Methods

I. Study System Lake Mývatn, Iceland (65°36 N, 17°0′ W) is a large (38 km²) shallow (4 m max depth) lake divided into two large basins that function mostly as independent hydrologic bodies (Ólafsson 1979). The number of non-biting midge (Diptera: Chironomidae) larvae on the lake bottom is high, but variable: midge production between 1972-74 ranged from 14-100 g ash-free dw m^-2 yr^-1, averaging 28 g dw m^-2 yr^-1 (Lindegaard and Jónasson 1979). The midge assemblage is mostly comprised of two species (> 90% of total individuals), Chironomus islandicus (Kieffer) and Tanytarsus gracilentus (Holmgren) that feed as larvae in the sediment in silken tubes by scraping diatoms, algae, and detritus off the lake bottom (Lindegaard and Jónasson 1979). At maturity (May-August) midge pupae float to the lake surface, emerge as adults, and fly to land, forming large mating swarms around the lake (Einarsson et al. 2004, Gratton et al. 2008). On land, midges are consumed by terrestrial predators (Dreyer et al. 2012, Gratton et al. 2008), or enter the detrital pool upon death (Gratton et al. 2008, Hoekman et al. 2012). Midge populations naturally cycle with 5-8 year periodicity, with abundances fluctuating by 3-4 orders of magnitude (Einarsson et al. 2002, Ives et al. 2008). II. Midge Emergence Measurement We used submerged conical traps to estimate midge emergence from Lake Mývatn. Traps were constructed of 2 mm clear polycarbonate plastic (Laird Plastics, Madison, WI) formed into a cone with large-diameter opening of 46 cm (0.17 m²). The tops of the cones were open to a diameter of 10 cm, with a clear jar affixed at the apex. The trap was weighted to approximately neutral buoyancy, with the jar at the top containing air to allow mature midges to emerge. Traps were suspended with a nylon line ~1 m below the surface of the lake from an anchored buoy. For sampling, traps were raised to the surface and rapidly inverted, preventing midges from escaping. Jars and traps were thoroughly rinsed with lake water to collect all trapped midges, including unmetamorphosed larvae and pupae, and scrubbed before being returned to the lake to prevent growth of epiphytic algae and colonization by midges. We assume that the emergence traps collect all potentially emerging midges from the sampling area, though it is likely an underestimate, since some midges initially captured could fall out of the trap. Thus, our results should be considered a conservative estimate of potential midge emergence from the surface of the lake.We sampled midge emergence throughout the south basin of Lake Mývatn. Emergence was sampled at six sites in 2008 and 2011 and ten sites in 2009 and 2010, with locations relocated using GPS and natural sightlines. Each site had two traps within 5 m of each other that were monitored during midge activity, approximately from the last week of May to the first week of August. Midge emergence outside of this time frame is extremely low (Lindegaard & Jónasson 1979) and we assume it to be zero. Traps were checked weekly during periods of high emergence (initial and final 2-3 weeks of the study), and bi-weekly during low emergence periods in the middle of the study (July). III. Identification, Counts, and Conversions Midges were counted and identified to morphospecies, small and large. The midge (Diptera,Chrionomidae) assemblage at Mývatn is dominated by two species, Chironomus islandicus (Kieffer)(large, 1.1 mg dw) and Tanytarsus gracilentus (Holmgren)(small, 0.1 mg dw), together comprising 90 percent of total midge abundance (Lindegaard and Jonasson 1979). First, the midges collected in the infall traps were spread out in trays, and counted if there were only a few. Some midges were only identified to the family level of Simuliidae, and other arthropods were counted and categorized as the group, others. Arthropods only identified to the family level Simuliidae or classified as others were not dually counted as Chironomus islandicus or Tanytarsus gracilentus . If there were many midges, generally if there were hundreds to thousands, in an infall trap, subsamples were taken. Subsampling was done using plastic rings that were dropped into the tray. The rings were relatively small compared to the tray, about 2 percent of the area of a tray was represented in a ring. The area inside a ring and the total area of the trays were also measured. Note that different sized rings and trays were used in subsample analysis. These are as follows, trays, small (area of 731 square centimeters), “large1” (area of 1862.40 square centimeters), and large2 (area of 1247 square centimeters). Rings, standard ring (diameter of 7.30 centimeters, subsample area is 41.85 square centimeters) and small ring (diameter of 6.5 centimeters, subsample area is 33.18 square centimeters). A small ring was only used to subsample trays classified as type “large2.”The fraction subsampled was then calculated depending on the size of the tray and ring used for the subsample analysis. If the entire tray was counted and no subsampling was done then the fraction subsampled was assigned a value of 1.0. If subsampling was done the fraction subsampled was calculated as the number of subsamples taken multiplied by the fraction of the tray that a subsample ring area covers (number of subsamples multiplied by (ring area divided by tray area)). Note that this is dependent on the tray and ring used for subsample analysis. Finally, the number of midges in an infall trap accounting for subsampling was calculated as the raw count of midges divided by the fraction subsampled (raw count divided by fraction subsampled).Other metrics such as total insects in meters squared per day, and total insect biomass in grams per meter squared day can be calculated with these data. In addition to the estimated average individual midge masses in grams, For 2008 through 2010 average midge masses were calculated as, Tanytarsus equal to .0001104 grams, Chironomus equal to .0010837 grams. For 2011 average midge masses were, Tanytarsus equal to .000182 grams, Chironomus equal to .001268 grams.

Methods

I. Field MethodsThe site where this manipulative field experiment was conducted on the Kalfastrond peninsula at Lake Myvatn is approximately 150 meters long and 75 meters wide. The vegetation consists of grasses Deschampsia spp., Poa spp., and Agrostis spp.), sedges (Carex spp.), and forbs (Ranunculus acris, Geum rivale,and Potentilla palustris). The experimental midge exclusions occurred from the middle or end of May to the beginning of August in each year, the period corresponding to the active growing season of plants and the flight activity of midges at this site. 2 by 2 meter plots were established in 6 blocks spread across the site. Each block included 3 treatment levels, an open control plot, a full exclusion cage and a partial exclusion cage, for a total of 18 experimental plots. Control plots were open to allow complete access to all arthropods. Experimental midge exclusion cages were 1 meter high and constructed from white PVC tubing affixed to rebar posts on each corner of the plot, Plate 1. Full exclusion cages were entirely covered with white polyester netting, 200 holes per square inch, Barre Army Navy Store, Barre VT, USA, to prevent midges from entering the plot. The mesh netting completely enclosed the 2 by 2 by 1 meter frame to prevent flying insects from entering, however the mesh was not secured to the ground in order to allow non flying,ground crawling, arthropods to freely enter and exit the cages. Partial exclusion cages had one 0.5 meter strip of mesh stretched around the outside of the frame and another 0.75 meter strip draped over the top. Partial cages were designed to examine any effects of cages themselves on terrestrial responses while minimally affecting midge inputs into the plots and arthropod movement.The partial exclusion treatment was discontinued in 2011. Each plot contains a pitfall and an infall trap that are continuously sampled during the summer, while the cages are up. Vacuum samples were taken from the plots about once per month in 2008 through 2010 and only once per summer for subsequent summers.Midge activity was measured in all plots to document changes in midge abundance over the course of a season and between years and to assess the degree to which cages excluded midges. Midge abundance in the plots was continuously measured using passive aerial infall traps consisting of a 1000 milliliter clear plastic cup, 95 square centimeter opening, attached to a post 0.5 meters high and filled with 250 milliliters of a 1 to 1 ethylene glycol to water solution and a small amount of unscented detergent to capture and kill insects that alighted upon the surface. Infall traps were emptied about every 10 days.II. AnalysisMidges were counted and identified to morphospecies, small and large. The midge (Diptera,Chrionomidae) assemblage at Myvatn is dominated by two species,Chironomus islandicus (Kieffer)(large, 1.1 mg dw) and Tanytarsus gracilentus(Holmgren)(small, 0.1 mg dw), together comprising 90 percent of total midge abundance (Lindegaard and Jonasson 1979). First, the midges collected in the infall traps were spread out in trays, and counted if there were only a few. Some midges were only identified to the family level of Simuliidae,and other arthropods were counted and categorized as the group, others. Arthropods only identified to the family level Simuliidae or classified as others were not dually counted as Chironomus islandicus or Tanytarsus gracilentus. If there were many midges, generally if there were hundreds to thousands, in an infall trap,subsamples were taken. Subsampling was done using plastic rings that were dropped into the tray. The rings were relatively small compared to the tray, about 2 percent of the area of a tray was represented in a ring. The area inside a ring and the total area of the trays were also measured. Note that different sized rings and trays were used in subsample analysis. These are as follows, Trays, small (area of 731 square centimeters), large1 (area of 1862.40 square centimeters), and large2 (area of 1247 square centimeters). Rings, standard ring (diameter of 7.30 centimeters, subsample area is 41.85 square centimeters) and small ring (diameter of 6.5 centimeters, subsample area is 33.18 square centimeters). A small ring was only used to subsample trays classified as type large2.The fraction subsampled was then calculated depending on the size of the tray and ring used for the subsample analysis. If the entire tray was counted and no subsampling was done then the fraction subsampled was assigned a value of 1.0. If subsampling was done the fraction subsampled was calculated as the number of subsamples taken multiplied by the fraction of the tray that a subsample ring area covers (number of subsamples multiplied by (ring area divided by tray area)). Note that this is dependent on the tray and ring used for subsample analysis. Finally, the number of midges in an infall trap accounting for subsampling was calculated as the raw count of midges divided by the fraction subsampled (raw count divided by fraction subsampled).Other metrics such as total insects in meters squared per day, and total insect biomass in grams per meter squared day can be calculated with these data. in addition to the estimated average individual midge masses in grams, For 2008 through 2010 average midge masses were calculated as, Tanytarsus equal to .0001104 grams, Chironomus equal to .0010837 grams. For 2011 average midge masses were, Tanytarsus equal to .000182 grams, Chironomus equal to .001268 grams.

Methods

BOOM SHOCKING1987:A standard WDNR electrofishing boat was used on Lake Mendota set at 300 volts and 2.5 amps (mean) DC, with a 20 % duty cycle and 60 pulses per second. On all sampling dates two people netted fish, the total electrofishing crew was three people. Shocking was divided into stations. For each station, the actual starting and ending time was recorded. Starting and ending points of each station were plotted on a nap. A 7.5 minute topographic map (published 1983) and a cartometer was used to develop a standardized shoreline mileage numbering scheme. Starting at the Yahara River outlet at Tenney Park and measuring counterclockwise, the shoreline was numbered according to the number of miles from the outlet. The length of shoreline shocked for each station was determined using the same maps. The objectives of the fall 1987 electrofishing was: to gather CPE data for comparison with previous surveys of the lake; develop a database for relating fall electroshocker CPE to predator density; collect fall predator diet data; make mark-recapture population estimates of YOY predators; and determine year-class-strength of some nonpredators (yellow perch, yellow bass, and white bass).1993: Electrofishing was used to continue marking largemouth and smallmouth bass (because of low CPE in fyke nets), to recapture fish marked in fyke netting, and to mark and recapture walleyes ( less than 11.0 in.) on Lake Mendota. Four person crews electrofished after sunset from May 05 to June 03, 1993. A standard WDNR electrofishing boat was used, set at about 300 volts and 15.0 amps (mean) DC, with a 20 % duty cycle at 60 pulses per second. On all sampling dates two people netted fish; thus, CPE data are given as catch per two netter hour or mile. Shocking was divided into stations. For each station the actual starting and ending time and the generator s meter times was recorded. Starting and ending points of each station were plotted on a map. 7.5 minute topographic maps (published in 1983) were used in addition to a cartometer to develop a standardized shoreline mileage numbering scheme. Starting at the Yahara River outlet at Tenney Park and measuring counterclockwise the shoreline was numbered according to the number of miles from the outlet. The length of shoreline shocked for each station was determined using these maps. The 4 person electroshocker crews were used again from September 20 to October 19. Fall shocking had several objectives: to gather CPE data for comparison with previous surveys of the lake; develop a database for relating fall electroshocker CPE to piscivore density; and make mark recapture population estimates of young of year (YOY) piscivores.1997:5/13/1997-5/20/1997: Electrofishing was completed at night on lakes: Mendota, Monona, and Waubesa. A standard WDNR electrofishing boat was used, set from 320-420 volts and 16-22 amps DC, with a 20 % duty cycle at 50 pulses per second. Two netters were used for each shocking event. At a particular station, starting and ending times where shocking took place were recorded. The location of the designated shocking stations is unknown.9/23/1997-10/14/1997: Electrofishing was completed at night on Mendota, Monona, Waubesa, and Wingra. A standard WDNR electrofishing boat was used, set from 315-400 volts and 16-24 amps DC, with a 20% duty cycle at 60 pulses per second. Two netters were used for each shocking event. Starting and ending time at each shocking station was listed. The location of the designated shocking stations is unknown.1998:Electrofishing was completed at night on Mendota, Monona, Wingra, and Waubesa from 5/12/1998- 10/28/1998. A standard WDNR electrofishing boat was used, set from 240-410 volts and 15-22 amps DC, with a 20% duty cycle at 50-100 pulses per second. Two netters were used for each shocking event. Starting and ending time at each shocking station was listed. The location of the designated shocking stations is unknown. FYKE NETTING1987:Fyke nets were fished daily from March 17 to April 24, 1987 on Lake Mendota. The nets were constructed of 1.25 inch (stretch) mesh with a lead length of 50 ft. (a few 25 ft. leads were used). The hoop diameter was 3 ft. and the frame measured 3 ft. by 6 ft. Total length of the net was 28 ft. plus the lead length. Nets were set in 48 unknown locations. Initially, effort was concentrated around traditional northern pike spawning sites (Cherokee Marsh, Sixmile Creek, Pheasant Branch Creek, and University Bay). As northern pike catch-per-effort (CPE) declined some nets were moved onto rocky shorelines of the lake to capture walleyes. All adult predators (northern pike, hybrid muskie, largemouth and smallmouth bass, walleye, gar, bowfin, and channel catfish) captured were tagged and scale sampled. Measurements on non-predator species captured in fyke nets were made one day per week. This sampling was used to index size structure and abundance, and to collect age and growth data. In each net, total length and weight of 20 fish of each species caught was measured, and the remaining caught were counted.1993:Same methods as 1987, except fyke nets were fished from 4/8/1993-4/29/1993 on Lake Mendota. The 1993 fyke net data also specifies the &ldquo;mile&rdquo; at which the fyke net was set. This is defined as the number of miles from the outlet of the Yahara River at Tenney Park, moving counterclockwise around the lake. In addition, abundance and lengths of non-gamefish species captured in fyke nets were recorded one day per week. Six nets were randomly selected to sample for non-gamefish data. This sampling was used to index size structure and abundance, and to collect age and growth data. In each randomly selected net, total length and weight was measured for 20 fish of each species, and the remaining caught were counted.1998:There is no formal documentation for the exact methods used for fyke netting from 3/3/1998-8/12/1998 on Lake Mendota. However, given that the data is similar to data collected in 1987 and 1993 it is speculated that the same methods were used.MINI-FYKE NETTING1989:There is no formal documentation for the exact methods used for mini-fyke netting on Lake Mendota and Lake Monona from 7/26/1989-8/25/1989. However, given that the data is similar to data collected from 1990-1993 it is speculated that the same methods were used. In the sampling year of 1989, mini-fyke nets were placed at 22 different unknown stations.1990-1993: Mini-fyke nets were fished on Lake Mendota and Lake Monona during July-September at 20, 29, 13, and 15 sites per month during 1990, 1991, 1992, and 1993, respectively to estimate year-class strength, relative abundance, and size structure of fishes in the littoral zone. Nets were constructed with 3/16 in. mesh, 2 ft. diameter hoops, 2 ft. x 3 ft. frame, and a 25 ft. lead. Sites were comparable to seine sites used in previous surveys. Sites included a variety of substrate types and macrophyte densities. To exclude turtles and large piscivores from minifyke nets, some nets were constructed with approximately 2 in. by 2 in. mesh at the entrance to the net. Thus, mini-fyke net data are most accurate for YOY fishes, and should not be used to make inferences about fishes larger than the exclusion mesh size. 1997:There is no formal documentation for the mini-fyke methods which were used on Lake Waubesa and Lake Wingra from 9/16/1997-9/18/1997. However, given that the data is similar to data collected in 1989, and 1990-1993, it is speculated that the methods used during 1997 are the same. SEINE NETTING1989, 1993: Monthly shoreline seining surveys were conducted on Lake Mendota and Lake Monona during June through September to estimate year class-strength, relative abundance, and size structure of the littoral zone fish community. Twenty sites were identified based on previous studies. Sites included a variety of substrate types and macrophyte densities. Seine hauls were made with a 25ft bag seine with 1/8 inch mesh pulled perpendicular to shore starting from a depth of 1 m. Twenty fish of each species were measured from each haul and any additional fish were counted. Gill Netting (1993)Experimental gill nets were fished in weekly periods during June through August, 1993. Gill nets were used to capture piscivores for population estimates of fish marked in fyke nets. All nets were constructed of five 2.5-4.5 in. mesh panels, and were 125 ft. long. Nets set in water shallower than 10 ft. were 3ft. high or less; all others were 6ft. high or less. Sampling locations were selected randomly from up to three strata: 1) offshore reef sets, 2) inshore sets, 6.0-9.9 ft. deep, and 3) mid-depth sets, 10-29.9 ft. deep. The exact location at which the gill nets were set on the lake is unknown because the latitude and longitude values which were recorded by the WDNR are invalid. Temperature and dissolved oxygen profiles were used to monitor the development of the thermocline and guide net placement during July and August. After the thermocline was established nets were set out to the 30 ft. contour or to the maximum depth with dissolved oxygen greater than 2 ppm. Walleye Age: Scale and Spine Analysis (1987) Scales were taken from walleye that were shocked during the fall of 1987 electrofishing events on Lake Mendota. Scales were taken from 10 fish per one-inch length increment. The scales were removed from behind the left pectoral fin, and from the nape on the left side on esocids. In addition, the second dorsal spine was removed from 10 walleyes per sex and inch increment (to age and compare with scale ages for fish over 20 inches). CREEL SURVEYS1989:Fishing pressure, catch rates, harvest, and exploitation rates were estimated from a randomized, access-point creel survey. The schedule was stratified into weekday and weekend/holiday day types. Shifts were selected randomly and were either 07:00-15:00 h or 15:00-23:00 h. In addition, two 23:00-03:00 h shifts and two 03:00-07:00 h shifts were sampled per month to estimate the same parameters during night time hours. During the ice fishing season (January-February) 22 access points around Lake Mendota and upstream to the Highway 113 bridge were sampled. The clerk counted the number of anglers starting and completing trips during the scheduled stop at each access point. During openwater (March-December) 13 access points were sampled; 10 were boat ramps and 3 were popular shore fishing sites<strong>. </strong>At each of these sites, an instantaneous count of shore anglers was made upon arrival at the site, continuous counts of anglers starting and completing trips at public and private access points were made. Boat occupants and ice fishing anglers were only interviewed if they were completing a trip. Both complete and incomplete interviews were made of shore anglers. Number caught and number kept of each species, and percent of time seeking a particular species were recorded. All predators possessed by anglers were measured, weighed, and inspected for finclips and tags. We measured a random sample of at least 20 fish of each non-predator species per day.1990-1993: Same as 1989, except 23 access points were used during the ice fishing season. In addition, 13 access points were sampled during the openwater (May-December) season; 9 sites were boat ramps and 4 sites were popular shore fishing sites. 1994-1999: No formal documentation exists, but given the similarity in the data and consistency through the years; it is speculated tha tthe methods are the same.