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Patterns of Interactive Segregation in Three Species of Sculpins (Cottus) in Western Oregon

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Patterns of Interactive Segregation in Three Species of Sculpins (Cottus) in Western Oregon AN ABSTRACT OF THE THESIS OF Terry Richard Finger for the degree of Doctor of Philosophy in Fisheries presented on May 22, 1979 Title: Patterns of Interactive Segregationin Three Species of Sculpins (Cottus) in WesternOregon. Abstract approved: Redacted for Privacy D / H'a1 1 Species interactions andtheir effect on theuse of habitat by benthic fishes were examined in a guild composed of Cottusperplexus, C, rhotheus, and C. beldingi in the Marys River abovePhilomath, Oregon, and in a constant environment provided byan artificial stream. In the Marys River system, the pattern of longitudinalsuccession in this guild was one of species additions downstreamrather than sequential longitudinal replacement. Cottus perplexus occurred throughout the river system, C. rhotheus was found only in themiddle and lower reaches, and C. beldingiwas restrictedtothe lower reaches within the study area. Longitudinal distributions appearedto be most strongly related to substrate characteristics in riffles. Cottus perplexus occurred on all substrates,C. rhotheus was found only where rock was present, and C. beldingi was captured onlyin riffles dominated by loose, cobble-sized rock with many interstitialspaces. Local use of habitatwas strongly related to the breadth of environmental tolerance of eachspecies arid thepresence or absence of other cottus species. The species broadly overlappedin their use of different substrates, depths, and distances fromshore, but segregated in an orderly manner withrespect to current velocity. Where it existed alone, C. perplexus utilizedboth pools and riffles and was found in greatest densitiesnear cover. Where it coexisted with C. rhotheus, C. perplexuswas primarily confined to pools and declined significantly in abundance. The degree to which C. perp].exus was excluded from riffles varied inrelation to seasons and the size composition of the C. rhotheuspopulation. In areas where there were low densities of large C. rhotheus,the exclusion of C. perplexuswas not as pronounced as in areas where therewere many large C. rhotheus, and was apparently due mainlyto competition. In these areas overlap between the species was alsogreater in spring, the time of peak food abundance, than in summeror winter. In areas where there were greater numbers of large C. rhotheus, the exclusionof C. perplexus from riffles was more pronounced and didnot vary seasonally. In these areas predation was apparently themost important mechanism causing the interactive segregation. Cottus rhotheus was generally confined to riffles, althoughsome large individuals did wander into pools during times of food scarcity. Cottus beldingi was exclusively found in riffles and hada greater maximum current velocity tolerance than C. rhotheus. In summer, all riffle areas in the MarysRiver flowed slowly enough for C. rhotheus, andC. beldingi apparently was forced, by predation and competitionexerted by C. rhotheus, to burrow into the substrate. In winter and spring, C. rhotheuswas excluded from areas of some riffles by thefaster currents. tn these areas C. beldingi had a refuge from competitionand predation andwas found atop the substrate to a greater degree. Experiments with the three speciesin the arti:Eicial stream supported the field data andsuggested that the competition contributing to the habitat shiftswas interference competition for space. Cottus perplexus selected riffleareas when alone, but was displaced partially to pools by C. beldingi throughinterference competition. The addition of C. rhotheus causedC. perplexus to be more rigidly confined to pools, butC. beldingi was able to coexist in riffles with C. rhotheusapparently by utilizing the interstitial spaces in the substrate. Comparison of these results with thoseof other studies suggests that segregation in stream fishesshould be more commonly basedon habitat than on food or time. A model was constructed that couldbe used to predict patterns of habitatutilization in other guilds of benthic fishes. Species interactions were also suggestedas having a major influence on the evolutionarypatterns of Cottus in western North America. Large, dominant species will obtaintheir preferred habitat throughout their geographicalrange. They will therefore not need to adapt to a widerange of conditions and thus may exhibit low morphological variability. Contrastingly, less dominant speciesmay be forced into many differenthabitats, depending on which dominant species are also present, andmay thus exhibit greater morphological variability. 1979 TERRY RICHARD FINGER ALL RIGHTS RESERVED Patterns of Interactive Segregation in Three Species of Sculpins (Cottus) in Western Oregon by Terry Richard Finger A THESIS submitted to Oregon State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy Completed May 22, 1979 Commencement June 1980 APPROVED: Redacted for Privacy ciate Professor of Fisheries In charge of major Redacted for Privacy ad of Department of Fiherie's andWildli Redacted for Privacy Dean of Graduate School Date thesis is presented May 22, 1979 Typed by researcher Terry Richard Finger ACKNOWLEDGEMENTS I wish to express my appreciationto my major professor and committee members, James D. Hall, Carl E.Bond, W. Scott Overton, and C. David Mclntire, for guidance throughoutthe course of this study and for their careful reviews ofthe manuscript. Many individuals unselfishly assisted in thefield work; I am particularly indebted to James Long, JefferyZiller, David Lundahi, and Russell Oates. I would also like to thank Wayne Seim ofthe Oak Creek Laboratory of Biology; withouthis collaboration the artificial stream experimental work wouldnot have been possible. Data analysis was made possible bya grant from the Oregon State University Computer Center. Field work expenses were paid in part by a Grant-in-Aid of Research fromSigma Xi. I would also like to add my thanksto Arthur J. Boucot, William J. Liss, and Charles E. Warren for themany informal discussions that heightened my enthusiasmand gave rise to the evolutionary framework of this thesis. TABLE OF CONTENTS P age INTRODUCTION 1 CONCEPTUAL FOUNDATION 5 Natural History of Cottus 5 Mechanisms Producing Community Structure 10 Guilds and Hierarchical Structure 13 Guild Structure in Stream Fishes 17 METHODS 20 The Study Site 20 Preliminary Survey 23 Local Patterns of Habitat Utilization 24 Artificial Stream Experiments 33 RESULTS AND INTERPRETATION 36 Composition of the Vertebrate Fauna 36 Longitudinal Distribution of Cottus species 38 Local Patterns of Habitat Utilization 40 Artificial Stream Experiments 79 DISCUSSION 85 General Patterns of Segregation in Freshwater Fishes 85 Mechanisms of Segregation in Stream Fishes 91 A Model of Interactive Segregation in Cottus 95 Species Interactions and the Evolution of Cottus in Western North America 103 LITERATURE CITED 109 APPENDICES 118 LIST OF FIGURES Figure Page 1 A portion of a hierarchically organizedsystem 16 based on the fauna of many of the streams of the Willamette River drainage inwestern Oregon. 2 The Marys River system, Benton and Lincoln 21 Counties,, Oregon. 3 Gradient map of the Marys River system, Benton 22 and Lincoln Counties, Oregon. 4 Numerical density of Cottus perplexus in the 43 upper Marys River, Oregon, in relation tocover provided by vegetation and wood debris. 5 Numerical densities of Cottus insummer in 53 relation to current velocity in riffles. 6 Numerical densities of Cottus in winter in 56 relation to current velocity in riffles. 7 Numerical densities of Cottus in spring in 59 relation to current velocity in riffles. S Numerical densities of Cottus insummer in 61 relation to mean substrate particle size in riffles. 9 Numerical densities of Cottus in winter in 62 relation to mean substrate particle size in riffles. 10 Numerical densities of Cottus in spring in 63 relation to mean substrate particle size in riffles. 11 Numerical densities of Cottus insummer in 64 relation to depth in riffles. 12 Numerical densities of Cottus in winter in 65 relation to depth in riffles. 13 Numerical densities of Cottus in spring in 66 relation to depth in rif1es. Figure Page 14 Numerical densities of Cottus insummer in 67 relation to distance to shore in riffles. 15 Numerical densities of Cottus in winter in 68 relation to distance to shore in riffles. 16 Numerical densities of Cottus in spring in 69 relation to distance to shore in riffles. 17 Within-riffle distribution of Cottus insummer 75 plotted in relation to the firsttwo principal coMponents of the environmental data. 18 Within-riffle distribution of Cottus in winter 76 plotted in relation to the first two principal components of the environmental data. 19 Within-riffle distribution of Cottus in spring 77 plotted in relation to the first two principal components of the environmental data. 20 Numerical densities of Cottus in each species 80 combination in relation to current velocity in the artificial streams. 21 Numerical densities of Cottus in each species 81 combination in relation to mean substrate particle size in the artificial streams. 22 Numerical densities of Cottus in each species 82 combination in relation to depth in the artificial streams. 23 A model of interactive segregation in the Cottus 97 guild in the Marys River. 24 Models of interactive segregation in several 101 guilds of benthic fishes. (a) Cottus asper, Cottus aleuticus inmany coastal Pacific slope streams. (b) Cottus beldingi, Cottus confusus inheadwater areas of Cascade streams. (c) Cottus beldingi, Cottus confusus in downstream areas of Cascade
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