nearly or quite naked," This is the 76 first recorded observation on prickling resorption and i t had been com- pletely overlooked by later authors. The suggestion of Robins and Miller (1957) that Cottus princeps is a long-standing derivative of asper is probably correct. The relationship is probably closest to the ancestral inland asper. Meristic evidence that the "asper species group" could have been de- rived from an ancestral inland form of Cottus asper is presented in Table VI. With few exceptions, the ranges of counts for a l l fin rays for a l l of the species f a l l within the range of counts typical of the present populations of inland Cottus asper. Such a<correlation may be interpreted to indicate that the genotype of the inland asper has a wide reaction range. A schematic representing the proposed second stage of the phylogeny of Cottus asper is presented in Fig. 20. Additional consideration of Cottus perplexus and C, gulosus will now be given. In its coastal distribution, Cottus perplexus is widespread in drainages of Washington and Oregon but does not cross the California border (Fig. 18). On the other hand, Cottus gulosus is found somewhat north and south of San Francisco, but only disjunctly in northern California and parts of the Oregon coast. When they occur*in the same stream, perplexus is generally, though not always, found in faster water, whereas gulosus occurs somewhat downstream in slightly slower velocity waters (Bond, 1963). Bond (personal communication) has stated that in areas where asper, gulosus, and perplexus occur in the same stream* a very thin slice of allopatry is cut! However, where upstream migration to the upper portion of streams has been blocked by impassable f a l l s , perplexus is often found above the falls where- as gulosus is always found below (Bond, 1963). This indicates, perhaps, that in streams where both species are present, gulosus is the most recent arrival. Both species have been found to be salt tolerant of at least 2H °/oo S. (Bond, 1963). Cottus asper is tolerant of long periods of TABLE VI. Summary of Meristic Data for Cottus asper and Other Species in the "asper species group,," Species D 1 D 2 D +D 1 2 A P 1 P 2 asper (coastal) 8-10 (11) 19-22 (23) (26) 27-32 15-18 \ 14-18 (3) 4 marginatus n 17-19 4 14-16 3 perplexus 2 7-8 2 18-20 2 13-16 (13) 1 14-15 (16) 14-16 2 4 klamathansis (6) 7-82 2 18-20 2 13-15 14-16 2 4 asper (inland) 7-10 (16) 18-22 (23) 26-32 15-18 (20) (13) 14-19 (19) (3) 4 princeps 2 6-8 2 19-23 26-302 15-182 14-162 2 4 qulosus pitensis (6) 5, 7- 9 8- 9 1 6 d o ) 1 (16) 3 16- 19 17- 19 6 17-18 (19) 1 . 14-163 13-156 12-145 (±2) 13-151 (14) (12) 1 15-16 (17) 14-166 1 13-15 (16) (3) 4 1 Superscript Notation; 1) Bailey and Bond, 1963; 2) Robins and-Miller, 1957; 3) Bond, 1961; 4 ) Schultz, 1936; 5) Snyder, 1905; 6) Snyder, 1913. A l l data on asper from this study. 78 Figure 20c=-Phylogeny of Cottus asper, Stage II, Evolution of "asper species group" During the Pleistocene, 79 immersion in higher salinities, i.e., greater than 31 °/oo S. (personal observation; and Bond, 1963). The salt tolerance of other "asper complex" species is unknown. Stage III. - By late Pleistocene, most of the speciation within the inland Cottus asper complex probably had occurred. But another surge of evolutionary activity was yet to* occur, primarily within the coastal popu- lation (Fig. 21). With the final retreat of the Puget and Juan de Fuca lobes of the Cordilleran ice sheet (late Vashon Stade) about 13,500 years before present (Armstrong, et a l , 1965), coastal Cottus asper began to move northward along the newly exposed coastline of British Columbia. With the more or less contemporaneous retreat of the Okanagan and Spokane lobes from the Columbia R. basin, inland Cottus asper from the lower Columbia advanced northward into Canada and, perhaps at that time, colonized the interior Fraser R. system through meltwater channels then in existence. Coastal Cottus asper now exists in three phenotypic forms in areas of the Pacific Slope formerly covered by the Cordilleran ice sheet: 1) The typically prickled*.estuarine spawning , coastal form with lower pectoral fin ray counts; 2) a typically prickled, fresh water spawning derivative form with high pectoral fin ray counts, found far inland in the Stikine, Nass, Peace, and parts of the Skeena R. drainages; and 3) a very weakly-prickled, fresh water spawning derivative form with high pectoral fin ray counts* found in certain mainland and is- land lakes. These "coastal derivative" forms probably followed the recent retreat of glacial lobes up narrow valleys in northern British Columbia. These forms are necessarily the most recent invaders, arriving, perhaps, even later than the upper Fraser R. forms of the inland Cottus asper from which 80 Figure 21„—Phylogeny of Cottus asper. Stage IIIo Evolution of Coastal Form Following Retreat of Cordilleran Glaciers During Recent Time, 81 they are distinct. Lindsey (1956) presented distributional evidence that the following primary fresh water species: Catostomus macrocheilus; Mylocheilus caurinum; Ptychocheilus oregonense; and Richardsonius balteatus; plus Cottus asper, a euryhaline species, had penetrated into the Peace R. system from the Skeena or Fraser R. system. " A l l are present in the Columbia, Fraser, and Skeena systems, a l l are absent from the Yukon, and a l l have been taken in the Mackenzie system only from the upper portion of the Peace drainage." The restricted range of these species in the Peace R. system indicates their relatively recent a r r i v a l . The present study indicates that the Peace R, representatives of Cottus asper are coastal derivatives which must have penetrated inland via the Stikine, Nass, or upper Skeena R. The inland form of Cottus asper has apparently failed to penetrate northward beyond the limits of the upper Fraser R, drainage. The exchange probably occurred and the connection be- came closed before the inland Cottus asper arrived on the scene. Evidence from the distribution of other primary freshwater fishes bears this out. Several species got only as far north as the inland asper. For example, the leopard dace, Rhinichthys falcatus, i s a Columbia R, form which has pene-. trated northward (presumably following the same routes as inland asper) as far as Middle River near Takla L. ,• about 120 miles northwest of Prince George, B.C. The bridgelip sucker, Catostomus columbianus, another Columbia R. form, has been taken in the Salmon-R, system just north of Prince George. Extensive collecting in the areas north of Prince George indicates that both species have failed to penetrate into the Peace or the Skeena R, systems. The prickly sculpin, Cottus asper, is interpreted as being a polytypic species which has become, and i s in the process of becoming, modified into several characteristic genotypes. The dichotomy between "inland" and "coastal" forms i s basically ? subspecific in nature. Classification of the 82 "asper species group" is beyond the limit of a system of binomial nomen- clature « Modifications in some of the earlier derivatives of the inland form have presumably become genetically fixed, and various authorities have assigned them ful l specific rank. More recently derived forms of the coastal Cottus asper , such as occur in the coastal lakes of British Columbia and Alaska, or in the British Columbia river drainage systems north of the Fraser R,, are modified sufficiently to perhaps warrant classification as subspecies, but the genetic nature of the modification has not been deter- mined. It is not known Whether the marked differences i n phenotype of the "coastal derivative" or*coastal lake forms are results of a phenomenon which occurs in one generation, or over a long period of time. If the changes are not environmentally induced, then genetic fixation can be assumed to have occurred within the past 10,000 years or less. 83 LITERATURE CITED Armstrong, E„, D„ R, Crandell, D, J , Easterbrook, and J , B, Noble, 1965, Late Pleistocene stratigraphy and chronology in southwestern British Columbia and northwestern Washington, Geol, Soc, Amer, Bull, 76:321-330, Bailey, R. M., and M. F, Dimick, 1949, Cottus hubbsi, a new cottid f i s h from the Columbia River system in Washington and Idaho, Occ. Pap, Mus, Zool, Univ. Mich. 513s1=18, Bailey, R. M., and C. E, Bond, 1963, Four new species of freshwater sculpins, genus Cottus, from western North America, Occ, Pap, Mus, Zool, . Univ, Mich, 634:1-27, Battle, H. I. 1929. 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Sci, Cong, 1967 2 54-58, 1962, Experimental study of meristic variation in a population of threespine sticklebacks, Gasterosteus aculeatus, Canad, J, Zool, 40(1962) % 272=312, Lord, J, K, 1866a, The naturalist in Vancouver Island and British Columbia, Richard Bentley, London, 1866, 121=358. 0 1866b, The naturalist in Vancouver Island and British Columbia, Richard Bentley, London, 1866, 2 21=375. McAllister, D, E. 1957, The systematics of the freshwater sculpins of British Columbia, M,A. Thesis, Univ, British Columbia Depto Zool, A p r i l , 1957sl=109, and C, C, Lindsey, 1959, Systematics of the freshwater sculpins (Cottus) of British Columbia, Nat, Mus, Can, Cont, Zool,, Bull. 172266-89, «,«..„«.0 i960. List of the marine fishes of Canada, Nat, Mus, Can, Bull. 168, Biol, Ser, 62 21=76. Meek, S„ E„ 1899. Notes on a collection of cold=blooded vertebrates from the Olympic Mountains. Field Mus. Nat, Hist, Publ. 31, Zool, Ser, (1897)1 (12)2225=236, Mottley, C. McC. 1934, The effect of temperature during the devedopment on the numbers of scales in the Kamloops trout, Salroo kamloops Jord. Contr, Canad, Biol,, N„ S. 8s254=263, Nichols, J. T. A small collection of Alaskan fishes, Proc, Biol, Soc, Wash, 21sl71-174, Pritchard, A, L. 1936, Stomach content analysis of fishes preying upon the young of Pacific Salmon during the fry migration at McClinton Creek, Masset Inlet, British Columbia. Canad. Field-Naturalist 50(6)2 104-105, Richardson, Sir J, 1836, Fauna Boreali-Americana, Part 3, The f i s h . Richard Bentley, London, Part 3 21-327, 88 =„ 1844, Ichthyology 0 Ins Voyage of the H, M. S, "Sulphur", under the command of Captain Sir Edward Belcher 9 during the years 1836=42, (R, B, Hinds, ed,), Not seen, Robeck, G, G,, C, Henderson, and R, C, Palange, 19540 Water quality Studies on the Columbia River, U, S, Dept. HEW, Pub, Health Svc,, R, A, Taft San. Engr. Center, Cincinatti 1954sl=99 plus 4 Appendices, Robins, C, R, and R. R, Miller, 1957, Classification, variation and dis- tribution of the sculpins, genus Cottus, inhabiting the Pacific Slope waters in California and southern Oregon, with a key to the species, Calif. Fish and Game 43(3)s213=233, Rutter, C, 1908, The fishes of the Sacramento=SanJoaquin Basin, with a study of their distribution and variation. Bull, U, S, Bur. Fish, 27(for 1907)sl03=152. Schultz, L o P. 1929, Check l i s t of the fresh=water fishes of Oregon and Washington, Publ. Fish,, Univ. Wash, Coll, Fish. 2(4)s43-50, -—-—, 1930, Notes on the species of Cottus in western Washington, Copeia 1930(1);14-15. ' and A. C, DeLacy, 1936a, Fishes of the American Northwest, A catalogue of the fishes of Washington and Oregon, with distributional records and a bibliography. Installment 3, Mid-Pacific Mag, (April-June) 1936sl27-142. ..._.( and =, 1936b, Fishes of the American Northwest. Installment 4, Mid-Pacific Mag, (July-Sept,) 1936S211-226. 1936. Keys to the fishes of Washington, Oregon and closely adjoining regions. Univ. Wash, Publ, Biol. 2(4)sl03=228. Seale, A. 1896. List of fresh-water fishes collected in the vici n i t y of Seattle, Washington, by Edwin C. Starks, Proc, Calif, Acad, Sci, (2)5 (for 1895) part 2s852-854. Seymour, A. 1959, Effects of temperature upon the formation of vertebrae and f i n rays in young chinook salmon. Trans, Am. Fish. Soc. 88J58-69, Shapovalov, L, and W. A, D i l l , 1950, A check l i s t of the fresh-water and anadromous fishes of California, Calif, Fish and Game 36(4)$382-391, -—-«-•, and A, C, Taft, 1954. The l i f e histories of the steel head rainbow trout (Salmo gairdneri gairdneri) and silver salmon (Onchorhychus kisutch), Calif. Dept. Fish and Game, Fish Bull. 98s1=375, - g w. A, D i l l , and A, J, Cordone, 1959, A revised check l i s t of the freshwater and anadromous fishes of California, Calif, Fish and Game H .. 45(3)sl59-180, Snyder, J. 0, 1905, Notes on the fishes of the streams flowing into San Francisco Bay. Rep, Bur. Fish, (for 1904), App. 5s327-338, 89 1908a, The fauna of the Russian River9 California, and its re-lation to that of the Sacramento, Science, N , S0 27s269-271, „ 1908b, The fishes of the coastal streams of Oregon and northern California, Bull, U. S, Bur. Fish, 27(for 1907).153-189. — — . .1913, The fishes of the streams tributary to Monterey Bay, California. Bull, U, S„ Bur. Fish, 32(for 1912)s47-72, 1916. Fishes of the streams tributary to Tomales Bay, Bull, U, S. Bur, Fish, 34(for 1914).375-381, Storer, D. H, 1846a, A synopsis of the fishes of North America, Mem, Amer. Acad. Arts and Sci., N, S, 2:253-550, —, 1846b, A synopsis of the fishes of North America, Metcalf and Co., Cambridge, 1846s1-298, Strawn, K, 1961. A comparison of meristic means and variances of wild and laboratory-raised samples of the fishes, Etheostoma grahami and E. lepidum (Percidae). Texas J. Sci. 13(2)sl27-159. ~° Suckley, G, 1859, Report upon the fishes collected on the Survey, Chap, lis Report upon the fishes exclusive of the Salmonidaes 350-368, Ins The natural history of Washington Territory, ^Cooper, J . G,, and Suckley, G, Baillere Brothers, New York, Part 111:399, 1960. Report upon the fishes collected on the Survey, Chap 2s Report upon the fishes exclusive of the Salmonidaw, U, S, Pac, R, R, Expl. and Surv,, (for 1853-55), 12(2) part 3 (5):350-368, Also appears as: H. Rep. Ex, Doc, 56, Wash., 1860:350-368. Sumner, F, H, 1942, A reconnaissance survey of Oregon coastal streams. Rep. Oregon St, Game Comm., 1-25 (mimeo). In_2 Bond, C, E, 1963:81. — — • . 1953. Migrations of salmonids in Sand Creek, Oregon, Trans, Am. Fish, Soc. 83(1952).139-150. Taft, A, C. 1943 California steelhead experiments. Trans, Am, Fish, Soc, 64:248-251, Taning, A, V, 1944, Experiments on meristic and other characters in fishes, I, Medd, Komm. Dan. Fisk. Havunders, ll(3)sl-66, , 1952, Experimental study of meristic characters in fishes, Biol, Rev, Cambridge Phil. Soc. 27:169-193, Wilimoysky, N. J. 1954. List of the fishes of Alaska, Stan, Ich, Bull, 4:279-294, — , 1958. Provisional keys to the fishes of Alaska, U, S, Fish and Wildlife Svc,, Fish, Res, Lab., Juneau. 1958:1-113. Appendix Table 1 0 List of Collections, Localities, and Other Pertinent Information. Pop, No. General Locale Collection No0 Specific Locality .60 mm S.L, :60 mm S.L, NoLato W.Long, COASTAL POPULATIONS Alaska "head of Chilkoot L," Lynn Canal, Favorite Channel:. Petersen Creek, 12 mi, N, Seward area: BC 64-311 NMC 61-133 Haines area: USNM 76408 Juneau area, AB 61-100 AB 61-489 " v.-,..-.-" " AB 61-490 " .. " . Loring area, Revillagigedo Id. s USNM 64130-49 Loring USNM 60801 Deep Bay USNM 60371 Mouth of River USNM 60316 Naha River Kasaan Bay, Prince of Wales Id. "ditch at airport" "lagoon at north edge of Seward" 1(56, 9(28, 0) 2-38,6) of Juneau 2(53,9-59,5) British 6. 7, Stream of Karta Bay Saltery -Cr, j Paul's Bight, Skowl Arm Saltery Cr, Saltery Cr,, 100 yds, up from mouth Is. . Graham Id.. Juskatla Inlet of Juskatla I I I I 9. 10, SU 23989 AB'62-76 AB 62-77 AB 63-167 Columbia Queen Charlotte BC 60-437 BC 60-439 BC 60-440 Queen Charlotte BC 60-420 BC 60-424 Princess Royal Id.: 1 BC 53-214 Bloomfield L,, N.E, side, Laredo Inlet King Id., Ocean Falls area: BC 54-434* Port John L. Rivers Inlet area: BC 56-629 Owikeno L.„ 2nd Narrows* Small stream about 2 mi. Mouth, of stream 3 mi, W, I t ; ... I I I I .'. I t I t I I Is, , Moresby Id, : Creek-entering Skidegate L, Shore of Skidegate L, W, of Juskatla 10(28,5-56,6) 59(26.4-59,6) . 38(32.7-59.5) 23(25, 12(25, 4(63 8( 62,9-125,0) 8( 73.2« 2( 70,1- 8( 78.0- 14(112,8= 2( 87,5= 3(115.4= 2( 68,9- 2(111.6- 11( 83.9< 2(149,5« 3( 94,3= 119.2) • 77.8) •142.1) •166.2) •f,'94.0) •135.0) • 76,0) •116.7) •139.0) •154.5) 112.1) ff-5?.&) 3-59,1) ,1-70,8?* 60°07' 149°24» 59°23» 58°22' 9( 60.2- 90.2) 11( 62,2-136,2) 6( 61,5- 83.3) 20( 61,0-132.2) 6( 62̂ .3- 98.5) 2(122.0-123,5) 16( 72.4- 85.7^* 10(none listed) 135°35« 134°36« 55°35' 131°40« 55°00« 35' 25' 53°00« 37' 36« 132°00e 35' 30* 132°00» 23* 25° 53°06» 132°00' 52°57' 52°06' 51°40' 128°50« 127°45' 126°40' *Counts taken by D.E. McAllister; **Lengths given are Total Lengths, Appendix Table 1, List of Collections, etc. (cont'd). 11. 12, 13. .14. 15, 16. 17. Vancouver Id BC 54-114 BC 64-312 BC 64-313 BC 64-314 Vancouver Id BC 54-39 Vancouver Id K61-13 Vancouver Id BC 63-1462 Gambler Id,, BC 58-410 White Rock area, Campbell R. area: Buttle L, Buttle L, Buttle L„ Upper Campbell L. Qualicum area: Horne L, Qualicuin area: Nile Cr. , 100 paces from mouth South ends Nitinat L, Howe Sounds Outlet Cr. 100 ft, from mouth Little Campbell R, K 60-52 Stn, C-22 K 60-40 I I K 60-71 n K 60-88 I I K 60-128 I I K 60-169 I I K 60.-27 I I c- 2 K 60-12 tt K 63-6 it K 60.-80 I I c-•3 K 60-101 ti K 60-116 it K 60-139 I I Saturna Id, BC 56-81 Strait of Georgia; Lye 11 Harbor Cr, 19. 20. 21. Clatsop Co. , UMMZ 93427 Cannon Beach areas Mouth of Elk Cr, Coos Co. 9 Lakeside, Ten Mile Lake outlet wier BC 63-1089 Ten Mile L. Curry Co.j Sixes areas SU 9270 Flores R„ Curry Co,, Winchuck Area, Winchuck R,s OSC 198 2 mi, upstream from U,S, Hwy 101 1(42.5) K 106.5) 49°47' 125°40' 2( 71.0- 76.0) 2( ca75-117.5) 1( 132.7 ) 49°55' 125°40' 49°20' 124°40' 11(108.1-132,0) 49°25' 124°40' 5(49,4-59.6) 17 ( 61,5- 91.0) 48°40' 124°50' 14(39,9-59.5) IK 63,0- 89.2) 49°25' 123°23' 17(23,0-59,9) 16 ( 60.5-129,3) 49°— 122°— 2( 89.7- 90,2) " 04' ii 1+0' 10( 77.5-107.7) 2( 92,8-127.5) 3( 93.0-102, 4) 2( 80,7-111,0) 2( 80,0- 89,5) 9( 60,6- 71, 8) " 00' " 46' 1( 39,5) 3( 64.0- 88.5) 6( 81,5-118,5) 33(13,0-58,5) 2( 70,1- 71.6) " 01' « 45' 5(55.0-58,6) 9( 60.5-97,3) 5(51,6-59,5) 7( 60,4- 73.3) 2(57,5-58,5) 9( 63,5- 83.2) 10(28,4-39,8)** 48°48' 123°10' 45°54' 123°58' 10(21.7-59,4) 5( 68,4-100.2 ) 43°35« 123°10' 33( 60.3-114,1) 42°55» 124°30« 3(44,2-55,3) 9( 77,0-154,0) 42°00« 124°15« 6(36,3-50.9) Appendix Table 1, List of Collections 9 e t c (cont'd),, California 22, ife Del Norte Co, 9 Crescent City.areas 41°50« 124°15« SU 23598 Point St. George K 62,3) I I it I I it CAS 18025 Wilson Cr., at bridge 1-05, U,S„ 101 K 84,7) " 55 • it I I CAS 18028 Smith R., at bridge on U,S, 199 3(31,7-397) 23, Del Norte Co.s 41°34e 124°10« SU 9272 Mouth of Klamath R,, at Requa 14(33,3=59,8) 16( 62,0=115,0) 24a, Mendocino Co,, Navarro area, Navarro R,s CAS 19066 Indian Cr,, bridge on Hwy, 128, Philo SU 35131=3 N, fork, o.2 mi, E, of Flynn Cr, SU 38016 Albion R., 2 mi. below Comptche 1(49,6) 1( 3( 63.0 ) 71.6=106.5) 39°09e 123°348 25a. Marin Co., Tomales Bay drain,, Lagunitas (Papermill) Crtt CAS 186I6A S,P„ Taylor St, Park, at Camp Taylor SU 40849 Near Point Reyes Station 6(22,7-44,4) 1( 93,0) 38°00e 122°45» 26. Santa•Cruz Co,, Waddell Cr.s CAS 20857 3 mi. S.W, of Ano Nuevo Point CAS 20862 " " " " " " " 24( 14( 96,0=143,2) 95,8=143.1) 37°069 122°17» 27, Santa Cruz Co., So quel areas CAS 19086 Soquel Cr,, at Soquel SU 4816 Soquel Cr, 1(54,7) 4( 64.9=105.2) 36°57« 12£°55« 28. San Luis Obispo Co., San Simeon areas I35°35» 121°128 SU 15183 San Simeon Cr, 4(22,9=42,5) 2(124,9=127,2) 29, Santa Barbara Co., Lompoc areas SU 40323 Santa Ynez R, 2( 76,8= 86.3) 34°41' 120? 35' 30. Santa Barbara Co,, Gaviotas UMMZ 131728 Gaviota Cr., at mouth 15(21,7-47.4) 34°24» 120°15' 31. Ventura Co., Ventura R*s 34°10' 119°20« UMMZ 13173ft 6.4 mi. N.E. of Ventura 1(113,5 ) UMMZ 132891 1/2 mi. from Foster Park, towards Ventura 2(119,0=151.1) Appendix Table 1. List of Collections, e t C o (cont'd),, COASTAL DERIVATIVE POPULATIONS British Columbia 32. Stikine R„ drainage s 56°409 128°40« BC 54=485 Cold Fish Lake BC 54=485 Cold Fish Lake* 2(77,7-^82,5) 3(48,6-66,7)^ \Wi 1(81,7 >*% ) 33. Nass R. drainage, Meziadin Lakes BC 55=221 At outlet BC 55=221=A At outlet BC 56=536 Off McLeod Creek 8(81,0=109.1) 5(66,2- 82.-0) 11(73,2= 87,2) 55°03" 129°10» 34. Skeena R, drainage, Babine Lake s BC 55=452 Wright Bay BC 55=453 Opposite Donald's Landing BC 56=455 At Topley Landing 9(15.8=34.4) 6(34.2=54.3) 1(60,6 ) 3(61,1= 96,9) 54°509 126°0=05« 35. Fraser R. drainage, Takla Lake outlets BC 54=249 Middle R. 12(27.4-35.2) 4(65,5= 83,5) 54°558 125°058 36. Mackenzie R. drainage. Nation Lakes areas BC 64=102 Tchentlo L. inlet stream mouth BC 64=101 Tchentlo L, in l e t end 3(41.2=54.7) 3(33.1-47.0) 55°12 9 125°005 37. Mackenzie R. drainage, Mansori Creek areas BC«56£458 Outlet of Wolverine L, 13(31.0=58,0) 11(65,0=133,3) 55°37 9 124°259 38,. Mackenzie R0 drainage, Parsnip R, watersheds BC-61=464 Tacheeda Lake 13(34,0=58.5) 2(65,6-666,2) 54°42 B 122°30° 39. Mackenzie R. , Crooked R. watersheds BC 54-477 McLeod Lake c 15(13,8-23,0) 55°009 123°009 40. Mackenzie R., Crooked R„ watersheds S-98 Bear L. N. of Summit L. 3(48,5=59,9) 4(61,5=141,0) 54 028 9 122°479 41. Mackenzie R,, Crooked R« watersheds BC 61-463 Hart L. off Hart Highway 2(79,5- 80.1) 42. Mackenzie R. , Crooked R, watersheds BC 56-362 Summit L, , N. of Prince George BC 54-459 Summit L., N„ of Prince George 1(25.9 ) 11(10.7=50,8) 54°159 122°40' 43. Dean R. drainage, Chilcotin areas BC 56-483 Nimpo L„ 15(12,5-34,4) 52°20« 125°109 Appendix Table l n List of Collections, etc. (cont'd),, INLAND POPULATIONS British Columbia Skeena R. Drainages . 44, ' 45„ Fraser 46. 47. 43. Fraser , 49. 50. Fraser 51„ 52, 53. 54, BC 56- BC 56- •411 #53 Lakelse R. , at Terrace Bulkley R0, 1 mi. below Bulkley R„, Nechako R. drainages BC 55-10 BC 54-162 BC 54-367 Thompson R. BC 54-242 BC 55-139 Fraser L., 2 mi Eulatazella L„ e Nadsilnich L., drainages Kamloops L. Pothole L from West end Deep Cr. St. Joseph's Cr. Fraser Valley drainages K mi s c, Squak urn7 L t outlet (Quilchena Cr.) outlet stream I I t i I I . I I ._ I I I I i t I t t i . I I K 62-19 i t r~ I I . t l . r I I BC 54-197 Squakum L, BC 54-196 I t BC 54-195 i i ... I I V-100 Vedder R. Stn. V-6, at end of Ford Rd, V-150 I I t i I I . . . . i t i t i t t t i t I I . V-203 n i t v-4, Vedder R, Canal BC 59-612 Vedder Canal, above highway bridge BC 54-43 Vedder R, BC 54-427 I I BC 59-504 i t BC 62-70 Cultus L. K misc. Cultus L. , outlet wier in Sweltzer Cr, K 63-16 I I t t - t i I I i t i t t t K 63-17 i t t i t t -511 I I I t t t SW-1 Sweltzer Cr., 1/4 mi, below Cultus L, BC 56-564 Hatzic L., Hatzic Slough BC 54-212 t t t i Swan Point BC 54-209 I I t t BC 56-120 I I i t Edwards Or, BC 55-313 i t t t " " mouth K misc. I I i t t i t t 14(40,8-55,9) 3(34,4-50,4) 15(16,8-55,6) 6(43,5-57,9) 9(53,6=67,0) 10(14,6-25,7) 12(25,3-58.Q) 11(35.0-59,0) 1(47.6 ) 2(33,5=43,0) 15(37.6-58,6) ft* 13(17,6-51,0) 6(24,2-32,2) 1(20,9 ) 13(32,2-50.5T 15( 60,5=153,6) 54°21« 128°35c 9( 67,1=130,7) 54°28( 126°30' l(468 !il ) 54°05' 124°50« 15( 62,2=105.5) 53°42' 123940' 53°45" 122°50» 50°45« 120°40' 10( 60,0- 88,7) 49°55» 120°30» 7( 73,8=104,8 9( 68,4=111,9 1( 68,8 7( 73,9-119,0 5 ( 1 0 3 3 ( 1 1 3 U3( 6 1 1 ( 97 2 ( 1 1 6 3 ( 1 1 3 3Q.09 2 ( 1 0 6 1 ( 1 0 4 5 ( 6 2 5 ( 3 ( 2 ( 2 ( 4 ( 9 ( 74, 72, 74, 70, 64, 75, 118,5 •143,8 •100,1 •129,0 •148,2 128,1 •140,0 •105,7 > 96,5 • 77,0 • 94,3 •116,0 • 70,1 •112.9 •130,2 3(104.5- 1(138,2 2( 66,7- 68.0 49°14' 122°00' 49°06' 122°008 49°04' 121°5-9» 49°10» 122°14' -F Appendix Table 1, List of Collections, etc, (cont'd). 55, Columbia R0 56, Columbia R„ 57„ Columbia R, 58, Washington- Columbia R„ 59 A-51 A-122 A-153 A-220 K 63-12 drainage, BC 54-369 BC 5 9-424 drainage, BC 54-124 BC 56-534 , Okanagan BC 53-187 Oregon S„ Alouette R, it it ti Stn, A-8, 14th Ave, A-6, 5th Ave, ti it tt tt it tt ti ti it Revel stoke areai Williamson L, A-7, tt ti 8th Ave, ti ti tt it off diving float Grand Forks area Christina L, ti it R„ drainage s Osoyoos L, 1 mio from S, end Columbia R, 61, drainage s USNM 104608 UW 14614 UW 14747 OSC misc, OSC " • OSC " CAS ( I o U c M , drainage, OSC misc. Moses L o , Grant Co,, Wash, McNary Dam, Benton Co,, Wash, Umatilla R, tt II mouth, Umatilla Co,, it II tt Ore, ti Portland and Vicinity? Abernathy Cr,, at mouth, Cowlitz Co, Kalama R 0 a 1/4 mi, above mouth Coweman R ti tt at mouth 100 yds, upstream " " Lewis R, j E.fork at U,S,'99, Clark Co, below U,S, 99 bridge it Washougal R, , 1 mi, above mouth " " " " , below old Hwy 30 bridge " Vista Slough, Big Rock, Multnomah Co, Sandy R, below old Hwy 30 bridge " Sandy R» Clackamas R„, near mouth, Clackamas Co, 3(30,0-54,2) 13(15,0-59,6) 14(27,8-39,8) 3(34,4-42,6) 11(19,8-34,4) 23(36,3-59*9) 3( 73,5- 82,6) 2( 95,4-103,4) 3(100,4-123,0) 5( 64,5-107,7) 8( 65,5- 97.0) 13( 86,0-123,6) 1( 73,3 ) 6( 60,5- 79.8) 49°14' 122°35 50°59' 118°11 49°09' 118°11 49°11' 119°30 20(18,8-26, 1(59,5 1(58,9 6) ) ) 3( 70,4-161,5 4( 76,8-100,3 2(125,4-169,2 1(149,5 3( 93,8-186,0 1( 62,2 2(118,8-119,5 1(113,5 1(105,6 3( 88,0-172,0 4(112,5-164,5 1(121,7 3( 94,6-106,0 1(146,5 1( 97,2 2(110.9-121.5 1( 91,4 1(175.0 47°06» 119°19' 45055" 119°20l 45°209-122°10( 46°028 122°52' Appendix Table 1 , List of Collections, etc, (cont'd), Columbia R o mouth, Washington Shore, Pacific Co,: 6 2 , Columbia 6 3 o Willamette 6 4 o UMMZ 9 3 4 2 5 Columbia R0 at end of Chinook Pier OSC misco Chinook R. OSC misCo Chinook R,, above tide gate SU 3 8 0 1 9 Columbia R, , 1 / 2 mi. W. of Megler Ferry 1 ( 2 4 , 6 ) , mouth, Oregon Shore, Clatsop Co,s SU 3 8 0 1 8 Columbia R, at Astoria SU 3 8 0 1 7 " " " " OSC 7 6 2 Walluski R., 6 mi, S.E. of Astoria OSC misco Big Creek, at mouth Ro drainage, Corvallis areas OSC 1 8 9 OSC misc. OSC 4 0 2 OSC 5 7 0 2 OSC misc. OSC misco Willamette R, drainage, 6 5 . OSC 7 4 4 California Sacramento R. drainage, 66a, CAS 1 8 0 8 1 UMMZ 9 3 4 2 8 6 6 b , USNM 2 9 5 Sacramento R, drainage, 6 7 , UMMZ 9 3 4 2 9 UMMZ 9 3 4 3 0 UMMZ 9 3 4 3 1 Sacramento R, drainage, 6 8 o OSC m£sc. Overflow slough at Corvallis Corvallis Sand 6 Gravel Pits, along R, Pothole, Willamette R, at Corvallis Stn, 5 , above Corvallis Stn, 6 , Harrisburg Stn. 7 , Peoria S o E „ of Eugene, Lane Co,; , Lookout Point Reservoir, Landax Shasta Co,s O'Brien Cr,,. emptying in Shasta L, Clear Cr„, a few mi, S, of Redding Fort Reading (on Cow Cr») Tehama and Glenn Co,s Sacramento R,, v a l l i t t l e below Red Bluff at Richfield ' Thomas Cr. . Stony Cr 0, near Orland Sacramento area s Sacramento Brickyard Pond, S, of city San Joaquin R, d r a i n a g e , San Joaquin Go,; 6 9 , CAS 2 0 8 8 7 San Joaquin R,(exact l o c a l i t y unknown) San Joaquin R, drainage, Stanislaus Co.; 7 0 , CAS 1 7 9 6 6 Irrigation ditch on Hwy 1 3 2 , 1 mi, E, CAS 1 9 0 9 8 of Empire. " " " " " " " San Joaquin R, drainage, Madera1and Feesno Co.; 7 1 , CAS 2 0 2 5 2 Madera Canal, presumably from Millerton 1 ( 5 3 , 4 ) 1 ( 3 4 , 5 ) 6 ( 9 7 , 5 - 1 5 0 , 6 ) 2 ( 1 1 6 , 5 - 1 1 8 , 0 ) 2 ( 8 6 , 5 - 1 2 8 , 2 ) 2 ( 6 0 , 1 - 1 1 2 , 7 ) 7 ( 1 2 7 , 5 - 1 5 4 , 0 ) 1 1 ( 1 0 0 , 8 = 1 3 1 , 5 ) 2 ( 8 7 , 8 = 8 9 , 8 ) 4 ( 1 1 9 , 3 - 1 3 4 , 5 ) 2 ( 1 5 0 , 3 = 1 5 0 , 7 ) 2 ( 1 2 3 , 5 - 1 3 3 , 5 ) K 7 3 , 3 ) 2 ( 6 6 . 8 = 7 4 . 6 ) 4 6 ° 1 7 « 1 2 3 ° 5 7 e 4 6 ° 1 1 ' 1 2 3 ° 5 0 s 4 4 ° 1 5 = 1 2 3 ° 1 0 = 3 0 " 1 5 " 1 1 ( 1 7 , 5 - 5 6 , 1 ) 1 0 ( 6 5 , 0 = 8 4 , 9 ) 2 ( 2 8 , 5 = 3 1 , 0 ) 1 ( 7 8 , 4 ) 4 3 ° 4 5 s 1 2 2 ° 4 0 ' 4 0 ° 4 8 8 1 2 2 ° 1 6 ! I I 3 0 « I I it 3( $6*7=105,2) ti 2 8 ' " 11*' 1( 69,7- ) i40°09' 122°13' 1( 99,0 ) 3 9 o 5 6 » i22°13' 1( 71,8 ) 3 9 0 4 5 s 122°139 2( 65,5-100,7) 38°30' 121°30' 10(31,5-56,5) 4 ( 6 1 , 0 - 8 4 , 0 ) ^ 3 7 0^59 121020' 37°35» 120°55' OSC misc. San Joaquin R, at Mendota 4 ( 2 7 , 5 - 4 4 , 9 ) 3 ( 2 5 , 0 - 5 0 . 5 ) 1 ( 3 0 , 0 1(120,9 )ca37°00' 1 1 9 ° 5 0 ' K 7 2 , 1 ) 37040* 120°20' to cn Appendix Table 1, List of Collections, e t c (cont'd),, San Joaquin R. drainage B Fresno and Tulare Co.s 72. CAS 25541 Kings R. , at Dutch John Cut CAS 20904 Kaweah R., near Vis a l i a INTERGRADE POPULATIONS Sacramento R. drainage. Contra Costa Co., Carquinez Straits 73. UMMZ 142373 ca 1/2 mi.E, of Eckley, S. of Dillon Pt San Pablo Bay area, Sonoma Co., Californias 74. USNM 296 Petaluma Additional Material From the Literature, Used Separately, or in Combination, with Populations Listed Above Sacramento R. drainages 75. Rutter, 1908sl45 Sacramento R. at Redding " " mouth of Battle Cr. " r " at Red Bluff " " at Jacinto • ••" " Arcade Cr., at Arcade Feather R, at Marysville 1(46.5 ) 3(34.5-58.3) 8(35.0-57.5) 20(60.0-183.4) 2(30.9-39.5) 2(71.8-109,5) 2 specimens 1 " 1 1 9 " 1 " San Francisco Bay, San Mateo Co.s 76. Snyder, 1905s338 San Francisquito Cr, Santa Cruz Co,, Watsonville areas 77. Snyder, 1913s72 Pajaro R„ Marin Co., Tomales Bay areas 25b, Snyder, 1908b$185 Papermill Gr. Mendocino Co., Point Arena areas 78. Snyder, 1908bsl85 Garcia R. Mendocino Co,, Navarro areas 24b. Snyder, 1908bsi85 Navarro R. Humboldt Co. , Areata areas 79. Snyder, 1908bs185 Mad:R„ Curry Co. Oregon, Gold Beach areas 80. Snyder, 1908bsl85 Rogue R,, near mouth Douglas Co., Reedsport areas 81. Snyder, 1908bsl85 Takenitch R. Tillamook Co., Nestucca Bay areas 82. Snyder, 1908bsl85 Nestucca R, King Co., Washington, Seattle areas 83. Snyder, 1908bsl85 Lake Washington 34 specimens 25 specimens 10 specimens 6 specimens 11 specimens 6 specimens 8 specimens 16 specimens 10 specimens 17 specimens ? i 36°15' 119°16" 38°00' 122°10 38°ir 122°40 40°30» 122°10 40°21» 122°09 40°09' 122°13 Additional Materials UMMZ 131771, Kern R,, California 1 specimen (not used) 39?059 121°35 37°24' 122'°0'9f aeoSO* i21°50 38°00* i22 u45 38°55» 123?45 39°09» 123°34 4Q°55' 124°08 42°26' 124°23 43°45' 124°07' 45°10« 123°57' 47°30» 122°15' 35°-- i i g o . _ Appendix Table 2. Weighted Mean Percentage Prickling and Resorption for Individual Populations of Cottus aSper Used in Compiling fable I and Eigure 4. Population No. Specimens £i60 mm S.L. Specimens S =60 mm S.L. % Resorption Coastal n Virtual Apparent n Virtual Apparent Relative Absolute 1 1* 37.9 34.2 „_ — — — 2 — 8 34.1 17,1 49,9 17.0 3 2 34.6 17.8 18 33.6 14.0 58.2 19.6 4 — 9 34.̂ 3 20,2 41.1 14.1 5 — 16 40.8 24,7 39.5 16.1 6 11 42.0 40.3 9 40,5 21,1 48.0 19.4 7 12 32.1 20.5 18 26.2 11,7 55,3 14.5 11 5 13.7 3.5 74,6 10.2 12 11 27.1 10,3 62.1 16.8 13 5 41.1 24.4 17 44.5 21,8 51.1 22.7 14 14 49.3 42.9 11 43.4 27.3 37.0 16.1 15 17 54.0 51.0 16, 43.0 24.8 42.3 18.2 16 16 44.2 41.4 29 40.3 27.5 31.8 12.8 18 10 49.7 41.6 5 31.0 15.4 50.1 15.6 19 — -«. 33 47.0 34.0 27,7 13.0 20-21 9 30.6 9 1* 16.5 — 25a 6 41.7 — ;j 24,2 — — 26 24 30,4 12.9 57,5 17.5 28 6 35.4 2 14,4 — —• - -29 2 -rs i7.6 — 30 15 44.8 43.8 — --?* - - — — 31 3 29.5 13.0 55.8 16.5 X 43.5 36.0 ~ 35.0 17.8 48.9 17.2 Coastal Derivative 32 2 15.0 8.9 40.3 6.1 33 — 37.8 25.4 32.9 12.4 34 5 49.5 48.6 1 35.4 22.3 37.0 13.1 35 12 49.0 46,2 4 43.8 38.6 11.8 5.2 36 6 42.4 38.4 — — — — 37 13 42.9 40.5 11 34.8 23.4 32, 8 11,4 38 13 34.5 29.1 2* 36.1 17.2 52.4 18.9 40 6 50.8 48,3 1 49.3 27.7 43.8 21.6 41 — — 2 40.5 31.6 21.8 8.9 X — 44 , 8 41,8 — 34.7 24.2 32.0 10,5 Appendix Table 2 (cont'd). Weighted Mean Percentage Prickling and Resorption for Individual Populations of Cottus asper Used in Compiling Table I and Figure 4, Population No, Specimens <C 60 mm S, L, Specimens I> 60 mm S.L, % Resorption Inland n Virtual Apparent n Virtual Apparent Relative Absolute 44 15 67,7 64, 8 14 64,1 53.9 16,0 10,2 45 3 69,3 65,8 9* 65,0 44,2 32,0 20, 8 46 4 65,1 61,6 1* 62,6 60,4 3,6 2,2 47 6 65,4 60, 7 15 59,9 54,3 9,4 5.2 50 12 61,4 56,4 10 59,6 49,9 16.3 9.7 51 11 65,4 60,6 31 55,3 46.1 16,6 9.2 52 15 56,4 53,5 25 59,9 52,6 12.1 7.3 53 13 63,1 61.0 15 58,3 49.7 14.7 8,6 54 20 77,6 73,3 21 73.5 68,0 7,5 5.5 55 12 69,8 67,0 20 67,4 53,5 20,7 13,9 56 14 52,3 51,1 13 51,0 45,0 11.8 6,0 57 8 66,9 66,3 1* 61,3 60,3 1,7 1,0 58 24 66,5 65.1 6 67,1 60,3 10,1 6.8 60 1* 84,2 81,2 7 69,5 57,8 16.9 11,7 61 == • — 15 67,3 55,8 17,1 11,5 62 2 67.0 6 58, 8 42,1 28,4 16,7 63 ™ 11 60,7 56,4 7,1 4,3 64 2 77,4 7 57,0 = = 65 11 81,9 80,6 10 76,7 65.8 14.2 10,9 66=68 2 74,5 74,5 6 63, 3 46,2 26.9 17,1 69=72 23 71,5 68,0 7 63,3 52,6 12,7 7„7 X == 67,2 65,3 == 63,2 53,2 16,1 10,0 Intergrade 73 8 60,1 57,1 20 52,9 35,6 32.8 17,3 Single specimens not included in means (X) Appendix Table 3. Non-weighted Mean % Virtual Prickling in Each of 14 Body Sections (Specimens> 60 mm S.L.) Population , No. Sample Lettered Body Sections (from Skin Map, Fig. 1) Size a b c d e f . g h I "f k 1 m n. Coastal 2 8 _ = 70.6 o o 35,6 72.5 o o v 32.5 55.6 3 18 — 65,9 1.1 2.8 31,1 60, 3r - - ' 42,8 53.6 4 ;8 — . . — 65.6 — 25,6 72,8 41,1 51,1 5 16 — 0.9 78.8 3.1 7.8 41.2 75.9 44.7 c66,6 „o OO 2.2 — _ 6 9 — 0^9 86.7 10.6 3.3 28.3 85,6 .41.1 66.1 a . — — 1.1 oo 7 18 — • ' 56.9 — 4.7 26.1 48,9 32.2 37.5 oo — — . • !»—•?"!• oo 11 5 2.4 = = " 18,0 25.0 25.0 17.0 oo o o o — ; ; o o 12 11 — 32.7 - = 14.1 47,7 ) = - 30.9 41.4 oo o o o o 13 17 ft 6.9 90.9 20.6 12.0 53.2 .88.8 11.8 43.5 69.7 1.2* oo oo o o 14 11 0.7 8.9 90.0 9.1 9.5 56,8 84, 5 2.3 42.3 67,3 = . = o o — — oo 15 16 4.7 90.3 20.0 14.7 50,3 86,6 2.1 43.4 64.4 o o ' — — 1.9 o o 16 6 — - = 85.8 ... ==w 5.8 40.8 8,0 J6 38.3 67.5 o o o — 5.8 — o 18 5 ft* 4.0 26.0 4.0 46.0 50', 0/ — - 41.0 47.0 OO oo 10.0 o o 19 33 3.0 16.5 92,3 8.9 27.4 65.9 87.0 3.5 43,2 74,1 OO oo 5.3 o o 20-21 9 - - (6.7) — (10.0) (39,4) (21,1) (35.6) (11.1) o = 25a 1 — o ~ • — 15,0 50.0 '•• — 35,0 50,0 -o o- OO . — 26 24 1.9 50,4 — - o 20,6 57.9 - - 36.2 52,3 28 2 — — — - - 17.5 — n 35.0 20.0 29 2 5.0 30.0 31 3 — — 53.3 5,0' 56.7 43.3 61.7 — _ » — Coastal Derivative 32 2 32.5 Oo 15.0 25.0 — 25.0 15.0 33 19 2.6 87.9 2.6 6.8 46.1 83.4 32.6 57.6 34 1 — ft oo 80.0 - - - - 65.0 65.0 35.0 50.0 ft 35 4 1.0 2.2 97.5 23.8 18.8 56.2 93.8 15,0 38.8 62.5 2.5 — - 5,0 — — 37 11 5.4 83.6 0.4 44,5 7,6.8 • — 32.3 48.2 38 2 _ « 90.0 20.0 37.5 80.0 — 30.0 55.0 — 2.5 41 < - . 2 — 7.0 90.0 — — 52.5 82.5 — — . 37.5 62.5 Appendix Table 3 (cont'd). Non-weighted Mean % Virtual Prickling in Each of 11 Body Sections (Specimens ̂ 60 mm S.L.) Population. Sample Lettered Body Sections (from Skin Map, Fig. 1) No. Size a b c d e f g h i j k 1 m n Inland. 11 11 33.3 68.9 99.6 56.8 100.0 98.6 98.9 37.5 57.5 91.8 10.7 21.8 3.6 15 9 31.6 52.8 100.0 68.3 60.6 86.1 100.0 60.6 56.7 97.2 22.8 1.2 32.2 19.1 16 i 75.0 70.0 100.0 65.0 100.0 100.0 100.0 25,0 55,0 85.0 =- -- 10.0 mm~ 17 15 23.7 51.0 99.0 59.3 57,3 89.0 99.0 53.0 50.3 90.7 25,3 17.3 <-,<» 5Q 10, 10.0 65.0 ibo.o 16.5 99.0 92,5 100.0 35,5 17.0 98.0 6,0 13.0 •»B 51 31 26.3 28.0 97.7 31.6 82.3 81.0 95,5 12.6 53.5 83.1 ==. - - 15.2 _» 52 25 15.8 65.6 99.0 .11. 0 91.H 91.8 97,2 29,8 55,0 87.2 6.6 - - 1.1 • » « 53 15 11.3 13.5 98.7 29.3 71.3 85.7 95.3 13.7 55,7 87.0 1.0 =-' 15,7 51 21 82.6 93,8 100.0 65.0 100. d 99.5 99.5 61.7 69,5 96.1 31.1 2.8 12.9 5.0 55 20 75.7 91,8 100.0 61,5 95.2 99.2 100.0 57.0 57.0 95,2 20,2 --• 21.8 5.5 56 13 6.3 15.3 98.8 38.8 62.3 72,7 91.9 69.6 45.8 80.0 0.8) Tt- : ), 2.8 57 1 60.0 75.0 100.0 70.0 100.0 100.0 100.0 35.0 50.0 85,0 10,0 10.0 58 6 77.5 91.2 100.0 72.5 100.0 100,0 99,2 53.3 51.2 93,3 20.8 27.2 5.0 60 7 68.3 92.1 99.3 50.7 97,1> 100,0 98.6 35.0 60.7 91.1 20.0 — 17.1 15,7 61 15 87.0 95.3 98.7 37.3 93.3! 100.0 96.0 33.0 60.3 88.7 10,0 0.7 30.3 1.7 62 6 26.7 13.3 100.0 15.0 75.0 89,2 98.3 26.7 52.5 83.3 1.7 20.8 63 11 51.8 + 89.5 91.1 27.7 95.1 90.9 22.7 60.0 78.6 1.1 i . i 11.8 61 7 (11.3) (78.6) (93.6) (25.7) (97.9) (95.0) (79,3) (7.1) (60.0) (73.6) (2.8) (0.7) (9.3) -65 10 91.5 100.0 100.0 69.5 100.0 100,0 100.0 19.5 72.0 96.0 . 30.5 7.0 59.0 8.0 66-68 6 61.0 91.2 100.0 57.5 97.5 100.0 100,0 19.2 50.8 89.2 11.7 — ' 11.7 — 69-72 7 17.1 82.8 95.0 31.1 80.0 98.6 93.3 33.6 5?.a 90.0 11.1 -- 3.6 — Intergrade ? 73 20 25.2 50.0 96.5 38.0 55.2 : . <_ 8.1,2 93.5 17,5 16.5 77.8 2.0 — — • 8. 8 — Represents only 1 or 2 prickles in this body section, '* One specimen in this sample was in a l l respects typically "inland" in i t s prickling, t Data enclosed by brackets indicate mean "apparent" prickling only. 1 These mean percentages have not been,weighted by multiplication of fractional constants appropriate to each body section. 102 Appendix Table 4. Mean Relative Prickling Intensity Index Values For A l l Populations of Cottus asper. Population Standard Length <60 mm Standard Length ^60 mm Sample Size Intensity Mean Range Sample Size Intensity Mean Range Coastal 1 2 3 4 5 6 7 8 11 12 13 14 15 16 18 19 20-21 25a 26 28 30 31 Range Coastal Derivative 32 33 34 35 36 37 38 39 40 41 42 43 2 2 10 12 5 14 17 28 10 9 6 6 15 15 12 6 13 13 14 6 11 13 1.0: 2.0 2,7 1.6 3.0 2.8 3.8 3>6 2.7 1.9 3.8 2.8 2.9 (1.0-3.8) Range 3.3 3.0 3.0 3.2 2.1 1.9 3.0 2.9 2.6 (1.9-3.3) e e e o o (2-3) (1-2) (2-4) (2-4) (3-5) (1-5) (1-4) (1-5) (3-5) (2-3) (2-3) (2-4) 0 0 0 0 0 0 0 0 0 0 (2-4) (1-3) (1-2) (2-4) (1-4) 8 18 10 16 9 18 2 2 11 17 11 16 26 5 33 6 1* 24, 2 19 4 11 2 2.9 2.3 2.4 3.1 2,9 1.2 3.0 1.0 1.5 2.6 2.6 3.8 3.2 1.6 3.4 3.2 4 2.0 3 2 3 (1,0-3,8) 1.5 3.0 3.0 4.0 2,9 2.5 3 3.0 (2.5-4.0) (2-3) (1-3) (1-3) (2-4) (2-4) (1-2) ooooo 0 0 0 6 0 (1-3) (1-5) (1-4) (2-5) (2-5) (1-3) (1-5) (2-4) 4 0 0 o 0 (1-3) (2-3) (1-2) (2-3) ooooo (3.5) (2-3) (2-3) ooo VW' ooooo 103 Appendix Table 4 (cont'd). Mean Relative Prickling Intensity Index Values For A l l Populations of Cottus asper. Population No. Standard Length <c60 mm Standard Length ̂ 6 0 mm Sample Size Intensity Sample Size Intensity Mean Range Mean Range Inland 44 15 3.7 U-4) 14 3.9 (3-5) 45 3 3.7 (3-4) 9* 4.0 • s o t s 46. 15 3.0 U-4) 1* 4 • • • 0 47 6 3.7 U-4) 15 3.7 (3-4) 50 12 3.3 (2-4) 10 3.4 (3-4) 51 11 5.0 0 4 0 0 6 31 4.6 (4-5) 5,2 15 3.1 (3-4) 25 4.2 (3-5) 53 13 3.8 (2-4) 15 4.3 (4-5) 54 20 4.3 (4-5) 21 4.9 (4-5) 55 16 3.9 (2-5) 20 3.9 (3-4) 56 14 3.2 t2-4) 1 3 * 3,3 (3-4) 57 14 3.6 (2-5) 1* .4 ooooo 58 24 4.5 (3-5) 6 4.7 (4-5) 59 2 0 * 3.6 (3-4) — m t a ~ 60 .1 5 ooooo 7 4.3 (4-5) 61 21 4.3 (3-5) 62 4.5 (4-5) 9 4,0 (3-5) 63 — - — - - 18 4.5 (4-5) 64 2 4.5 (4-5) 7 4.9 (4-5) 65 11 4.2 (4-5) 10 4.2 (4-5) 66-68 2 4.5 (4-5) 7 4.0 (3-5) 69-72 23 4.2 (4-5) 6 3.6 (3-4) Range (3.0-5.0) (3.3-4,9) Intergrade 73 8 3.8 (3-5) 20 3.4 (2-4) Single specimens not used in computing means Appendix Table 5,. Pectoral Fin Asymmetry in Cottus asper, by Population. Population No. Sample Size (Pairs) Symmetrical No. % Pairs Asymmetrical No, % Pairs Total Ray Count Greater Greater on Left on Right Direction of Asymmetry Within Population Si n i s t r a l Dextral Equal Coastal ,1 10 9 90.0 i 10.0 1 — _ _ 2 8 7 87.5 l 12.5 1 - . + - 3 20 14 70.0 6 30.0 5 1 + - - 4 21 18 85.7 3 14.3 3 - + - - 5 18 16 88.9 2 11.1 1 1 - - + 6 19 15 78.9 4 21.1 2 2 - + 7 30 20 66.7 10 33.3 9 1 + - - 8 2 2 100,0 0 — - - - - 11 5 3 60.0 2 40.0 - 2 . - 12 11 9 81,8 2 18.2 1 1 - + 13 22 15 68.2 7 31, 8 5 2 - - 14 24 17 70.8 7 29.2 5 2 + - 15 33 29 87.8 4 12.2 3 1 + - - 16 72 60 83.3 12 16.7 7 5 + - - 18 15 13 86.7 2 13.3 - 2 - + - 19 33 30 90,9 3 9.1 2 1 + - 20 12 8 67.7 4 33.3 3 1 +: - - 21 kS 4 80.0 1 20.0 1 - +. - - 23 30 26 86„7 4 13.3 3 1 + • - - 25a 7 4 57.1 3 42.9 1 2 - + - 26 35 28 80.0 7 20.0 5 2 + - - 28 8 4 50.0 4 50.0 3 1 + - - 29,31 5 2 40.0 3 60.0 2 1 t - -30 15 14 93.3 1 6.7 — 1 — + — Totals 46.0 367 79.8 93 20.2 62 31 15(65.2) 5(21.7) a (13.1) Appendix Table 5 (cont'd). Pectoral Fin Asymmetry in Cottus asper, by Population Population No. Sample Size (Pairs) Symmetrical No. % Pairs Asymmetrical No. % Pairs Total Ray Count Greater Greater on Left on Right Direction of Asymmetry Within Populations Sinistral Dextral Equal Inland 44 26 26 89.6 3 10.4 2 1 + 45 12 9 75.0 3 25,0 3 - + - - 46 16 10 62.5 6 37.5 4 2 + — - 47 21 17 81.0 4 19.0 - (4 - + - 50 22 9 40.9 13 59.1 5 8 - + - 51 42 32 <76.2 10 23.8 6 ;4 + - - 52 40 34 85.0 6 15.0 5 1 + - - 53 27 20 74.1 7 25.1 6 1 + - - 54 41 30 73.2 11 26.8 § 5 + - 55 37 29 78.4 8 21.6 6 2 + - - 56 27 17 63.0 10 37.0 4 '6 + - 57 15 7 46.7 8 53.3 5 3 + - 58 30 22 73.3 8 26.7 3 5 - • 59 18 9 50.0 9 50.0 6 3 + - - 60 16 11 68.8 5 21.2 <:8 2 + - - 61 21 15 71.4 6 28.6 6 : r - - 62 19 19 100.0 0 — •- - - — 63 19 14 73.7 5 26.3 3 2 + - - 64 8 7 87.5 1 12.5 1 - + - - 65 22 17 77.3 5 22.7 2 3 - + - 66-68 11 % 63.6 4 36.4 2 2 - - + 69-72 30 .26 86.6 4 13.4 4 - + - - 73 28 .'21 75.0 7 25.0 5 2 + — — Totals 5 5 1 -.408 74.0 143 26.0 87 56 16(72.7) 5(22.7) 1(4.6) Appendix Table 5 (cont'd). Pectoral Fin Asymmetry in Cottus asper, by Population. Population Sample . Symmetrical Asymmetrical Total Ray Count Direction of Asymmetry Ho. Size No. % No. % Greater Greater Hi thin Population*; (Pairs) Pairs Pairs on Left on Right Sinistral Dextral Equal Coastal Derivative 32 2 0 2 100.0 2 + 33 22 18 81.8 4 18.2 4 - + 34 19 14 73.7 5 26.3 2 3 - + 35 16 11 68.8 5 21.2 3 2 + - - 36 6 4 66.7 2 -33.0 1 1 - + 37 23 16 69.6 7 30.'4 4 3 + - - 38 15 11 73.3 4 26. 7 3 1 + - — 39 15 12 80.O 3 20.0 3 - + - - 40 7 S 71.4 2 28.6 2 - + 41 2 2 100.0 0 - — _ _ _ 42 11 10 90.9 1 9.1 1 - + - 43. . w- 10 . 71.4 4 28.6 2 2 - + Totals 152 113 74.4 39 25.6 27 12 8(72.7) 1( 9.1) 2(18.2) Grand Totals 1163 888 76.4 275 23.6 176 99 39(69.6) 11(19.6) 6(10.7 Appendix Table 6. Summary of Meristic Counts (Means and Ranges) for a l l Populations of Cottus asper,- Pop. No. No,Specs. Dl D2 A P i l e f t 1. 10 8,80 (8-9.) 20,70 (20=22) 16.70 (16=18) 15,30 (15-16) <& • 8 9.88 (9=11) 20,50 (20-21) 16,88 (16-17) 15,75 (15-16) 3. 20 8.95 (9-10) 19,85 (19-21) 16,35 (16-17) 15,50 (15-16) 4. 21 8.95 (8-10) 20,38 (20-21) 16,67 (15-18) 15,95 (15=17) 5. 18 9,06 (9=10) 20,50 (20-21) 17,11 (16-18) 15.22 (15-16) 6. 19 8.79 (8=9 ) 19, Nt (19=21) 16,37 (16-17) 15,63 (15-16) 7, 30 9.13 (9-10) 20,60 (20=21) 16,40 (15=17) 16,63 (16-17) 8. 2 9.00 (9 ) 20.50 (20-21) 17,00 (17 ) 16,00 (16 ) 9. 20 9,10 (9=10) 20.70 (20=22) 17,00 (16=18) 15.84 (15-17) 10. 10 9,00 (9 ) 21,00 (20-22) 17.00 (16=18) 16,60 (16-18) l i e 6 9.00 (9 ) 20,83 (20=21) 16,50 (16-17) 16,50 (16=17) 12. 11 8.82 (8-9 ) 20,27 (20=21) 16,64 (16=17) 17,00 (16=18) 13, 22 8,95 (8=9 j 20,73 (20-22) 17,18 (16=18) 15,95 (15-17) 14. 25 9,08 (8=10) 20.32 (19=21) 16,44 (15=18) 15,38 (15-16) 15, 33 9,06 (8=10) 20,76 (20=21) 16,88 (16=18) 15,55 (15-16) 16 o 72 8.99 (8=10) 20,53 (19=21) 16,80 (16-18) 15,89 (15-17) 17, 10 8,90 (8=9 ) 20,80 (20-21) 17,00 (16-18) 16,20 (16-17) 18. 15 9.00 (9 ) 20,67 (20=21) 16,87 (16=18) 15,93 (15-16) 19, 33 8.82 (8=10) 20,27 (18-21) 16,76 (16-18) 16,45 (16-17) 20. 12 9,00 (9 ) 20,42 (20=21) 16,75 (16=18) 16,75 (16-18) 21. 6 9,00 (9 ) 20.83 (20-21) 17,00 (16=18) 16.20 (16-17) 22, 30 9.00 (9 ) 20,47 (20=21) 16,87 (16=18) 16,10 (15-17) 23. 5 9,00 (9 7 20,60 (20=21) 17,20 (17-18) 16,40 (16-17) 24a. 5 9.00 (9 ) 20,80 (20-22) 16,80 (16=18) 15,60 (15-16) 25a S b. 17 9.00 (9 ) 20,14 (20-21) 16,00 (15-17)^ 16,14 (15-17) 26. 35 9,03 (8-10) 20.23 (19-21) 16,49 (15=18) 16,46 (16-17) 27. 5 8.60 (8-9 ) 20.00 (20 ) 16,60 (16-17) 16,20 (16-17) 28, 8 8,75 (8-9 ) 19.50 (19-20) 16.50 (16-17) 16,38 (15-17) 29.31. 5 8.80 (8-10) 20.00 (20 ) 16,80 (16-17) 15,60 (15-16) 30. 15 8.67 (8=9 ) 19,73 (19-20) 16,47 (15-17) 16,00 (15-17) 32. 6 9.00 (8-10) 22.17 (22-23) 17,67 (17-18) 16,67 (16-17) 33. 24 9.46 (9-11) 20,70 (20-21) 16,75 (16-18) 16,09 (16-17) 34. 19 9.05 (9-10) 20.21 (20-21) 16.42 (15-17) 16,53 (16-17) 35. 16 9,19 (9-10) 20,25 (20-21) 17,00 (16-18) 16,25 (16-17) 36, 6 -- . . . < . . . . . . 16,50 (16-17) 37. 24 9.42 (9-10) 20.54 (20-22) 16.96 (16-18) 16,39 (15-17) Pj_ right D l + D2 15,20 15,88 15,30 15,81 15,22 15,63 16,37 16,00 16,83 17,00 15,82 15,24 15,48 15,86 16,07 16,42 16,58 16,00 16,00 16,20 15,60 16,29 16,37 16,00 16,12 15,40 16,07 15,91 16.58 16.19 16,50 16,38 15=16) 15=16) 15=16) 15=16) 15=16) 15=16) 16=17) 16 ) 16=17) 17 ) 14=16) 15=16) 15=16) 15=17) 16=17) 16=17) 16=18) 16 ) 14=17) 16-17) l S = i & ) 16=17) 15-17) 15-17) 15=17) 15=16) 15-17) 15- 17) 16=18) 16- 17) 16-17) 15-17) 29,50 30,38 28,80 29,33 29,56 28,63 29.73 29,50 2 9, 80 30,00 29,83 29,09 29,68 29,40 29,82 29<,f2 2 9.70 29,67 29,09 29.42 29,83 2 9. ,45 29,60 29,80 29,14 29,26 28,60 28,25 28.67 28.40 31.17 30.17 29,26 29.44 (29-31) (2 9=32) (28=30) (28=30) (29=30) (28=30) (29=31) (29=30) (29-32) (29,31) (29-30) (28=30) (29=31) (28,31) (2 9=31) (28=31) (29=30) (29=30) (26=30) (29=30) (29=30) (29=30) (29=30) (29=31) (29=30) (28=30) (28=29) (27-29) (28=30) (27=29) (30=32) (29=31) (29-31) (29=31) 29,96 (29-32) Appendix Table 6, Summary of Meristic Counts (Means and Ranges) for a l l Populations of Cottus asper p(cont' d). Pop. No, No, Specs, P l l e f t ?l right Dl + D2 38, 15 9,13 (9-10) 20.47 (20-21) 16,93 (16-18) 16,731 (16KL7) 39, 15 9.27 (9-10) 20.40 (20-21) 16.80 (16-18) 16,27 (15-17) 40, 7 -- —_ 17.43 (17-18) 41, 2 9.50 (9-10) 20.00 (20 ) 16.50 (16-17) 17,50 (17-18) 42, 12 9,17 (9-10) 20,17 (20-21) 16,42 (15-18) 16,17 (16-17) 43, 15 9.61 (8-10) 20,31 (20-21) 16,27 (16-18) 16,67 (16-17) 44, 29 9.03 (9-10) 20,27 (19-21) 16,59 (15-17) 16,86 (16-18) 45, 12 9.00 (9 ) 2;0,17 (20-21) 16,83 (16-18) 16,75 (15-18) 46, 16 9.25 (9-10) 20,31 (20-21) 16,25 (15-17) 16,50 (15-18) 47, 21 9.10 (8-10) 20,71 (20-22) 16,52 (15-17) 16,29 (15-17) 48. 9 9,22 (9-10) 20.00 (19-21) 16,12 (16-17) 49. 10- 8,80 (8-9)) 20,70 (20-22) 17,10 (16-18) 16,90 (16-17) 50. 22 8.52 (7-10) 20,64 (16-23) 16,67 (15-17) 17,14 (13-18) 51. 42 8,98 (8-10) 20.76 (20-21) 17,31 (16-20) 16,93 (14-18) 52. 40 9.00 (9 ) 20.82 (20-22) . 17,02 (16-18) 16.60 (16-18) 53. 28 9,00 (8-10) 20,82 (20-21) 16, t i (1-6-.13) 16,93 (16-18) 54. 41 8.98 (8-9 ) 20.90 (19-22) 16,98 (15-18)® 16,44 (15-18) 55. 37 8,95 (8-9 ) 20,70 (19-22) 16,89 (16-18) 16,43 (16-17) 56. 27 8.89 (8-9 ) 20.89 (20-22) 17,18 (16-18) 16,30 (15=17) 57. 15 8.80 (8-9 ) 20.93 (20-22) 17,00 (16-18) 16,67 (14-18) 58. 30 8.87 (8-9 ) 20.13 (19-22) 16,27 (15-17) 16,73 (16=18) 59. 20 9.06 (8-10) 20.31 (19-21) 16,81 (16-18) 16,67 (14-18) 60. 16 8.69 (8-9 ) 20.87 (2® £2 2) 16,81 (15-18) 16,44 (16-17) 61. 21 8.81 (8-10) 20.69 (19-22) 17,24 (16-19) 16,29 (15-18) 62. 13 9.08 (9-10)s? 20*69) (20-22) 17.15 (16-18) 16,00 (15-17) 63. 24 9.00 (8-10) 20.29 (19-21) 16.58 (15-18) 16,25 (16-17) 64, 9 8.89 (8-9 ) 20.56 (20-21) 17.00 (16-18) 1.6,00 (15-17) 65. 22 8.68 (8-9 ) 20,23 (19-21) 17,18 (16-18) 16,14 (15-17) 66a. 3 8.00 (8 ) 19.67 (19-20) 16.67 (16-17) 16.00 (15-17) 66b. 3 8.00 (8 ) 20,00 (20 ) 17,00 (17 ) 16.00 (16 ) &V1 o 3 8.00 (8 ) 19.67 (18-21) 16.33 (16-17) 15.67 (14-17) ?-2 8.00 (8 ) 20.00 (20 ) 17,50 (17-18) 16.00 (16 ) 6*f.c 14 8.21 (7-9 ) 19.00 (19 ) 16,64 (16-18) 15,64 (15-16) 70?'= 8 8.00 (8 ) 19.12 (18-20) 17,00 (16-18) 16,50 (16-17) 7i|p 3 8.00 (8 ) 19.33 (19-20) 16,67 (16-17) 16,33 (16-17) 72|* 4 8.00 (8 - ) 18,75 (18-20) 16,50 (16-17) 16.00 (15-17) 73> 28 8.82 (8-10) 19.93 (19-21) 16,54 (15-18) 16,21 (15-17) 16,67 16.07 17,14 17.50 16,00 16,64 16,83 16,50 16,38 16,45 17,50 16,86 16,50 .16,74 16,39 16.32 16,41 16,53 16,80 16,16 16,38 15,86 16,00 -16,21 15.89 16,18 15.67 16,33 16.00 15,00 15,50 16,25 16,33 16,00 16,11 16- 18) 15- 17) 17- 18) 17-18) 16 ) 16- 17) 16-17/) 15-18) 15=18) 15=17) 17=18) 14=19) 16=17) 16-18) 15=17) 15=17) 16=17) 14=17) 16=18) 14=18) 15=17) 15=17) 15=17) 15=17) 15=16) 16=17) 15- 16) 16- 17) 15- 17) 14-16) 14- 16) 16- 17) 16-17) 15- 17) 15-17) 29,60 29.67 29,50 29.34 29/92 29,31 29.18 29,56 29,81 29,22 29,50 29.14 29,74 29,83 29,82 29.88 29,65 29,78 29.73 29,00 29f.45 29,53 29.50 29,77 29,29 29,45 28.91 27,67 2 8.00 27,67 28,00 27,21 27,12 27,33 26,75 28,75 29-31) 29-31) 29-30) 29-31) 29-31) 28=30) 29-30) 29-31) 28=31) 29=30) 29=30) 28- 32) 2 9-32) 29- 31) 28=31) 28=31) 28=31) 28=31) 29=31) 27- 31) 28=31) 28- 31) 28=31) 2 9=31) 2 8-31) 28=30) 28-30) 27-28) 28 ) 26-28) 28 ) 26-28) 26- 28) 27- 28) 26- 28) 27- 31) Appendix Table 7, Monthly Mean Temperature in °C for Selected Coastal and Inland Localities,, Locality Latitude J F M A M J J A S 0 N D Annual Range Coastal Valdez 61° 07' N . - 7.1 - 5,9 - 2,6 + 1,4 6,3 10,2 11,3 10,9 7,8 3,0 - 3,9 - 7,5 18,8 Prince Rupert 54° 17' N . 1.7 ,2,9 4,2 6,3 9,8 12,0 13.4 13,9 12,0 8,7 5,1 2,7 12,2 Victoria 48° 25' N o 3.6 5,7 7,2 9,6 12,4 14,3 15,9 15,7 14,4 10.9 7.3 •>5.:2 12,3 San Francisco 37° 47' N o 10.0 11,0 11.9 12,5 13,3 14,4 14,3 14,7 15.9 15,3 13.6 10,9 5.9 Santa Barbara 11,8 — _= 19,1 — — — . ='= — 7,3 San Diego 32° 44' N . 12,3 13,4 14,8 16,2 17,8 19,0 21,0 21,7 20.9 18.6 15.6 13,2 9,4 N-S Range 19,4 19.3 17,4 14,8 l i , 5 8,8 9,7 11,8 13,1 15,6 -19,5 20.7 Inland Watson L o 60° 07' -22.0 18,4 - 8,9 - 0,4 8,0 13,6 14,8 12,8 9,0 1.1 -13,7 -22,1 .36.8 Prince George 53° 54' - 9,9 6,6 - 1.4 + 4,6 9.9 13.4 14,9 13,7 9,9 4,8 - 2.7 - 7,5 24.8 Spokane 47° 37' - 4.4 +0.4 3.8 8,7 13,1 16,3 20,9 20,0 15,4 9,2 2,7 -1,1 25.3 Red Bluff 40° 09" 6,9 9,2 11.4 14,6 19,8 23,0 27,1 25,4 23,5 17,4 11.6 7,7 20.2 Sacramento 38° 35' 7.6 10.0 12,2 16,3 18,0 21,3 23.9 23,2 22,3 17,6 12,2 8,3 16,3 Fresno 7,5 27,4 — - — — 19.9 N-S Range 29.6 28.4 21.1 15,9 11,8 9,6. 12.6 12,6 14,5 16,5 25,9 30,4 — *10 year-averages, 1941-1950 (World Weather Records, U„S, Weather Bur, Wash., D,C, 1959.1-1361)

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