FooD INTERRELATIONSHIPS OF THEſom en scULPIN.CŒTUS BĀ ------… RAINBOW TROUT, SALĀTO GARDNERI, ÎN A TRIBUTARY OF LAKE SUPERIOR - PAUL R. HANNUKSELA Raeſ, AſS OF THE . NATURAL SCIENCE LIBRARY i-- " --- *.*.* . Piº P B R ry on s : º: . . . ... 3-3 - ; :=s. zººs ETFFT, VERTRs Srº- -º- * ABSTRACT FOOD INTERRELATIONSHIPS OF THE MOTTLED SCULPIN, COTTUS BAIRDI, AND JUVENILES OF THE RAINBOW TROUT, SALMO GAIRDNERI, IN A TRIBUTARY OF LAKE SUPERIOR By Paul R. Hannuksela Young rainbow trout and sculpins occupy similar habitat in many Michigan streams and thus, may compete for available food. To examine this possibility, I investigated the production and food habits of coexisting populations of the mottled sculpin and juveniles of the rainbow trout in relation to the standing crop of bottom fauna in a small tributary of Lake Superior. Age 0 and I rainbow trout production from June to August 1970 was 2.8 g/ m”. Mottled sculpins, age I and II, produced 1.5 g/m” during the same period. The bottom fauna was composed mostly of Tendipedidae, Baetidae, Hydropsychidae, Rhyacophilidae, Limnephilidae, and Gastropoda. The mean standing crop of bottom fauna was 9.4 g/ m*. The rainbow trout and the mottled sculpin had similar diets; bottom fauna was eaten in approximate proportion to their abundance in the stream. - No mutual predation occurred between the two fishes. The low total food consumption of 1.9 times the mean standing crop of benthos indicated that detrimental food competition probably did not occur between rainbow trout and mottled sculpins. FOOD INTERRELATIONSHIPS OF THE MOTTLED SCULPIN, COTTUS BAIRDI, AND JUVENILES OF THE RAINBOW TROUT, SALMO GAIRDNERL IN A TRIBUTARY OF LAKE SUPERIOR By Paul R. Hannuksela A thesis submitted in partial fulfillment of the requirements for the degree Of Master of Science 1973 School of Natural Resources Department of Wildlife and Fisheries The University of Michigan Committee members: - Dr. Karl F. Lagler, Chairman Dr. William C. Latta ACKNOWLEDGMENTS Drs. Karl F. Lagler and William C. Latta made up my graduate studies committee and offered suggestions. Personnel from the Marquette Fisheries Research Station, Michigan Depart- ment of Natural Resources, assisted with the field work. James R. Ryckman advised on statistical procedures. Thomas M. Stauffer reviewed the manuscript. - - The study was supported in part by funds from the Federal Aid in Wildlife Restoration Act under Dingell–Johnson Project F-31-R, Michigan. ii TABLE OF CONTENTS ACKNOWLEDGMENTS . . . . . . . . . . LIST OF TABLES . . . . . . . . . . . . . . . . . . . . . INTRODUCTION . . . . . . . . . . . . . STUDY SITE . . . . . . . . . . . . . . . METHODS . . . . . . . . . . . . . . . . Fish Populations and Production Bottom Fauna. . . . . . . . . . . Fish Food Habits . . . . . . . . Fish Food Consumption . . . . Stream Measurements . . . . . RESULTS . . . . . . . . . . . . • • * ~ * Fish Populations and Production Bottom Fauna. . . . . . . . . . . Comparison of Food Habits . . . Food Consumption . . . . . . . . DISCUSSION . . . . . . . . . . . . . . . . . . . . . . . . . . CONCLUSIONS . . . . . . . . . . . . . LITERATURE CITED . . . . . . . . . . iii iv. Table LIST OF TABLES - Page Estimated numbers of the rainbow trout and the mottled sculpin and their mean length and weight in the Little Garlic River, June–August 1970 . . . 9 Mean standing crop, production and food consump- tion of the rainbow trout and the mottled sculpin in the Little Garlic River, June–August 1970 . . . . 10 Taxa and number of Organisms in 10 bottom samples each month, June and August 1970, Little Garlic River . . . . . . . . . . . . . . . . 12 Percentages of total number and weight Of organisms found in bottom samples and in the stormachs of the rainbow trout and the mottled sculpin, Little Garlic River, June and August 1970 . . . . . . . . . . . . . • • * * * * * * * * * * * 14 Analysis of variance of the total diet of the rainbow trout and the mottled sculpin in the Little Garlic River, June and August 1970 . . . . 15 Production of trout, salmon, and sculpins in various streams, 15 June– 15 August 18 iv. INTRODUCTION Many Michigan tributaries of Lake Superior are nursery streams that support substantial populations of juveniles of the rainbow trout (Salmo gairdneri). Most of these streams also contain large populations of sculpins (Cottus bairdi and/or C. cognatus). During the first 1 to 3 years of life, young rainbow trout are in close association with sculpins. Thus, trout and sculpins may compete for available food and prey upon each other. Prior investigations of food interrelationships have been done on various species of salmonids and cottids. However, to my knowl- edge, none has been done on the rainbow trout and mottled sculpin (C. . bairdi) combination. Dineen (1951) found that the brook trout (Salvelinus fontinalis), brown trout (Salmo trutta) and mottled sculpin eat the same food. But he, as did Koster (1939), contended that food competition was lessened by the ability of trout to feed in all planes of the water from top to bottom, whereas sculpins are essentially bottom feeders. Brocksen, Davis, and Warren (1968) concluded that the reticulate sculpin (C. perplexus) could influence the food consump- tion and production of the cutthroat trout (Salmo clarki) by cropping the benthic food supply. They concluded that this reduced the supply of drift organisms (which are important to trout [Hunt, 1965]), as well as the benthos of the substrate. The evidence regarding sculpin predation on juvenile trout and salmon is conflicting. Koster (1939) and Patten (1962, 1971) concluded that trout and salmon fry make up a negligible portion of the diet of several species of sculpins. Conversely, significant predation by the reticulate sculpin on rainbow trout sac fry (Phillips and Claire, 1966) and by slimy sculpins (C. cognatus) on brown trout sac fry (Clary, 1972) has been demonstrated in aquaria. Juvenile rainbow trout in streams reportedly do not prey upon fish (McAfee, 1966). To provide information on food interrelationships of popula- tions of the rainbow trout and mottled sculpin coexisting in one stream system, I investigated their food habits and production in relation to the standing crop of bottom fauna. STUDY SITE This study was conducted on the Little Garlic River, a rain- bow trout nursery stream tributary to Lake Superior in Marquette County, Michigan. Field data were collected during 10–13 June, 8-9 July, and 5-7 August 1970, from a 300-m long stream section some 2 km upstream from the river mouth. The Little Garlic River flows for most of its length through rugged terrain characterized by steep hills forested mainly with mature hardwoods and some conifers. However, the study Site was in that portion of the stream which flowed through an area of relatively low relief. The stream banks were lined with alder (Alnus), aspen (Populus), and willow (Salix). Emergent vegetation was lacking in the study section. Submerged vegetation was primarily green and blue- green filamentous algae sparsely covering the stream bottom. - The stream bottom in the study section was estimated to be 47% rubble, 35% gravel, and 18% sand. Seventy per cent was riffle area with pools making up the remaining 30%. Between June and August, the stream became narrow and shallow because of receding water levels. Mean width decreased from 6.5 to 4.8 m, mean depth from 26 to 17 crin, mean volume of flow from 0.4 to 0. 1 m*/sec, and the area of submersed bottom from 1,950 m” in June, to 1, 770 m? in July, and 1,440 m” in August. Conductivity was 153 umho/cm3 in August at 18 C. Mean water temperature increased from 13 C in June to 18 C in August. Age 0 and I rainbow trout and mottled sculpins were abundant and adults of the longnose dace (Rhinichthys cataractae) Welſ’ e COIY) IO, OI). in the study section. The brook trout, young (age 0) of the coho salmon (Oncorhynchus kisutch), age II and older of the rainbow trout, and the mottled sculpin were rare. Young-of-the-year of the rainbow trout and of the mottled sculpin began to emerge in early June. The other age groups of both species were present during the entire study. - Aquatic insect nymphs and 1arvae, snails, and Oligochaetes were abundant in the stream substrate. METHODS Fish Populations and Production Population estimates of age 0 (August only) and age I rainbow trout and age I and II mottled sculpins were made in June, July, and August using the methods described by Shetter (1957). The fish were captured, marked by excising part of a fin, and released one day. The following day the ratio of marked to unmarked fish was determined. Calculations followed those of Bailey (1951). All fish captured for the population estimate were measured to the nearest millimeter, total length, and 10 from each 10-mm length group were weighed to the nearest 0.1 g to obtain average weights of the age groups for calculating production. TWO random samples of unmarked rainbow trout and mottled sculpins were preserved in 10% formalin for determining age and growth and food habits--35 rainbow trout and 53 mottled sculpins in June, and 78 rainbow trout and 54 mottled sculpins in August. These preserved fish were measured to the nearest millimeter, total length, and weighed to 0.1 g. Scales were removed from the rainbow trout at an area between the origin of the dorsal fin and the lateral line and otoliths were excised from the labyrinths of the mottled sculpin for age assessment. Otoliths, cleared in hot xylene (Larsen and Skud, 1960), and scales, impressed on plastic slides, were enlarged with a microprojector to a magnifica- tion of 107X to determine the age of the fishes (Lagler, 1956). Production of fish flesh was determined graphically by plotting standing crops of rainbow trout and mottled sculpins in numbers on the Ordinate and corresponding mean weights on the abscissa (Allen, 1951; Chapman, 1965). Areas under the resulting 6 production curves were measured for the period 11 June to 6 August 1970. Estimated production of age 0 rainbow trout was low because the August population estimate was used as the estimate (minimal) of their numbers in June and July. BOttom Fauna Ten randomly selected samples of benthos (Hildebrand, 1971) were collected on 12 and 13 June and 10 on 6 and 7 August with a 0.1 m” modified Hess sampler having size-30 mesh netting (Waters and Knapp, 1961) and preserved in 40% isopropyl alcohol. In collecting, the sampler was set on the sample site and forced into the substrate as deeply as possible. Large rocks within the sample site were individually cleaned of fauna. Fine substrate was sifted by hand, causing the fauna to drift into the mesh bag of the sampler. - The sugar flotation method of separating fauna from inadvertently collected substrate and debris (Anderson, 1959) was attempted, but filamentous algae hampered the technique. Manual sorting of the bottom samples proved most feasible. Identification of the Organisms followed Pennak (1953), with individuals being sorted into the appropriate taxonomic group and counted. To get standing crop, average weights of individuals in each taxon were calculated for both sampling periods in the following manner. Four bottom fauna samples were randomly selected from each of the two sets of 10 bottom samples collected in June and August. The known number of individuals in each taxon were centrifuged for 2 minutes at 2500 rpm and weighed to 0.001 g. Trichoptera were weighed without cases. An average weight for individuals in each taxon was calculated. This average was used to calculate weight of each taxon present in the stream and the weight of food at the time of ingestion. Fish FOOd Habits Stomach contents of the random sample of preserved rainbow trout and mottled sculpins were usually identified to family (Pennak, 1953; Ross, 1965), sorted into appropriate taxonomic groups, and counted. A three-way analysis of variance (Freund, 1952) was used to detect any differences between food habits of juvenile rainbow trout and mottled sculpins. I tested the hypothesis that there was no difference between the diets of the two fishes. Sources of variation were the sampling dates, six food categories (classes and Orders) with families as replicates, and the two fish species. Fish Food Consumption Total food consumption during my study was calculated by summing the daily maintenance and the growth ration. The maintenance and growth factors were adapted from other investigations. Allen (1951) and Hopkins (1970) used a daily maintenance ration for brown trout of 1.2% of the standing crop of fish and a growth ration of 4.2 times brown trout production. Bonham (1949) found the growth ration for rainbow trout to be three times rainbow trout production. For purposes of this study, I used 1.2% of the average standing crop of the rainbow trout and of the mottled sculpin as the daily maintenance ration for each, and three times the production of these fishes as the growth ration. Stream Measurements Stream width in meters, depth in centimeters, rate of current flow in meters per second (No. 622 Gurley current meter) were measured at each randomly selected bottom sampling site. Also, the substrate was classified as rubble, gravel or sand and the area of each type was estimated as per cent of the total area. RESULTS Fish Populations and Production Several changes occurred in the populations of the rainbow trout and the mottled sculpin during the course of this study (Table 1). Young-of-the-year rainbow trout were just beginning to emerge from their redds in June; by August all had emerged and their population amounted to 2,845 individuals (2.0/m”). Age I rainbow trout, which numbered 724 (0.4/m”) in June and 464 (0.3/m”) in July, decreased significantly to 263 (0. 2/m2) in August. This was probably due to downstream, 1akeward migration (Stauffer, 1972). In June, age I rainbow trout averaged 91 mm in length and weighed 6.9 g. They increased to 99 mm and 9.7 g in July, and to 108 mm long and 11. 7 g in August. Age 0 rainbow trout averaged 44 mm long and weighed 1.0 g in August. Biomass production of rainbow trout from June to August was 2, 292 g (1.3 g/m”) for age 0 fish and 2, 480 g (1.5 g/m”) for age I fish (Table 2). Although only a few very small young-of-the-year mottled sculpins were observed in June, many were present in July and August, but they were too small for me to estimate their numbers. Age I mottled sculpins numbered 1,930 (0. 9/m”) in June; 2, 879 (1.6/m”) in July; and 1,770 (1.2/m2) in August. Although the point estimates differed among the three population estimates, the 95% confidence limits overlapped widely (Table 1), which indicated that there was no significant change in age I mottled sculpin numbers. Age II mottled sculpins numbered 70 in June, 26 in July, and 24 in August. Again, 95% confidence limits overlapped widely. Age I mottled sculpins grew from means of 49 mm and 1.6 g in June, to 55 mm and 2.0 g in July, and to 61 mm and 2.6 g in August. Age II 8 Table 1. --Estimated numbers of the rainbow trout and the mottled sculpin and their mean length and weight in the Little Garlic River, June–August 1970 Population e © Den- HiO – Mean Mean Species, age estimate and & ge Of e Sit mass length weight and month 95% confidence (m.4) ( /m2) (mm) (g) limits g g Rainbow trout Age 0 June 2, 845& 1.5 0.3 23 0.2 July 2, 845a. 1. 6 0. 7 35 0.4 August 2, 845 + 320 2. 0 2. 0 44 1. 0 Age I June 724 + 206 0.4 2. 6 91 6. 9 July 464 + 112 0.3 2. 6 99 9. 7 August 263 + 45 0. 2 2. 1 108 11. 7 Mottled sculpin Age I June 1, 930 + 624 0. 9 1. 6 ºf $3 1. 6 July 2, 879 + 955 1. 6 3. 3 55 2. 0 August 1, 770 + 440 1.2 3. 2 61 2. 6 Age II June 70 + 74 <0. 1 0.4 92 10. 9 July 26 + 16 <0. 1 0.2 93 12. 3 August 24 + 29 <0. 1 0.2 95 13. 1 * These values were derived from the August population estimate and applied to measurements of a sample of young-of-the-year in June and July to derive the matching values in each of the four columns to the right. 10 Table 2. --Mean standing crop, production, and food consumption of the rainbow trout and the mottled sculpin in the Little Garlic River, June–August 1970 Species Mean e FOOd e and standing crop Production consumption age g g/mº g g/mº/ g g/mº/ - day day Rainbow trout Age 0 1, 517 0.9 2, 292 0.02 7, 914 0.1 Age I 4, 191 2.5 2,480 0.03 10,307 0.1 Total 5, 708 3.4 4, 772 0.05 18, 221 0. 2 Mottled sculpins Age I 4, 483 2.6 2, 460 0.03 10, 446 0.1 Age II 466 0.3 - 80 <0.01 559 30. 1 Total 4, 948 2.9 2, 540 0.03 11, 005 0. 1 11 mottled sculpins attained means of 92 mm and 10.9 g in June, 93 mm and 12.3 g in July, and 95 mm and 13. 1 g in August. Change in ichthyomass of the mottled sculpin production from June to August was 2,460 g (1.4 g/m2) for age I fish, plus 80 g (0.1 g/m2) for age II fish (Table 2). - BOttom Fauna Thirty-five taxa were found in the bottom samples (Table 3). The most abundant groups were Tendipedidae, Baetidae, Hydro- psychidae, Rhyacophilidae, Limnephilidae, and Gastropoda. These taxa made up 85% of the total number of organisms and 82% of the total weight. Biomass of bottom fauna was 16.1 kg (8.2 g/m2) in June and 15.3 kg (10.7 g/m2) in August, with a mean of 15.7 kg (9.4 g/m2). There was little change in biomass and species composi- tion of the benthos between June and August. Comparison of Food Habits The 35 taxonomic groups were apportioned into 11 major food categories to test relationships between numbers and weight of various groups in the bottom fauna and the food habits of the rainbow trout and the mottled sculpin (Table 4). Those taxa with an abundance of individuals formed their own food item categories, usually families, while taxa with few members were combined within appropriate higher taxonomic groupings. The frequency of Occurrence and weight of bottom organisms in the stomachs of both fishes was in approximate proportion to bottom fauna abundance in the stream, with few excep- tions. The rainbow trout fed slightly more frequently on baetid nymphs and the mottled sculpin fed a little more often on Trichoptera naiads. Adult stages of insects made up a small portion (9.5%) of the rainbow trout diet, whereas, mottled sculpins ate none. No significant difference existed between the diets of young rainbow trout and mottled sculpins (Table 5). All observed values of F for sources of variation and their interactions were not significant. 12 Table 3. --Taxa and number of organisms in 10 bottom samples each month, June and August 1970, Little Garlic River June August TOtal Taxa Num-- Per Nunn - Per Num- Per ber cent ber cent ber cent Diptera Tendipedidae 3, 443 59 2, 069 39 5, 512 49 Ceratopogonidae 145 2 258 5 403 4 Tipulidae 101 2 30 tr 131 1 Rhagionidae 16 tr 15 tr 31 tr Simulidae 19 tr 3 tr . 22 tr Tabanidae 2 tr 5 tr 7 tr Empidae 5 tr 1 tr 6 tr Cecidomyidae 2 tr 0 0 2 tr Drosophilidae 1 tr 0 O 1 tr Unidentified 4 tr 3 tr 7 tr Ephemeroptera Baetidae 1, 000 17 523 10 1, 523 14 Heptageniidae 57 1 168 3 225 2 Ephemeridae 16 tr 1 tr 17 tr Unidentified 2 tr 0 0 2 tr Trichoptera Hydropsychidae 197 3 55.2 10 749 7 Rhyacophilidae 43 tr 398 8 441 4 Limnephilidae 265 4 129 2 394 4 Leptoceridae 4 tr 128 2 132 1 Hydroptilidae 100 2 O 0 100 1 Psychomyiidae 13 tr 50 1 63 1 Phryganeidae 2 tr 5 tr 7 tr Unidentified 17 tr 13 tr 30 t]." Coleoptera - Finnidae 63 1 126 2 189 2 Unidentified 17 tr 7 tr 24 t;" Plecoptera 86 1 90 2 176 2 Odonata 8 tr 4 tr 12 tr Hemiptera 1 tr 6 tr 7 tr Collermbola 3 trº 1 tr 4 tr (continued, next page) 13 Table 3. --concluded June August Total Taxa Num- Per Nurm — Per Num — Per ber cent ber Cent ber cent Megaloptera 3 tr O 0 3 tr Lepidoptera 1 tr 0 0 1 tr Hymenoptera 1 tr 0 O 1 tr Hydrac arina - 97 2 149 3 246 2 Araneae - 2 tr O 0 2 tr Gastropoda Physidae - 2 tr 498 9 500 4 Pelecypoda - 2 tr 7 tr 9 tr Oligochaeta 90 1 61 1 151 1 Hirudinea 3 tr 11 tr 14 tr Nematoda 11 tr 7 tr 18 tr Fish" 6 tr 2 tr 8 tr Unidentified 0 0 1 tr 1 tr Totals 5, 850 5, 321 11, 171 Includes sac fry of Catostomidae and Cyprinidae. 14 Table 4. --Percentages of total number and weight of organisms found in bottom samples and in the stomachs of the rainbow trout and the mottled sculpin, Little Garlic River, June and August 1970 Percentage of total Percentage of total Taxa number weight Bottom Rainbow Mottled Bottom Rainbow Mottled samples trout sculpins samples trout sculpins Diptera Tendipedidae 49. 2 37. 2 47. 1 4.4 5. 8 4. 2 Other 5. 6 10. 0 7.4 1. 8 6.4 3. 8 Trichoptera - Hydropsychidae 6.7 4. 7 14. 7 62. 5 32. 1 55. 0 Limnephilidae 3. 5 1. 8 2. 2 2. 6 1.0 1. 7 Rhyacophilidae 3. 8 0. 5 0. 0 6. 0 3. 8 0. 0 Other 3. 1 2.4 5.4 1. 8 2.8 5. 5 Ephemeroptera Baetidae 13. 6 28. 3 8.0 5. 8 24. 5 3. 2 Other - 2. 3 1.2 4. 2 1.9 1.4 6, 7 Other insecta 6. 1 5. 6 2.9 9. 5 10. 0 8. 0 Gastropoda 4. 5 7. O 6.4 1. 5 7.2 5. 9 Oligochaeta 1. 6 1. 3 1. 7 2. 2 5. 0 6. 0 15 Table 5. --Analysis of variance of the total diet of the rainbow trout and the mottled sculpin in the Little Garlic River, June and August 1970 . Degrees Sums - F Source of variation Of Of Mean freedom squares square Observed Expected" Months 1 2. 14 2. 14 0. 72 4. 35 Food groups * 5 19. 38 3. 88 1. 32 2. 71 Fish species 1 5. 28 5, 28 1. 79 4. 35 Months X Orders 5 9. 76 1. 95 0.66 2. 71 Months X species 1 5.36 5.36 1. 82 4.35 Orders X species 5 10. 81 2. 16 0. 73 2. 71 Months X Orders X species 5 29. 56 5. 91 2. 00 2. 71 Error 20 59. 06 2. 95 Total 43 141. 55 * p = 0.05. See Table 4 for food groups with families as replicates. 16 Even though rainbow trout fry were present in ever- increasing numbers in the study section of the stream during June to August, no fry were found in stomachs of either rainbow trout Or mottled sculpins. The only fish encountered as a food item of another fish was a young-of-the-year mottled sculpin in an age II mottled sculpin. Food Consumption Total food consumption during the 57 days of the study was 29.2 kg, of which rainbow trout ate 18.2 kg, while mottled sculpins ate 11.0 kg (Table 2). Rainbow trout consumed 319.7 g of food per day (0.2 g/m2) which was 2% by weight of the mean standing crop of benthos. Mottled sculpins ate 193. 1 g of benthos per day (0. 1 g/m2) or 1.2% of the mean benthic biomass. DISCUSSION Production of the rainbow trout in the Little Garlic River was within the range of values found elsewhere by others, particularly in salmonid nursery streams (Table 6). Only one report (Allen, 1951) had substantially greater salmonid production, but he was able to compute values for more age groups than I did. Production values for salmonids of corresponding age groups (Goodnight and Bjornn, 1971; Chapman, 1965) are similar to my findings. Information on sculpin production is scant, but Goodnight and Bjornn (1971) reported much lower production values for Cottus sp. than I found for the mottled sculpin (Table 6). A comparison of food habits and bottom fauna abundance revealed that young rainbow trout and mottled sculpins fed on the various bottom fauna in approximate proportion to their abundance in the stream. An analysis of variance test confirmed that the diets of these two fishes were similar. No occurrences of mutual predation between rainbow trout and mottled sculpins were found during this study. Even though large numbers of the fry of both fishes were available for predation, only one young mottled sculpin was eaten by a larger sculpin. This supports the contentions by Koster (1939) and Patten (1962, 1971) that sculpin predation on trout and salmon fry in streams can be negligible. My finding of an absence of predation by the rainbow trout on young of the mottled sculpin is augmented by McAfee (1966) who contended that in general, juveniles of the rainbow trout in freshwater eat mainly aquatic and terrestrial insects. I could not accurately measure the extent of food competition. My experimental design provided me only with data on standing crop of bottom fauna. Total food consumption by the rainbow trout and 17 18 Table 6. --Production of trout, salmon, and sculpins in various Streams, 15 June – 15 August Fish e d Produc – Investigators Stream lS sº º tion - - ge g D (g/m2) Present study Little Garlic R., Rainbow trout 0–I 2. 8 Michigan Mottled sculpin I-II 1. 5 Goodnight and Big Springs Cr. , Rainbow trout 0-I 3. 8 Bjornn (1971) Idaho Cottus sp. I-II 0. 5 Lemhi R. , Rainbow trout 0–I 0. 6 Idaho Cottus sp. I-II 0.3 Chinook salmon 0-I 1. 7 Chapman (1965) Deer Cr., Coho salmon 0-I 1.5 Oregon Flyn Cr. , Coho salmon O-I 1. 3 Oregon Needle Branch, Coho salmon 0 — I 1. 2 Oregon Hunt (1966) Lawrence Cr. , Brook trout 0-IV 1.6 Wisconsin Section A Lawrence Cr. , Brook trout 0-IV 2. 9 Wisconsin . Section B Allen (1951) Horokiwi Stream, Brown trout A11 9. 0 New Zealand 19 mottled sculpin for 57 days was 1.9 times the mean standing crop of bottom fauna. Bottom fauna production values reported by other investigators indicate that this level of consumption probably was much less than the amount of benthos produced in the Little Garlic River. Allen (1951) estimated that the total annual production of bottom fauna in the Horokiwi Stream must have been 40–150 times the average standing crop, based on fish production. Allen's production estimates are regarded as being too high by Mann (1967). Mann reported that in the littoral zone of lakes and in streams where the fish population is at the maximum level permitted by food resources, annual food production may be 10 or more times the average standing crop of food. In support of this view, annual productivity of bottom fauna living on a rock-outcrop in a southern Piedmont stream was reported by Nelson and Scott (1962) to be 11-12 times the mean standing crop. Daily food consumption by the rainbow trout was 2.0% of the mean biomass of bottom fauna; the mottled sculpin consumed 1.2%. This rate of cropping was substantially lower than that which Hopkins (1970) found for fish in a brown trout nursery stream. He concluded that a 5 to 20% daily consumption of the standing benthic population represented a high rate of cropping. Based on the low total consumption and low daily cropping rate, food competition between rainbow trout and mottled sculpins in the Little Garlic River probably was not detrimental to either because more food was produced than the fishes could consume. CONCLUSIONS Populations of the rainbow trout and the mottled sculpin in the Little Garlic River exhibited production comparable to salmonids in other waters, particularly salmonid nursery streams. With minor exceptions, the food habits of the trout and sculpins in the Little Garlic River were similar. Mutual predation did not occur between or among the age and size groups studied. Detrimental food competition probably did not occur because of the low rate of food consumption compared to the abundance of food organisms. 20 LITERATURE CITED Allen, K. R. 1951. The Horokiwi Stream. New Zealand Mar. Dep. , Fish. Bull. No. 10, 231 p. Anderson, R. O. 1959. A modified flotation technique for sorting bottom fauna samples. Limnol. Oceanogr., 4 (2): 223-225. Bailey, N. J. J. 1951. On estimating the size of mobile populations from recapture data. Biometrika, 38: 293–306. Bonham, K. 1949. Some tests with experimental groups of fingerling rainbow trout, Salmo gairdnerii, on uniformity and rate Of growth, diet and photographic size recording. Trans. Amer. Fish. Soc. , 79: 94-104. Brocksen, R. W., G. E. Davis, and C. E. Warren. 1968. Competition, food consumption, and production of sculpins and trout in laboratory stream communities. J. Wildl. Manage. , 32(1): 51-75. Chapman, D. W. 1965. Net production of juvenile coho salmon in three Oregon streams. Trans. Amer. Fish. Soc. , 94 (1): 40–52. Clary, J. R. 1972. Predation on the brown trout by the slimy sculpin. Prog. Fish-Cult. , 34(2): 91-95. Dineen, C. F. 1951. A comparative study of the food habits of Cottus bairdii and associated species of Salmonidae. Amer. Midl. Nat. , 46 (3): 640-645. Freund, J. E. 1952. Modern elementary statistics. 3rd ed. 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Techniques for studying herring scales and otoliths. Prog. Fish-Cult. , 22(2): 84-86. Mann, K. H. 1967. The cropping of the food supply, p. 243-257. In Blackwell Scientific Publications, The biological basis of freshwater fish production [Shelby D. Gerking (ed.)], Oxford and Edinburgh. McAfee, W. R. 1966. Rainbow trout. p. 192–215. In Calif. Dep. Fish Game, Inland fisheries management [Alex Calhoun (ed.)], Sacramento. Nelson, D. J., and D. C. Scott. 1962. Role of detritus in the productivity of a rock-Outcrop community in a Piedmont stream. Limnol. Oceanogr., 7: 396-4 13. Patten, B. G. 1962. Cottid predation upon salmon fry in a Washington stream. Trans. Amer. Fish. Soc., 91: 427-429. Patten, B. G. 1971. Predation by sculpins on fall chinook salmon, Oncorhynchus tshawytscha, fry of hatchery origin. Nat. Mar. Fish. Serv. , Spec. Sci. Rep.--Fish No. 62, 13 p. Pennak, R. W. 1953. Freshwater invertebrates of the United States. The Ronald Press Co., New York, 769 p. Phillips, R. W., and E. 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