U.S. Department of the Interior

Bureau of Land Management

BLM Alaska Open File Report 107

BLM/AK/ST-06/020+6700+040

August 2006

Waterfowl Use of Wetlands 

in Relation to Limnological Variables 

in the Kvichak River Area, Alaska

A fi nal report of the 1992 and 1993 waterfowl habitat evaluation 

effort in the Kvichak and Alagnak block of the Anchorage Field Offi ce

Bruce E. Seppi

���

Alaska

U.S. DEPARTMENT OF THE INTERIOR BUREAU OF LAND MANAGEMMENT
Mission Statement

The Bureau of Land Management (BLM) sustains the health, diversity and productivity of the public lands 

for the use and enjoyment of present and future generations.

Authors

Bruce E. Seppi is a wildlife biologist with the Anchorage Field Offi ce, Anchorage, Alaska.

Cover

BACKGROUND PHOTO: Aerial view of the study area between the west shores of Lake Iliamna and 

Alagnak River. TOP LEFT: the author conducts alkalinity testing on wetlands in the study area. LOWER 

RIGHT: a fl oatplane lands at a small lake to collect researchers. Wetlands too small for fl oatplane access 

required helicopters for transporting researchers and their equipment.

Open File Reports

Open fi le reports present the results of inventories or other investigations published outside the formal 

BLM-Alaska technical publication series. These reports can include preliminary or incomplete data and 

are not published and distributed in quantity. 

Reports are available while supplies last from BLM External Affairs, 222 West 7th Avenue, #13, 

Anchorage, Alaska 99513 (907) 271-5555 and from the Juneau Minerals Information Center, 100 Savikko 

Road, Mayfl ower Island, Douglas, AK 99824, (907) 364-1553. Copies are also available for inspection 

at the Alaska Resource Library and Information Service (Anchorage), the United States Department of 

the Interior Resources Library in Washington D.C., various libraries of the University of Alaska, the BLM 

National Business Center Library (Denver), and other selected locations. 

A complete bibliography of all BLM-Alaska scientifi c reports can be found on the Internet at:

http://www.blm.gov/ak/affairs/sci_rpts.html.

Related publications are also listed at: http://www.blm.gov/ak/jrmic/index.html.
Waterfowl Use of Wetlands 

in Relation to Limnological Variables 

in the Kvichak River Area, Alaska

A fi nal report of the 1992 and 1993 waterfowl habitat evaluation 

effort in the Kvichak and Alagnak block of the Anchorage Field Offi ce

By

Bruce E. Seppi

BLM Open Report 107

August 2006

U.S. Department of the Interior

Bureau of Land Management

Alaska State Offi ce

222 W. 7th Ave., #13

Anchorage AK 99513
i 

TABLE OF CONTENTS 

List of Tables ......................................................................... iii 

List of Figures ........................................................................ iv 

List of Appendices ................................................................... iv 

Acknowledgements ................................................................... v 

Abstract ................................................................................. 1 

Introduction 

Related Research .............................................................. 2 

Objectives ...................................................................... 4 

Study Area Description .............................................................. 5 

Geology and Topography .................................................... 7 

Soils ............................................................................. 7 

Climate .......................................................................... 7 

Methods 

Selection of Wetlands ......................................................... 9 

Brood Surveys ................................................................ 10 

Limnology .................................................................. 10 

Invertebrate Sampling ....................................................... 11 

Vegetation Sampling ........................................................ .12 

Statistical Analysis ........................................................... 12 

Results 

Limnological Variables ..................................................... 16 

Biological Variables .........................................................19 

Brood Use/Overall Waterbird Use ......................................... 19 

Statistical Analysis .......................................................... 21 

Principal Component Analysis ............................................. 24 

Regression Models .......................................................... 28 
ii 

Discussion 

Hydrology .....................................................................32 

Limnological/Wetland Traits ............................................... 33 

Aquatic Invertebrates ........................................................ 36 

Brood Use ......................................................................37 

Management Implications ................................................... 39 

Literature Cited ......................................................................40 

Appendices ............................................................................ 45 
iii 

LIST OF TABLES 

Table 1. Number of wetlands surveyed in each habitat stratum in the Kvichak study area 

in 1992 and 1993 .......................................................................... 9 

Table 2. Number and species of duck broods observed in the Kvichak study area in 1992 

and 1993 ...................................................................................13 

Table 3. Number and species of waterbirds observed on wetlands surveyed in the 

Kvichak study area in 1992 and 1993 .................................................14 

Table 4. Descriptive statistics of 20 variables measured within the Kvichak study area in 

1992 and 1993 ............................................................................17 

Table 5. Mean and standard error of wetland characteristics within each habitat stratum 

in the Kvichak study area in 1992 and 1993 

............................................. 18 

Table 6. Number of ducklings of each species observed within each habitat type in the 

Kvichak study area in 1992 .............................................................20 

Table 7. Number of ducklings of each species observed within each habitat type in the 

Kvichak study area in 1993 .............................................................20 

Table 8. Number of adult waterbirds of each species observed within each habitat type in 

the Kvichak study area in 1992 .........................................................22 

Table 9. Number of adult waterbirds of each species observed within each habitat type in 

the Kvichak study area in 1993 .........................................................23 

Table 10. Factor loadings for each variable along principal component axes 1, 2 & 3.......24 

Table 11. Factor loadings for each variable along principal component axes 1, 2, 3 with 

cations and anions combined ........................................................... 25 

Table 12. Correlation matrix for 20 habitat variables ...........................................29 

Table 13. Stepwise multiple regression equations showing the relationship between all 

species of duck broods and 15 habitat variables on wetlands within the Kvichak 

study area ..................................................................................30 

Table 14. Stepwise multiple regression equations showing the relationship between all 

species of adult ducks and 15 habitat variables on wetlands within the Kvichak 

study area ..................................................................................30 

Table 15. Stepwise multiple regression equations showing the relationship between all 

species of waterbirds and 15 habitat variables on wetlands within the Kvichak 

study area ..................................................................................31 
iv 

LIST OF FIGURES 

Figure 1. Map showing location of study area .................................................... 6 

Figure 2. The 95% confidence ellipses of wetlands with at least one brood and wetlands 

with no broods along the first 2 principal component axes .........................26 

Figure 3. The 95% confidence ellipses of wetlands with at least 1 species of adult duck 

and wetlands with no ducks along the first 2 principal component axes 

.........26 

Figure 4. The 95% confidence ellipses of wetlands with at least 1 species of waterbirds 

and wetlands with no waterbirds along the first 2 principal component axes ....27 

Figure 5. The 95% confidence ellipses of wetlands with at least 1 swan and wetlands with 

no swans along the first 2 principal component axes ................................27 

LIST OF APPENDICES 

Appendix 1. Explanation of variable abbreviations .................................................45 

Appendix 2. Raw data of 20 habitat variables for each wetland .................................. 46 

Appendix 3. The confidence ellipses of the first 3 principal components for wetlands with 

and without broods (3a), adult ducks (3b), all waterbird (3c), and swans (3d) 

....... 53 

Appendix 4. Occurrence of all invertebrate organisms found in 3 wetland habitat strata within 

the Kvichak study area ..................................................................55 

Appendix 5. List of aquatic and upland plant species recorded from wetlands surveyed in the 

Kvichak study area in 1992 and 1993 .................................................57 
v 

Acknowledgements

This project was funded entirely by the Anchorage Field Office of the 

Bureau of Land Management. 

Many people were involved in this project over the last 2 field seasons, 

and several volunteered their summers and came to Alaska to help collect the data 

from the field. I’d like to thank Dan Simpson, Dina Arel and Steve Fellman in 

1992 and Sybille Klenzendorf, Dave Van Rijn and Lee Gowey in 1993 for 

volunteering their time. Their help was greatly appreciated. 

In King Salmon, I’d like to thank U.S. Fish and Wildlife Service refuge 

manager Ron Hood and the staff of the Alaska Peninsula/Becharof National 

Wildlife Refuge for the use of a vehicle, the bunkhouse, floatplane dock, and 

aircraft fueling facilities in 1992 and the heliport in both years. Their cooperation 

reduced costs and simplified logistics while working in King Salmon. Thanks 

also goes to Dick Russell of the Alaska Department of Fish and Game in King 

Salmon who generously offered us the use of his personal vehicle, a place to pitch 

camp, and the use of cooking and shower facilities in 1993. 

I’d also like to thank pilots Ross Carswell of Country Lakes Air, Jack 

Gordon and Lester of Trans Alaska Helicopters, John Penny and mechanic Brad 

Craig of Alaska Helicopters, and the pilots of Katmai Air for their skills and 

patience while flying in and out of the numerous and often very shallow and 

muddy wetlands we sampled. 

I’d like to thank Dr. Jacqueline LaPerriere of the Alaska Cooperative Fish 

and Wildlife Research Unit, University of Alaska Fairbanks for the use of the 

flourometer and university laboratory facilities to analyze chlorophyll a samples. 

I thank Dr. John “Jack” Jones of the University of Missouri Columbia, 

Department of Fisheries and Wildlife for providing low-cost, accurate and precise 

water chemistry analysis results for both years. 
1 

Abstract 

The relationship between 20 habitat variables and use by duck broods and 

adult waterbirds of all species was investigated in 1992 and 1993 on 156 wetlands 

in the Kvichak River area, Alaska. The study examined whether wetland size and 

shape, water chemistry characteristics, primary productivity and aquatic 

invertebrate biomass affected the density of waterfowl on wetlands. Biases in the 

visibility and detectability of birds, especially duck broods, were recognized. 

Results suggest brood use by all species of ducks was influenced by 

wetland cation concentrations, particularly sodium. Use by adult ducks and all 

species of waterbirds was related to total alkalinity and shoreline length. 

Limnological variables associated with swan use of wetlands were inconclusive, 

but general trends indicate productivity and salinity may be factors. On a density 

basis (birds/m shoreline), sodium, chlorophyll a, pH and total alkalinity most 

strongly influenced use by duck broods, adults and all species of waterbirds. 

These results suggest that large, relatively productive wetlands with low to 

moderate salinity were used most by waterfowl. 
2 

Introduction 

Waterfowl populations in North America have declined to historically low 

levels due to increasing losses of breeding, wintering and migration habitat (N. 

Am. Waterfowl Management Plan 1986). These declines are primarily a result of 

the continued conversion of wetland habitat to agricultural use in the midcontinent prairie-parkland breeding areas (Tiner 1984, Sanderson 1980), as well 

as in the Central Valley of California (Graziano and Cross 1993). Industrial and 

urban land development has also contributed to loss of wetlands nationwide (Dahl 

and Johnson 1991). Although legislation exists that attempts to slow or reverse 

these trends (Section 404-1B of the Clean Water Act), habitat loss remains a 

substantial threat to waterfowl production. The production potential of pristine 

high latitude areas is therefore increasingly important, as these areas now contain 

substantial portions of breeding populations of species of significant management 

interest (e.g. northern pintail) (Derksen and Eldridge 1980, Smith 1970, Hansen 

and McKnight 1964). 

Alaska contains millions of acres of wetland habitat representing a 

significant production area for many species of waterfowl. While much of this 

habitat is managed by the U.S. Fish and Wildlife Service, important breeding 

areas exist on Bureau of Land Management (BLM) lands. Some of these lands 

may be more important as waterfowl breeding habitat than others, however the 

Kvichak and Alagnak block within the Anchorage Field Office contains 

substantial wetland habitat, representing significant waterfowl production 

potential. Aerial breeding pair surveys in 1992 and 1993 revealed breeding 

waterfowl densities of 35.9 and 29.3 ducks per square mile in stratum 8 (Bristol 

Bay) in 1992 and 1993 respectively (Conant and Groves 1992, 1993). While 

these densities are low relative to other BLM lands in Alaska, it is crucial to 

understand habitat features important to breeding, nesting and brooding waterfowl 

using the Kvichak area so comparisons of its waterfowl production potential can 

be made with other areas. 

Related Research 

BLM has funded a similar investigation of waterfowl and wetland habitat 

quality relationships in the Iditarod George block of the Anchorage Field Office. 

The relationship between 20 habitat variables and use by waterfowl broods was 

investigated in the Lower Innoko River area, Alaska. In that area, brood use was 

most strongly associated with shoreline length, potassium, and aquatic 

invertebrates, or temperature, color, chlorophyll a and shoreline length, depending 

on an analysis using separated or combined cations and anions (Seppi 1993). 

When limnological variables were compared with overall duckling density 

(broods/m shoreline length), stepwise regression linked chlorophyll a, 

temperature, color and total phosphorous concentrations with higher brood 

densities (Seppi 1993). 

Other research has assessed waterfowl use patterns and hydrochemical 
3 

characteristics of Interior Alaska wetlands. Murphy et al. (1984) showed 

phosphate and nitrate concentrations in the water column to be correlated with 

duck use. High concentrations of these nutrients where linked to both high duck 

density and species richness. Hydrological connection to flowing water was also 

associated with higher densities of ducks when compared to isolated waterbodies, 

because of higher nutrient levels associated with the inflow of water (Kessel et al. 

1980, Murphy et al. 1984, Heglund 1988, 1992). Breeding densities of waterbirds 

(ducks, geese, swans, loons, grebes) were found to be positively correlated to total 

phosphorous concentrations by Nilsson and Nilsson (1978) as well. 

Inorganic nutrients, particularly phosphorous and nitrogen, are considered 

limiting factors in the productivity of wetlands (Richardson et al. 1978). Growth 

of aquatic plants is often limited by nitrogen and phosphorous availability, 

therefore the rates at which these nutrients are supplied, cycled and removed from 

the wetland environment are critical to plant growth (Kadlec 1979). Ducks in turn 

may use nitrogen and phosphorous rich lakes in a response to higher productivity 

levels. On Interior Alaska wetlands, Heglund (1988) reported a positive 

correlation between waterbird density (ducks, geese, swans, loons, grebes) and 

total phosphorous and nitrogen concentration, cation salinity (calcium, sodium 

and potassium concentration) and shoreline length. Large, productive, slightly 

brackish lakes supported the highest densities of waterbirds. Further work in 

boreal forest wetlands by Heglund (1992) suggests brackish water and wetmeadow wetlands with sedge and grass vegetation, which provided cover and 

abundant invertebrate and plant foods were preferred by some species of 

waterfowl and grebes. Differences in visibility of broods among wetland types 

may have influenced these results, which are therefore offered with caution. 

Habitat requirements for duck broods depend both on the amount of 

surface water available and wetland fertility. Patterson (1976) found the highest 

density of duck broods on fertile, hard water ponds with abundant submerged 

macrophytes because these ponds provided both escape cover and food. 

Courcelles and Bedard (1979) found that broods, as well as adult birds, selected 

habitats of open cattail (Typha augustifolia) with open areas dominated by 

submerged hydrophytes (Lemna, Myriophylum, Ceratophylum and Utricularia

species). 

Submerged aquatic plants and their associated invertebrates are important 

factors in waterfowl use of ponds. Duck use of wetlands is positively correlated 

with invertebrate numbers (Joyner 1980), although invertebrate numbers depend 

largely on the species and abundance of submerged aquatic plants (Krull 1970). 

Aquatic vegetation alone can be an important waterfowl food, however even those 

plants of low food value are indirectly important to waterfowl production as they 

provide habitat for macroinvertebrates which are an important source of animal 

protein for ducks (Krull 1970, Serie and Swanson 1976). Dabbling ducks 

concentrate their foraging in areas with the highest abundance and biomass of 

invertebrates and often seek out areas inhabited by prey items of high nutritive 

quality (Kaminski and Prince 1981). High invertebrate populations often coincide 

with peak hatching of waterfowl (Bergman et al. 1977) and constitute a major 

portion of the diet for many species of ducklings (Sudgen 1973). 
4 

Wetland salinity may also be a factor in use by waterfowl broods. Brood 

use of saline lakes is determined principally by salt concentration, the availability 

of freshwater, and aquatic foods, although ducklings cannot tolerate high 

concentrations of sodium chloride (Swanson et al. 1984). In addition, high 

concentrations of sodium chloride can cause changes in the species structure of 

invertebrates and plants attractive to ducks, thus affecting their use of saline 

wetlands (Swanson et al. 1984). 

The physical, chemical and biological characteristics of wetlands directly 

and indirectly affect use patterns by waterfowl. These characteristics can 

therefore be used as indicators in evaluating wetlands as potential habitat for 

waterfowl. Before management to increase or maintain waterfowl production is 

undertaken, it is essential that the relationship between habitat factors and brood 

production be understood. Limited data exist that relate characteristics of Alaskan 

wetlands to their use by waterfowl broods (Murphy et al. 1984, Heglund 1988, 

1992). The Lower Innoko/Iditarod George study is the only data that exists for 

BLM lands in Alaska (Seppi 1993). The purpose of this study is therefore to 

examine if any relationships exist between wetland characteristics and waterfowl 

use on BLM lands in the Kvichak area. These results can in turn be compared to 

other waterfowl habitat on public lands under BLM management and used to 

recognize those areas most important to waterfowl production. 

Objectives 

The overall objective of this project was to evaluate the Kvichak area for 

its waterfowl production potential. To do this, the project attempted to determine 

if a correlation exists between a wetlands water chemistry, morphological 

measurements and selected biological factors and its use by waterfowl broods, 

adult birds, (ducks, geese and swans) and other water-loving birds (loons, grebes). 

This involved identifying those wetlands most used by waterbirds and used to 

produce waterfowl broods and surveying and describing their: basic 

morphometric characteristics; water chemistry; aquatic vegetation and associated 

macroinvertebrate fauna. These parameters were then statistically compared to 

waterfowl production and waterbird use within each wetland to identify possible 

habitat\use relationships. 
5 

Study Area Description 

The study area is located at the base of the Alaska Peninsula near King 

Salmon, Alaska, approximately 465 km (290 miles) southwest of Anchorage. The 

lands are BLM-administered public lands, however large portions are in the 

process of conveyance to the State of Alaska. The area (59o

 00’N, 156o 30’W), 

covers approximately 8,300 square kilometers and extends east to the western 

shore of Lake Iliamna, west to the Nushagak River, north to 59o 45’N latitude, 

and south to the village of King Salmon (Figure 1). At present, 490 square 

kilometers of BLM-administered lands are interspersed with private and stateowned lands (ADNR December 1992). 

The major drainages for the area are the Nushagak, Alagnak and Kvichak 

Rivers. The villages of Levelock and New Stuyahok lay within or on the 

boundaries of the study area. Several private hunting and fishing guide/outfitters 

operate within or near the region. The area is roadless, with the exception of a 

road between King Salmon and Naknek, a small coastal community largely 

influenced by the Bristol Bay salmon fishery. 

The area lies on a glacial outwash of shallow sandy/gravel soils with 

discontinuous and isolated masses of permafrost (Selkregg 1976). The dominant 

vegetation is a transition between the forest/tundra plant communities found 

farther north and the treeless grass/sedge/dwarf shrub/lichen tundra typical of the 

Alaska Peninsula. Most of the area is rolling dwarf shrub tundra with numerous 

small pothole lakes and larger, more shallow wetlands. A band of spruce forest 

(Picea spp.) extends in a narrow strip across the center of the area, approximately 

along the path of the Alagnak and Kvichak Rivers. 

Aerial view of the study area between the west shores of Lake Iliamna and the Alagnak 

River. 
6 
7 

A unique band of wetlands exists along a glacial moraine in the northeast 

portion of the study area. This band of wetlands is situated west of Lake Iliamna 

and extends for 72 kilometers (45 miles) in an arc which begins northwest of the 

lake and ends southwest of the lake. These wetlands are of variable size and 

shoreline development, yet all are deep, clear and virtually void of sediment and 

aquatic vegetation. These pools typically have a deep emerald green color that is 

apparent from the air and when using a Secchi disk to measure transparency. 

They have deep, bowl-shaped basins with rock, gravel or sand bottoms that drop 

off quickly in depth with minimal shallow areas along the shoreline. Those 

moraine lakes surveyed averaged 4.4 m in overall depth and 8.2 m maximum 

depth, including one wetland with a maximum depth which exceeded 14 meters 

(waterbody 1002, appendix 2). These wetlands have closed hydrological regimes 

with snowmelt, rain and ground water as their only source of water replenishment. 

Such wetlands obtain their nutrient flux mainly from rain and ground water, 

depending on the substrate (Gosselink and Turner 1978), thus have a low nutrient 

flux and may be less desirable to waterfowl (Murphy et al. 1984). 

Larger wetlands in the other parts of the study area had larger surface 

areas, were shallower, more turbid and have more aquatic plant life than the 

moraine lakes. These non-moraine wetlands had both closed and open 

hydrological regimes, however even those with closed hydrologies were relatively 

more productive because of their shallow depth (< 1 m) which provides favorable 

conditions for aquatic plant growth. The higher surface area, shallow depths and 

subsequent higher productivity may make these wetlands more desirable to 

breeding waterfowl (Heglund 1988, 1992, Seppi 1993). 

Geology and Topography 

The geology of the area is dominated by glacial moraine and drift, or 

glacialfluvial outwash deposits bordering older moraines, or alluvial flood plain 

deposits associated with rivers and streams (Selkregg 1984). 

Soils 

Soils are well drained, shallow and gravelly, and are formed in well 

drained course volcanic ash deposits in the inland areas to poorly drained, 

fiberous peat soils over a shallow, discontinuous permafrost table near Lake 

Iliamna and the coast (Selkregg 1984). 

Climate 

Climate in the Kvichak area is considered transitional between the 

Interior’s continental climate and the maritime climate of the southern coastline of 

the Aleutian Islands (Selkregg 1984). Average maximum temperatures range 

from 63 0F (17 0C) to 38 0F (3 0C) in July and 29 0F (-2 0C) to 4 0F (-160

C) in 

January. Extremes of -42 0F (-410

C) and 880F (310C) have been recorded at King 

Salmon. Mean annual precipitation is 20 in (51 cm), which includes 45 in (114 
8 

cm) of snowfall (Selkregg 1984). Winds average 9.6 knots (17.8 kph), with an 

extreme of 115 kph. Average freeze up of the Nushagak River is November 20th, 

the river is ice free by May 5th. An average growing season of 97 days is 

estimated for Dillingham (Selkregg 1984). 
9 

Methods 

Selection of Wetlands 

Wetlands were selected using 1:63,360 topographical maps and 1:21,120 

black and white aerial photographs. Wetland habitats were stratified according to 

the following criteria: connected wetlands (wetlands that were hydrologically 

connected to flowing water); closed wetlands (wetlands that were not connected 

to flowing water); moraine wetlands (the deep green pools along the glacial 

moraine described earlier); rivers (the major river channels of the Alagnak and 

Kvichak Rivers). To select individual wetlands for brood and waterfowl use and 

limnological analysis, each 1 mi2

 section marked on the topographical maps was 

assigned a number. After single sections within the study area were selected with 

the use of a random numbers table, a second random number was used to pick a 

wetland from that section. If sections without wetlands were chosen, they were 

rejected and a new section chosen. This was done repeatedly until 200 individual 

wetlands had been selected. 

Thirty-two connected wetlands, 61 closed wetlands, 5 moraine wetlands 

and 3 rivers were surveyed in 1992; 35, 58, 14, and 1 of these respective wetlands 

were surveyed in 1993, resulting in samples from 156 different wetlands in the 2 

years (Table 1). No wetlands were repeatedly surveyed within each year, 

however I returned to 53 ponds in 1993. Forty-eight wetlands were sampled in 

1992 only and 55 in 1993 only (Table 1). 

Table 1. Number of wetlands surveyed in each habitat stratum in the Kvichak 

study area 1992 and 1993. 

YEAR SURVEYED 

Wetland 1992 1993 Both 

Type 1992 1993 only only years 

Connected 32 35 15 18 17 

Closed 61 58 28 25 33 

Moraine 5 14 2 11 3 

River 3 1 3 1 0 

Total 101 108 48 55 53 

Total different wetlands surveyed in 2 years = 156 
10 

Brood Surveys 

Use of individual wetlands by waterfowl was estimated by aerial and 

ground surveys. Larger, deeper wetlands were accessed by float plane. Smaller, 

shallower ponds were accessed by helicopter. Complexes of wetlands of various 

sizes were also accessed by float plane by landing on the largest lake and walking 

to the nearby, smaller ponds. All wetlands accessed by helicopter were surveyed 

for broods from the air as they were first approached. The shoreline perimeter of 

each selected wetland was flown at low altitude (<50 feet) and broods recorded as 

they flushed. Larger wetlands accessed by floatplane were surveyed from the 

ground. A collapsible pack canoe was also used on wetlands accessed by 

floatplane. For ground surveys, two people walked, or when possible canoed in 

opposite directions along the shoreline of each wetland and recorded the species, 

age class (Gollop and Marshall 1954) and number of all waterfowl broods 

observed. Adult, non-breeding waterbirds without young of all species were also 

recorded on the wetlands surveyed for broods. 

Ideally, brood surveys were to be timed as to occur midway between the 

peak hatch and occurrence of dabbler and diver broods in the area. I did not feel 

this was accomplished. Brood surveys were initiated on 25 June and 19 June in 

1992 and 1993 respectively. The brood survey continued until 17 July in 1992 

and 5 July in 1993. Only one survey was conducted for each wetland. The 

estimated peak hatch of broods for the Kvichak/Alagnak river drainage area is 

between 7-15 to 7-21 (Hodges and Conant 1990), therefore the survey may be an 

underestimation of broods as it was too early in both years and did not account for 

those broods hatching in July, particularly diving broods. 

Limnology 

Limnological analysis of wetlands was carried out on the same day 

wetlands were surveyed for waterfowl use. Water samples were collected at the 

surface of each wetland in plastic containers for total nitrogen, total phosphorous, 

major cations (sodium, calcium, potassium, magnesium) and major anions other 

that carbonates (chloride, sulfate). Water samples for chlorophyll a were 

collected and filtered and preserved on site for subsequent laboratory analysis. 

Physical characteristics (transparency, surface temperature, average depth) and 

those chemical characteristics that could be determined on site (conductivity, total 

alkalinity, water color, pH) were also measured at each water body. I estimated 

shoreline length and surface area for all wetlands by digitizing 1:21,120 black and 

white aerial photographs using Arc/Info (1991). A shoreline development index, 

which compares shoreline irregularity, was calculated from the ratio of shoreline 

length to the circumference of a circle that has the same area as the lake (Wetzel 

1983). Water transparency was measured using a Secchi disk attached to a 

graduated cord and depth was recorded to the nearest 0.25 meter. Conductivity 

(recorded in μS/cm) and pH were measured using a Corning Check-Mate 90 

handheld meter with detachable conductivity and pH sensors. Water temperature 

was measured at the surface with a digital thermometer built into each detachable 
11 

sensor. Filtered water samples were visually compared with a Hach color 

comparitor kit to estimate water color and recorded in platinum cobalt units. 

Bicarbonate alkalinity was determined by titration with sulfuric acid using a Hach 

digital titrator. Phenolphthalein and bromcresol green-methyl red were used as 

titration endpoint indicators and total alkalinity recorded as mg/L CaCO3. 

Chlorophyll a samples were collected using a hand operated vacuum pump 

to filter up to 1 liter of surface water through a 0.45μm Gelman A/E glass fiber 

filter. The glass filters were then wrapped in paper filters and placed in airtight, 

darkened containers filled with desiccant for preservation by drying. Chlorophyll 

a concentrations were later determined flourometrically (Knowlton 1984) using 

acetone extraction (APHA 1985) and recorded in mg/L. I used laboratory 

facilities in the Department of Biology and Wildlife at the University of Alaska, 

Fairbanks to determine chlorophyll a concentrations from field samples. 

Water samples collected to determine concentrations of total nitrogen, 

total phosphorous, major cations and major anions were sent to the University of 

Missouri at Columbia for analysis. Samples for total phosphorous and total 

nitrogen were collected in 500 ml plastic bottles and subsamples of 10 ml each 

were pipetted into clean acid washed test tubes at the end of each day. Total 

nitrogen samples were preserved with one drop of concentrated sulfuric acid. 

Total phosphorous samples were kept in a cool place and preserved by 

refrigeration upon return from the field. A single filtered sample of 50 ml was 

collected for calcium, magnesium, potassium and sodium and preserved with one 

drop of concentrated sulfuric acid. A single water sample of 125 ml was collected 

for sulfate and chloride analysis and also reserved by refrigeration. 

Invertebrate Sampling 

Aquatic invertebrate samples were collected only in 1993 on a limited 

number of wetlands within each stratum in conjunction with limnological 

sampling. At each wetland, aquatic invertebrate abundance in the benthos and 

water column was measured using a sampling device made of a graduated 

Plexiglass tube (Pederson and Pederson 1983) with a 500 micron screen that 

could be inserted into a slot at the bottom. Samples where collected by quickly 

lowering the tube through the water column and 3 cm into the benthos. The water 

column and benthos enclosed by the sampler was then agitated to suspend benthic 

organisms. The screen was then slid into place to collect a combined sample of 

organisms from the sediments and water column. Water was allowed to drain 

from the sampler, and material on the screen was then washed into plastic jars 

with 95% ethanol and saved for later sorting. Samples were taken at 3 random 

sites around the perimeter of the wetland in water no deeper than 40 cm. In the 

laboratory, the 3 samples from each wetland were combined, then sorted and 

identified to family when possible using Pennak (1978), and Thorp and Covich 

(1991). After weighing the samples wet, I dried them for 24 hrs at 100o

C and 

reweighed them to determine total dry biomass (g/cm2) of invertebrates per 

sample. The mass of invertebrates was determined on the basis of area and not 

volume as the sampling device used was probably biased toward benthic 
12 

invertebrates and did not capture many of the more mobile organisms in the water 

column. 

Vegetation Sampling 

I sampled aquatic vegetation in each wetland and collected field notes on 

terrestrial vegetation to enhance the descriptions of wetland habitat 

characteristics, but I did not directly compare vegetation data with use by 

waterfowl. Aquatic vegetation types were recorded once for all wetlands during 

limnological sampling. Two transects across each wetland were used to 

determine the species of aquatic plants present. On each wetland, transect lines 

were established across the center of the wetland in east-west and north-south 

directions perpendicular to the shoreline. Using the canoe and a compass bearing, 

each transect line was followed from shoreline to shoreline. At approximately 10 

m intervals, all aquatic plant species encountered were recorded. Those species 

that could not be identified on site were collected and placed in a plant press for 

later identification using Welsh (1974) and Hulten (1968). 

Depth measurements were also taken along transect lines used to sample 

aquatic vegetation. Depth was determined using a weighed measuring line with 

0.25 m graduations. Soundings were taken at approximately 10 m intervals for 

transects >50 m and every 5 m for shorter transects. Depth measures from both 

transects combined were used to calculate mean depth (m) for each wetland. 

Statistical Analysis 

Waterfowl use of wetlands was divided into 4 categories for statistical 

analysis: ducklings of all species (total ducklings), including dabbling ducklings 

(tribe Anatini) and diving ducklings (tribes Aythyini, Mergini); adult dabbling 

and diving ducks; all waterfowl/waterbirds, including adults and juveniles of 

ducks, geese and swans (tribes Anatini, Aythyini, Mergini, and Anserini) and 

loons (genus Gavia); and swans only (trumpeter swans (Cygnus buccinator) 

and/or tundra swans (Cygnus columbianus). Six species of duck broods (Table 

2), 12 species of adult ducks, 2 species of loons and 2 species of swans (Table 3), 

occurred in surveys and were used in the analysis. 
13 

Table 2. Number and species of duck broods observed in the Kvichak study area 

in 1992 and 1993a

. 

 1992 1993 

Species # Broods # Ducklings # Broods #Ducklings 

Scaup1 1(5)3 1(2)4 1(17)3 9(30)4

Black Scoter 1(5) 2(3) 0(0) 0(0) 

White-winged Scoter 0(0) 0(0) 0(0) 0(0) 

Mallard 3(15) 11(17) 2(33) 11(37) 

Goldeneye2 0(0) 0(0) 0(0) 0(0) 

Pintail 2(10) 5(8) 1(17) 2(7) 

Oldsquaw 0(0) 0(0) 0(0) 0(0) 

Green-winged Teal 2(10) 10(16) 0(0) 0(0) 

Wigeon 4(20) 12(19) 0(0) 0(0) 

Hooded Merganser 0(0) 0(0) 0(0) 0(0) 

Bufflehead 0(0) 0(0) 0(0) 0(0) 

Unidentified 7(35) 22(35) 2(33) 8(27) 

Total 20(100) 63(100) 6(100) 30(100) 

a

 adult birds were seen of all species listed 

1 includes both greater and lesser scaup 

2

 includes both common and barrow’s goldeneye 

3

 percent of total broods in parenthesis 

4

 percent of total ducklings in parenthesis 
14 

Table 3. Number and species of waterbirds observed on wetlands surveyed in the 

Kvichak study area in 1992 and 1993. 

Species 1992 1993 Total 

Swans1 54(11)a 69(9)b 123(10)c

Scaup2

 208(42) 202(27) 410(33) 

Black Scoter 129(26) 361(48) 490(39) 

White-winged Scoter 16(3) 4(0.5) 20(2) 

Red Breasted Merganser 1(0.2) 4(0.5) 5(0.3) 

Common Loon 7(1) 15(2) 22(2) 

Arctic Loon 21(4) 8(1) 29(2) 

Mallard 10(2) 5(0.7) 15(1) 

Goldeneye3

 6(1) 2(0.3) 8(0.6) 

Pintail 9(2) 2(0.3) 11(0.9) 

Oldsquaw 1(0.2) 4(0.5) 5(0.4) 

Green-winged Teal 11(2) 3(0.4) 14(1) 

Wigeon 5(1) 1(0.1) 6(0.5) 

Hooded Merganser 6(1) 0(0) 6(0.5) 

Bufflehead 3(0.6) 0(0) 3(0.2) 

Unidentified 11(2) 77(10) 88(7) 

Total 498(100) 757(100) 1255(100) 

1

 includes tundra swans adult and cygnets 

2

 includes both greater and lesser scaup 

3

 includes both common and barrow’s goldeneye 

a

 percent of total for 1992 

b

 percent of total for 1993 

c

 percent of total for both years 
15 

To reduce dimensionality of wetland variables I calculated scores for each 

wetland along the first 3 principal components derived from wetland variables. 

Transparency and aquatic invertebrates where excluded from all multivariate 

analysis. Transparency exceeded wetland depth in most wetlands, and aquatic 

invertebrates were collected on a limited number of wetlands in only 1993, 

resulting in too few data points for each variable. Factor score coefficients were 

based on the correlation matrix of the original variables (Johnson and Wichern 

1988). To more easily recognize the relationship between wetland characteristics 

and brood use of wetlands, 95% confidence ellipses of the first 2 principal 

component scores for wetlands used by duck broods of all species, adult ducks, all 

species of waterbirds and swans were plotted separately for the first 2 principal 

components. The confidence ellipses of the first 3 principal components are also 

included in appendix 3 for a 3 dimensional comparison. 

Stepwise multiple regression was used to determine which wetland habitat 

variables were most strongly related to waterfowl use (Draper and Smith 1980). 

All species of duck broods combined, adult ducks, total waterbirds and swans 

were used as the dependent variables in 4 separate analysis. Before regression 

analysis, variables were tested for multicollinearity by examining a correlation 

matrix of all variables and eliminating 1 variable from each with a correlation 

0.70 (Bowyer et al. 1988, table 6). The final regression model was tested for 

aptness by plotting residuals against fitted dependent variables (Neter et al. 1985). 

Finally, the Durbin-Watson test for autocorrelation was performed on the final 

model to ensure that no correlation existed among error terms (Neter et al. 1985). 

 This allowed me to determine if any important variables were not included in the 

final model or mistakenly eliminated because of autocorrelation with other 

variables. All statistical analysis were performed using StatSoft Statistica 

software packages (StatSoft 1994). 

The conductivity of natural waters is closely related to concentrations of 

the major ions present (Wetzel 1983). Conductivity of wetlands ranged from 5 to 

468 μS/cm, with lowest average conductivity in moraine wetlands and highest 

conductivity in rivers and connected wetlands (Table 5). Salinity concentrations 

from 6 major ions reflect similar patterns as conductivity measures in the 4 strata; 

lowest average concentrations in moraine wetlands and higher in rivers and closed 

and connected wetlands (Table 5). 
16 

Results 

Limnological Variables 

Wetlands ranged in surface area from <0.5 to 668 ha (Table 4). 

Connected wetlands had the largest sampled average surface area (60 ha), 

followed in order by rivers, moraine and closed wetlands (Table 5). Wetlands 

averaged 1.32 m deep overall (Table 4). Moraine wetlands had the highest mean 

depth (3.97 m), while mean depths of closed wetlands were 1.0 m (Table 5). No 

depth measurements were taken on the rivers because of swift currents, but 

estimated depths probably did not exceed 2 m (personal observation). Shoreline 

length, which was measured from aerial photos taken in mid summer, ranged 

from 187 to 12,425 m (Table 4), and was greatest in rivers and lowest in closed 

wetlands (Table 5). Shoreline development indices ranged from 1.02 (nearly 

circular closed wetlands) to 3.57 (very long irregular shorelines of the rivers, 

Tables 4,5). 

Wetland surface water color ranged from zero (very clear, unmeasurable 

color) to 275 PtU (heavily stained), with a mean value of 47 PtU (Table 4). Color 

varied somewhat across strata; rivers and moraine wetlands were clear while 

connected and closed tundra ponds were more highly stained (Table 4). 

Secchi disk transparencies exceeded wetland depth in 78% of the wetlands 

in 1992 and 68% in 1993 (Appendix 2). Even heavily stained wetlands were 

often too shallow to measure transparency. In those wetlands were it could be 

measured, Secchi transparency averaged 2.0 m across all strata (Tables 4, 5). 

Surface water temperature averaged 16 oC across all wetlands (Table 4), 

rivers were coolest while moraine wetlands were most warm (Table 5). Wetland 

pH was near neutral in each stratum and averaged 7.12 overall (Tables 4,5). 

Total alkalinity ranged from 0.0 (unmeasurable with field kits) to 47 mg/L 

CaCO3 (Table 4), with highest alkalinities in rivers and connected wetlands and 

lowest alkalinities in moraine wetlands (Table 5). 

Total phosphorous and total nitrogen levels ranged from 4.0 to 409 μg/L 

and 0.03 to 1.55 mg/L respectively (Table 4). River strata had higher average 

total phosphorous concentrations and moraine wetlands lower average 

phosphorous concentrations than other strata (Table 5). Connected wetlands 

averaged higher in total nitrogen while rivers averaged lowest (Table 5), although 

wetlands in all strata were relatively close to the 0.37 mg/L average overall (Table 

4). 
17 

Table 4. Descriptive statistics of 20 variables measured within the Kvichak study 

area in 1992 and 1993. 

Variablea N Mean S.E. Minimum Maximum 

Invert 31 0.140 0.071 0.004 2.223 

Trans 60 2.02 0.34 0.25 11.0 

Temp 204 5.58 0.21 9.1 26.16 

pH 192 7.12 0.04 5.5 9.45 

Cond 208 29.50 2.46 4.65 467.5 

T. Alk 206 8.18 0.53 0.0 46.5 

Color 204 47.28 3.27 0.0 275.0 

TP 174 36.09 3.39 4.0 409.0 

TN 174 0.37 0.02 0.03 1.55 

Ca 174 1.04 0.08 0.03 7.68 

Mg 174 0.67 0.04 0.06 3.54 

K 174 0.64 0.15 0.08 25.0 

Na 174 2.72 0.15 0.76 14.1 

Cl 174 2.19 0.68 0.20 119.5 

SO4 173 0.42 0.04 0.05 3.3 

Chloroa 208 4.58 0.44 0.052 63.52 

Surface 209 51.70 6.24 0.32 667.9 

SLL 209 2531.6 142.8 187.2 12425.4 

SLD 209 1.30 0.03 1.02 3.57 

Ave Depth 183 1.32 0.11 0.10 8.61 

a

 Explanation of variable abbreviations in Appendix 1. 
18 

Table 5. Mean and standard error of wetland characteristics within each 

habitat stratum in the Kvichak study area in 1992 and 1993.

WETLAND TYPE 

 Connected Closed 

Variablea

 Wetland Wetland Moraine River 

Invert 0.3074±(0.4073) 0.0676±(0.0372) 0.0193±(0.0116) - 

Trans 0.82±(0.24) 1.34±(0.57) 6.20±(1079) - 

Temp 15.36±(0.70) 15.57±(0.55) 16.52±(1.37) 14.51±(2.14) 

pH 7.20±(0.12) 7.06±(0.12) 7.25±(0.23) 7.11±(0.22) 

Cond 31.94±(4.07) 30.31±(7.73) 13.40±(5.24) 44.62±(5.10) 

T. Alk 9.97±(1.86) 7.98±(1.33) 2.23±(1.68) 13.80±(1.87) 

Color 56.5±(11.5) 46.3±(7.9) 20.8±(15.1) 13.3±(15.8) 

TP 39.29±(6.34) 38.78±(10.56) 9.64±(2.30) 48.67±(74.42) 

TN 0.4014±(0.0418) 0.3949±(0.0462) 0.1748±(0.0356) 0.1107±(0.0215) 

Ca 1.29±(0.24) 0.91±(0.15) 0.73±(0.79) 2.64±(0.09) 

Mg 0.82±(0.13) 0.66±(0.12) 0.25±(0.12) 0.93±(0.36) 

K 0.53±(0.07) 0.76±(0.49) 0.36±(0.07) 0.47±(0.15) 

Na 3.06±(0.41) 2.78±(0.44) 1.43±(0.25) 2.19±(0.33) 

Cl 1.55±(0.31) 2.77±(2.30) 1.29±(0.23) 0.73±(0.13) 

SO4 0.45±(0.12) 0.42±(0.10) 0.29±(0.13) 1.07±(0.44) 

Chloroa 5.28±(1.17) 4.83±(1.32) 1.11±(0.34) 1.03±(0.55) 

Surface 60.46±(12.70) 46.39±(17.89) 53.1±(49.32) 56.15±(67.68) 

SLL 3039.3±(417.9) 2061.5±(301.4) 3273.3±(1578.8) 4488.1±(2007.4) 

SLD 1.27±(0.06) 1.22±(0.04) 1.73±(0.32) 2.24±(0.46) 

Ave Depth 2.72±(3.28) 1.02±(0.20) 3.97±(0.90) - 

a

 Explanation of variable abbreviations in Appendix 1. 
19 

Biological Variables

Chlorophyll a concentrations averaged higher in connected wetlands (5.28 

mg/L) than the other strata (Table 5). 

Seven classes, 5 orders, 5 suborders, and 20 families of aquatic 

invertebrates from 8 phyla occurred in sampled wetlands (Appendix 4). No 

aquatic invertebrate samples were taken in the river strata. Moraine wetlands had 

fewer invertebrate taxa (Appendix 4), and lower biomass than the other strata, 

while connected wetlands had the highest biomass of invertebrates (Table 5). 

Predacious diving beetle larva (dytiscidae), mosquito larva (culicidae), black fly 

larva (simuliidae), hydra (hydridae), lymnaeid snails, nemtodes and gordian 

worms (gordiidae) were found only in closed wetlands (Appendix 4). Midge 

larva (ceratopogonidae) and leptophlebiid may flies were found only in connected 

wetlands while caddis flies (leptoceridae) were found only in moraine wetlands 

(Appendix 4). Aquatic earth worms (oligochaeta), chrimomid midge larva, 

limnephilid caddis flies, planorbid snails and fingernail clams (sphaeriidae) were 

found in all strata (Appendix 4). 

Thirty species of aquatic emergent, submergent and upland plants where 

recorded in wetlands or near shorelines (Appendix 5). The river and moraine 

wetland strata were devoid of aquatic vegetation. No truly aquatic plant species 

were found in the river stratum; the main species recorded where willows along 

the riverbank. Most moraine wetlands had no aquatic plants, although bureed, 

water shield, and pond lily occurred in some moraine wetlands that had shallower 

areas. Most aquatic plant species occurred in both connected and closed wetlands 

with the exception of manna grass, cotton grass, labrador tea, marsh fivefinger, 

lingonberry, sweet gale, spike rush, water hemlock and pondweed (Potomogeton 

natans) found only in and around hydrologically closed wetlands (Appendix 5). 

A fern species (probably Thelypteris phegopteris) occurred only in the elevated 

upland areas around moraine wetlands. 

Brood Use/Overall Waterbird Use 

Duck broods of 6 species were recorded within the study area. American 

wigeon (Anas americana), green-winged teal (Anas crecca), mallard (Anas

platyrhynchos) and northern pintail (Anas acuta) were the most abundant species 

of broods in both years (Table 2). Even though 6% more wetlands were surveyed 

in 1993 than in 1992, 52% fewer ducklings were recorded in 1993 compared to 

1992 (Table 2). In 1992 and 1993, 100% and 93%, respectively, of all ducklings 

observed were seen on connected or closed wetlands (Tables 6, 7). No broods 

were recorded on the river strata in either year (Tables 6, 7). Scaup broods 

(Aythya spp.) were seen only on closed wetlands in both years while greenwinged teal (Anas crecca), were seen only on connected wetlands. 
20 

Table 6. Number of ducklings of each species observed within each habitat 

type in the Kvichak study area in 1992. 

 WETLAND TYPE 

 Connected Closed Moraine 

Species Wetlands Wetlands Wetlands River Total 

Scaupa

 0(0)b 1(6) 0(0) 0(0) 1(2) 

Black Scoter 0(0) 0(0) 0(0) 0(0) 0(0) 

Mallard 0(0) 2(12) 0(0) 0(0) 2(3) 

Pintail 0(0) 2(12) 0(0) 0(0) 2(3) 

Green-winged Teal 10(24) 0(0) 0(0) 0(0) 10(17) 

Wigeon 5(12) 7(41) 0(0) 0(0) 12(21) 

Unidentified 26(64) 5(29) 0(0) 0(0) 31(54) 

Total 41(100) 17(100) 0(0) 0(0) 58(100) 

a

 includes both greater and lesser scaup 

b

 number in parenthesis equal percent of total ducklings within each stratum 

Table 7. Number of ducklings of each species observed within each habitat 

type in the Kvichak study area in 1993. 

 WETLAND TYPE 

 Connected Closed Moraine 

Species Wetlands Wetlands Wetlands River Total 

Scaupa

 0(0)b 9(53) 0(0) 0(0) 9(30) 

Black Scoter 0(0) 0(0) 0(0) 0(0) 0(0) 

Mallard 7(64) 4(24) 0(0) 0(0) 11(37) 

Pintail 0(0) 0(0) 2(100) 0(0) 2(7) 

Green-winged Teal 0(0) 0(0) 0(0) 0(0) 0(0) 

Wigeon 0(0) 0(0) 0(0) 0(0) 0(0) 

Unidentified 4(36) 4(23) 0(0) 0(0) 8(26) 

Total 11(100) 17(100) 2(100) 0(0) 30(100) 

a

 includes both greater and lesser scaup 

b

 number in parenthesis equal percent of total ducklings within each stratum 
21 

Fifteen species of adult waterbirds were also seen on the study area. 

Swans (Cygnus buccinator, C. columbianus), scaup (Aythya spp.) and black 

scoter (Melanitta nigra) were the most abundant species seen in both years (Table 

3). Twenty percent more waterbird species were seen in 1993 than in 1992 (Table 

3). In both 1992 and 1993, 99% of all waterbirds seen were observed on 

connected and closed wetlands (Tables 8,9). Connected wetlands were most used 

overall with 65% of the total birds observed in both years (Tables 8,9). No 

waterbirds were recorded on the river stratum in either year. 

Statistical Analysis 

All habitat variables were originally included in principal component 

analysis and multiple stepwise regression. The results from these analysis 

indicated shoreline length, sodium and magnesium were the dominant factors 

associated with brood use of wetlands. It appeared that sodium and magnesium 

were an overall measure of cation concentration. At the same time, shoreline 

length was highly correlated with adult ducks and total waterbirds, but the 

analysis did not account for the density of birds using wetlands. Therefore, to 

allow other variables to enter the analysis, salinity variables were combined in a 

second analysis that used major cations and anions other than carbonates in place 

of each ion separately. A third analysis also used density (ducklings or adult 

ducks or waterbirds/shoreline length) as the dependent variable in multiple 

stepwise regression. The results of all 3 analysis are reported. 
22 

Table 8. Number of adult waterbirds of each species observed within each 

habitat type in the Kvichak study area in 1992. 

 WETLAND TYPE 

 Connected Closed Moraine 

Species Wetlands Wetlands Wetlands River Total 

Swansa

 33(11)b 21(13) 0(0) 0(0) 54(54) 

Scaupc

 159(52) 46(29) 0(0) 0(0) 205(43) 

Black Scoter 64(21) 61(38) 3(50) 0(0) 128(27) 

White-winged Scoter 10(3) 6(4) 0(0) 0(0) 16(3) 

Red-breasted Merganser 0(0) 1(4) 0(0) 0(0) 1(<1) 

Common Loon 0(0) 5(3) 2(33) 0(0) 7(1) 

Arctic Loon 17(6) 3(2) 1(17) 0(0) 21(4) 

Mallard 3(4) 4(3) 0(0) 0(0) 7(1) 

Goldeneyed

 2(4) 4(3) 0(0) 0(0) 6(1) 

Pintail 2(<1) 5(3) 0(0) 0(0) 1(<1) 

Oldsquaw 0(0) 1(<1) 0(0) 0(0) 1(<1) 

Green-winged Teal 7(2) 2(1) 0(0) 0(0) 9(2) 

Wigeon 1(<1) 0(0) 0(0) 0(0) 1(<1) 

Hooded Merganser 6(2) 0(0) 0(0) 0(0) 6(1) 

Bufflehead 3(1) 0(0) 0(0) 0(0) 3(1) 

Unidentified 0(0) 0(0) 0(0) 0(0) 0(0) 

Total 307(100) 159(100) 6(100) 0(0) 472(100) 

a

 tundra swans 

b

 number in parenthesis equals percent of each species in each stratum 

c

 includes both greater and lesser scaup 

d

 includes both common and barrow's goldeneye 
23 

Table 9. Number of adult waterbirds of each species observed within each 

habitat type in the Kvichak study area in 1993. 

 WETLAND TYPE 

 Connected Closed Moraine 

Species Wetlands Wetlands Wetlands River Total 

Swansa

 37(8)b 32(13) 0(0) 0(0) 69(9) 

Scaupc

 140(29) 55(22) 1(10) 0(0) 195(26) 

Black Scoter 237(49) 124(50) 0(0) 0(0) 361(48) 

White-winged Scoter 2(<1) 2(<1) 0(0) 0(0) 4(<1) 

Red-breasted Merganser 0(0) 0(0) 4(40) 0(0) 4(<1) 

Common Loon 2(4) 8(3) 5(50) 0(0) 15(2) 

Arctic Loon 0(0) 8(3) 0(0) 0(0) 8(1) 

Mallard 0(0) 3(1) 0(0) 0(0) 3(<1) 

Goldeneyed

 1(<1) 1(<1) 0(0) 0(0) 2(<1) 

Pintail 0(0) 1(<1) 0(0) 0(0) 1(<1) 

Oldsquaw 1(<1) 3(1) 0(0) 0(0) 4(1) 

Green-winged Teal 3(<1) 0(0) 0(0) 0(0) 3(<1) 

Wigeon 0(0) 1(<1) 0(0) 0(0) 1(<1) 

Hooded Merganser 0(0) 0(0) 0(0) 0(0) 0(0) 

Bufflehead 0(0) 0(0) 0(0) 0(0) 0(0) 

Unidentified 64(13) 12(5) 0(0) 0(0) 76(11) 

Total 487(100) 250(100) 10(100) 0(0) 746(100) 

a

 tundra swans 

b

 number in parenthesis equals percent of each species in each stratum 

c

 includes both greater and lesser scaup 

d

 includes both common and barrow's goldeneye 
24 

Principal Component Analysis 

Three biologically interpretable principal components (PC) were 

developed from the data set of 18 variables. The first 3 principal components 

accounted for 51% of the variance in the original data (Table 10). Magnesium, 

sodium, total nitrogen, total alkalinity, conductivity, chlorophyll a, total 

phosphorous, color and calcium loaded positively on the first principal component 

while temperature and shoreline development loaded negatively (Table 10). PC2 

described a gradient of variables where shoreline length, surface area, and 

shoreline development varied inversely with temperature, pH, total phosphorous 

and chlorophyll a. On PC3, chlorophyll a, total nitrogen and total phosphorous 

load positively, and chloride and conductivity load negatively. 

Table 10. Factor loadings for each variable along principal component axes 

1, 2 and 3. 

Habitat Variable PC1 PC2 PC3 

Total Nitrogen 0.738 -0.056 0.331 

Total Phosphorous 0.590 -0.204 0.325 

Chloride 0.482 -0.063 -0.778 

Sulfate 0.478 -0.118 -0.466 

Calcium 0.550 0.164 0.082 

Magnesium 0.824 0.007 -0.178 

Sodium 0.804 -0.116 -0.194 

Potassium 0.129 -0.051 -0.246 

Average Depth 0.070 0.041 0.025 

Surface Area 0.184 0.738 0.106 

Shoreline Length 0.159 0.920 0.045 

Shoreline Development -0.132 0.511 -0.124 

Temperature -0.316 -0.466 -0.013 

pH 0.387 -0.294 0.236 

Conductivity 0.697 -0.012 -0.665 

Total Alkalinity 0.712 0.059 0.203 

Color 0.581 0.091 0.089 

Chlorophyll a 0.691 -0.112 0.386 

Cumulative variance accounted for 50.7% 
25 

When cations and anions other than carbonates where each combined, the 

first three principal components accounted for 55% of the variance in the original 

data (Table 11). Total nitrogen, chlorophyll a, cations, total alkalinity, 

conductivity, total phosphorous and color loaded positively on PC1, while 

temperature and shoreline development loaded negatively (Table 11). On PC2, 

shoreline length, surface area and shoreline development loaded positively as 

temperature, pH, and total phosphorous were negative. On PC3, chlorophyll a, 

total phosphorous and total nitrogen were positive while anions and conductivity 

where negative (Table 11). 

Table 11. Factor loadings for each variable along principal component axes 

1, 2 and 3 with cations and anions each combined. 

Habitat Variable PC1 PC2 PC3 

Total Nitrogen 0.779 -0.073 0.290 

Total Phosphorous 0.628 -0.242 0.302 

Total Anions 0.469 0.009 -0.816 

Total Cations 0.750 -0.088 -0.117 

Average Depth 0.071 0.025 0.001 

Surface Area 0.224 0.731 0.154 

Shoreline Length 0.178 0.905 0.112 

Shoreline Development -0.143 0.511 -0.065 

Temperature -0.373 -0.492 -0.05 

pH 0.407 -0.311 0.183 

Conductivity 0.636 0.021 -0.748 

Total Alkalinity 0.660 -0.028 0.080 

Color 0.620 0.116 0.076 

Chlorophyll a 0.764 -0.140 0.355 

Cumulative variance accounted for 54.9% 

Wetlands with ducklings of all species had significantly higher PC scores 

along PC1 (t137=-2.99) than wetlands with no page ducklings (Figure 2). 

Wetlands with adult ducks of all species had significantly higher PC scores along 

PC1 (t137=-3.79, Figure 3). Wetlands with waterbirds of any species (t137=-3.46) 

as well as wetlands with swans (t137=-3.11) also had significantly higher PC 

scores along PC1 (Figures 4,5). 
26 

Explanation of variable abbreviations in Appendix 1. 

Explanation of variable abbreviations in Appendix 1. 
27 

Explanation of variable abbreviations in Appendix 1. 

Explanation of variable abbreviations in Appendix 1. 
28 

Regression Models 

In all regression models where cations and anions where analyzed 

separately, aquatic invertebrates, transparency, magnesium, chloride and total 

nitrogen were removed from the analysis because of high correlations with other 

variables (Table 12). When salinity variables where combined, and when the 

density (birds/meter of shoreline) was used as the dependent variable, aquatic 

invertebrates, transparency and total nitrogen were excluded. 
29 
30 

I conducted stepwise multiple regression for all species of adult ducks 

combined, for all species of ducklings combined, for all waterbird species, and for 

swans. Using 15 variables, stepwise regression models identified 6 habitat 

variables that most strongly influence use of wetlands by ducklings, adult ducks 

or all waterbirds. For all species of duck broods with ions separated, sodium was 

the only significant variable that contributed to the variation in habitat use 

(R2

=0.11, Table 13). When ions were combined, total cations explained 8% of 

the variation in use (Table 13). When density of ducklings was used, sodium 

(R2

=0.07), chlorophyll a (R2=0.03) and pH (R2=0.03) accounted for 15% of the 

variation in use by all species of ducklings (Table 13). 

Table 13. Stepwise multiple regression equations showing the relationship between all 

species of duck broods and 15 habitat variables on wetlands within the Kvichak study area. 

WITH IONS SEPARATED

TOTAL # DUCKLINGS = -0.423 + 0.325 (sodium) 

R2=0.11 p=0.0001 

WITH IONS COMBINED

TOTAL # DUCKLINGS = -0.312 + 0.289 (Total Cations) 

R2=0.08 p=0.0006 

AS DUCKLINGS/SHORELINE LENGTH

TOTAL # DUCKLINGS = -0.002 + 0.323 (sodium) - 0.25 (chlorophyll a) + 0.204 (pH) 

R2=0.13 p=0.0002 

For adult ducks analyzed with ions separated and with ions combined, 

total alkalinity (R2=0.12) and shoreline length (R2=0.03) explained 15% of duck 

use (Table 14). When analyzed by density, chlorophyll a explained 19% of the 

adult duck use of wetlands (Table 14). 

Table 14. Stepwise multiple regression equations showing the relationship between all 

species of adult ducks and 15 habitat variables on wetlands within the Kvichak study area. 

WITH IONS SEPARATED

TOTAL # DUCKS = -0.997 + 0.319 (total alkalinity) + 0.176 (shoreline length) 

R2=0.15 p=0.00011 

WITH IONS COMBINED

TOTAL # DUCKS = -0.997 + 0.319 (total alkaklinity) + 0.176 (shoreline length) 

R2=0.15 p=0.00002 

AS DUCKS/SHORELINE LENGTH

TOTAL # DUCKS = 0.0009 + 0.438 (chlorophyll a) 

R2=0.19 p=0.00000 
31 

For waterbirds of all species, total alkalinity (R2

=0.14) and shoreline 

length (R2

=0.03) explained 17% of the waterbird use when cations and anions 

where either separated or combined (Table 15). Chlorophyll a (R2

=0.18) and total 

alkalinity (R2=0.03) accounted for 21% of the variation in use when the density of 

waterbirds was used as the dependent variable (Table 15). 

When stepwise regression analysis was run for swans, no variables were 

significant, suggesting that there was no relationship between the 15 habitat 

variables and use of wetlands by swans. 

Table 15. Stepwise multiple regression equations showing the relationship between all 

species of waterbirds and 15 habitat variables on wetlands within the Kvichak study area. 

WITH IONS SEPARATED

TOTAL # WATERBIRDS = -0.905 + 0.346 (total alkalinity) + 0.189 (shoreline length) 

R2

=0.18 p=0.00002 

WITH IONS COMBINED

TOTAL # WATERBIRDS = -0.905 + 0.346 (total alkalinity) + 0.189 (shoreline length) 

R2=0.18 p=0.000002 

AS WATERBIRDS/SHORELINE LENGTH

TOTAL # WATERBIRDS = 0.0007 + 0.353 (chlorophyll a) + 0.199 (total alkalinity) 

R2=0.21 p=0.00000 
32 

Discussion 

Hydrology 

Murphy et al. (1984) suggested that hydrologic connection to flowing 

water is an important factor in the nutrient dynamics of Interior Alaska wetlands 

and is reflected in the patterns of habitat use by ducks. In the Iditarod George 

block of the Lower Innoko River area, morphological variables and some 

chemical and biological variables were significantly different between 

hydrologically closed and connected wetlands suggesting hydrology affected the 

chemical composition of the wetlands there (Seppi 1993). In the Kvichak area, I 

found no significant difference in limnological characteristics between closed and 

connected wetlands when all variables were compared with a multivariate T-test 

(Hotelling’s T2=27.42, p<0.1788). However T-tests for 4 individual variables 

were significant (calcium p= 0.0027, magnesium p=0.0159, shoreline length 

p=0.0029, total alkalinity p=0.0086) suggesting hydrology may influence the 

chemical composition of wetlands in the study area. 

The wetlands in the Kvichak area were not as hydrologically distinct as 

those in Lower Innoko River area. In fact, the hydrology of many wetlands was 

apparent only from aerial photos, and was often difficult to determine. Some 

wetlands were connected to ephemeral drainages and were difficult to classify as 

hydrologically connected or closed. Hydrology is important in controlling the 

chemical and biological characteristics of aquatic systems (Gosselink and Turner 

1978), and the results of this study indicate that wetland hydrology does influence 

the nutrient and cation concentrations in ponds. Connected wetlands were higher 

in total nitrogen, total phosphorous, calcium, magnesium, sodium, sulfate and 

chlorophyll a than closed wetlands (Table 5). More broods were found in 

connected wetlands in 1992 (Table 6), and 37% of connected wetlands were used 

by broods in 1993 (Table 7). Waterbirds of all species were most likely to be 

found on connected wetlands than on closed wetlands, although a substantial 

number were seen on closed tundra ponds in both years (Tables 8,9). 

Brood use data should be interpreted cautiously as it is likely that many of 

the broods present were missed because they were difficult to see when they took 

cover in surrounding vegetation, particularly on large wetlands where the float 

plane was used for access. 

Additional evidence of a correlation between water chemistry and use by 

adult ducks, broods and waterbirds in general is that multiple stepwise regression 

showed that birds were associated with large, shallow, more productive wetlands 

with higher cation (sodium) and chlorophyll a concentrations, higher alkalinity 

and longer shorelines (Tables 13,14,15). The combination of these characteristics 

was typical of all hydrologically connected and some closed tundra ponds in the 

study area. 
33 

Limnology/Wetland Traits 

Cation concentration contributes to salinity and is positively correlated 

with conductivity (Wetzel 1983, Heglund 1988, 1992, Seppi 1993). Data from 

this project shows a high correlation between sodium and chloride and 

conductivity, suggesting that NaCl is the predominant source of ions (Table 12), 

and probably is influenced by the marine environment, especially in wetlands 

close to the coast. Conductivity and individual cations loaded positively on PC1 

(especially magnesium and sodium, Table 10), as did combined cations (Table 

11). PC1 scores imply conductivity and salinity from cation concentrations had a 

substantial influence on use of wetlands by duck broods and waterbirds of all 

species (Figures 2,3,4,5). The importance of salinity to duck broods is also 

suggested by the results from multiple stepwise regression, as sodium emerges as 

1 of 2 variables with a major influence on duck brood use when ions are analyzed 

separately and on a density basis (Table 13). 

Swanson et al. (1984) has shown that highly saline water can be toxic to 

ducklings. Moderate salinity levels have been related to the density of waterfowl 

broods in boreal forest wetlands (Heglund 1988, 1992, Seppi 1993), and on the 

prairies (Swanson et al. 1988, Swanson and Duebbert 1989). Moderately saline 

wetlands have also been associated with adult waterbirds (Heglund 1992). Levels 

of salinity in the Lower Innoko River area are less than 500 μS/cm and considered 

fresh (mean=67.5 μS/cm, Seppi 1993), as are salinity levels in the Kvichak area 

(mean=29.5 μS/cm, Table 4), and both are far lower than average salinities on 

prairie wetlands (Stewart and Kantrud 1971). The positive correlation between 

brood use in the Innoko River area and duck brood/waterbird use I found in the 

Kvichak area seems to apply to slightly or moderately saline wetlands (Seppi 

1993), but becomes lethal to duck broods at very high levels (Swanson et al. 

1984). 

Total alkalinity is a measure of the buffering capacity of water resulting 

from the alkaline contributions of the negative ions of salts of weak acids (APHA 

1985). Alkalinity was measured in this study as mg/L as CaCO3. Total alkalinity 

loaded strongly on PC1 (Tables 10,11), and comparisons of wetlands with and 

without use by broods, adult ducks, waterbirds and swans with principal 

component axes 1 and 2 imply that all categories of birds were using wetlands 

with higher total alkalinities than wetlands were no birds were found (Figures 

2,3,4,5). In addition, stepwise multiple regression implies a correlation with total 

alkalinity and adult duck use with ions separated (R2=0.12, p=0.00011) and 

combined R2=0.12, p=0.00002, Table 14), as well as with use by all waterbirds 

(ions separated and combined R2=0.14, p=0.00002), and by density of waterbirds 

(R2=0.03, p=0.00000, Table 15). Total alkalinity may indirectly influence 

waterfowl use by providing a higher buffering capacity and thereby providing 

more favorable conditions to plants and animals that are attractive to waterfowl. 

This scenario was suggested for the Innoko River area where wetlands with little 

or no brood use had very low alkalinity and low aquatic invertebrate biomass 

(Seppi 1993). This relationship between alkalinity and waterfowl use was also 

found in the Kvichak area, as the majority of duck brood and waterfowl use was 
34 

on wetlands with the higher alkalinities (Tables 5,6,7,8,9). There are probably 

other factors involved in this relationship, however, as very little waterfowl use 

was seen on rivers in the Kvichak area, yet total alkalinity was highest in rivers 

compared to other strata (Table 5,6,7,8,9). 

pH was included in the multiple stepwise regression for 

ducklings/shoreline length, although the relationship was weak (R2

=0.03, 

p=0.0002, Table 14). pH loaded moderately on PC1 (Tables 10,11), and factor 

scores for wetlands with no waterfowl were low, suggesting waterfowl use 

increased on wetlands with higher pH (Figure 2,3,4,5). Low pH (<6.0) has a 

negative affect on aquatic invertebrates (Eilers et al. 1984, Bell 1971), including 

benthic organisms (Haines 1981), although some species are more sensitive than 

others. McAuley and Longcore (1988) found diets of ring-necked ducklings from 

high pH wetlands in maine (>6.0) to be more diverse than those from low pH 

wetlands (<6.0), and suggested retarded growth and lower survival rates could 

result if ducklings were forced to feed on less nutritious and more mobile species 

when invertebrate diversity was low. Low pH may therefore indirectly affects 

wetland use by reducing abundance of invertebrate species important to broods, as 

well as adult birds. Heglund (1992), found broods of all species of ducklings used 

wetlands higher in pH, with the exception of deep water divers (ring-necked duck 

Aythya collaris, barrow’s bucephala islandica and common goldeneye Bucephala

clangula, bufflehead, Bucephala albeola) which were found in slightly more 

acidic water. On the Yukon Flats, Alaska, most species of adult waterbirds used 

wetlands with moderate to high pH, although deep water divers and loons tended 

to use wetlands with pH 7.0 (Heglund 1992). In the Kvichak area, all strata had 

pH values very near 7.0 (Table 5). The relationship between pH and waterfowl 

use of wetlands I found is weak, but this may be attributed to the narrow range of 

pH of wetlands in the Kvichak area (5.5 to 9.5, Table 4). In addition many 

species adult waterfowl may be habitat generalists, especially post-breeding males 

and unmated pairs, making relationships with any habitat variable hard to detect. 

Temperature loaded negatively on PC1 (Tables 10,11), and PC1 scores for 

wetlands used by waterfowl suggest birds preferred cooler wetlands (Figures 

2,3,4,5). This variable is hard to interpret, because surface water temperature may 

vary depending on weather conditions. Therefore the relationship may actually be 

artificial and related to other habitat variables, since temperature is not highly 

negative on PC1 (-0.316 Table 10, -0.373 Table 11). Moraine wetlands tended to 

be slightly warmer than other strata, yet they did not have any of the 

characteristics desirable to waterfowl (i.e. shallow depth, cover, plant and 

invertebrate foods). Therefore I think birds were not actually using wetlands that 

were cooler, but were using cooler wetlands that provided them with the food and 

cover which the warmer, moraine wetlands did not provide. 

Shoreline development also loaded negatively on PC1 (Tables 10,11), and 

PC1 scores for wetlands used by waterfowl suggest birds use wetlands with more 

round, less convoluted shorelines (Figures 2,3,4,5). At the same time, shoreline 

length emerged as an important factor for use by adult ducks and all species of 

waterbirds (Table 14,15). Multiple stepwise regressions imply birds occur every 

1700 to 1800 meters of shore length on average (Tables 13,14,15). Longer 
35 

shorelines provide more shallow areas, emergent vegetation and escape cover 

important to broods and molting waterfowl. Surface area and shoreline 

irregularity of man-made ponds in Quebec influenced broods use; however those 

same ponds had abundant emergent vegetation and high densities of aquatic 

invertebrates which provided broods with adequate cover and food (Belanger and 

Couture 1988). Small stock ponds in Montana were likely ignored by broods 

because of the lack of escape cover and food, not because of their small size 

(Hudson 1983). Shoreline length was also associated with dabbler brood use in 

South Dakota stock ponds, but wetlands with long shorelines provided more 

shallow water and emergent shoreline vegetation than wetlands with smaller shore 

lengths (Mack and Flake 1980). Results from Heglund (1992) again imply a 

strong relationship between shoreline length and use of wetlands by 6 species of 

dabbling ducks, 11 species of diving ducks and 6 other species of waterbirds. 

Shoreline length and shoreline development have both been related to waterbird 

use in this study, although the two variables are correlated (0.51, Table 12). I 

believe the relationship with wetland shore length and shape in this study is also a 

result of birds using wetlands that provide them with the most food and cover, and 

not of them using wetlands because of there morphological characteristics. Many 

of the wetlands in the Kvichak area are tundra ponds, which tend to be more 

circular than the convoluted shorelines of moraine wetlands which lay between 

the small hills and valleys created by the moraine itself. Large tundra ponds 

provide waterfowl with shallow depths, and relatively more cover and plant and 

invertebrate foods than moraine wetlands with long, convoluted shorelines (see 

shoreline development, Table 5). I believe birds are using the connected and 

closed tundra ponds more because they have more food and cover yet are more 

round, not because they have a lower shoreline development than moraine 

wetlands, which provide lesser amounts of aquatic invertebrate food and cover 

(Table 5). 

Phosphorous and nitrogen are the two most important nutrients governing 

wetland primary productivity (Kadlec 1979, Wetzel 1983), although phosphorous 

is more often limiting in aquatic systems and far more important to wetland 

productivity in comparison to nitrogen (Schindler 1974). Phosphorous 

concentrations have also been related to annual rates of primary productivity of 

algae in many lakes world wide (Vollenweider 1979, in Wetzel 1983). Total 

phosphorous and total nitrogen loaded strongly positively on PC1 (Tables 10,11). 

The plotted factor scores of wetlands with brood, waterfowl and general waterbird 

use suggests birds used wetlands higher in total nitrogen and total phosphorous 

concentrations (Figures 2,3,4,5). 

Eutrophication status (level of primary productivity) has been related to 

concentrations of total phosphorous (Vollenweider 1968); lakes are considered 

eutrophic if total phosphorous concentrations exceed 20 μg/L. Across all 

wetlands, total phosphorous concentrations averaged 36 μg/L (Table 4), and 

wetlands in all habitat strata were above 20 μg/L, except for moraine wetlands 

(9.64 μg/L, Table 5). This would indicate that all wetlands could be considered in 

or near a eutrophic state, with the exception of Moraine wetlands. Total 

phosphorous concentrations were positively correlated with dabbling and diving 
36 

duck use of oligotrophic lakes in Sweden (Nilsson and Nilsson 1978). This 

association between phosphorous and waterfowl use was related to higher levels 

of productivity in more eutrophic lakes. Increases in total phosphorous levels due 

to pollution also caused higher productivity, but did not lead to higher waterbird 

species richness in formerly oligotrophic lakes. Similarly, Nilsson (1978) 

reported breeding waterfowl used naturally more eutrophic lakes more often. 

However when oligotrophic lakes where polluted with phosphorous, diving 

species disappeared, and dabbling species first increased then declined with 

increasing eutrophication (Nilsson 1978). Duck density was directly related to 

natural levels of phosphate and nitrite in boreal forest wetlands in Alaska 

(Murphy et al. 1984). Heglund (1988) showed waterbird density, including adult 

dabbling and diving ducks, to be related to total phosphorous and total nitrogen. 

Heglund (1992) found associations between omnivorous dabbling broods and 

total phosphorous concentrations. 

A positive relationship between annual average chlorophyll a

concentrations and total phosphorous has been found for many lakes 

(Vollenweider 1979, cited in Wetzel 1983). I found that as chlorophyll a

concentrations increased, duck brood density decreased (Table 13), while adult 

duck and waterbird density increased (Tables 14,15). In addition, total 

phosphorous and chlorophyll a loaded strongly positively on PC1, and the plotted 

factor scores of wetlands with duck broods, adult ducks, swans and waterbirds of 

all species shows a relationship between bird use and wetlands with higher levels 

of total phosphorous and chlorophyll a (Figures 2,3,4,5), suggesting productive 

wetlands with higher nutrient levels are attractive to waterbirds. This relationship 

was apparent in moraine wetlands where total phosphorous and chlorophyll a

levels were low, and waterfowl use was low (Tables 5, 6,7,8,9). The negative 

correlation between chlorophyll a and broods may not be real but a result of an 

inadequate count of the broods that were actually present. Many large, productive 

wetlands were used by broods but were not seen during surveys, and suggests a 

relationship that is probably false. 

Aquatic Invertebrates 

Aquatic invertebrates are an important source of food for ducklings (Chura 

1960, Bartonek and Hickey 1969, Bengston 1971, Sudgen 1973, Krapu and 

Swanson 1977, Street 1978, Eriksson 1978, Joyner 1980, Talent et al. 1982, 

Hohman 1985, Jarvis and Noyes 1986). In the prairie pothole region fresh (<500 

μS/cm) to slightly brackish (500-2000 μS/cm) wetlands (as defined by Stewart 

and Kantrud 1971) support the greatest abundance and diversity of aquatic 

invertebrates (LaBaugh and Swanson 1988, Kantrud et al. 1989a). Submerged 

aquatic plants play a crucial role in providing habitat for macroinvertebrates 

(Krull 1970) and are therefore also a major factor in determining their abundance 

and availability. Aquatic plant communities are themselves important for broods 

as they provide escape cover (Patterson 1976) and food for some species, 

particularly in the later stages of development before fledging (Sudgen 1973, 

Swanson and Duebbert 1989). In the prairie pothole region, Stewart and Kantrud 
37 

(1972) found that plant community composition was related to wetland salinity, 

and that many plant species were intolerant of highly saline conditions. In 

general, species diversity of aquatic vegetation decreases with increasing salinity 

(Kantrud et al. 1989b), which in turn reduces potential habitats for aquatic 

invertebrates. Therefore salinity indirectly influences brood use of wetlands by 

affecting the potential of wetlands to support the species composition and 

abundance of aquatic plants and invertebrates that are attractive to waterfowl 

(Joyner 1980, Swanson et al. 1988, Heglund 1992). Results of this study show 

waterfowl use was highest on closed and connected wetlands (Tables 6,7,8,9). 

These wetlands had higher cation and anion concentrations and the highest 

invertebrate biomass (Table 5), which apparently provided a richer food source 

for birds. The aquatic invertebrate data is limited, however (31 wetlands in 1993, 

Appendix 2), and for this reason was not included in the multivariate statistical 

analysis. Simple linear regression with aquatic invertebrates as the independent 

variable reveals a correlation with all waterbirds (r2

=0.26, p=0.003) and adult 

ducks (r2

=0.26, p=0.003). 

Brood Use 

The use of wetlands by waterfowl and other waterbirds was most strongly 

associated with sodium, total alkalinity, and shoreline length or total cations, total 

alkalinity and shoreline length, depending on the separation or combination of 

cations and anions (Tables 13,14,15). These results suggest that sodium is a 

measure of overall ionic concentration, which is a logical result from an area that 

is in close proximity to a marine environment. When the density of waterfowl 

(birds/m of shoreline) is compared with limnological variables, stepwise 

regression equations imply and association of sodium, chlorophyll a and total 

alkalinity and use by ducklings, adult ducks, and waterbirds (Tables 13,14,15). 

No waterfowl were seen on river strata during surveys or at any other 

time, although the two rivers (Kvichak and Alagnak) were surveyed for chemistry 

and waterfowl only 4 times over the two years. Both rivers were extremely clear 

with swift currents and had no visible aquatic vegetation and were probably not 

attractive to broods, but use was undoubtedly underestimated because of small 

sample sizes in the river stratum. 

The greatest total number of waterfowl of all species and the highest 

biomass of aquatic invertebrates were found on connected wetlands (Tables 

5,6,7,8,9). This suggests that birds were using connected wetlands to feed where 

invertebrates were more abundant. Connected wetlands provided conditions of 

higher total alkalinity and higher water color. Darker water color was associated 

with brood use in the Innoko River area (Seppi 1993), but wetlands were very 

clear in the Kvichak and there was no apparent relationship with waterfowl use 

and water color. 

Connected and some closed wetlands provided large areas of shallow 

water and long shorelines lined with emergent vegetation which were important to 

broods for cover and food. I found an association between shoreline length and 

use by broods and all types of waterbirds (Tables 13,14,15). I did not relate use 
38 

directly with the amount of shoreline cover, nor did I relate waterfowl use to 

individual species of plants. Shoreline length may be an indirect method of 

evaluating plant cover and the extent of shallow areas of a wetland. Many other 

studies have reported relationships between shoreline cover and brood use (Mack 

and Flake 1980, Hudson 1883, Belanger and Couture 1988, Heglund 1992) and 

indirect associations were found for broods in the Lower Innoko River area (Seppi 

1993), and on the Kvichak area for all waterbirds. 

Nitrogen and phosphorous have been recognized as limiting factors in 

productivity in boreal forest wetlands in Alaska (Alexander and Barsdate 1971, 

1974, Barsdate and Alexander 1971). Moraine wetlands had lower chlorophyll a

concentrations and lower aquatic invertebrate biomass associated with lower 

phosphorous levels, making them less attractive to waterfowl. In the Kvichak 

area, I believe broods used wetlands that provided them with maximum cover and 

food, which was found on ponds with increased biotic activity resulting from 

higher nutrient levels. The river strata had the highest total phosphorous 

concentrations, but no waterfowl use. Some species of waterfowl do use rivers 

for brooding (i.e. common and red-breasted merganser, harlequin), although most 

species of dabbler and diver prefer ponds and are not able to take advantage of 

deep or swift current rivers. 

All habitat variables notwithstanding, I believe the ability of field crews to 

spot and identify broods and differences in the visibility of broods among wetland 

types may have influenced the results. Small, deep moraine wetlands and small 

closed or connected tundra ponds were easy to survey for broods and there was no 

doubt all birds were accounted for. However, larger wetlands with long, 

convoluted shorelines and thick shoreline cover were very difficult to survey and 

often held higher densities of broods and waterfowl, which were momentarily 

visible on approach by plane. But after landing and canoeing or walking the 

shorelines, most birds particularly broods, could not be flushed to be counted, 

resulting in vast underestimations of use. This is apparent in the low total 

numbers of broods seen in each year and in the large percentage of birds that were 

classified as unidentified. More substantial statistical associations could be 

determined for adult ducks and other waterbirds. This is probably a result of the 

greater visibility of these birds as well as the probability that they are habitat 

generalists and do not require abundant food and cover in specific wetlands when 

compared to broods. Adult birds are mobile and may have been seen on wetlands 

that provided them with loafing sites but not food gathering habitat. It is possible 

that I never observed pre-breeding and non-breeding adult birds in the habitats 

that are important to them for food (i.e. coastal marine environments). 

In conclusion, wetland use by broods and waterfowl is related to several 

limnological and biological factors. Productive lentic wetlands with hydrologic 

connection to flowing water, moderate salinities and relatively high alkalinities 

are used most by duck broods, adult ducks, swans and other waterbirds. As with 

all biological systems, these factors do not function independently of one another, 

and may be indirect determinants of the physical and biological habitat variables 

that actually attract waterfowl. 
39 

Management Implications 

The vast size and remote location of BLM lands in Alaska creates unique 

management problems requiring decisions that must account for pending land 

selection procedures, subsistence use of wildlife resources, public use of federal 

lands and use of other natural resources, as well as long-term wildlife 

management goals. Therefore, to be cost effective and appropriate, management 

decisions on important wetlands and waterfowl habitat require an understanding 

of the ecology and habitat requirements of nesting, breeding and migrating 

waterfowl. 

The Kvichak area is a mosaic of private, federal, state and Native 

corporation lands, a situation common to other important waterfowl habitat on 

BLM lands in Alaska. There are increasing economic incentives to develop 

natural resources (i.e. minerals development) in the Bristol Bay river drainages. 

At the same time, sport hunting, sport and commercial fishing and subsistence 

hunting remain an important way of life for the region. These lifestyles and the 

desire to maintain them may persuade Native and hunting and fishing groups, and 

the general public, to protect these lands and allow this largely pristine ecosystem 

to remain intact, and thereby protect vital wetlands important to staging and 

breeding waterfowl. 

Due to its geographic location and size, management to improve or create 

waterfowl habitat in the Kvichak area is impractical. The production of 

waterfowl in the area is lower than other areas of the state (Hodges and Conant 

1990) with the exception of swans (Groves et al. 1990, Groves and Conant 1991). 

Yet, the Kvichak area may be an important staging ground for waterfowl bound 

for the Yukon Kuskokwim Delta and the North Slope, particularly in the spring 

before breeding ground ponds are ice free. Protection of the area is the best 

management practice by insuring that needed migration and breeding habitat for 

waterfowl, as well as many other species of wildlife that depend on the Kvichak 

and Alagnak river drainages, will remain for future production. 
40 

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45 

Appendix 1. Explanation of variable abbreviations.

Invert = aquatic invertebrates (mg/cm2) 

Trans = transparency (m) 

Temp = temperature (oC) 

pH = pH (no units) 

Cond = conductivity (μS/cm) 

T. Alk = total alkalinity (mg/L) 

Color = water color (platinum cobalt units) 

TP = total phosphorous (μg/L) 

TN = total nitrogen (mg/L) 

Ca = calcium (mg/L) 

Mg = magnesium (mg/L) 

K = potassium (mg/L) 

Na = sodium (mg/L) 

Cl = chloride (mg/L) 

SO4 = sulfate (mg/L) 

Chloroa = chlorophyll a (mg/L) 

Surface = surface area (hectares) 

SLL = shoreline length (m) 

SLD = shoreline development index (no units) 

Ave Depth = average depth (m) 
46 

Appendix 2. Raw data of 20 habitat variables for each wetland. 

Date WB TN TP Cl SO4 Tot-An Ca Mg Na K Tot-Cat Ave-Depth 

6-25-92 299 0.293 20.3 5.00 0.15 5.15 1.10 0.95 2.81 0.64 5.50 0.41 

6-25-92 301 0.360 22.7 1.70 1.40 3.10 1.53 1.07 2.29 0.54 5.43 0.67 

6-25-92 302 0.236 21.0 .020 0.15 0.35 0.24 0.24 1.15 0.30 1.93 0.38 

6-25-92 303 0.220 19.7 0.40 0.15 0.55 0.52 0.53 1.69 0.23 2.97 1.20 

6-26-92 298 0.64 

6-26-92 304 0.40 0.15 0.55 0.12 0.17 1.08 0.26 1.63 1.45 

6-26-92 305 0.25 

6-28-92 274 0.25 

6-28-92 276 0.400 32.0 1.40 0.50 1.90 0.52 0.32 2.00 0.33 3.17 0.25 

6-28-92 277 0.38 

6-28-92 300 0.47 

6-28-92 290 0.266 20.3 1.30 0.60 1.90 1.69 1.02 2.99 0.46 6.16 

6-28-92 295 0.116 7.00 .080 0.154 0.95 0.16 0.15 0.77 0.31 1.39 4.31 

6-28-92 296 0.143 6.00 1.40 0.15 1.55 0.12 0.17 0.87 0.29 1.45 3.41 

6-28-92 297 1.99 

6-29-92 280 0.246 10.0 0.90 0.15 1.05 0.76 0.31 1.65 0.27 2.99 3.98 

6-29-92 281 1.24 

6-29-92 284 0.216 12.0 0.50 0.15 0.65 1.16 0.54 2.27 0.32 4.29 0.25 

6-29-92 285 0.50 

6-29-92 288 0.38 

6-29-92 289 0.45 

6-29-92 291 0.75 

6-29-92 294 1.38 

6-30-92 149 1.88 

6-30-92 150 0.343 70.0 2.90 0.15 3.05 2.04 1.54 14.1 0.63 18.31 0.42 

6-30-92 153 0.263 15.0 0.60 0.15 0.75 0.68 0.34 1.39 0.29 2.70 0.50 

6-30-92 156 0.42 

6-30-92 158 0.310 13.7 0.50 0.15 0.65 0.92 0.62 2.12 0.34 4.00 1.84 

6-30-92 160 5.33 

6-30-92 161 0.353 22.3 0.60 0.15 0.75 0.22 0.20 1.00 0.13 1.55 0.50 

7-1-92 164 1.00 

7-1-92 207 0.266 8.70 0.40 0.15 0.55 0.18 0.24 1.24 0.56 2.22 2.77 

7-1-92 208 0.296 8.70 0.60 0.15 0.75 0.58 0.29 1.57 0.23 2.67 

7-1-92 209 

7-1-92 214 0.290 18.3 0.30 0.15 0.45 0.53 0.51 1.66 0.38 3.08 0.38 

7-1-92 216 0.490 54.7 0.70 0.40 1.10 1.10 1.04 2.73 0.35 5.22 0.94 

7-1-92 218 0.50 

7-1-92 222 0.46 

7-1-92 223 0.413 29.0 0.30 0.15 0.45 1.03 .095 1.86 0.38 4.22 0.71 

7-2-92 254 1.09 

7-2-92 255 0.323 27.0 0.30 0.30 0.60 0.30 0.28 1.57 0.24 2.39 0.44 

7-2-92 257 2.45 

7-2-92 258 0.46 

7-2-92 259 0.253 20.3 0.80 0.15 0.95 2.60 1.49 3.56 0.79 8.44 1.35 

7-2-92 140 0.63 

7-2-92 143 0.376 33.7 2.90 .015 3.05 1.34 1.52 3.92 0.78 7.56 0.57 

7-2-92 145 0.220 20.3 1.50 0.40 1.90 0.36 0.44 2.17 0.24 3.21 0.46 

7-2-92 146 0.63 

7-3-92 8B 0.69 

7-3-92 8C 0.326 30.0 0.60 0.15 0.75 1.13 1.33 3.66 0.46 6.58 1.22 

7-3-92 8D 0.25 

7-3-92 145A 0.340 38.3 1.00 0.30 1.30 0.44 0.36 2.06 0.17 3.03 0.25 

7-3-92 145E 0.340 19.0 1.20 0.15 1.35 3.67 2.59 3.60 0.56 10.42 0.38 

7-3-92 183 0.56 

7-3-92 136 0.246 10.7 0.40 0.15 0.55 0.22 0.14 1.09 0.16 1.61 0.25 

7-3-92 137 1.28 

7-4-92 183 0.133 10.0 0.80 0.70 1.50 2.68 1.08 2.27 0.38 6.41 

7-4-92 188 0.223 7.30 1.80 0.30 2.10 7.68 0.29 1.46 0.40 9.83 3.63 

7-4-92 191 0.443 11.7 .040 0.15 0.55 0.20 0.21 1.19 0.82 2.42 0.63 

7-4-92 198 0.320 19.0 0.50 0.15 0.65 0.24 0.26 1.37 0.43 2.30 0.39 
47 

Date WB TN TP Cl SO4 Tot-An Ca Mg Na K Tot-Cat Ave-Depth 

7-7-92 4E 

7-7-92 22 0.300 25.70 0.70 0.15 0.85 0.95 0.17 1.23 0.15 2.50 0.50 

7-7-92 23 0.406 84.70 0.60 0.15 0.75 1.28 1.10 2.43 0.48 5.29 0.82 

7-7-92 30 0.686 56.00 1.70 0.15 1.85 1.24 1.00 3.42 0.50 6.16 0.70 

7-7-92 32 0.77 

7-7-92 4C 0.386 51.30 1.00 0.15 1.15 2.07 1.32 2.37 0.39 6.15 1.35 

7-8-92 96A 0.146 6.7 1.2 0.15 1.35 0.15 0.15 0.95 0.15 1.40 

7-8-92 90A 0.156 11.0 0.7 0.15 0.85 0.70 0.51 1.65 0.25 3.11 

7-8-92 100C 0.446 41.30 2.80 0.90 3.70 2.58 1.32 4.62 0.60 9.12 2.31 

7-9-92 86 0.83 

7-9-92 88 0.433 41.00 1.60 0.15 1.75 1.35 0.85 2.02 0.39 4.61 0.52 

7-9-92 94 0.363 38.70 0.40 1.10 1.50 4.11 1.62 2.65 0.58 8.96 0.82 

7-10-92 15 0.110 93.00 0.30 0.15 0.45 0.22 0.19 1.12 0.15 1.68 1.89 

7-10-92 41 0.166 7.300 0.80 0.15 0.95 0.86 0.50 1.66 0.38 3.40 7.06 

7-10-92 61 0.276 15.50 0.60 0.15 0.75 0.33 0.28 1.11 0.17 1.89 0.42 

7-10-92 81 0.373 30.00 1.20 0.15 1.35 1.85 1.44 2.87 0.48 6.64 1.25 

7-11-92 186 0.490 31.7 0.70 0.15 0.85 2.03 1.38 2.42 0.51 6.34 0.25 

7-11-92 183 0.096 127.0 0.60 1.00 1.60 2.69 1.16 2.44 0.40 6.69 

7-11-92 kvic 

7-11-92 39 0.366 23.30 1.10 0.15 1.25 2.18 1.65 2.20 0.31 6.34 0.75 

7-11-92 68 0.380 36.30 0.80 0.10 0.90 0.51 0.55 3.00 0.37 4.43 0.50 

7-13-92 12 0.466 27.30 0.70 2.50 3.20 0.83 0.70 1.79 0.30 3.62 1.40 

7-13-92 14 0.356 28.00 1.20 0.15 1.35 1.69 1.17 2.55 0.44 5.85 3.84 

7-13-92 54 0.330 25.70 1.30 0.15 1.45 1.59 1.21 2.27 0.39 5.46 0.44 

7-14-92 6 0.426 42.00 1.10 0.15 1.25 0.87 0.65 2.04 0.31 3.87 0.75 

7-14-92 310 0.803 85.0 0.60 0.15 0.75 2.42 1.74 2.84 0.82 7.82 

7-14-92 312 1.196 138.0 1.50 0.15 1.65 2.71 2.41 14.1 2.22 21.44 0.28 

7-14-92 20 0.380 61.00 0.40 0.15 0.55 0.45 0.28 1.95 0.24 2.92 0.25 

7-14-92 21 0.256 24.30 0.40 0.15 0.55 0.38 0.20 1.40 0.16 2.14 0.25 

7-14-92 47 0.343 72.00 2.60 0.15 2.75 1.21 0.86 5.06 0.54 7.67 0.85 

7-15-92 2 0.293 16.70 0.50 .015 0.65 0.49 0.16 1.15 0.19 1.99 0.25 

7-15-92 66 0.123 18.00 0.50 1.10 1.60 4.27 2.36 2.95 0.89 10.47 3.19 

7-15-92 67 0.653 42.70 1.70 0.60 2.30 2.91 2.70 9.60 1.05 16.26 0.42 

7-15-92 70 0.380 22.70 0.70 0.15 0.85 1.10 0.70 2.09 0.30 4.20 0.75 

7-16-92 8 0.566 118.3 2.70 0.15 2.85 4.38 1.57 6.10 1.24 13.29 0.80 

7-16-92 73 0.33 

7-16-92 79 0.233 14.00 0.40 0.15 0.55 0.48 0.20 1.07 0.80 1.83 1.08 

7-16-92 84 0.333 26.70 1.50 0.15 1.65 0.75 0.64 2.10 0.55 4.04 0.25 

7-17-92 69 0.370 39.70 1.10 0.30 1.40 0.34 0.30 1.43 0.19 2.26 0.50 

7-17-92 71 0.65 

7-17-92 85 0.413 35.00 0.70 0.40 1.10 1.30 0.81 2.95 0.32 5.38 0.63 

6-19-93 302 0.310 28.00 1.20 0.70 1.90 0.16 0.14 1.78 0.28 2.36 0.67 

6-19-93 303 0.353 16.00 1.02 0.30 1.50 0.43 0.34 1.85 0.26 2.85 1.23 

6-19-93 305 0.347 30.00 1.60 1.70 3.30 0.88 0.60 3.11 0.71 5.30 0.30 

6-19-93 299 0.540 85.00 1.20 0.60 1.80 0.76 0.40 2.56 0.44 4.16 0.88 

6-19-93 301 0.373 35.00 1.60 0.20 1.80 1.52 0.79 2.81 0.61 5.73 0.5 

6-20-93 276 0.276 32.00 1.60 0.80 2.40 0.30 0.20 2.16 0.47 3.13 0.33 

6-20-93 277 0.210 14.00 0.80 0.05 0.85 0.18 0.12 1.20 0.48 1.99 0.34 

6-20-93 274 0.207 12.00 1.20 0.05 1.25 0.16 0.10 1.15 0.49 1.90 0.38 

6-20-93 304 0.103 12.00 1.60 0.05 1.65 0.10 0.10 0.79 0.62 1.61 

6-20-93 296 0.030 6.00 0.80 0.05 0.85 0.11 0.11 1.09 0.26 1.57 3.28 

6-20-93 295 0.053 12.00 0.800 0.200 1.00 0.28 0.06 0.93 0.32 1.59 5.18 

6-20-93 298 0.167 11.00 0.80 0.10 0.90 0.36 0.24 1.27 0.16 2.03 0.89 

6-20-93 297 0.373 49.00 1.20 0.30 1.50 1.50 1.22 2.38 0.57 5.67 1.49 

6-20-93 300 0.630 8.00 0.80 0.05 0.85 0.03 0.07 0.76 0.17 1.03 0.71 

6-21-93 294 0.103 10.00 0.80 0.05 0.85 0.06 0.08 0.90 0.18 1.22 1.22 

6-21-93 290 0.230 17.00 2.10 0.10 2.20 0.90 0.45 2.40 0.28 4.03 1.04 

6-21-93 291 0.277 28.00 0.80 1.30 2.10 1.14 0.54 1.86 0.34 3.88 0.83 

6-21-93 288 0.293 33.00 2.30 0.30 2.60 1.80 0.89 3.41 0.85 6.95 0.66 

6-21-93 281 0.430 29.00 0.80 0.10 0.90 1.12 0.82 2.54 0.38 4.86 1.58 

6-21-93 280 0.300 13.00 2.00 0.80 2.80 0.48 0.25 1.81 0.30 2.84 6.69 

6-21-93 285 0.190 19.00 0.80 0.30 1.10 0.15 0.18 2.09 0.37 2.79 1.33 
48 

Date WB TN TP Cl SO4 Tot-An Ca Mg Na K Tot-Cat Ave-Depth 

6-21-93 289 0.330 47.00 0.80 0.70 1.50 1.22 0.88 3.30 0.67 6.07 

6-22-93 161 0.427 29.00 0.80 0.50 1.30 0.17 0.11 1.14 0.18 1.60 0.33 

6-22-93 164 0.623 245.00 0.80 0.40 1.20 1.12 0.65 2.10 0.42 4.29 1.00 

6-22-93 158 0.253 17.00 1.20 0.10 1.30 1.04 0.43 1.82 .036 3.65 2.96 

6-22-93 156 0.267 13.00 0.80 0.05 0.85 1.56 0.14 1.23 0.26 3.19 0.63 

6-22-93 284 0.260 13.00 1.20 0.05 1.25 0.84 0.32 2.52 0.42 4.10 0.81 

6-22-93 150 1.440 211.0 4.60 0.80 5.40 2.04 1.15 12.00 1.33 16.52 0.68 

6-22-93 149 0.177 11.00 1.20 0.30 1.50 0.18 0.17 1.67 0.30 2.32 3.95 

6-22-93 153 0.400 18.00 0.80 0.05 0.85 0.34 0.14 1.40 0.28 2.16 0.50 

6-23-93 223 0.493 44.00 0.80 1.30 2.10 0.70 0.59 2.16 0.47 3.92 1.50 

6-23-93 207 0.317 10.00 2.00 0.05 2.05 0.47 0.14 1.49 0.58 2.68 3.11 

6-23-93 208 0.273 14.00 1.20 0.40 1.60 1.34 0.26 1.64 0.38 3.62 

6-23-93 214 0.407 35.00 1.20 1.40 2.60 0.66 0.26 2.57 0.44 3.93 

6-23-93 209 0.377 15.00 3.00 0.20 3.20 0.96 0.64 4.86 0.52 9.68 0.38 

6-23-93 222 0.370 30.00 1.20 1.10 2.30 0.68 0.48 2.46 .034 3.96 0.58 

6-23-93 216 0.657 88.00 1.20 1.00 2.20 0.83 0.76 2.68 0.63 4.90 1.31 

6-23-93 218 0.490 409.00 1.60 1.30 2.90 2.43 1.78 3.52 1.06 8.79 0.53 

6-24-93 1010 0.143 4.00 0.80 0.40 1.20 0.23 0.09 0.95 0.36 1.63 3.00 

6-24-93 1011 0.133 6.00 1.02 0.05 1.25 0.32 0.18 1.82 0.26 2.58 3.31 

6-24-93 1012 0.110 7.00 1.20 0.30 1.50 0.18 0.18 1.23 0.30 1.89 4.90 

6-24-93 1013 0.293 24.00 2.00 0.10 2.10 0.40 0.30 1.84 0.42 2.96 1.98 

6-24-93 1014 0.103 9.00 0.80 1.50 2.30 2.54 0.56 1.86 0.62 5.58 

6-24-93 1000 0.153 9.00 0.80 0.20 1.00 0.09 0.11 0.96 0.22 1.38 4.96 

6-24-93 1001 0.233 8.00 1.70 0.30 2.00 0.14 0.12 1.02 0.40 1.68 6.65 

6-24-93 1002 0.140 5.00 1.60 0.05 1.65 0.14 0.14 1.01 0.29 1.58 8.61 

6-24-93 1003 0.247 9.00 1.20 0.40 1.60 0.25 0.16 1.48 0.36 2.25 5.40 

6-24-93 1004 0.323 19.00 1.20 1.20 2.40 0.35 0.34 1.91 0.31 2.90 0.90 

6-27-93 259 0.337 43.00 3.10 0.40 3.50 1.58 0.82 2.88 0.44 5.75 

6-27-93 257 0.280 8.00 2.50 0.30 2.80 0.52 0.34 1.82 0.46 3.14 

6-27-93 258 0.317 55.00 3.60 0.40 4.00 0.36 0.20 2.35 0.29 3.20 

6-27-93 140 1.007 193.00 5.10 0.70 5.80 1.35 2.24 4.68 2.21 13.48 

6-27-93 143 0.500 60.00 3.00 1.80 4.80 1.18 1.10 3.98 0.520 6.78 

6-28-93 105A 0.540 31.00 1.70 0.30 2.00 1.45 0.95 2.94 0.74 6.08 2.56 

6-28-93 100A 0.553 41.00 1.60 0.60 2.20 1.51 0.55 1.04 0.94 7.37 0.97 

6-28-93 100B 0.657 43.00 2.00 0.20 2.20 1.53 0.78 3.08 0.74 6.13 0.72 

6-28-93 96 0.620 28.00 1.60 0.10 1.70 1.66 0.89 3.38 0.58 6.51 1.00 

6-28-93 105A 0.303 20.00 2.50 0.05 2.55 1.14 0.92 2.17 0.53 4.76 4.84 

6-28-93 100 0.570 45.00 2.00 1.00 3.00 3.14 1.57 5.18 1.02 10.91 1.04 

6-28-93 96A 0.577 38.00 1.20 0.10 1.30 1.81 0.96 3.50 0.64 6.91 

6-29-93 1014 0.527 49.00 2.60 0.10 2.70 3.03 0.56 4.99 1.02 9.60 4.79 

6-29-93 105B 0.310 27.00 1.20 0.10 1.30 1.82 0.74 3.80 0.64 7.01 2.33 

6-29-93 111B 0.177 23.00 1.20 0.05 1.25 0.76 0.39 1.74 0.32 3.21 1.60 

6-29-93 100C 0.330 44.00 3.10 0.80 3.90 1.94 1.02 4.74 0.77 8.47 

6-29-93 111B 0.207 14.00 1.60 0.05 1.65 1.45 0.56 2.00 0.28 4.29 1.54 

6-29-93 105B 0.733 38.00 3.00 0.40 3.40 1.02 0.50 4.12 0.52 6.16 0.69 

6-29-93 100C 0.157 19.00 0.80 0.10 0.90 0.97 0.64 2.48 0.18 4.27 

6-30-93 95A 0.210 17.00 1.20 0.20 1.40 0.58 0.24 1.50 0.32 2.64 0.78 

6-30-93 88 0.653 35.00 1.60 0.05 1.65 1.13 0.54 2.12 0.43 4.22 1.00 

6-30-93 12 0.297 32.00 1.20 0.40 1.60 0.70 0.56 1.71 0.39 3.60 

6-30-93 15 0.120 13.00 1.60 0.05 1.65 0.24 0.80 1.14 0.20 1.66 1.38 

6-30-93 90A 0.260 19.00 1.20 0.05 1.25 0.24 0.10 1.40 0.22 1.96 0.68 

6-30-93 90 0.403 37.00 0.80 0.40 1.20 1.37 0.94 2.47 0.80 5.58 0.38 

7-1-93 311A 0.463 44.00 0.80 0.20 1.00 1.21 0.94 1.72 0.76 4.63 0.95 

7-1-93 311B 0.763 79.00 0.80 1.00 1.80 1.36 1.30 3.31 0.55 6.52 0.79 

7-1-93 311C 0.183 14.00 1.20 0.05 1.25 0.37 0.26 1.26 25.00 26.89 2.19 

7-1-93 4B 0.397 20.00 1.20 0.50 1.70 0.28 0.14 1.80 0.25 2.47 0.75 

7-1-93 311 0.560 44.00 0.88 0.60 1.40 1.92 1.20 5.01 1.32 9.45 0.938 

7-1-93 4 0.377 90.00 1.60 0.60 2.20 0.43 0.28 2.40 0.34 3.45 0.58 

7-1-93 6 0.350 51.00 7.00 0.80 7.80 0.49 0.30 3.20 0.54 4.55 0.58 

7-1-93 310 0.940 133.00 0.80 0.30 1.10 1.58 1.44 4.85 1.09 8.96 0.88 

7-2-93 4A 0.423 33.00 2.00 0.20 2.20 0.29 0.26 2.97 0.25 3.77 1.00 

7-2-93 4C 0.395 17.00 1.20 0.40 1.60 0.40 0.44 2.71 0.28 3.83 0.69 
49 

Date WB TN TP Cl SO4 Tot-An Ca Mg Na K Tot-Cat Ave-Depth 

7-2-93 4E 0.340 34.00 0.80 0.60 1.40 0.43 0.35 2.12 0.33 3.23 0.79 

7-2-93 32 0.567 37.00 1.20 0.50 1.70 0.92 0.69 4.33 0.39 6.33 0.88 

7-2-93 30 1.550 77.00 1.60 0.80 2.40 0.86 0.71 5.73 0.49 7.79 0.58 

7-2-93 44A 0.587 37.00 1.20 1.20 2.40 0.74 0.54 3.30 0.52 5.10 0.58 

7-2-93 44 0.430 27.00 1.60 0.10 1.70 0.20 0.21 3.21 0.26 3.88 0.75 

7-3-93 270A 0.730 85.00 119.5 3.30 122.8 1.56 3.54 7.22 5.17 17.49 0.75 

7-3-93 1020 0.180 14.00 2.70 0.20 2.90 0.19 0.27 2.72 0.29 3.47 0.81 

7-3-93 85 0.375 41.00 3.30 0.40 3.70 0.68 0.60 2.58 0.26 4.12 0.75 

7-3-93 85A 0.365 21.00 4.00 0.20 4.20 0.38 0.42 2.74 0.60 4.14 0.75 

7-3-93 81 0.350 33.00 1.20 0.70 1.90 1.60 1.30 4.46 0.60 7.96 0.81 

7-3-93 84 0.307 23.00 1.20 0.40 1.60 0.56 0.55 3.42 0.49 5.02 0.56 

7-3-93 8A 0.460 49.00 2.50 1.10 3.60 0.34 0.28 2.76 0.30 3.68 0.50 

7-3-93 8B 0.340 19.00 3.60 0.40 4.00 0.26 0.41 2.96 0.16 3.97 0.75 

7-3-93 8C 0.180 11.00 3.00 0.80 3.80 0.50 

7-3-93 8E 0.297 21.00 3.00 0.80 3.80 0.25 0.40 3.34 0.39 4.38 0.46 

7-3-93 8D 0.263 23.00 0.29 0.24 2.22 0.44 3.19 0.46 

7-5-93 137B 0.350 15.00 0.80 1.20 2.00 0.28 0.22 2.80 1.70 3.47 0.58 

7-5-93 137A 0.467 18.00 1.60 0.10 1.70 1.12 0.86 2.78 0.23 4.99 0.58 

7-5-93 1023 0.283 12.00 1.20 0.80 2.00 1.54 1.22 2.38 0.94 6.08 2.25 

7-5-93 66 0.303 8.00 2.00 0.30 2.30 1.92 1.40 2.88 0.94 7.14 4.50 

7-5-93 145A 0.530 42.00 4.00 0.50 4.50 1.20 1.46 7.84 0.97 11.47 

7-5-93 145B 0.403 16.00 3.90 0.50 4.40 0.24 0.39 2.40 0.29 3.32 

7-5-93 145C 0.723 25.00 5.00 1.70 6.70 0.74 0.84 4052 0.62 6.72 0.25 

7-5-93 145D 0.517 16.00 4.00 1.50 5.50 0.47 0.70 5.01 0.14 6.32 0.96 

7-5-93 145E 0.680 22.00 8.20 0.30 8.50 0.54 0.82 4.91 0.66 6.93 0.10 

Date WB Surface SLL SLD Trans Temp pH Cond Alk Color Chloro Invert 

6-25-92 299 32.30 2453.40 1.22 15.83 7.18 28.23 8.10 20 2.502 

6-28-92 300 4.80 862.20 1.11 9.10 57.90 0.80 0 0.958 

6-25-92 301 2.80 604.80 1.02 14.33 6.90 34.00 10.80 20 2.857 

6-25-92 302 11.10 1446.50 1.22 16.25 7.80 8.97 5.00 25 1.006 

6-25-92 303 4.60 899.10 1.18 15.66 7.38 15.04 3.00 20 1.602 

6-26-92 304 0.97 356.60 1.02 10.96 8.12 1.90 10 0.525 

6-26-92 305 1.80 684.20 1.44 12.06 30.10 8.50 160 5.769 

6-26-92 298 9.30 1479.40 1.37 11.70 7.80 14.16 6.50 5 0.970 

6-28-92 295 1.40 461.30 1.10 12.46 9.79 0.80 0 0.587 

6-28-92 296 3.30 921.30 1.43 11.30 8.88 1.10 0 0.548 

6-28-92 297 3.50 736.00 1.11 0.75 12.36 7.00 48.00 15.20 40 16.458 

6-28-92 290 6.90 980.70 1.05 12.66 7.67 37.40 12.30 5 3.035 

6-28-92 274 0.37 222.20 1.03 13.50 7.90 20.86 50 9.632 

6-28-92 276 6.70 1286.90 1.40 9.80 7.74 18.25 2.50 53 2.568 

6-28-92 277 3.10 711.30 1.14 11.15 8.01 16.74 1.40 5 1.00 

6-29-92 291 17.10 1598.00 1.09 16.73 38.50 6.80 8 2.763 

6-29-92 294 25.30 2267.20 1.27 7.31 18.25 1.90 0 0.649 

6-29-92 280 2.30 551.90 1.03 6.98 24.30 3.80 0 1.414 

6-29-92 281 11.20 1877.60 1.58 7.19 31.36 10.60 25 6.822 

6-29-92 284 6.20 1156.90 1.31 14.33 36.30 4.00 25 1.347 

6-29-92 285 4.70 1023.40 1.33 14.50 10.64 9.50 30 1.413 

6-29-92 288 8.50 1078.90 1.04 14.40 7.53 63.16 17.20 25 4.895 

6-29-92 289 11.90 1345.40 1.10 7.29 61.83 5.20 43 8.462 

6-30-92 149 9.70 1626.50 1.47 15.73 7.83 13.47 4.90 2 1.457 

6-30-92 150 1.20 443.30 1.14 13.26 7.57 83.26 20.00 70 5.068 

6-30-92 153 18.10 1762.40 1.17 13.55 12.87 7.00 5 2.647 

6-30-92 156 7.80 1122.10 1.13 16.36 9.75 0.50 10 0.613 

6-30-92 158 6.70 1084.60 1.18 15.16 20.83 6.60 15 2.037 

6-30-92 160 11.80 1861.20 1.53 2.00 14.63 38.56 17.50 30 4.511 

6-30-92 161 3.10 730.60 1.17 16.46 7.80 11.67 2.30 25 2.539 
50 

Date WB Surface SLL SLD Trans Temp pH Cond Alk Color Chloro Invert 

7-1-92 164 16.90 2070.60 1.42 0.75 15.63 7.27 35.00 18.50 37 12.284 

7-1-92 207 2.30 692.90 1.29 26.16 7.84 12.33 8.50 10 0.686 

7-1-92 208 0.86 341.60 1.04 19.10 6.75 14.61 8.60 10 1.150 

7-1-92 209 3.50 714.10 1.08 20.03 7.00 45.16 10.90 45 1.447 

7-1-92 214 17.40 1717.40 1.16 19.26 6.50 15.19 3.00 65 1.174 

7-1-92 216 6.80 1181.70 1.28 0.50 18.73 7.54 14.80 11.60 80 9.981 

7-1-92 218 6.10 921.40 1.05 23.80 6.83 69.06 26.60 72 8.821 

7-1-92 222 0.83 348.50 1.08 16.80 6.50 20.43 4.70 80 1.954 

7-1-92 223 8.90 1104.60 1.04 13.66 6.75 20.90 9.60 70 3.854 

7-2-92 254 7.30 1371.10 1.43 14.93 17.79 8.00 10 0.699 

7-2-92 255 3.20 810.20 1.28 15.40 7.75 34.36 1.90 40 1.472 

7-2-92 257 10.70 1384.20 1.19 14.80 6.25 23.56 5.10 10 1.041 

7-2-92 258 14.70 1585.50 1.17 1.25 15.36 6.25 14.13 1.80 50 1.756 

7-2-92 259 12.10 1307.60 1.06 15.80 7.00 40.10 12.60 70 5.861 

7-2-92 140 22.20 1836.20 1.10 15.22 6.25 72.70 28.90 100 3.321 

7-2-92 143 4.40 760.50 1.02 14.76 6.25 41.80 125 2.735 

7-2-92 145 6.90 1438.80 1.55 15.50 11.41 2.40 45 0.962 

7-2-92 146 5.90 1063.20 1.23 14.00 27.63 3.60 80 1.118 

7-3-92 8B 1.90 589.40 1.21 17.10 6.25 26.03 7.00 200 2.437 

7-3-92 8C 4.90 2150.00 2.74 17.60 6.50 37.45 15.00 40 5.110 

7-3-92 8D 1.00 382.60 1.08 20.70 5.50 15.10 5.90 75 1.460 

7-3-92 145A 7.40 1060.10 1.10 18.13 5.50 20.90 2.50 42 2.845 

7-3-92 145E 49.00 2978.80 1.20 15.30 7.25 64.85 23.70 50 2.632 

7-3-92 183 22.90 2413.50 1.42 16.25 6.00 11.39 3.50 45 2.013 

7-3-92 136 15.00 1440.10 1.05 13.76 6.00 9.62 6.30 2 0.660 

7-3-92 137 6.20 1132.90 1.28 1.50 14.86 6.50 23.76 7.80 58 1.756 

7-4-92 188 34.90 6030.40 2.89 7.00 17.40 6.50 11.65 3.90 3 0.596 

7-4-92 191 1.60 570.90 1.27 18.53 6.50 16.31 2.30 10 0.512 

7-4-92 198 0.46 264.80 1.10 20.30 6.50 13.26 8.20 5 2.091 

7-4-92 183 8.90 2701.30 2.55 12.63 7.00 42.66 12.20 5 1.588 

7-7-92 4E 58.60 3501.4 1.29 15.93 7.00 28.26 7.20 35 4.237 

7-7-92 22 62.30 3097.60 1.11 15.27 6.25 7.36 1.70 20 2.377 

7-7-92 23 64.40 3058.10 1.07 0.50 15.67 7.00 46.25 9.10 150 9.623 

7-7-92 30 372.30 7997.20 1.17 0.25 15.36 7.00 29.90 17.20 40 7.123 

7-7-92 32 104.40 3903.90 1.08 16.80 7.00 28.26 11.20 10 3.020 

7-7-92 4C 64.60 3792.60 1.33 0.50 13.67 7.25 38.35 13.50 30 12.778 

7-8-92 96A 90.80 3795.5 1.12 16.10 6.00 14.27 5.00 5 0.266 

7-8-92 90A 52.10 2738.40 1.07 13.90 21.20 4.40 15 1.028 

7-8-92 100C 115.50 6045.80 1.59 1.25 15.82 8.00 55.95 18.50 20 8.406 

7-9-92 86 62.40 3043.00 1.09 16.90 6.00 10.43 5.20 20 1.546 

7-9-92 88 194.50 7505.40 1.52 16.33 7.25 25.43 11.20 55 11.805 

7-9-92 94 45.50 2773.10 1.16 0.75 15.63 7.50 60.70 20.80 45 4.818 

7-10-92 15 181.90 5458.90 1.14 15.63 6.25 13.51 5.00 1 0.562 

7-10-92 41 58.50 5510.30 2.03 5.00 15.60 7.00 25.93 4.30 5 732 

7-10-92 61 59.60 3129.70 1.14 14.93 6.25 11.74 9.50 12 3.501 

7-10-92 81 124.20 4529.60 1.15 1.00 16.52 7.50 38.30 18.50 20 4.317 

7-11-92 39 41.30 2531.60 1.11 19.03 6.75 41.10 19.90 10 2.617 

7-11-92 186 18.20 1857.10 1.23 17.70 8.00 35.56 11.20 55 1.627 

7-11-92 183 8.90 2701.30 2.55 14.37 7.00 49.92 15.60 30 0.906 

7-11-92 kvic 159.50 6834.10 1.53 

7-11-92 68 57.60 3076.10 1.14 19.73 6.50 23.30 9.80 40 2.945 

7-13-92 12 97.30 5133.80 1.47 17.23 6.25 38.76 6.60 40 0.632 

7-13-92 14 33.00 3295.30 1.62 1.25 17.13 7.50 49.11 13.10 35 6.275 

7-13-92 54 173.50 5428.20 1.16 20.16 7.00 41.70 9.10 40 1.353 

7-14-92 20 52.90 2689.50 1.04 0.25 16.23 6.50 14.00 2.80 50 3.154 

7-14-92 21 34.50 2498.70 1.20 15.60 6.25 11.63 3.00 20 0.722 

7-14-92 310 76.20 3397.20 1.10 15.70 7.50 47.54 46.50 55 4.563 

7-14-92 312 76.50 3438.20 1.11 16.45 9.00 98.63 44.70 150 12.917 

7-14-92 6 164.90 6187.80 1.36 0.37 15.16 6.50 18.73 8.20 50 2.065 

7-14-92 47 78.00 3348.70 1.07 0.50 15.80 7.00 39.34 13.10 150 3.980 

7-15-92 2 60.20 2967.40 1.08 16.10 6.25 11.52 1.80 1 0.498 

7-15-92 66 67.60 4656.90 1.60 2.50 14.92 7.50 68.12 27.20 5 1.493 
51 

Date WB Surface SLL SLD Trans Temp pH Cond Alk Color Chloro Invert 

7-15-92 67 76.10 3635.30 1.18 0.25 17.32 8.50 82.24 3.90 125 9.188 

7-15-92 70 69.70 5706.50 1.93 18.13 6.80 22.33 12.50 5 1.443 

7-16-92 8 43.80 2439.80 1.04 0.50 18.13 7.25 45.93 15.30 95 9.653 

7-16-92 73 25.40 2076.40 1.16 18.26 7.50 29.10 8.60 125 15.628 

7-16-92 79 42.40 2796.30 1.21 17.43 6.00 8.11 1.90 20 0.642 

7-16-92 84 42.70 3209.70 1.39 16.90 6.40 21.73 9.30 45 0.733 

7-17-92 69 113.80 4327.00 1.14 14.20 6.25 11.39 3.00 5 2.958 

7-17-92 71 73.80 4537.70 1.50 0.50 14.86 6.70 37.53 20.80 35 8.344 

7-17-92 85 69.40 3772.40 1.28 14.26 6.70 25.10 10.00 60 3.608 

6-19-93 299 32.30 2453.40 1.22 21.00 7.93 40.00 2.70 30 16.775 0.226 

6-19-93 301 2.80 604.80 1.02 19.53 7.71 41.20 13.40 10 4.805 0.167 

6-19-93 302 11.10 1446.50 1.22 21.51 7.67 13.41 0.00 11 0.906 

6-19-93 303 4.60 899.10 1.18 19.26 7.49 22.05 5.20 10 0.509 

6-19-93 305 1.80 684.20 1.44 22.40 7.58 28.00 14.10 15 0.990 

6-20-93 296 3.30 921.30 1.43 16.83 8.18 6.55 1.00 5 0.052 

6-20-93 295 1.40 461.30 1.10 8.50 17.07 7.77 7.02 1.50 15 0.530 

6-20-93 298 9.30 1479.40 1.37 20.01 6.99 12.68 2.10 15 0.972 0.004 

6-20-93 297 3.50 736.00 1.11 20.70 6.79 39.10 13.80 30 2.978 

6-20-93 300 4.80 862.20 1.11 20.05 6.42 4.65 0.70 10 .0615 

6-20-93 276 6.70 1286.90 1.40 19.75 6.98 13.57 0.60 5 2.319 0.034 

6-20-93 277 3.10 711.30 1.14 31.60 6.99 12.21 7.40 5 0.646 

6-20-93 274 0.37 222.20 1.03 23.30 6.91 8.45 1.90 5 1.210 

6-20-93 304 0.97 356.60 1.02 19.50 6.94 8.06 0.50 5 0.643 0.007 

6-21-93 294 25.30 2267.20 1.27 17.70 7.04 9.48 1.00 5 0.911 0.009 

6-21-93 290 6.90 980.70 1.05 19.10 6.93 30.60 9.50 5 2.123 0.044 

6-21-93 291 17.10 1598.00 1.09 20.55 6.92 26.73 4.90 10 1.409 

6-21-93 288 8.50 1078.90 1.04 20.70 6.88 45.30 5.00 40 4.398 0.021 

6-22-93 28 46.20 1156.90 1.31 17.67 8.08 23.43 6.00 100 1.950 

6-22-93 150 1.20 443.30 1.14 0.25 17.53 9.45 5.55 24.20 175 63.515 

6-22-93 149 9.70 1626.50 1.47 3.50 18.33 8.45 9.70 0.80 50 1.805 

6-22-93 153 18.10 1762.40 1.17 19.00 7.72 12.33 3.70 10 3.805 

6-22-93 161 3.10 730.60 1.17 13.90 6.98 8.34 0.00 15 2.212 0.005 

6-22-93 164 16.90 2070.60 1.42 15.70 6.92 31.70 17.30 30 20.186 

6-22-93 158 6.70 1084.60 1.18 2.00 17.60 7.03 20.05 0.80 10 2.662 0.006 

6-22-93 156 7.80 1122.10 18.40 6.99 8.39 2.50 10 0.749 

6-23-93 207 2.30 692.90 1.29 5.00 16.93 7.63 10.93 1.00 30 2.038 0.010 

6-23-93 208 0.86 341.60 1.04 17.47 7.26 15.58 3.10 30 1.194 

6-23-93 214 17.40 1717.40 1.16 0.50 17.90 7.02 16.35 3.80 70 4.070 0.191 

6-23-93 209 3.50 714.10 1.08 0.50 19.75 6.97 40.60 11.60 50 1.577 

6-23-93 223 8.90 1104.60 1.04 1.50 13.76 8.67 17.20 8.10 15 9.536 

6-23-93 222 0.84 348.50 1.08 14.90 7.88 20.40 3.80 30 2.398 

6-23-93 216 6.80 1181.70 1.28 1.00 15.75 8.63 22.40 5.20 25 22.727 

6-23-93 218 6.10 921.40 1.05 15.98 7.34 56.80 20.00 25 13.623 

6-24-93 1010 5.10 871.70 1.09 16.80 7.11 5.30 0.00 5 0.386 0.021 

6-24-93 1011 23.30 3940.40 2.30 17.20 7.08 7.95 0.00 5 2.661 0.008 

6-24-93 1012 69.30 12425.40 3.57 7.00 18.80 7.09 9.27 0.90 15 0.621 

6-24-93 1013 6.60 1676.80 1.84 2.75 19.00 7.31 14.64 1.50 20 1.688 0.029 

6-24-93 1014 47.30 5715.50 2034 16.53 7.34 41.27 13.60 5 0.610 

6-24-93 1000 106.40 11471.40 3.14 9.50 16.40 7.10 7.90 0.50 1 0.592 

6-24-93 1001 1.50 474.90 1.09 8.00 17.50 7.00 8.90 0.20 5 0.906 

6-24-93 1002 4.90 1198.30 1.53 11.00 18.50 7.00 9.10 0.00 5 0.587 

6-24-93 1003 3.50 896.40 1.35 5.50 18.90 7.00 11.00 1.80 10 0.929 

6-24-93 1004 9.20 1777.80 1.65 23.60 7.00 15.60 1.90 100 1.723 

6-27-93 259 12.10 1307.60 1.19 12.70 7.02 33.00 11.50 75 7.415 0.121 

6-27-93 257 10.70 1384.20 1.19 13.80 7.00 20.50 4.70 10 1.487 0.040 

6-27-93 258 14.60 1585.50 1.17 11.40 7.00 14.40 1.30 100 8.199 

6-27-93 140 22.20 1836.20 1.10 11.40 6.97 68.30 25.20 125 26.427 0.103 

6-27-93 143 4.40 760.50 1.02 11.30 6.97 39.40 9.30 125 10.718 

6-28-93 105A 480.00 6547.80 1.20 1.25 15.30 7.30 42.40 15.80 50 11.743 

6-28-93 100A 85.10 4945.70 1.51 0.75 14.50 7.30 50.90 19.50 75 6.192 

6-28-93 100B 78.90 2578.30 1.17 13.90 7.30 32.90 10.80 75 10.289 0.075 

6-28-93 96 144.60 4591.60 1.08 0.50 14.80 7.09 36.90 10.40 50 5.320 
52 

Date WB Surface SLL SLD Trans Temp pH Cond Alk Color Chloro Invert 

6-28-93 105A 73.80 3718.90 1.22 15.70 7.12 37.40 13.10 15 3.414 

6-28-93 100 95.60 3606.50 1.04 0.75 14.40 7.11 76.10 22.50 100 10.702 

6-28-93 96A 51.90 2716.00 1.06 0.50 14.60 7.11 43.60 14.00 75 9.536 

6-29-93 100C 115.50 6045.80 1.59 2.00 11.80 7.10 55.40 1.20 50 5.961 2.223 

6-29-93 111B 577.30 8927.20 1.50 2.50 11.90 7.10 28.40 0.60 50 1.267 0.338 

6-29-93 105B 114.50 5377.50 2.03 0.25 11.90 7.06 30.10 5.80 150 13.631 

6-29-93 100C 59.70 2914.20 1.52 14.10 7.00 29.30 9.50 5 2.601 0.016 

6-29-93 1014 97.90 3958.10 1.13 2.00 12.70 7.21 62.60 26.00 50 5.984 

6-29-93 105B 118.40 5532.30 1.43 13.00 7.27 42.90 15.60 35 1.727 

6-29-93 111B 667.90 3397.60 1.16 13.60 7.44 19.40 8.80 50 0.681 

6-30-93 95A 56.30 3237.90 1.23 12.30 7.20 11.20 6.20 2.114 0.028 

6-30-93 88 194.50 7505.40 1.52 12.50 7.18 31.50 13.40 25 7.761 0.122 

6-30-93 12 97.90 5313.60 1.47 13.30 6.80 19.10 6.30 30 0.872 0.020 

6-30-93 15 181.90 5458.90 1.14 13.60 7.20 6.90 0.60 5 1.144 

6-30-93 90A 57.30 2898.30 1.08 13.20 7.05 10.37 1.50 15 1.649 

6-30-93 90 125.70 4844.70 1.22 13.10 7.00 49.60 18.20 75 1.661 

7-1-93 311A 28.70 2034.90 1.07 13.60 6.90 26.90 9.50 125 2.359 0.080 

7-1-93 311B 36.30 2284.10 1.07 14.40 7.14 33.00 11.80 100 6.207 

7-1-93 311C 10.50 1810.40 1.58 3.50 14.60 7.30 11.50 0.80 20 1.711 

7-1-93 4B 69.30 4517.80 1.53 0.50 10.70 7.10 9.40 0.10 75 3.062 

7-1-93 311 54.70 2759.00 1.05 14.00 7.03 56.40 20.90 100 10.847 

7-1-93 4 165.20 5386.80 1.18 12.30 7.01 12.72 1.90 275 13.725 

7-1-93 6 164.90 6187.80 1.36 11.70 7.00 15.99 1.60 225 24.775 

7-1-93 310 76.20 3397.20 1.10 13.20 7.02 46.40 18.80 60 20.328 0.174 

7-2-93 4A 40.50 3279.90 1.45 0.50 10.60 7.48 8.96 1.50 80 4.921 

7-2-93 4C 10.30 1276.00 1.12 11.40 8.20 13.50 4.20 75 3.203 0.044 

7-2-93 4E 49.20 2811.40 1.13 0.50 12.20 7.90 13.40 4.90 75 4.697 

7-2-93 32 104.40 3903.90 1.08 12.10 7.03 27.60 7.20 25 6.635 

7-2-93 30 372.30 7997.20 1.17 0.25 11.90 6.98 24.60 6.30 25 18.630 

7-2-93 44A 64.50 3201.40 1.12 11.90 7.03 24.50 4.20 100 7.599 

7-2-93 44 34.60 2208.80 1.06 12.60 7.04 12.31 1.00 50 4.487 

7-3-93 270A 76.50 2783.50 1.28 0.25 11.50 7.40 467.50 14.00 150 16.551 

7-3-93 1020 142.50 3239.30 1.09 11.20 6.76 12.50 0.20 25 2.037 

7-3-93 85 69.40 3772.40 1.28 11.00 7.00 22.00 4.70 25 5.746 

7-3-93 85A 12.50 2333.50 1.86 11.50 8.34 21.50 3.50 50 2.806 

7-3-93 81 124.20 4529.60 1.15 0.75 11.50 7.17 41.90 15.60 50 4.793 

7-3-93 84 42.70 3209.70 1.39 11.40 7.10 18.70 3.90 100 4.988 

7-3-93 8A 51.30 4285.10 1.69 0.25 10.50 7.03 14.27 6.50 125 7.268 

7-3-93 8B 2.10 550.50 1.07 11.20 7.05 16.71 1.30 100 1.721 0.011 

7-3-93 8C 0.85 338.90 1.04 14.40 7.05 24.07 2.60 1.342 

7-3-93 8E 2.10 525.40 1.02 13.65 7.05 19.14 0.60 1.762 

7-3-93 8D 35.80 2398.50 1.13 0.33 12.27 7.02 15.22 0.10 1.818 

7-5-93 137B 47.60 3658.30 1.50 0.50 11.10 6.90 16.10 3.30 125 5.230 

7-5-93 137A 44.40 4239.40 1.79 11.60 7.00 50.60 16.00 75 3.869 

7-5-93 1023 487.40 6661.90 1.22 0.50 11.30 7.10 59.40 17.00 125 2.604 

7-5-93 66 67.60 4656.90 1.60 14.60 7.30 84.30 24.00 50 2.840 

7-5-93 145A 73.80 2628.70 1.23 11.35 7.03 4.68 11.90 70 8.187 

7-5-93 145B 5.30 670.30 1.17 0.50 13.15 7.10 18.97 3.20 70 1.196 0.128 

7-5-93 145C 67.50 3195.20 1.57 15.43 7.03 38.63 3.00 150 5.073 0.044 

7-5-93 145D 23.70 1449.40 1.34 15.85 7.01 25.57 1.20 100 1.770 

7-5-93 145E 0.32 187.20 1.20 17.25 7.01 40.07 1.80 60 3.813 
53 
54 
55 

Appendix 4. Occurrence of all invertebrate organisms found in 3 wetland 

habitat strata within the Kvichak study area.

 Connected Closed Moraine 

Annelida

 Oligochaeta X X X 

 Hirudinea 

 Glossiphoniidae X X 

Arthropoda

 Insecta 

 Coleoptera 

 Dytiscidae X 

 Gyrinidae X X 

 Diptera 

 Ceratopogonidae X 

 Chironomidae X X X 

 Culicidae X 

 Simuliidae X 

 Ephemeroptera 

 Leptophlebiidae X 

 Hemiptera 

 Corixidae X X 

 Trichoptera 

 Leptoceridae X 

 Limnephilidae X X X 

 Phryganeidae X 

Cnidaria

 Hydrozoa 

 Hydridae X 

Crustacea

 Amphipoda 

 Gammaridae X X 

 Cladocera 

 Chydoridae X X 

 Daphnidae X X 

 Copepoda 

 Cyclopoida X X 

Arachnida

 Hydracarina X X 
56 

Mollusca

 Gastropoda 

 Lymnaeidae X 

 Planorbidae X X X 

 Pelecypoda 

 Sphaeriidae X X X 

Nemetoda X 

Nemetomorpha

 Gordiodea 

 Gordiidae X 
57 

Appendix 5. List of aquatic and upland plant species recorded from 

wetlands surveyed in the Kvichak study area in 1992 and 1993.

 Connected Closed Moraine 

 Wetland Wetland Wetland River 

UPLAND/SHORELINE 

Bluejoint Grass (Calamagrostis canadensis) X X 

Manna Grass (Glyceria borealis) X 

Sedge (Carex aquatilus) X X 

 (Carex rostrata) X X X 

Horsetail (Equisetum spp.) X 

Cotton Grass (Eriophorum augustifolium) X X

Labrador Tea (Ledum palustre) X X

Cassandra (Chamaedaphne calyculata) X

Dwarf Birch (Betula nana) X X

Shrub Birch (Betula glandulosa) X X

Willow (Salix spp.) X X X

Marsh Fivefinger (Potentilla palustris) X

Lingonberry (Vaccinium vitis-idaea) X

Sweet Gale (Myrica gale) X

Alder (Alnus crispa) X X X

Black Spruce (Picea marianna) X X

White Spruce (Picea glauca) X X

Moss (Sphagnum spp.) X X

Fern (Thelypteris phegopteris?) X

EMERGENTS 

Mares Tail (Hippuris vulgaris) X X

Spike Rush (Eleocharis acicularis) X

Water Hemlock (Cicuta mackenzieana) X

Buckbean (Menyanthes trifoliata) X X

SUBMERGENTS 

Pondweed (Potomogeton praelongus) X X

 (Potomogeton natans) X

Water Milfoil (Myriophyllum spicatum) X X 

Bureed (Sparganium augustifolium) X X X

Water Shield (Brasenia schreberi) X X

Pond Lilly (Nuphar polysepalum) X X X

Hornwort (Ceratophyllum demersum) X X