Diet of the Round Goby in the St. Clair River and Lake St. Clair, 1993

Similar documents
Charter Boat Catch and Effort from the Michigan Waters of the Great Lakes, 1994

STUDY PERFORMANCE REPORT

Summary of 2012 DNR and Partners Fisheries Surveys in St. Clair/Lake Erie and Fishery Forecast for 2013

The Round Goby Botulism Connection. Renea A. Ruffing Graduate Research Assistant Penn State University

Charter Boat Catch and Effort from the Michigan Waters of the Great Lakes, 1998

Zooplankton Availability to. Larval Walleye (Sander vitreus) in Black Lake, MI, USA

INDEX WORDS: Round goby, zebra mussels, sphaeriid clams, Great Lakes.

JadEco, LLC PO BOX 445 Shannon, IL 61078

Lake St. Clair Fish Community and Fishery

STUDY PERFORMANCE REPORT

Changes in Mottled Sculpin and Johnny Darter Trawl Catches after. the Appearance of Round Gobies in the Indiana Waters of Lake.

Introduction: JadEco, LLC PO BOX 445 Shannon, IL 61078

Status of Lake Erie s Western Basin Fish Populations: Trends and Environmental Conditions

STUDY PERFORMANCE REPORT

Exploring the relationship between native smallmouth bass and invasive mussels in the Huron Erie Corridor

DNR. Manual of Fisheries Survey Methods II: with Periodic Updates STATE OF MICHIGAN DEPARTMENT OF NATURAL RESOURCES FISHERIES DIVISION SPECIAL REPORT

Cedar Lake Comprehensive Survey Report Steve Hogler and Steve Surendonk WDNR-Mishicot

Charter Boat Catch and Effort from the Michigan Waters of the Great Lakes, 1996

MINNESOTA DEPARTMENT OF NATURAL RESOURCES DIVISION OF FISHERIES AND WILDLIFE. Gamefish Assessment Report

STUDY PERFORMANCE REPORT

Arizona Game and Fish Department Region VI Fisheries Program

During the mid-to-late 1980s

LAKE DIANE Hillsdale County (T8-9S, R3W, Sections 34, 3, 4) Surveyed May Jeffrey J. Braunscheidel

STUDY PERFORMANCE REPORT

Charter Boat Catch and Effort from the Michigan Waters of the Great Lakes, 2001

Lake Monitoring Program: Lesser Slave Lake Stock Assessment

Tahquamenon River Assessment Appendix

Status of the Lake St. Clair Fish Community and Sport Fishery,

Lake Winnibigoshish Fisheries Information Newsletter

Impact of Round Goby Predation on Zebra Mussel Size Distribution at Calumet Harbor, Lake Michigan

In each summer issue of Lake

Results of the 2015 nontidal Potomac River watershed Smallmouth Bass Young of Year Survey

Comparative Survival of Pellet-Reared Muskellunge Stocked As Fingerlings In Bluegill Ponds With and Without Largemouth Bass

Crawford Reservoir. FISH SURVEY AND MANAGEMENT INFORMATION Eric Gardunio, Fish Biologist Montrose Service Center

FOX AND COYOTE TRAPPING SURVEY

TABLE ROCK LAKE 2014 ANNUAL LAKE REPORT. Shane Bush Fisheries Management Biologist Missouri Department of Conservation Southwest Region

Michigan Dept. of Natural Resources Status of the Fishery Resource Report Page 1. Weber Lake Cheboygan County, T34N, R3W, Sec.

Nearshore Fish Community of Lake Michigan: 2017 Summer Harbor Assessment

STUDY PERFORMANCE REPORT

Arizona Game and Fish Department Region I Fisheries Program. Chevelon Canyon Lake Fish Survey Report Trip Report April 2015

Predation on Zebra Mussels by Freshwater Drum and Yellow Perch in Western Lake Erie

Tittabawassee River Assessment. Miles. Gladwin Smallwood Impoundment. Harrison. Clare. Midland. Mt. Pleasant. St. Louis. Saginaw.

STUDY PERFORMANCE REPORT

Behavioral Interactions Between Round Gobies (Neogobius melanostomus) and Mottled Sculpins (Cottus bairdi)

The presence of exotic species, such as zebra mussels, can often become problematic

Hooked on. Tagging study. Inside this issue: Volume: #2 July Tagging study

BIG TWIN LAKE Kalkaska County (T28N, R05W, Section 18, and T28N, R06W, Section 13) Surveyed May 1999

Caro Impoundment, Tuscola County

2017 Glad Lake Pike Transfer Summary of Activities

CORRESPONDENCE/MEMORANDUM

feeding - clear moderate-sized shallow streams with moderate vegetation spawning - nests in gravel, sand, or hard rock substrate

Rainy Lake Open-water Creel Survey:

Status of the Lake Erie Fish Community Jeff Tyson, Ohio Division of Wildlife

The Walleye Fishery of the Detroit River, Spring 2000

2011 Haha Lake Northern Pike Control

Non-indigenous Fish Management Information Fisheries Division 2005

MARTINDALE POND Wayne County 2004 Fish Management Report. Christopher C. Long Assistant Fisheries Biologist

Protection Measures Against Exotic Species

Ohio Lake Erie Fisheries Report For 2002

Diet Study of Walleye in Black Lake Using Isotope Analysis of d15n and d13c

Laurentian Great Lakes and their invasive species. Jeff Gunderson Minnesota Sea Grant College Program

Mille Lacs Lake Creel Survey Report for. Open Water Season of and. Winter Season of

STUDY PERFORMANCE REPORT

2011 Status of Major Stocks. Lake Erie Management Unit March 2012

Aquatic Invasive Species

Countermeasures against Alien Fishes (Largemouth Bass and Bluegill) in Lake Biwa

Dauphin Lake Fishery. Status of Walleye Stocks and Conservation Measures

Lake Butte des Morts Commercial Seining Project Report

Student Worksheet: River Health and Indicator Species

FISH COMMUNITIES AND FISHERIES OF THE THOUSAND ISLANDS AND MIDDLE CORRIDOR

Minnesota Department of Natural Resources Division of Fisheries and Wildlife. Winter Creel Survey Report for Leech Lake

Current Status and Management Recommendations for the Fishery in the Cloverleaf Chain of Lakes

Susquehanna River Walleye Fishery

Status of Lake St. Clair Submerged Plants, Fish Community, and Sport Fishery

Summary of and Initial Response to public comments on MN Department of Natural Resources proposal to manage new waters for Muskellunge

SKIATOOK LAKE MANAGEMENT PLAN

Onondaga Lake Fishery: 2011 Fact Sheet

Relative Size Selectivity of Trap Nets for Eight Species of Fish'

Geology. Key Factors. Overfishing. Great Lakes Fishes. Historical Fishing. About 10,000 years since last glacial retreat very young ecologically

Greetings By Rick Bruesewitz MNDNR Aitkin Area Fisheries Supervisor

NEVADA DEPARTMENT OF WILDLIFE STATEWIDE FISHERIES MANAGEMENT

Recovery and Decline of Lake Whitefish in U.S. Waters of Eastern Lake Ontario,

The impact of fish predation on invasive rusty crayfish (Orconectes rusticus) populations in two northern temperate lakes

STUDY PERFORMANCE REPORT

Lake LeAnn 2018 Fish Survey

HUBBARD LAKE Alcona County (T27N, R7E; T28N, R7E) Surveyed May and September Tim A. Cwalinski

Fish Survey of Arctic Lake (ID # ), Scott County, Minnesota in 2012

Cast Net Report. As required by Minnesota Statutes, section 97C.345, subdivision 3a, paragraph (c) 03/01/2018. Cast Net Report 1

BENSON PARK POND FISH SPECIES

Michigan Department of Natural Resources Status of the Fishery Resource Report Page 1

Mille Lacs Lake Creel Survey Report for. Open Water Season of and. Winter Season of

Largemouth Bass. Max 30 inches Average 4-12 inches EATS

MIDDLE FORK RESERVOIR Wayne County 2004 Fish Management Report. Christopher C. Long Assistant Fisheries Biologist

Aquatic Plant Management and Importance to Sport Fisheries

Appendix C Bill Cross Rapids Freshwater Mussel and Fishery Study Plan

The relationship between the spatial distribution of common carp and their environmental DNA in a small lake

Lake information report

Costs of Rearing and Stocking Walleye in Michigan During 1994

MISSISSIPPI MAKEOVER A Plan for Restoration, Just Around the Bend

STUDY PERFORMANCE REPORT

Transcription:

Michigan DNR STATE OF MICHIGAN DEPARTMENT OF NATURAL RESOURCES Number 96-2 April 30, 1997 Diet of the Round Goby in the St. Clair River and Lake St. Clair, 1993 Michael V. Thomas www.dnr.state.mi.us FISHERIES DIVISION TECHNICAL REPORT

MICHIGAN DEPARTMENT OF NATURAL RESOURCES FISHERIES DIVISION Fisheries Technical Report 96-2 April 30, 1997 DIET OF THE ROUND GOBY IN THE ST. CLAIR RIVER AND LAKE ST. CLAIR, 1993 Michael V. Thomas The Michigan Department of Natural Resources (MDNR), provides equal opportunities for employment and access to Michigan s natural resources. Both State and Federal laws prohibit discrimination on the basis of race, color, national origin, religion, disability, age, sex, height, weight or marital status under the Civil Rights Acts of 1964, as amended, (1976 MI P.A. 453 and 1976 MI P.A. 220, Title V of the Rehabilitation Act of 1973, as amended, and the Americans with Disabilities Act). If you believe that you have been discriminated against in any program, activity or facility, or if you desire additional information, please write the MDNR Office of Legal Services, P.O. Box 30028, Lansing, MI 48909; or the Michigan Department of Civil Rights, State of Michigan, Plaza Building, 1200 6 th Ave., Detroit, MI 48226 or the Office of Human Resources, U. S. Fish and Wildlife Service, Office for Diversity and Civil Rights Programs, 4040 North Fairfax Drive, Arlington, VA. 22203. For information or assistance on this publication, contact the Michigan Department of Natural Resources, Fisheries Division, Box 30446, Lansing, MI 48909, or call 517-373-1280. This publication is available in alternative formats. Michigan DNR Printed under authority of Michigan Department of Natural Resources Total number of copies printed 200 Total cost $223.36 Cost per copy $1.11

Michigan Department of Natural Resources Fisheries Technical Report No. 96-2, 1997 Diet of The Round Goby in the St. Clair River and Lake St. Clair, 1993 Michael V. Thomas Michigan Department of Natural Resources Lake St. Clair Fisheries Research Station 33135 South River Road Mt. Clemens, MI 48045 Abstract. The round goby Neogobius melanostomus, a species indigenous to the Black and Caspian Sea region of Europe, was first discovered in the St. Clair River in 1990. Since that time, they have expanded into Lake St. Clair and have also been collected from Lake Michigan and Lake Erie. Food habits of the round goby in the St. Clair River and Lake St. Clair have not been previously described. The stomach and intestine of round gobies obtained in 1993 from St. Clair River anglers (84 fish) and Lake St. Clair survey trawls (39 fish) were examined. Only one of the 123 fish examined was empty. The most common food item consumed for both locations was the non-indigenous zebra mussel Dreissena polymorpha. The only other important component of the diet for St. Clair River fish was caddisfly larvae. Zebra mussels were found in the diet of 96% of all St. Clair River round gobies; in contrast, the diet for Lake St. Clair round gobies consisted of amphipods, snails, ostracods, fingernail clams, caddisfly and chironomid larvae, in addition to zebra mussels. Although zebra mussels were found in 67% of all Lake St. Clair round gobies, the frequency of occurrence for several other food items, such as amphipods (63%), was relatively high. The study revealed little evidence of piscivory in round gobies from either area. Only one fish, a 28mm brook stickleback Culaea inconstans was found in the 123 round gobies examined. Two species of gobies, the tubenose goby Proterorhinus marmoratus and round goby Neogobious melanostomus, were discovered in the St. Clair River in 1990. Jude et al. (1992) described the initial discovery of these two species and briefly discussed their possible effects on the native benthic fish community. Since 1990, round gobies have become a nuisance species for walleye Stizostedion vitreum anglers in the St. Clair River. Round gobies aggressively attack earthworms, the preferred bait for walleye in the St. Clair River. In fact, many long-time walleye fishermen believe the abundance of round gobies and their appetite for earthworms is impeding walleye fishing success in the river. Tubenose gobies have not been caught by anglers. Observations by recreational divers in the St. Clair River also suggest that the abundance of round gobies has increased dramatically since 1990. These same anecdotal reports indicate that abundance of previously common species such as logperch Percina caprodes and sculpins Cottus spp. declined sharply during the same time period. We enlisted the help of walleye anglers in collecting round gobies from the St. Clair River. We also incidentally caught round and tubenose gobies in trawls in Lake St. Clair, while collecting yellow perch Perca flavescens for a 1

separate research project. The objective of this study was to document the food habits of the round goby in the St. Clair River and Lake St. Clair during summer 1993. Methods Round gobies were collected from recreational anglers targeting walleye in the St. Clair River during July and August, 1993, near Marine City and Algonac, Michigan (Figure 1). All were taken using earthworms as bait. Most were placed on ice immediately upon capture and frozen whole within 6 h for later diet analysis. Round gobies from Lake St. Clair were collected with a bottom trawl. Trawl collections were made in August, September, and October, 1993. The trawl used was a bottom trawl with a 10-m headrope and 9.5-mm stretched mesh (cod end), and was towed by the R/V Channel Cat at approximately 2.0 mile per hour. Round gobies were immediately placed on ice and frozen whole within 8 hours for later diet analysis. Round gobies were thawed in the lab, and individual length, weight, and sex recorded. Scale samples were taken for age analysis. The anterior portion of the digestive tract was slightly enlarged, but a distinct stomach was not apparent. Therefore, the entire digestive tract was removed and all contents were placed in 70% ethyl alcohol. Food items were later identified to the lowest possible taxon. The diet was analyzed for "percentage composition by number" and "frequency of occurrence" as described by Windell and Bowen (1978). Percent composition by number is the percentage of the total number of food items examined accounted for by a selected taxa. Frequency of occurrence describes the percentage of fish with non-empty stomachs that contained at least one of a selected food item. Length of all intact zebra mussel shells were measured to the nearest millimeter for a subsample of 24 St. Clair River fish. Biological Data Results A total of 84 round gobies were collected from walleye anglers fishing the St. Clair River. The fish ranged from 61 to 133 mm in total length, with an average length of 110 mm. Average weight for round gobies from St. Clair River was 24.3 g. Aging by scales was possible in 79 of the fish (Table 1). Ages ranged from 1 to 3 yr, with a mean age of 2. Males accounted for 74% of the sample. Thirty-nine round gobies were collected by bottom trawl in Lake St. Clair. These fish ranged from 32 to 117 mm in total length, with an average length of 83 mm. Average weight for round gobies from Lake St. Clair was 10.8 g. Only 16 fish could be aged by scales (Table 1). Ages ranged from young of the year to 2 yr, with a mean age of 0.7 yr. Males made up 79% of the sample. Diet Analysis Percent Composition of Diet Zebra mussels Dreissena polymorpha dominated the diet of fish from St. Clair River, accounting for 62% of the 1,409 food items examined (Table 1). Other numerically important components of the diet included caddisfly cases and larvae, as well as rocks. Zebra mussel byssal threads were attached to many of the rocks examined, suggesting that they were inadvertently consumed while foraging on zebra mussels. The largest number of zebra mussels found in a single fish was 25; the fish was 110 mm and 23.94 g. The mean number of zebra mussels per non-empty stomach was 11. No significant relationship was evident between round goby weight and the number of zebra mussels found in the gut (Figure 2). No fish were found in stomachs of round gobies from the St. Clair River. Zebra mussels were also the most abundant component of the diet for round gobies from Lake St. Clair, accounting for 34% of the total of 370 food items examined (Table 2). Round 2

gobies from Lake St. Clair consumed a more varied diet, with amphipods, ostracods, snails, fingernail clams, caddisfly larvae, and chironomid larvae also numerically important in the diet. The highest number of zebra mussel shells found in a single fish was 16 (the fish was 101 mm and 15.22 g) with a mean number of 5 mussels per non-empty stomach. The relationship between number of zebra mussels in the gut and weight of round gobies (Figure 2) was significant (F = 0.0001, R 2 = 0.32) with smaller gobies containing fewer zebra mussels. One fish, a 28 mm brook stickleback Culaea inconstans, was found in a Lake St. Clair round goby (110 mm, 21.8 g). Frequency of Occurrence Zebra mussels were found in 96% of the non-empty St. Clair River round gobies examined (Table 1). Rocks were the next most frequently occurring item found. Other taxa identified in more than 20% of the non-empty stomachs included caddisfly larvae and cases, fingernail clams, and snails. For Lake St. Clair, zebra mussels occurred in 68% of the non-empty stomachs examined (Table 2). Amphipods were nearly as frequent (63%). Other taxa occurring in more than 20% of the non-empty stomachs included fingernail clams, caddisfly larvae, snails, ostracods, chironomid larvae, and caddisfly cases. Lengths of Zebra Mussels Consumed A total of 139 whole zebra mussel shells were found in the stomach and intestines of a subsample of 24 round gobies from the St. Clair River. Shell lengths ranged from 3 to 12 mm, with a mean of 6.9 mm. Regression analysis indicated no significant relationship between goby total length and mean length of whole zebra mussel shells found in the digestive tract (Figure 3). Discussion Collections made for this study indicate that round gobies are well established in the St. Clair River, and are successfully colonizing Lake St. Clair. Fish of three age-groups were collected from both areas, with young of the year included in trawl catches from Lake St. Clair. If these fish are capable of colonizing Lake St. Clair, with its abundant populations of Great Lakes muskellunge Esox masquinongy, walleye, smallmouth bass Micropterus dolomieu, largemouth bass Micropterus salmoides, northern pike Esox lucius, rock bass Ambloplites rupestris, yellow perch, and channel catfish Ictalurus punctatus, I believe they will have no problem colonizing most inshore waters of the lower Great Lakes. Round gobies collected from the St. Clair River were larger and older than those collected from Lake St. Clair. It is probable that the difference in collection methods between the two areas is the main reason for these differences. Fishing with hook and line, the method of collection for the St. Clair River gobies, should select for larger individuals than trawling. Perhaps younger and smaller fish are actually more common in Lake St. Clair, since it was colonized more recently than the St. Clair River. However, this study does not provide the data needed to draw that conclusion. It is clear that the round goby, an exotic fish species, forages heavily on another exotic species, the zebra mussel, in the St. Clair River and Lake St. Clair. As many as 25 zebra mussels, ranging in length from 3 to 12 mm, were found in a single round goby during this field study. In a laboratory study, Ghedotti et al. (1995) found round gobies consumed an average of 5.4 zebra mussels per hour, ranging in length from 4 to 13 mm. Few native fish species are expected to consume zebra mussels (French 1993). Thus, round gobies may have a competitive advantage over native species which may be competing with each other for shared food resources. Although, direct competition for food resources between round gobies and native benthic fishes in the St. Clair River may not be intense, round gobies may still interact in other ways with those fish. Jude et al. (1992) 3

suggested that round gobies may influence darters Etheostoma spp., sculpins Cottus cognatus, and logperch Percina caprodes through predation on eggs, young and yearlings. I found little evidence of piscivory in round gobies collected from either the St. Clair River or Lake St. Clair in 1993. In fact, only 1 fish was found among the nearly 1800 food items in the 123 round gobies examined. However, soft tissued food items such as fish eggs or larvae, are infrequently encountered in diet studies due to their quick digestion time. As a result, predation by round gobies on fish eggs or young of the year may be underestimated by this study. Further, even low levels of piscivory by round gobies could be important at high population densities. The St. Clair River has a higher taxonomic diversity of macrozoobenthos than Lake St. Clair (Edsall et al. 1988). Despite this, round gobies from Lake St. Clair consumed a more diverse diet than those collected from the St. Clair River. The reason for this is not clear. Perhaps areas in the St. Clair River where round gobies are routinely captured by anglers are characterized by benthic diversity much lower than the rest of the St. Clair River. Additionally, differences in the ages and sizes of gobies collected in the two areas may be involved. Finally, zebra mussel densities may be much higher at the collection sites in the St. Clair River than in Lake St. Clair, allowing the fish to forage more selectively. Zebra mussels have been viewed as rechanneling energy flow from algae and zooplankton to the benthic macroinvertebrate community (Griffiths 1992). If round gobies are preyed upon by walleye or other game fish species, they offer a direct energy link from zebra mussels to piscivorous game fish. Unfortunately, there is little evidence from sport anglers to suggest that game fish species feed consistently on round gobies, and no concentrated survey effort targeting potential goby predators has been conducted. This study was based on round gobies caught incidentally by survey bottom trawls in Lake St. Clair and sport anglers in the St. Clair River. While this small sample has provided a brief glimpse into the diet of the round goby in these waters, many unanswered questions surround the round goby and its effects on the fish community. As this species continues to expand its range and population densities across the Great Lakes, increased public awareness and concern over this newest of exotics is inevitable. A thorough, wellplanned study to investigate the effects of both goby species on the Lake St. Clair and St. Clair River fish community would be a prudent effort. 4

Lake Huron St. Clair River Marine City Algonac Lake St. Clair Detroit River Figure 1. Map of St. Clair River and Lake St. Clair showing areas of round goby collections by angling (cross hatched) and trawling (gray stippled). 5

25 St. Clair River Number of mussels in gut 20 15 10 5 0 0 10 20 30 40 50 Fish weight (g) 16 Lake St. Clair Number of mussels in gut 12 8 4 0 0 5 10 15 20 25 30 Fish weight (g) Figure 2. Relationship between the number of zebra mussels found in the digestive tract and the weight of round gobies from the St. Clair River (above) and Lake St. Clair (below). 6

10 Mean zebra mussel length (mm) 8 6 4 80 90 100 110 120 130 140 Fish length (mm) Figure 3. Mean length of zebra mussels consumed by 24 round gobies in the St. Clair River. 7

Table 1. Mean length and weight at age for round gobies Neogobius melanostomus collected in 1993. One SE is given in parentheses. Age 0 I II III St. Clair River Mean length (mm) 90 (17) 113 (10) 118 (10) Mean weight (g) 11.9 (6.1) 25.4 (7.9) 28.6 (7.9) Percent males 60 79 69 Number 0 10 56 13 Lake St. Clair Mean length (mm) 53 (12) 97 (12) 110 Mean weight (g) 2.6 (1.4) 14.6 (4.6) 21.8 Percent males 43 78 0 0 Number 7 9 1 0 8

Table 2. Taxa accounting for >5% by number of total stomach contents of round gobies collected in 1993 from the St. Clair River. n = 84. Zebra Caddisfly Caddisfly mussel Rocks cases larvae Percent composition of diet: Total 874 104 192 113 Mean 11 2 6 3 Range 1-25 1-13 1-11 1-44 % of diet 62.0 7.4 13.6 8.0 Frequency of occurrence: Stomachs containing at least 1 81 44 34 34 % of non-empty stomachs 96.4 52.4 40.5 40.5 9

Table 3. Taxa accounting for >5% by number of total stomach contents of round gobies collected in 1993 from Lake St. Clair. n = 39. Zebra Fingernail Caddisfly Chironomid Seed mussel clam larvae Snail Scuds larvae shrimp Percent composition of diet Total 127 29 22 42 58 20 36 Mean 5 2 2 3 2 2 3 Range 1-16 1-4 1-5 1-7 1-9 1-7 1-7 % of diet 34.3 7.8 5.9 11.4 15.7 5.4 9.7 Stomachs containing at least 1 % of non-empty stomachs Frequency of occurrence: 26 14 13 13 24 10 12 68.4 36.8 34.2 34.2 63.2 26.3 31.6 10

Literature Cited Edsall, T.A., B.A. Manny, and C.N. Raphael. 1988. The St. Clair River and Lake St. Clair, Michigan: An ecological profile. U. S. Fish and Wildlife Service Biological Report 85 (7.3), 130 p. French, J.R.P. 1993. How well can fishes prey on zebra mussels in eastern North America? Fisheries 18(6):13-19. Ghedotti, M.J., J.C. Smihula, and G.R. Smith. 1995. Zebra mussel predation by round gobies in the laboratory. Journal of Great Lakes Research 21:665-669. Griffiths, R.W. 1992. Effects of zebra mussels (Dreissena polymorpha) on the benthic fauna of Lake St. Clair. Pages 415-438 in T.F. Nalepa and D.W. Schloesser, editors. Zebra mussels: biology, impact, and control. CRC Press, Boca Raton, Florida. Jude, D.J., R.H. Reider, and G.R. Smith. 1992. Establishment of Gobiidae in the Great Lakes basin. Canadian Journal of Fisheries and Aquatic Sciences 49:416-421. Windell, J.T. and S.H. Bowen. 1978. Methods for study of fish diets based on analysis of stomach contents. Pages 219-226 in T. Bagnel, editor. Methods for the assessment of fish production in fresh waters, 3rd edition. Blackwell Scientific Publications, Oxford, England. Report approved by Paul W. Seelbach James S. Diana, Editor James E. Breck, Editorial Board Reviewer Alan D. Sutton, Graphics Barbara A. Diana, Word Processor 11