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thesis entitled
LAND-COVER AND COARSE PARTICULATE ORGANIC

INFLUXES TO A SMALL STREAM

presented by

David Craig Mahan

has been accepted towards fulfillment
of the requirements for

Philosophy

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Doctoral

 

 

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LAND-COVER AND COARSE PARTICULATE ORGANIC

INFLUXES TO A SMALL STREAM

By

David Craig Mahan

A DISSERTATION

Submitted to
Michigan State University
in partial fulfillment of the requirements

for the degree of

DOCTOR OF PHILOSOPHY

W. K. Kellogg Biological Station
Department of Fisheries and Wildlife

1980

J /"/o'2 41¢? 5/

ABSTRACT
LAND-COVER AND COARSE PARTICULATE ORGANIC INFLUXES
TO A SMALL STREAM
By

David Craig Mahan

Effects of variation in streamside vegetation on allochthonous
coarse particulate organic (CPOM) influx were determined for Augusta
Creek, a third-order stream in a multiple-land-use catchment in
southwestern Michigan. Three sampling sections were in common woody
riparian associations; another had undergone selective vegetation
removal. CPOM contributions were estimated with direct infall and
lateral transport samplers for two years.

Significant differences in allochthonous CPOM were found, with
litter totals correlated with vegetation and channel width changes.
Litterfall was greatest to a first-order reach in dense forest (647
g/m2) and lowest in the selectively cut section (108 g/mz). As channel
width increased, exogenous CPOM declined on an areal basis, due to
decreased bank to channel area ratio and reduced canopy. Since most
North American watersheds are multiple-land-use, these results suggest
that CPOM influx variations should characterize most headwater streams.

Because of differences in channel width and vegetation,
allochthonous CPOM was unevenly distributed among streamrorders. First—
and second-orders made up 49% of the stream's surface, yet accounted
for over 62% of the litter entering Augusta Creek. These totals
underscore the role of small tributaries in CPOM collection and

indicate that food resources for coarse particle detritivores may be

inversely correlated with stream-order in response to increased
channel width.

Variations in patterns of CPOM influxes may also have a critical
influence on in-stream detritus levels. In both years, CPOM totals
and types were similar within each location. Leaves (70%) and wood
(18%) were the dominant litter types, with more rapidly degraded
fruits and herbs found in small amounts. Most litter entered the
stream in autumn (70%), however, the remainder was quite evenly
distributed among the other seasons, affording a low, but constant
influx at these times. Several features of the CPOM influx pattern,
its annual similarity, seasonal continuity and predominance of leaves
and wood, should lend stability to the detrital resource base in
Augusta Creek.

The mode and distance of CPOM transport to the stream are also
important, since they determine the area of undisturbed vegetation
required for normal litterfall. Most CPOM was transported by direct
infall (78%), with low relief affording little lateral movement.
Concurrent vegetation analyses indicated that most allochthonous
CPOM originated within 10 m of the channel. Nearstream shrubs were
particularly important litter sources, as suggested by correlations of
shrub basal area and stand density with CPOM influxes. Thus, a ZO-m
corridor of natural vegetation should maintain typical litterfall
levels to this stream.

While the catchment is multiple-land-use, the riparian zone is
generally undisturbed. This natural "greenbelt" has probably been a
leading factor in the maintenance of normal headwater stream structure

and function in Augusta Creek.

ACKNOWLEDGEMENTS

This dissertation is dedicated to my wife, Barb, whose love and
support never faltered. In addition, my parents, Paul and Mildred
Mahan, were a continual source of encouragement. Abby and Meegan,
who joined our family during this time period, were understanding when
daddy had to "go do his work".

I wish to thank Dr. Kenneth Cummins, committee chairman, for his
support and encouragement in all aspects of the study, and for the
fishing trips, which taught me much about streams. I would also like
to thank the other committee members, Drs. Michael Klug, Richard
Merritt and Ray White, for their contributions to this study. Their
insistence on high standards made this a more rewarding educational
experience.

I am very grateful to Dr. George Lauff for providing facilities
at the W. K. Kellogg Biological Station and for his continual
encouragement. Technical assistance was provided by others on the
station staff, Dolores Haire, Carolyn Hammerskjold, Marilyn Jacobs,
Charlotte Seeley and Art Weist. Leah Jackson typed the dissertation,
and Anita Johnson and Red Mahan did the illustrations.

Numerous friends and colleagues contributed to the study in
many ways, especially with their ideas. They include: George Spengler,
Robert King, Donna King, Roger Ovink, Milton Ward, Charley Dewberry,
Dan Lawson, and David Hart.

Financial support was provided by the Department of Energy through
Grant No. E(11-1)2002), National Science Foundation Grant Number
GB-36069X1 and Office of Water Resources Research Grant Number
A-OBl-Mich.

ii

TABLE OF CONTENTS

INTRODUCTION ....... . .............. . .......... .... ............... 1

DESCRIPTION OF STUDY AREA ....................................... 6
Drainage BaSin0....0.000000000000000.......OOOOOOOOOOOOOOOO 6

Methods(Temperature and Discharge) ..... ..... . ............. . 10
Sampling Sites ........ ........ .... ......................... 10
METHODS AND MATERIALS ......................................... .. 18
Land-Cover ... ...... . ............... . ....................... 18
Vegetation Analyses ........................................ l9
CPOM Influx Estimates . ..... .......... ...... .. .......... .... 20
RESULTS AND DISCUSSION ......... ................................. 23
Land-Cover ......................... ..... . ................. . 23
Vegetation Analyses ..... ...... .......... . .................. 31
CPOM Influxes at the Sampling Sites .. ...................... 45
CPOM Influxes Among the Sampling Sites ..... . ............... 61
Literature Comparisons of Litterfall Estimates .. ........... 72

syntheSj-S ......OOOOOOOOOOOO......OOOIOOO.......OOOOOOOO.... 77
Allochthonous CPOM Resources .......................... 77

Other Estimates of Detrital Resources ........... ..... . 82
Biotic Responses to Changes in Allochthonous CPOM ..... 83
Allochthonous CPOM and the Riparian Zone .............. 89
SUMMARY AND CONCLUSIONS ............... . ......................... 91
LIST OF REFERENCES ................... ..... ......... ............. 94
APPENDIX ........................................................ 109

iii

LIST OF TABLES
TABLE PAGE

1. Summary of temperature values from Augusta Creek sites,
1976-1977 .....000000.0.000.00.000.000000000000000000000000. 14

2. Summary of discharge values from Augusta Creek sites,
1976-1977 0000-0000000oooooooooooooooooo0000000000000 ooooooo 16

3. Land-use patterns in the Augusta Creek Drainage Basin ...... 24
4. Summary of riparian cover along Augusta Creek .............. 28
5. Summary of vegetation transects at the Smith site .......... 33
6. Summary of vegetation transects at the 43rd Street site .... 34
7. Summary of vegetation transects at the Nagel site .......... 36

8. Summary of vegetation transects at the Kellogg Forest
Site ........ 0.00......0.0000 000000000 00.000.00.000 0000000 0.0 38

9. Dominant trees and shrubs and quadrat summaries at
the sites ................................... ............... 40

10. A comparison of selected stands of woody riparian
vegetation from the Midwestern U.S.A. .. ........ . ........... 44

11. Seasonal and total CPOM influxes at the Augusta Creek
sites by sampler types ............................ ...... ... 46

12. Categories of CPOM and total CPOM influxes at the
Augusta Creek sites .................. ...................... 47

13. Combined seasonal CPOM totals and rates at the Augusta

Creek sites ......................... ........... ..... ..... .. 48
14. Leaf influx at the Smith site .... ....... . .................. 50
15. Leaf influx at the 43rd Street site ............... ......... 54
16. Leaf influx at the Nagel site . ............................. 57
17. Leaf influx at the Nagel Forest site ..... ............... ... 6O

18. Results of Kruskal-Wallis and Dunn's multiple comparison
tests for the combined trap totals ..... .. ........ . ......... 62

iv

 

TABLE PAGE

19. Results of Kruskal—Wallis and Dunn's multiple comparison
test for the combined trap totals (leaves only) ............. 66

20. Summary of leaf collections expressed on the basis of
processing rates .......... . ................. .. .............. 7O

21. Comparison of annual CPOM inputs to streams with
different riparian vegetation ...................... . ........ 73

22. Annual CPOM influxes to Augusta Creek based on

litterfall and land-cover estimates ......................... 78
23. Annual CPOM influxes by categories to Augusta Creek ......... 80
A1. Woody vegetation along Augusta Creek... ................. .... 109

A2. Total CPOM input by category and season at the Smith
site ........................................................ 110

A3. Total CPOM input by category and season at the 43rd
Street site..... .......................... .......... ........ 111

A4. Total CPOM input by category and season at the Nagel
Site 0000000 0 00000000000000000 0 000000000 0 0000000 0000 000000000 112

AS. Total CPOM input by category and season at the Kellogg
ForeSt Site0.0..0 ..... 0000000... 0000000000000000000000000000 113

A6. Average CPOM inputs for the 28—day sampling periods ......... 114

A7. Leaf species collected in the litter traps arrayed by
processing types..... ....................................... 115

A8. Annual totals of combined direct infall and lateral
transport samplers in g/m of bank ........................... 116

A9. Variables used in rank correlations and the correlation
results.....00........... 00000000000000000000000000000000000 117

A10. Total fruit collections at the sites by species and
trap type ....... ... ......................... . ..... . ......... 118

A11. Seasonal and total CPOM inputs at the sampling locations
by channel surface area ..................................... 119

A12. Seasonal production/respiration ratios (P/R) from selected
riffle sections of Augusta Creek, 1974-1975 (King, 1980).... 120

A13. Riffle invertebrate community functional group analysis
at the Augusta Creek sites from the River Continuum
Project, Summer, 1976 (Vannote, Cummins, Minshall, Sedell).. 120

V

 

TABLE PAGE
A14. Particulate organic matter (POM) in Augusta Creek

sediments from the River Continuum Project (Vannote,
Cummins, Minshall, Sedell, 1976)..... ............... . ....... 121

vi

 

LIST OF FIGURES

FIGURE

1.

10.

11.

12.

13.

14.

Augusta Creek Drainage Basin (cf. Figures 2 and 3 for

sampling locations and scal'e)0.0.00.000..0..0....0000 000000

Augusta Creek sampling sites: 1=Smith; 2=43rd Street;
3=Nagel; 4=Kellogg Forest

Representation of litterfall samplers, with direct
infall traps shown at the top and lateral transport
traps shown at the bottom of the figure

Land—use in the Augusta Creek Drainage Basin.. ..... . .......

Average basal area of the woody vegetation (>2.0 cm DBH)
at the sites .......... ....... .......... ... ...... .. .........

Seasonal CPOM influx in g/m of streambank at the
Smith site. .......... ... ............. . ..... .... ............

Seasonal CPOM influx in g/m of streambank at the
43rd Street site... ............. ..... ......................

Seasonal CPOM influx in g/m of streambank at the
Nagel Site............... ...... . ......... .. ................

Seasonal CPOM influx in g/m of streambank at the
Kellogg Forest site......................... ...............

Annual CPOM totals by litter types at the sites ............

Annual CPOM influx, with CPOM types expressed as a

percentage of the total.......... ..... ...... ...............
Annual CPOM totals by sampler type at the sites ............
Seasonal CPOM totals at the sites ....... ..... ..............

Annual leaf influxes by processing categories at
the Sites.000...0 0000000000 0 000000 000 000000 0 00000 0 000000000

vii

PAGE

26

52

63

64

67

71

INTRODUCTION

Headwater streams are the primary interface between aquatic and
terrestrial environments (Sedell, et al., 1973). Their importance is
underlined by the estimate that they account for over 85% of the total
length of North American running waters. (Leopold, et al., 1964). Small
streams are detritus-based ecosystems (Cummins, 1974; Mdnshall, 1978),
and often depend on particulate organic material (POM; >O,5pm) from the
terrestrial environment for a sizable portion of their annual energy
budget (Hynes, 1963; Minshall, 1967; Cummins, 1974). In forested
catchments, POM, most of which is greater than 1 mm in diameter (CPOM),
may account for as much as 50 to 60% of the allochthonous organic influx
to these lotic systems (Fisher and Likens, 1973; Sedel, et al., 1974).

Allochthonous CPOM usually takes the form of leaves, wood, fruit
and herbaceous matter in forested areas (Gosz, et al., 1972). Autumnal-
shed leaves are the dominant litter type in the deciduous forest biome
(Kaushilk, 1969; Liston, 1972; Bell and Sipp, 1975), since over 90% of
the foliage biomass is typically in the tree layer (Ovington, 1965;
Whittaker, 1966; Monk, et al., 1970). While these losses are relatively
insignificant to the terrestrial ecosystem (Fisher and Likens, 1973),
they represent a tremendous resource to the aquatic system.

Detrital use by shredders (coarse particle feeders), collectors
(fine particle feeders), and the microflora (Suberkropp and Klug, 1976)

is the dominant community activity in small, forested streams (Anderson

INTRODUCTION

Headwater streams are the primary interface between aquatic and
terrestrial environments (Sedell, et al., 1973). Their importance is
underlined by the estimate that they account for over 85% of the total
length of North American running waters. (Leopold, et al., 1964). Small
streams are detritus-based ecosystems (Cummins, 1974; Minshall, 1978),
and often depend on particulate organic material (POM; >0,5um) from the
terrestrial environment for a sizable portion of their annual energy
budget (Hynes, 1963; Minshall, 1967; Cummins, 1974). In forested
catchments, POM, most of which is greater than 1 mm in diameter (CPOM),
may account for as much as 50 to 60% of the allochthonous organic influx
to these lotic systems (Fisher and Likens, 1973; Sedel, et al., 1974).

Allochthonous CPOM usually takes the form of leaves, wood, fruit
and herbaceous matter in forested areas (Gosz, et al., 1972). Autumnal-
shed leaves are the dominant litter type in the deciduous forest biome
(Kaushilk, 1969; Liston, 1972; Bell and Sipp, 1975), since over 90% of
the foliage biomass is typically in the tree layer (Ovington, 1965;
Whittaker, 1966; Monk, et al., 1970). While these losses are relatively
insignificant to the terrestrial ecosystem (Fisher and Likens, 1973),
they represent a tremendous resource to the aquatic system.

Detrital use by shredders (coarse particle feeders), collectors
(fine particle feeders), and the microflora (Suberkropp and Klug, 1976)

is the dominant community activity in small, forested streams (Anderson

and Cummins, 1979). Shredder life cycles appear to be timed to coincide
with normal allochthonous CPOM inputs. Their growth is greatest from
late fall to early spring, which corresponds with the maximum
availability of deciduous leaf litter (Mackay and Kalff, 1973; Cummins
and Klug, 1979). In turn, fine particulate organic material (FPOM)
generated by shredders, and microbial and physical processes, may be
utilized by collectors (Cummins, 1973). Shredders and collectors may
serve as food for community members who function as predators.

The rate of detrital degradation by the biotic community is
correlated with several characteristics of the detritus, including
nutrient levels (particularly nitrogen - Hynes, 1975), structural
components, rate of microbial colonization, and water temperature
(Kaushik and Hynes, 1971; Barlocher and Kendrick, 1973; Anderson and
Sedell, 1979). Due to seasonal, local and stream-order differences in
detrital inputs, production and storage, detritivores in running waters
are subject to spatially and temporally variable food resources (Cummins
and Klug, 1979). These variations cause important changes in food
quality (King, 1978; Ward and Cummins, 1978), which may affect growth
rates, duration of life cycles, fecundity and survivorship (Anderson and
Cummins, 1979).

The biota of small streams is well suited to make use of external
organic influxes which contain a high percentage of CPOM (Cummins, 1974).
In fact, up to 80% of this CPOM may be retained long enough to be used
by aquatic macro- and microorganisms (Anderson and Sedell, 1979).
Because the organisms of headwaters are adapted to these resource
conditions (Ross, 1963), allochthonous CPOM differences due to

nearstream vegetation changes should be reflected by variations in the

species complex of the aquatic community (Vannote and Sweeney, 1980).

The area of the drainage basin which most influences running waters
is the riparian zone (Meehan, et al., 1977). The riparian forest
maintains normal structure and function of the stream ecosystem,
especially due to shading and organic inputs which it provides (Fisher
and Likens, 1973). Large wood debris physically stabilizes the channel
and, by its retention of POM, increases POM availability to detritivores
(Swanson, et al., 1976; Marzolf, 1978). Habitat and cover for aquatic
organisms is another function of wood debris (Hynes, 1970; Marzolf,
1978). Riparian vegetation also reduces sediment and nutrient transport
to streams (Mannering and Johnson, 1974; Sommers, et al., 1975; Ohlander,
1976) and is important in temperature control (Swiff and Messer, 1971).

When disruption of this streamside vegetation occurs, significant
physical, chemical and biological changes in the aquatic environment may
result (Karr and Schlosser, 1978). Clearcutting can greatly increase
nutrient and temperature ranges in the water (Brown, 1970; Sedell, et al.,
1973; Likens, et al., 1977). Serious silt buildup on stream beds, with
concommitant reductions in fish and invertebrate populations, has also
resulted from poor logging practices (Tebo, 1955; Chapman, 1962). In
heavily agriculturalized areas, running water communities have
experienced pronounced faunal changes (Thompson and Hunt, 1930;
Trautman, 1939; Larimore and Smith, 1963). Urbanization has also
affected many drainage basins, and the hydrologic problems which
followed have been summarized by Coughlin and Hammer (1973).

Of particular interest to this study are the effects which changes
in land-cover within a catchment, especially in the riparian zone, can

have on allochthonous CPOM influxes to streams. Differences in detritus

 

within small watercourses have resulted from changes in nearstream
woody species and/or their abundance (Minshall, 1968; Liston, 1972).
When deciduous woodlands were converted to conifers, differences in
food quality of external CPOM occurred (Woodall and Wallace, 1972;
Webster and Patten, 1979). More subtle changes in allochthonous
influxes may result from natural vegetation variations along a stream
due to altitudinal or other environmental differences (Meehan, et al.,
1977). Introduction of plant diseases or alien species may also cause
changes in organic influx patterns. For example, all the effects of the
introduction and management of saltcedar (Tamarix) on the watercourses
of the American Southwest will probably never be known (Robinson, 1965;
Graf, 1978).

Previous measurements of CPOM influxes to running waters were in
single—land-use drainage basins (Fisher and Likens, 1973; Sedell, et al.,
1974; McDowell and Fisher, 1976; Webster and Patten, 1979), or at
selected locations of one type of riparian vegetation (Otto, 1975;
Dawson, 1976; Winterbourn, 1976). However, a majority of contemporary
North American catchments encompass a number of land-use and vegetation
types. Therefore, these results are of limited use in identifying
exogenous CPOM contributions to multiple-land-use drainage basins.

This study was an attempt to more completely describe CPOM influxes.
The objectives were: 1) to quantify distributional variation in woody
streamside vegetation and its influence on the influx of allochthonous
CPOM to a stream; 2) to identify the area of maximal CPOM contribution
within the riparian zone; 3) to evaluate CPOM influxes in relation to
other ecological measurements (detrital standing crOp, P/R ratio,

invertebrate standing crop, leaf processing rates). Because Augusta

 

Creek is a multiple-land-use drainage basin, these results should more
accurately identify allochthonous CPOM contributions which are

characteristic of most stream systems.

DESCRIPTION OF STUDY AREA

Drainage Basin

 

Augusta Creek is a small stream located in Barry and Kalamazoo
Counties, Michigan (Figure l). The total length of the watercourse,
including its tributaries, is 63.3 Km, encompassed by a catchment of
72.3 sz. At its mouth, it is a third-order (Strahler, 1957) stream
with an average discharge of 1.20 m3/s (42.2 cfs) (U.S.G.S., 1978).

The stream arises at an elevation of approximately 286.7 m and falls
to 241.0 m at its terminus, with an average gradient of 2.0 m/Km (0.2%)

Most of the drainage basin is glacial outwash, with some moraines
occurring near Augusta and in the Gilkey Lakes - Fair Lake area of
Barry County (Schmalz, 1978). The land is moderate to slightly
rolling with slopes usually less than 12%, except in morainic areas.
Upland soils are generally well drained, with moderately permeable
upper layers overlying permeable sand and gravel to loamy sand
subsoils. Areas of organic soils (Adrian and Houghton muck) occupy
some extinct glacial lake beds and lowlands along the stream (Konwinski,
1978).

Chemically, one of the most characteristic features of the stream
is the high total hardness, averaging nearly 280 mg/l. This is a result
of the calcareous sand and gravel which underlies the upper soils.

With exception of the rather high levels of inorganic nitrate nitrogen
(2-5 mg/l), which enters via the groundwater, other chemicals are
present in amounts indicative of few disturbances along the stream

(e.g. P as PO

4 of 10-40 ug/l). In a random survey of 10 similar sized

tributaries of the Kalamazoo River, Augusta Creek was rated second

Figure 1. Augusta Creek Drainage Basin (cf. Figures 2 and 3 for
sampling locations and scale).

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in terms of overall water quality (U.S.D.A., 1977).

Prior to 1840 most of the basin was in forest, predominantly the
oak-hickory association. Along the stream the southern lowland forest
as described by Curtis (1959) was common (Kenoyer, 1930). Most of the
uplands were cleared for agricultural purposes in the mid to late 1800's
and have remained under this use. The lowland forests, while also cut
for lumber in the same period, have regrown in many areas into some
resemblance of the stands which once existed (Thompson, 1972). One
notable difference in the floodplain forests is the lack of large elms
(Ulmus spp.), which were eliminated in the 1950's and 60's by the Dutch
elm disease (Thompson, 1972). A characteristic feature along most of
the stream's length is a dense layer of streamside shrubs. The woody
riparian communities are important in maintaining the relatively natural
condition of the stream.

Today the catchment is a multiple—land-use area, about half covered
with lakes and natural vegetation such as forests, shrub-lands and
marshes, with most of the remaining land in agriculture. Much of the
more natural landscape lies adjacent to the stream, serving to buffer
the creek from human activities.

The climate of southwestern lower Michigan is primarily continental,
although it is somewhat modified by the proximity of Lake Michigan.
Since the area is latitudinally at 42° North, strong seasonal
temperature contrasts prevail (Eichenlaub 1978). The mean annual
temperature of the study area, based on data from 1940-1969, is 9.1°C with
a frost free growing season of 151 days. The coldest month of the year
is January with a mean of -4.7°C whereas July, the warmest month,

averages 22.2°C (Strommen, 1971). Mean annual precipitation is 87.1 cm

 

10

and is rather evenly distributed throughout the year. June is the
wettest month, with 10.5 cm of rain, while February, with a 3.8 cm
average, is the driest month. Winds prevail from the southwest and
average 16.1 kpm (Strommen, 1971). Because of frequent low pressure
systems during the late winter and early spring, this period is often
marked by storm activity, much cloudiness and occasional strong winds

(Eichenlaub, 1978).

Methods

Stream temperature was monitored weekly at the four sites for two
years with Bristol continuous temperature recording instruments (Model
636, Bristol Corporation, Waterburry, CN.). From the recording charts,
mean daily, maximum and minimum weekly temperatures were determined.
Seasonal degree days were estimated from weekly averages.

Discharge was monitored every Wednesday at the sites for the same
time period. Mean velocities were estimated with a Beauvert Midget
Current Meter (Nerpic Corporation, Grenoble, France), and cross-sectional
areas were determined. Discharge calculations followed the method
described by Hynes (1970) and were summarized as seasonal means and

extremes .

Sampling78ites

 

Four locations were chosen along Augusta Creek which represented
typical woody riparian plant communities (Figure 2). Each sampling area
was named after the landowner (Smith, Nagel) or a nearby landmark (43rd
Street, Kellogg Forest). Complete vegetative descriptions will follow
in a later section.

The Smith site is typical of most first-order tributaries in the

11

Figure 2. Augusta Creek sampling‘sites: 1 = Smith; 2 = 43rd Street;
3 = Nagel; 4 = Kellogg Forest.

12

Figure 2

 

13

drainage basin. Streamside vegetation is dense, lowland forest, which
provides a continuous canOpy over the 1.0-2.0 m wide stream. This
shade, plus relatively large groundwater contributions, serve to keep
the water cooler than the other sites in warm periods and relatively
warmer in the winter (Table 1). Although discharge is quite stable,
variability was high because its small volume (i = .014 m3/s,.494 cfs)
was readily changed by rainstorms and snow melt (Table 2). The upper
banks lepe at 6% from a narrow floodplain of 4-20 m in width. The
floodplain soil is Adrian muck, which is replaced by Kalamazoo sandy
loam on the adjacent uplands.

At 43rd Street, a representative shrub-carr vegetation association
is found. Highest summer water temperatures were found at this location
(Table 1), probably due to its proximity to several lake outflows, and
a reduced vegetative canopy upstream from the site. The average
elevational gradient was 7%, ‘with one bank being moderately steep
(10%). The 15-50 m wide floodplain is Adrian muck, with uplands of
Sunfield sandy loam. Augusta Creek is a second-order stream at this
location, averaging about 7 m in width and 0.668 m3/s (23.60 cfs) in
discharge (Table 2).

Vegetation at the Nagel site is predominantly shrub-carr, with
some lowland forest. It was selected as an example of a disturbed
location, since on one side of the channel, only a 5-10-m corridor of
natural vegetation has been retained. A lawn, with scattered trees,
lies beyond this point. Annual degree days are higher here than the
downstream Kellogg Forest site, probably due to a reduction in the
canOpy (Table 1). Upper bank slope is 2.3%, with a relatively wide

(35-170 m) area of Adrian muck along the stream. The creek is

14

 

 

 

 

 

 

 

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15

 

 

 

 

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000mm3 x 000mm x
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16

 

 

 

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17

 

 

 

 

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18

third-order at this location, and averages 7 m in width and 0.873 m3/s
(30.83 cfs) in discharge (Table 2).

The Kellogg Forest area illustrates a mature, lowland forest.
Because of increased canopy and greater groundwater contributions from
nearby porous soils (U.S.D.A., 1979), water temperature is cooler in
the summer and warmer in the winter than the upstream Nagel site (Table
1). Bank slope is low (1-2Z), with the stream flowing through a
90-115 m wide floodplain of Adrian muck. This third-order channel is
6-10 m wide and has a mean discharge of 1.010 m3/s (35.65 cfs) (Table
2).

Continuous discharge records have been collected just below the
Kellogg Forest since 1964 by the U. S. Geological Survey. Recurrence
intervals for maximum and minimum discharge during the study period
were estimated from these data. Highest discharge occurred in 1976
with a five-year flood, while 1977 was a time of drought with a 14-year

low.

METHODS AND MATERIALS

Land-Cover

 

Information on land-cover (use) patterns within the Augusta Creek
drainage basin was obtained from three sources: aerial photographs;
U.S.G.S. 7.5 Minute (1:15,000) Series Topographic Maps; direct
observation on the land surface (ground truthing). In order to get a
thorough knowledge of the riparian environment I walked along most of
the mainstream and the accessible tributaries.

The aerial photography and general information on land-cover was

supplied by the Remote Sensing Project of Michigan State University.

19

This group was contracted to delineate land-cover in the Kalamazoo
River Basin (e.g. Richason and Enslin, 1973) as part of a land and
water resource planning project by the Soil Conservation Service
(U.S.D.A., 1977). Color infrared imagery was obtained for the Augusta
Creek basin and predominant land-cover was identified within all six
acre sections by using a grid overlay. Coverage types followed the
classification scheme used by the Remote Sensing Project.

In addition to the land-use analysis, land-cover was more closely
identified within a 100 m wide zone along the stream. The categories
were chosen to be compatible with vegetation and organic input analyses

which followed.

Vegetation Analyses

Riparian woody vegetation was inventoried by the transect method
discussed by Cain and Castro (1959). At each end of the 30-m section
of channel where terrestrial particulate organic influxes were
monitored, two 10 x 30-m belt transects running perpendicular to the
watercourse were established on the two opposing 310pes. Gradient of
these slopes was determined with a Brunton pocket transit. Within
each 300-m2 transect, three IOO-m2 quadrats were outlined with stakes,
representing O-lO-m, lO-ZO-m and 20-30-m distances from the channel.
Therefore, at each sampling location twelve 100-m2 areas were censused,
with four of these quadrats combined for each IO-m distance from the
stream. The 100-1112 samples represent plot sizes normally selected for
woody vegetation analyses (Costing, 1956; Cain and Castro, 1959).

Within the quadrats all woody vegetation greater than 2 cm in

diameter at breast height (DBH) was identified, measured to the nearest

20

0.5 cm and enumerated. With these data, relative density, relative
dominance and relative frequency for each species were calculated, and
the results combined to determine importance values, following the
method described by Cox (1967). Taxonomic nomenclature followed

Gleason and Cronquist (1963), with plants classified as tree or shrub

based on the criteria of Harlow (1957) and Braun (1961).

CPOM Influx Estimates

 

The movement of organic material greater than 1 mm in diameter
(CPOM) into the stream was estimated at the four sampling locations,
within 30-m sections of creek channel. The channel area in m2 at the
sites was: Smith - 53.7; 43rd Street - 213.3; Nagel - 209.1; Kellogg
Forest - 250.2. In the designated 30-m section, the sampling apparatus
was randomly placed within two distinct zones, depending on sampler
type. Two kinds of samplers were used in order to measure CPOM
falling directly into the stream and material transported laterally down
the bank (Figure 3). The direct litterfall (infall) traps consisted of
1.0 m2 wooden frames mounted on steel rods and lined with 1 mm mesh
nylon bags. They were 0.4 m deep and suspended just above the water
level at bankfull discharge. The infall traps were positioned at the
stream edge and projected out over the water surface. The lateral
transport (lateral) samplers were 0.25 m wide and 0.2 m high. They
were three-sided boxes, with corregated steel bottoms, and had wooden
covers to eliminate direct fall-in. They were each located on the bank
adjacent to an infall trap, just above the bankfull discharge level,
with their open ends positioned upslope. Such trap placement assured

the collection of CPOM destined to enter the stream along its margin.

 

 

22

Six sampling units of each type were placed at each site (in 1977 at
Smith, n=12), which allowed for six independent litterfall estimates
per meter of bank.

Litter collections were made every 28 days, from November 5, 1975
through November 1, 1977. Samples were oven-dried at 50° C, sorted
and then weighed to the nearest 0.01 g on a top-loading balance
(Mettler P163). Particles were sorted into four categories: leaves;
wood (branches and twigs); fruits; herbs (herbaceous plant parts).
Leaves and fruits were identified to species (or genera), in order to
facilitate comparisons with the vegetation transects. After inspection
of the data, the 28-day amounts were combined into four seasonal totals
in a manner similar to that of Grigal and Grizzard (1975). Based on
input trends, the periods chosen were: winter - November through
February (112 days); spring - March to late May (84 days); summer -
late May to mid August (84 days); autumn - mid August through October
(84 days).

The small number of replications confounded determination of the
frequency distribution and statistical analysis of the samples. In
addition, variability was inherently large due to the seasonal
nature of the litter movement, and the uneven pattern of
distribution of the vegetation (Gosz, et al., 1972; Post and delaCruz,
1977). When the raw data were examined, variances were usually larger
than the mean and proportional to it, indicating a negative binomial
distribution (Elliot, 1971). Following logarithmic transformation,
however, variances were still often not homogeneous. Therefore,
appropriate nonparametric statistics, as described by Gibbons (1976),

were used. These tests are particularly suitable for small sample

23

sizes with contagious distributions, and may be almost as efficient as
their parametric equivalents (Sokal and Rohlf, 1969). Initially, the
Kruskal - Wallis test, a nonparametric alternative to the one way
analysis of variance (Gibbons, 1976). was used. If significant
differences in the ranking of totals were detected, the multiple
comparison test of Dunn (1964) was utilized in order to detect sample
differences. Where correlation analysis between variables was desired,
Kendall's coefficient of rank correlation was determined (Sokal and

Rohlf, 1969).

RESULTS AND DISCUSSION
Land-Cover

The general pattern of land-cover in the catchment is summarized
in Table 3 and shown in Figure 4. Almost half of the area is in a
more natural state, being covered with lakes, wetlands or native
terrestrial plant communities. Wetlands are predominantly of the type
classified as shrub-swamp (Jeglum, et al., 1974) or shrub-carr (Curtis,
1959), with some herbaceous areas (true marsh - Jeglum, et al., 1974).
The major associations in the forested areas were previously noted,
except for conifer plantings which are found in scattered upland areas.
The native stands are all second or third growth, with no virgin
remnants.

Most of the remaining land is under agricultural use, underscoring
the rural nature of the area. Because of porous soils and moderate
slope, there is little prime agricultural land, and pasturing is common

(U.S.D.A., 1977). Nevertheless, overgrazing, streambank erosion and

Table 3. land use patterns in the Augusta Creek Drainage Basin.

 

 

 

Use Categories Acres Hectare (Ha) [— 7.
Water (Laks ard Streams) 504.7 204.2 2.8
Agriculture 44.7
Cultivated crcplarrl
(row creps, grairs) 3495.6 1414.6 19.6
"Pemanert" pasture, hay, crop
rotation fields (inactive agriculture) 4317.0 1747.0 24.2
Fruits lCB.3 43.8
Tree 39.4 15.9 0.2
Bush arrl vineyards 68.9 27.9 0.4
Feel lots 49.2 19.9 0.3
Rsidential 314.9 127.4 1.8
Transportation, Camunication,
Utilitie 297.0 120.2 1.6
Ccmnercial (services ani iIstimtional) 78.7 31.8 0.4
Otter urban-outcbor recreation,
cemetery 131.2 53.1 0.7
Industrial 19.7 8.0 0.1
Extractive (gravel pits) 19.7 8.0 0.1
Woodlarrl 8.3
Broadleaf forest 2219.2 898.1 12.4
Coniferous 816.7 330.5 4.6
Mixed 464.2 188.0 2.6
Wang) forest (forested wetlards) 1730.8 711.0 9.7
Brush (less than 25% tree) 1053.0 4%.1 5.9
Nonforested Wetlandsa 12.6
emergent vegetation 297.1 120.2 1.7
vegetated open water 191.6 77.5 1.1
shrub-cart 1756.8 711.0 9.8
Totals 17 ,865.4 7,23 .9 100
(2.9012) (72.2102)

 

aIn Figure 4, nonforested wetlands are called narshlarrls.

25

Figure 4. Land-use in the Augusta Creek Drainage Basin.

AREA: 2732 «W 71.54 lun’

       

 

, ....n _ .
. .. ... » 1. .. . .
. I . ., H ...; . .... . a y ./ . I
s . . . 0
1. 0 . . ..I. a _ . .-
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.1 . . .... . /// u. .. /
. _ . . n . I . I .. w. v.
~ .1
203.... I

_ ....m.....w.., .....é I”
. . /.

26

KAI. All)” C0

Figure 4

27

livestock wastes do not presently create significant water quality
problems along the creek. Areas classified as brush are principally
inactive agricultural lands which are being invaded by woody vegetation.

Residential, commercial and industrial uses occupy only an
estimated 2.5% of the total land area. However, within a 48 Km radius
of the town of Augusta, there are over 250,000 people in the Kalamazoo-
Battle Creek metropolitan area (Heller, 1978). If current trends
continue (Ross Twp. Planning Com., 1974; Heller, 1978), the impact of
these people on the drainage basin in terms of housing, recreation and
other uses will increase.

The pattern of land-cover (uses) in the basin is significant
(Figure 4). Nearly all of the non-forested wetlands (marshlands in
Figure 4), the lowland forests, and roughly one-third of the upland
forest stands lie adjacent to the stream. Almost two-thirds of the
more natural landscape is closely associated with the creek, which
should be a critical factor in maintaining high water quality.

In respect to land-cover, the Augusta Creek basin appears to be
fairly typical of most rural basins of similar size in southwestern
Michigan (U.S.D.A., 1977). However, profound changes in land use are
taking place in many areas of this region (U.S.D.A., 1977; Heller, 1978)
and Augusta Creek may one day be affected.

As noted earlier, riparian cover within a 100-m corridor along
the stream was determined (Table 4). The general categories are
descriptive of vegetation types, exclusive of lakes and human impacted
areas, along the watercourse. A 100-m zone was chosen, because it
usually included the entire floodplain and some adjoining uplands. By

the criteria of Trimble and Sartz (1957), this width was more than

28

Table 4. Summary of riparian cover along Augusta Creek.

 

 

Cover Type Channel Length (Km) % of Total
Marsh 3.1 4.9
Shrub-Carrac 15.1 23.9
Shrub-Carrbc 6.4 10.1
Lowland Foresta 22.5 35.6
Lowland Forestb 6.9 10.9
Lake 8.1 12.8
Human Impacted 1.1 1.8
Totalsd 63.2 100.0

 

 

8First- and second-order channels.
Third-order channels.
CIncludes some wet-meadow where shrub cover is less than 25%.

dAlmost 77% of the length of the stream was first- and second-order.

29

double the vegetative corridor required for effective treatment of
surface runoff when upper bank slopes do not exceed 12%.

The most common streamside plant community was lowland forest
(Table 4). This ash-basswood-elm (Fraxinus - T}lia_- Ulmug) dominated
forest resembles the southern lowland forest described by Curtis (1959).
A distinction was made between lowland forest (and shrub-carr) along
first- and second-order channels and third-order reaches, in order to
estimate their relative abundance in conjunction with allochthonous
CPOM measurements. Due to their width, the vegetative canopy was
closed over first- and second-order tributaries. Along third-order
sections, occasional gaps in vegetative cover were found. Considering
that first- and second-order tributaries account for about 77% of the
total length of stream (Table 4), the distribution of lowland forest
along these smaller channels was almost the same (46%) as that along
third-order reaches (47%).

While lowland forests were widespread on river and stream valleys
and old glacial lake plains (Stearns and Kobriger, 1975), the extent
of their present coverage (47%, Table 4) as compared to presettlement
times is not known. The lack of remaining uncut (i.e. virgin) stands
probably indicates a reduced contemporary coverage of this vegetation
type. Relatively large areas (70%) of the original acreage is still
occupied by these forests in Wisconsin, because of frequent flooding
and poor drainage (Curtis, 1959).

Shrub-carr was the next most abundant nearstream plant association
(Table 4). This type of cover consists of a moderate, to thick growth

of one to four meter tall shrubs (Curtis, 1959). The stems of these

shrubs are usually clumped, so that a vigorous growth of sedges,

30

grasses, forbs, and low shrubs covers the ground (Stearns and
Kobriger, 1975). In some areas, shrub cover may have been sparse
enough (<25%) so that a wet-meadow association (Curtis, 1959) existed.
The shrub-carr is a common vegetation type along streams of this
geographical region (White, 1965), and may be more commonplace today
than at time of settlement (Stearns and Kobriger, 1975).

Marshes are not very common plant communities along Augusta Creek.
Distinctive marsh flora such as cattails (Eypha spp.) and arrowheads

(Sagittaria spp.) are frequent in a narrow zone at the channel margin,

 

but extensive stands of marsh species are not widespread. Most of the
estimated riparian marsh occurs in the extensive wetlands between 43rd
and 45th Streets which is an extinct glacial lake bed (Figures 1 and 4).

The Augusta Creek catchment contains one of the largest
concentrations of lakes in the Kalamazoo River Basin (U.S.D.A., 1977)
(Figure 1). Lakes are a prominent feature in the upper part of the
drainage basin, where they serve as the primary source of the two
first-order tributaries which form the main branch of the stream.
Several low-head dams along the creek have created small flooded areas.
They are not so deep as to be considered lakes, but rather fall under
the vegetative types mentioned above.

The final riparian cover recognized was that land which showed
obvious signs of human alteration. This includes bridges and culverts,
livestock crossings, channelized sections and residentially influenced
areas. Although the entire basin was previously affected by cultural
activities, most of it now shows a more natural aspect and was not

included in this class (Table 4).

31

While the Augusta Creek catchment encompassed a number of different
land-cover types, a natural assemblage of nearstream vegetation was
characteristic along the stream. Especially common were plant
communities dominated by lowland species of trees and shrubs. This
undisturbed riparian cover should aid in maintaining and protecting the

associated aquatic environment (Karr and Schlosser, 1978).

Vegetation Analyses

 

Woody vegetation was described in order to assist in the
analysis of CPOM influxes. Approximately 50 species of trees and
shrubs were observed at the sites along Augusta Creek (Table A1).
Those types found in the vegetation transects are shown in Tables 5-8,
with species listed in descending order of dominance (basal area).
Importance values are useful for the descriptive analysis of plant
communities (Curtis and McIntosh, 1957; Curtis, 1959), however, in
situations where some species differ significantly in size from others,
trends in importance value may diverge from density and basal area
(Reiners, 1972). In addition, basal area is a fundamental measure of
forest structure, roughly paralleling biomass and production (Reiners,
1972) and showing a higher degree of correlation with litterfall
(Crosby, 1961; Bray and Gorham, 1964). The transects were oriented
perpendicular to the stream channel in order to parallel the major
axis of environmental gradients (cf. Bell, 1974; Killingbeck and Wali,
1978). This placement should.have given a more complete analysis of
stand composition (Bormann, 1953) and, in combination with litterfall
measurements, allowed some rough estimate of overland transport of

allochthonous particulate organics.

32

Except for the shrub corridor and black walnut (Juglans nigra) and

 

American elm (Ulmus americana) at the stream margin, the vegetation is

 

largely upland in nature at the Smith site (Table 5). The dominant

trees are mesic forest species such as sugar maple (Acer saccharum),

 

black cherry (Prunus serotina) and shagbark hickory (Carya ovata)

 

 

(Braun, 1961) (Table 9). As indicated by basal coverage and importance
values, the leading species from 10-30 m is sugar maple. Stand basal
area increased along the t0pographic gradient back from the stream,
while density declined due to the presence of fewer, but larger,
individual stems. Based on overstory coverage and composition (Curtis,
1959; Bell and Del Mbral, 1977), soil type, and the extent of flooding
during extreme discharges, the forest community beyond 10 m from the
channel is a young, mesic upland association.

0f the 22 woody species at Smith, 15 were shrubs and seven were
trees (Tables 5 and A1). Shrubs formed an important component in all

samples, but were greatest in coverage near the stream. Gray dogwood

(Cornus racemosa) was the leading woody species in all descriptive

 

categories in the first 10 m, creating an almost inpenetrable thicket.
A comparable predominance of dogwoods was noted by Liston (1972) along
a small Kentucky stream. Some other dominant shrubs at the Smith site

were musclewood (Carpinus caroliniana) on the floodplain, and staghorn

 

sumac (RhUS typhina) on well-drained soils.

At 43rd Street, shrubs accounted for almost 30% of the total basal
area, and when all vegetation discriptors are included (i.e. importance
value), they were found to be the leading woody components at the site

(Table 6). Hawthorn (Crataegus spp.), black willow (Salix nigra) and

 

red-osier dogwood (C; stolonifera), all common along southern Michigan

 

33

Table 5. SunnagV of vegetation transects at the Snith site.

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Den. Rel. Dom. Rel. Rel. Imp.
Species #/ 1031112 Den. @0317? Dom. Freq. Freq. Value

0-100
Cornus racamsa 820.0 66.9 @855 26.7 4 10.5 104.1
Prunus serotina 17.5 1.4 3284.42 21.4 2 5.3 28.1
Jug1ans nigra 15.0 1.2 2172.60 14.2 3 7.9 23.3
Carpinus caroliniana 60.0 4.9 2050.00 13.4 2 5.3 23.6
Ulnus americana 52.5 4.3 1789.50 11.7 4 10.5 26.5
Populus tramloides 5.0 0.4 465.82 3.0 2 5.3 8.7
Corylus arericana 65.0 5.3 319.15 2.1 2 5.3 12.7
Salix nigra 57.5 4.7 82.32 1.8 2 5.3 11.8
Sarbucus canadensis 37.5 3.1 184.12 1.2 3 7.9 12.2
Viburnum lentago 15.0 1.2 143.75 1.0 2 5.3 7.5
Crataegus spp. 15.0 1.2 121.25 0.8 2 5.3 7.3
Acer saccharun 7.5 0.6 96.48 0.6 3 7.9 9.1
Rosa palustris 17.5 1.4 85.92 0.6 2 5.3 7.3
Rhus gyphina 17.5 1.4 85.92 0.6 1 2.6 4.6
Ionicera xylosteun 7.5 0.6 73.15 0.5 1 2.6 3.7
bees anericanun 10.0 0.8 49.10 ‘ 0.3 2 5.3 6.4
Vitfs aestivalfs 5.0 0.4 24.55 0.2 1 2.6 3.2
TOTALS 1225.0 99.8 f 15,316.60 100.1 38 100.2 300.1

10-2011
Acer saccharun 160.0 21.7 5498.08 27.8 4 14.3 63.8
EFF serotina 22.5 3.1 5478.65 27.7 3 10.7 41.5
Carya ovata 7.5 1.0 2374.38 12.0 1 3.6 16.6
mus typhma 72.5 9.8 1427.53 . 7.2 1 3.6 20.6
Carpinus caroliniana 32.5 4.4 1416.90 7 2 1 3.6 15.2
Cornus racamsa 227.5 30.8 1292.75 6 5 4 14.3 51.6
Ulnus anerfcana 37.5 5.1 1164.68 5.9 4 14.3 25.3
Ilex verticillata 50.0 6.8 245.50 1.2 1 3.6 11.6
Satbucus canadensis 37.5 5.1 184.13 0 9 1 3.6 9.6
Corylus arericana 35.0 4.7 171.85 0 9 2 7.1 12.7
Popqus tramloides 15.0 2.0 133.30 0.7 1 3.6 6.3
Quercus boreaIis 12.5 1.7 126.43 0.6 1 3.6 5.9
Vitis aestivalis 5.0 0.7 107.00 0.5 1 3.6 4.8
Mmlus 10.0 1.4 85.90 0.4 1 3.6 5.4
Salix nigra 7.5 1.0 $.83 0.2 l 3.6 4.8
Crataegus spp. 5.0 0.7 24.55 0.1 1 #36 4.4
TOTALS 737 .5 100.0 19 ,768.46 99.8 28 100.3 300.1

20-3011
Acer saccharun 160.0 49.2 9398.98 54.7 4 21.1 ' 15.0
Carya ovata 2.5 0.8 3141.60 18.9 1 5.3 5.0
Prunus serotina 7.5 2.3 1455.93 8.7 2 10.5 . 21.5
Ulnus arericana 32.5 . 10.0 1198.20 7.2 3 15.8 33.0
mercus borealis 35.0 10.8 659.75 4.0 2 10.5 25.3
Pbras rubra 2.5 0.8 601.33 3.6 1 5.3 9.7
Rhus typhina 57.5 17.7 332.38 2.0 2 10.5 30.2
Cornus racemsa 12.5 3.8 61.38 0.4 1 5.3 9.5
Crataegus spp. 7.5 2.3 36.82 0.2 1 5.3 7.8
. POpulus tremloides 2 5 0.8 31.42 0.2 1 5.3 6.3
Corylus arericana 5.0 1.5 24.55 0.1 1 5.3 6.9
TOTALS .33 .0 100T 16 ,3232 100.0 19 100.2 300. 2

 

34

Table 6. Sumary of vegegtion transects at the 43rd Street site.

 

 

 

 

 

 

 

 

 

Den. Rel. Dom. Rel. Rel. Imp.
Species (per 1000112) Den. (cmz/ 1000n2) Dom. Freq. Freq. Value
O-lOm
Salix % 5.0 0.7 5,987 42.2 1 3.7 46.6
Salix $5.0 35.2 1,892 13.4 4 14.8 63.4
Prums serotina 17.5 2.3 1,68) 11.8 2 7.4 21.5
Robinia psuedoacacia 62.5 8.3 1,371 9.7 1 3.7 21.7
(mafia spp. 45.0 6.0 997 7.0 3 11.1 24.1
(brnus stolonifera 155.0 20.6 784 5.5 3 11.1 37.2
M oElifolius 75.0 10.0 $5 2.7 2 7.4 20.1
Zantl'flcylun americarun 55.0 7.3 299 2.1 2 7.4 16.8
Viburnun 131% 32.5 4.3 $7 1.9 2 7.4 13.6
Mg 7.5 1.0 255 1.8 2 7.4 10.2
Ulnus americana 10.0 1.3 150 1.1 2 7.4 9.8
wgustrls 22.5 3.0 110 0.8 3 11.1 14.9
TOTALS 752. 5 100.0 14,177 100.0 27 99.9 299.9
10-20n
Prums serotina 22.5 3.6 11,385 59.5 3 14.3 77.4
m spp. 60.0 9.6 2,618 13.7 3 14.3 37 .6
Robinia 185.0 29.7 2,290 12.0 1 4.8 46.5
Cornus stoloniferaa 125.0 20.0 686 3.6 3 14.3 27.9
gpliifolius 93.0 14.4 442 2.3 2 9. 5 26.2
Viburnun lentag 47.5 7.6 $1 2.0 2 9.5 19.1
Ulms americana 5.0 0.8 374 2.0 1 4.8 7.6
.Mw 10.0 1.6 $4 1.9 2 9.5 13.0
Salix % 52.5 8.4 317 1.7 1 4.8 14.9
Malus <_:_____oronaria 2.5 0.4 142 0.7 1 4.8 5.9
Zantllxylun— americanun 12.5 2.0 61 0.3 1 4. 7.1
Sanbucuscanader_1s—is 10.0 _1.6 60 0.3 1 4.8 6.7
TOTALS— 622.5 99.7 19,120 100.0 21 100.2 299.9
20-3011
Robinia W 35.0 19.4 1,0% 56.7 1 16.7 92.8
Vihmnm 131% 50.0 27.8 319 17.5 1 16.7 62.0
Oornus stoloniferaa 52.5 29.2 263 14.4 2 33.3 76.9
W flaws 32.5 18.0 160 8.7 1 16.7 43.4
Sanbucus canadensis 10.0 5.6 49 2.7 1 16.7 25.0
TOTALS 180.0 100.0 1,827 100.0 6 100.0 3(1). 1

 

 

aMost Oomus was fistolonifera, with sane scattered C_.aumum arrl C_.racanosa also

 

present .

35

stream borders (Otis, 1950), were the dominant shrubs, and accounted for‘
82% of the shrub basal area. Vegetation varied in respect to composition
and coverage between the sample quadrats, reflecting the properties of
the sample location. One transect fell along a moderately steep bank
and accounted for most of the black cherry and black locust (Robinia

pseudoacacia) found at this site. In comparison to the other locations

 

(Table 10), this site was low in woody species (16), with three of
these confined to one transect (steep bank). The other transects were
more representative of the shrub-carr vegetation type found along
Augusta Creek. Stem density declined markedly back away from the
stream, not only due to a decline in shrubs, but also due to the
general lack of woody vegetation.

Disturbance at the Nagel site is indicated by its vegetative
description (Table 7). Of the 19 species found at Nagel, 11 were trees
and nine were shrubs. Most of the woody plants here are characteristic
of swamp forests of the region (Otis, 1950; Braun, 1961). Red maple
Q5; rubrum) and musclewood dominate the trees and shrubs, respectively,
together comprising almost 60% of the basal area of the stand (Table 9).
Interpretation of the transects was difficult at this location because
of disturbance. Basal area was very low, at least in part, because of
vegetation removal. For example, only scattered red maples were
present in most disturbed area due to selective cutting by the
landowner. Shrub cover has been reduced in comparison to the other
sites, as shown by the low density and basal coverage, particularly at
the creek margin (Table 9).

In the Kellogg Forest, Augusta Creek transverses a well developed

(50-60-year—old, Walt Lemmien, pers. com.) bottomland forest. Three

Table 7 .

Summary of vegetation transects at the Nagel site.

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Den. Rel. Dom. Rel. Rel. Imp.
Species (per 100062) Den. (mg/1000112) 06m. Freq. Freq. Value
O-10m
Prunus serotina 22.5 15.8 783 40.6 3 17.6 74.0
Salix nigra 42.5 29.8 601 31.1 4 23.5 84.4
Physocarpus opulifolius 25.0 17.5 123 6.4 3 17 .6 41.5
Carpinus ccaroliniana 12.5 8.8 122 6.3 1 5.9 21.0
Salix discolor 15.0 10.5 102 5.3 2 11.8 27.6
Cornus racemosa 12.5 8.8 97 5.0 1 5.9 19.7
Francinus nigra 2.5 1.8 49 2.5 1 5.9 10.2
Vibumm lentago 5.0 3.5 30 1.6 1 5.9 11.0
Cornus stoloniferaa 5.0 3.5 25 1.3 1 5.9 10.7
TOTALS 142.? 100.0 1, 932 100.? 17 100.6 300.1
10-20m
Betula lutea 12.5 7.9 1,857 11.3 2 18.2 62.4
Carpinus caroliniana 57.5 8.5 1,724 33.7 1 9.1 79.3
Prums serotina 2.5 1.6 567 11.1 1 9.1 21.8
Acer rubrun 5.0 3.2 300 5.9 2 18.2 27.3
1335 rubra 27.5 17.5 212 4.1 1 9.1 30.7
Populus trenuloides 35.0 22.2 204 4.0 1 9.1 35.3
Robinia psuedoacacia 2.5 1.6 158 3.1 1 9.1 13.8
Acer saccharun 7.5 4.8 53 1.0 1 9.1 14.9
Physocarpus opulifolius 7.5 4.8 37. 0.7 1 9.1 14.6
TOTALS 157.5 100.1 5,112 99.9 11 100.1 300.1
20-3111
Acer rubrun 17.5 12.5 8,644 70.3 3 25.0 107.8
Tilia americam 2.5 1.8 1,431 11.6 1 8.3 21.7
Carpims caroliniana 15.0 10.7 554 4.5 1 8.3 23.5
Oornus racemosa 67.5 48.2 492 4.0 2 16.7 68.9
Ulmus americana 5.0 3.6 403 3.3 1 8.3 15.2
Horas rubra 22.5 16.1 269 2.2 1 8.3 26.6
Betula lutea 2.5 1.8 213 1.9 l 8.3 12.0
Prunus aviun 2.5 1.8 216 1.8 1 8.3 11.9
Populus tremloides 5.0 3.0 42 0.3 1 8.3 12.2
TOTALS 140.0 100.1 12,289 99.9 12 99.8 89.8

 

 

alncluchs some C_. anrnun.

37

canopy species, basswood (Tilia americana), red ash (Fraxinus

 

pennsylvanica) and black ash (F; nigra), account for almost 90% of the

 

basal area (Tables 8 and 9). The two ashes predominated in the first

20 m while basswood was the leading species in the following 10 m.

Total basal area was highest in the O-lO-m plots where red ash attained
greatest size. This species is very tolerant to high (>3%) flooding
frequencies (Bell, 1974), and was apparently best suited to the
environment conditions at streamside. Basswood, which requires more
well-drained soils (Otis, 1950), increased as the terrace was approached.
The canopy association at the Kellogg Forest site is much like the type
dominating the most strongly developed seral stages on the floodplain

of Hickory Creek, Illinois (Bell and Del Meral, 1977).

Shrub cover, which constituted less than 10% of total basal
coverage, was dominated at this site by musclewood and gray dogwood.
Similar reduced shrub cover was noted by Johnson, et al., (1976) under
mature, dense-canopied ash-elm floodplain forests along the Missouri
River. Species richness among shrubs was high, since they accounted
for 15 of the 22 woody species at the site. As with the trees, shrubs
were predominantly those types associated with lowland habitats.

When comparing the different locations in this study and other
riparian associations, it is important to consider controlling
environmental factors. Variations in the structure of stands of
streamside vegetation have been associated with various physical
gradients, such as relative flooding severity (Bell, 1974; Franz and
Bazzaz, 1977), slope aspect and slope angle (Wikum and Wali, 1974;
Killingbeck and Wali, 1978). Due to microtopographic changes,

floodplains exhibit a flood frequency gradient. Along this gradient

38

Summary of vegetation transects at the Kellogg Forest site.

Table 8 .

 

Imp.
Value

Rel.
Freq. Freq.

Rel

Rel. Dom.

Den.

61891660070

.594.7971734

.8
.Am MN... 111

um

82.1
32.8
21.1

889889988999999

812882588225n/fl22

11.8

34.133112331121111

llSnU../.3299w6l.ml43331

60.442111000000000

3%..25
1

160
149
1 10

500.0nU.nwnU.550550.0.50.

W0808557flm328825

89.7

34 99.7

8 ,668 100.0

100.0

 

495.0

3 .l. 2 5 5
3 1y82 On ..I.9 3
[45541535111311

O.SnU.SSSfiU.SfiU.50.50.5

002253076262

32 100.0 89.8

99.8

21,690

100.0

 

410 .0

1930.7.18rnw346536wnwo.

(msss
91Ml4

613r0.rO.3—/.661rnwr0.r0.66

37.4.3340.337.33333

10.7

1.2/4114311211111

763698065532111

765322100000030.

67.8

94 760 020/... 9
mflmmfimmnmmsmamsu
12:.)2n/flll

6r0.l4..3.6—/.90.3065933

O~lo620pnfinlo8702111u

16 .6

snwnU.0.55555550.50.nw

2msoz7 my3n20755

65.0

.1

28 100.2

100.0

32,245

99.8

 

395 .2

 

manna Mmoflmm¥)um

Species
0- 10m

 

Fraxinus permsylvanica
Tilia americana
SPP-

Fraxinus nigra

 

Carpinus caroliniana

Corylus anericana

Cra

 

Physocarpus opulifolius

 

 

Populus treruloides
Corms racemosa

 

Rhamnus cathartica

 

 

Vitis aestivalis

 

 

Ulnus americana

 

Rosa palustris

 

Viburnun lentago
Viburrun opulus
Cornus 31mm

 

 

 

 

10-20m

Spp-

Fraxims pennsylvanica

Fraidnmnigra

 

W
W
0

 

 

 

Populus traruloides
Coqlus americana

Vibumun lentago
Sanbucus canadensis

Rhanrms cathart icus

Pmnus virginiana

 

 

 

 

Ulnus americana
Corrus ammun

 

 

 

Tilia americana
Corms racemosa

 

Crata

 

20-30n

spp-

 

 

 

Corylus americana

Cra
Sanbucus canadensis

Cornus anrmm
Rhannus cathartica

 

 

Prunus serotina

 

W opulifolius

Fraxims pennsylvanica

Acer rubrun

 

Carpinus caroliniana

Cornus racemosa

 

 

Populus tremloides

 

Vibtmm lentago

 

 

Tilia americana
Pims strobus

 

Fraximsniga

 

 

 

 

39

the best predictor of the field distribution of a species is its
relative level of flood tolerance (Gill, 1970; Franz and Bazzaz,
1977). Another consideration important to stand composition is its
successional development (Hosner and Minckler, 1963; Bell and Del
Moral, 1977). Generally, old, compositionally stable floodplain
forests characteristically have the highest basal area and a reduced
density (Curtis, 1959; Johnson, et al., 1976).

The coverage of woody vegetation, particularly the tree species,
varied considerably between the four Augusta Creek locations. Total
tree basal area was highest in the Kellogg Forest by a factor of two,
three and five as compared to the Smith, 43rd and Nagel sites,
respectively (Table 9; Figure 5). Furthermore, there was little
species overlap between the stands. Black cherry was among the leading
species at Smith, 43rd and Nagel, while Kellogg Forest and Nagel shared
basswood. Even at a location, there were often considerable changes in
canopy species along the transect, probably reflecting the topographic
gradient back from the channel. Within transect differences were
particularly evident at Smith, where the floodplain was most narrow
and an upland association prevailed. The other three sampling areas
were predominantly lowland communities and reflected their differences
in wetness, succession and disturbance.

The most striking vegetational differences were observed at the
Nagel and Kellogg Forest sites (Figure 5). Nagel has undergone the
most alteration, and all stand descriptors have been greatly reduced
(Table 7). The canopy species which remained were representative of
the wet-mesic southern lowland forest (Curtis, 1959; Catana, 1967),

and indicated lower soil moisture than that at the Kellogg Forest. The

Table 9. Dominant trees and shrubs and quadrat summaries at the sites.

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Basala Z of X of Quadras Totals

Species Site Area Type m /ha

Tree Total Smith 2.9 Trees
Acer saccharum 4.9 0-10m 7.8
Prunus serotina 3.4 10-20m 14.8
Carya ovata 1.8 20-30m 16.2
Ulmus americana 1.4

Tree Total 43rd 8.3
Prunus serotina 4.4 0-10m 9.4
Salix alba 2.0 10-20m 14.4
Robinia pseudoacacia 1.6 20-30m 1.0
Juglans nigra 0.2

Tree Total Nagel 5.0
Acer rubrum 3.0 0-10m 0.8
Betula lutea 0.7 10—20m 3.1
Tilia americana 0.5 20-30m 11.0
Prunus serotina 0.4

Tree Total Kel. For. 27.9
Tilia americana 12.9 0-10m 33.8
Fraxinus pennsylvanica 11.2 10-20m 19.6
Fraxinus nigra 2.1 20-30m 30.4
Pinus strobus 0.8

Shrub Total Smith 4.3 Shrubs
Cornus racemosa 1.8 0-10m 7.5
Carpinus caroliniana 1.2 10-20m 5.0
Rhus typhina 0.6 20-30m 5.0
Corylus americana 0.2

Shrub Total 43rd 3.4
Crataegus spp. 1.2 0-10m 4.7
Salix nigra 0.7 10-20m 4.7
Cornus stolonifera 0.6 20-30m 0.8
Physocarpus opulifolius 0.3

Shrub Total Nagel 1.5
Carpinus caroliniana 0.8 O-lOm 1,1
Salix nigra 0.2 10-20m 2.0
Cornus racemosa 0.2 20-30m 1.3
Mbras rubra 0.2

Shrub Total Kel. For. 2.3
Carpinus caroliniana 0.6 0-10m 2.9
Cornus racemosa 0.5 10-20m 2.1
Viburnum lentago 0.2 20-30m 1.9

0 2

Corylus americana

 

 

 

aMean mZ/ha based on the quadrat totals.

41

40

 

Vegetation sham

D UNDERSI’ORY
- CANOPY

30.—

   
    

‘ ‘ I
'7. (v..‘ .&

  

m2/ ho
8
fi?”?§¥§

A 7‘ A. {I’M-7".” HMJ 5” J5. .tit‘ fi.’!flm-l PPM'; I , ‘v:
r - .7 J7 -' . A 4 :2 l 1:; a. a" 5:1 ‘ . , - ;.
l I ”7777‘. ‘I' I .7- w_, .‘ . 5‘. ,' ~.‘ ».-.,- ,

 

 

 

KF

Figure 5. Average basal area of the woody vegetation (>2.0 cm DBH)
at the sites.

42

community in Kellogg Forest experienced more complete inundation

(i.e. across the entire transect), and resembles the wet, southern
lowland forest type (Curtis, 1959). Because of the most uniform
environment along the transects, the Kellogg Forest stand showed the
greatest homogeneity in vegetative cover among the sites (Table 8).

The increased basal area, reduced density and greater coverage of trees
(90%, Table 9) all suggested that this was also the oldest stand
surveyed in this study (cf. Johnson, et al., 1976).

Relative to the trees, shrub basal area was much more uniform
between locations, and between quadrats within a location (Table 9;
Figure 5). Greatest coverage was found at the Smith site, with its
young forest association. The lowest total was at Nagel, reflecting
its disturbance. Dominant shrub species were similar at the sites,
with musclewood and gray dogwood shared among three locations and black
willow between two of them. Except for hawthorn at 43rd Street, the
shrub layer was largely made up of dogwoods, musclewood and black
willow. From other regional studies (White, 1965; Liston, 1972;
Sytsema and Pippen, 1980), these are common riparian species.

Shrubs appeared to be well adapted to the edaphic and moisture
conditions next to the stream, since they reached their greatest
density and basal coverage there (Table 9). Ice damage, erosion,
flooding and siltation, which are most prevalent at the channel margin
(Sigafoos, 1961; Lindsay, et al., 1961), are normal effects of the
hydrologic regime of north-temperate running waters (Cooke and
Doornkamp, 1971). These environmental attributes may serve to maintain
an earlier successional stand dominated by a few small species (i.e.

shrubs - Nicholson and Monk, 1975). Once established, many streambank

43

shrubs form dense colonies from underground, horizontal rootstalks,
which tend to persist in a relatively stable condition for a long period
of time (White, 1965). Ware and Penfound (1949) noted that shrubs were
of primary importance in stabilizing the sides of the channel of an
Oklahoma river and, in flood prone areas, these species tended to remain
dominant.

Published results from other deciduous, riparian woodlands are quite
variable, in part because of the different locations and stand histories,
but also due to dissimilar methods of analysis (Table 10). Most
investigations of woody vegetation along watercourses have been carried
out in areas of relatively undisturbed or mature second growth forests.
In addition, as is evident by the paucity of data on basal coverage of
understory woody vegetation, these surveys concentrated on canopy
species. Such differences make comparisons with the Augusta Creek sites
more difficult.

Among riparian forests in the Midwest, the Augusta Creek stands
appear to be younger than most which have been studied, but comparable
in species numbers (Table 10). Density was very high at the sites in
this study, while basal area ranged from moderate (Kellogg Forest) to
low (other sites) levels. Because of the relationship between biomass
and litterfall (Bray and Gorham, 1964), the age of the stands is
important to the level of CPOM influx to the aquatic system. With
normal seral develOpment, forests along Augusta Creek should accumulate

biomass and decrease in density (Bormann, et al., 1970).

44

A mg: 385 node :Sumoafimmflou
£me 5 3.8 Hogan
.3}. 5 >39QO

 

 

mm a 5 gm m2 3 as ea 8: .88.. 80:3.
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abate segues. meouom on 7 7 7 7 7 s8 Nam m8 m5 .4. u. .836“
£7.88. Seamus... 5.3.38 8 7 7 7 7 7 mm 98 2.. £2 ..E .... .comuflom
Etta EBB... J... .3 : w m mam 38 7 7 7 7 82 £83.. a swim
.25 53%.... Qua. B m 889 ...... 58 . .... .89 as; mean 8.8m

8 m €53 Hoop 888:: E95

 

.<.m.: 58ng m5 Sum cowumumwg 54.83.“ 683 MO mud—3m vmuooaow mo 83.8950 < .9 03mm.

45

CPOM Influxes at the Sampling Sites

 

The total amount of allochthonous CPOM entering the stream per
meter of bank at the Smith location was an estimated 714 g/m in 1976
and 547 g/m in 1977 (Table 11). Of these totals, almost 80% was direct
litterfall, with the rest downslope movement (Figure 13). Generally,
leaves and woody material were the most common components, comprising
80% or more of the litter (Table 12; Figure 6). Fruit totals were
especially large in 1977, with this being a good mast year for black
walnut (Table A10). The high amount of wood in 1976 (45% of total)
resulted from several severe storms in the winter and early spring
(Figure 6; Table A2).

Autumn was the season of largest particulate influx and accounted
for 52% of the annual total in 1976 and 76% in 1977 (Table 13; Figure
6). Inputs were quite similar in the fall between the two years and
were mostly leaves. Consistently lower totals were recorded for the
other three seasons, with variability at these times due to influxes of
woody material as a result of inclement weather patterns. Except for
these wood pulses, winter appears to be the time of lowest litter input
to the aquatic system (Table 13).

Leaf inputs were quite constant during the two years, totalling
366 g/m and 348 g/m in 1976 and 1977, respectively (Table 14). An
estimated 65% of the incoming leaves were from species characteristic
of the medium processing (decomposition rate) range, predominantly
black walnut and American elm (Table 20). Species which break down
rather rapidly in streams ("fast") averaged 20% of the total and were
mostly gray dogwood. "Slow" leaves made up 15% of the leaf litter and

were mostly quaking aspen (Populus tremuloides).

 

6
l4

cu mac mum o<l~< mam

.xcmn mo E\w u Hmuop m.E\w cw uuoamcmuu Hmumuma u a ”~E\w cw Hanna“ uomuwv u Ho
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: cos: mnmfi :ufism you unmoxm mouwm mam mcommmm Ham now 0 u an
.muouum mcaocsou

RHINH mean ca omosu mam magma mficu :H mHmuou mzu :mosumn mmfiocmamuomfiv unwwam ones

 

 

 

N©~.moq nno.oaa wmm.mmc mmm.mqm A<Hoe nz<mu
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Hmao.m cmm.mm amom.qm owo.do fimow.mm nwm.¢- unoc.m mun.mo Hamm
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mmdo.c onm.¢~ mmcm.n Hoc.oH mmm¢.m~ mmm.~q oodm.m ooo.~m umassm H
«mmo.~ oqc.- mama.“ ooc.m nmma.m oao.o~ N-~.c omn.qm magnum
omm~.~ 909.5 ~Hoq.o m~n.m omqm.q ~m~.- fimoo.~ moc.w nous“:
dom.omm <oc.~a~ ww~.ohm nw~.q- A<HQH oz<mu
ndo.~o mN¢.o- mac.om~ awm.am~ A<HOH
on~¢.~ o~o.o< NNNN.¢¢ N¢~.mn ooww.m~ -¢.Ho~ q¢m~.~H cho.am Hamm
wmdq.m o~o.c co-.m om~.m~ w-o.q No~.w~ monn.~ ~m¢.o~ Hoseam A
mcmw.o «am.m «Nom.m omm.N~ won~.¢ woo.o~ Hmmq.q m~m.cm wcfiuam
«mow.o cae.m m-~.~ mnn.m~ mNow.m Nom.m~ ano.¢ www.mm noun“: e~¢~
qwm.<o~ mmo.mn mn~.q~m won.¢mm A<HOH
mom¢.mfi omc.mom omwo.mm www.mm a~om.mm msq.o¢~ comm.nm mm~.mm~ Hamm
mNNm.m cww.mm quo.~ ~o~.¢ o~wo.m w-.om «Hom.m wuo.nm umaasm pH
omm©.m oqn.q~ ~woo.~ omo.o homo.n ~mc.o~ omfin.om www.mma magnum
n<©m.~ wo~.- nmmm.~ omq.H~ fiomm.~ mum.o~ ¢o~m.co mma.oo umucas
.m.m cmm: .m.m cmmz .m.m coo: o.m.m sumo: commwm
amouom wwoaamx Hmwmz oomuum vume Luwam
m.mmmmu umHmEmm zn mmuam xwmpo mumsws< mnu um moxSchH zomu HmuOu cam Hmcommmm .HH manme

47

.0000 000000000 H00000H u A "0000 HH000H 0000H0 u H

m

 

 

 

 

 

 

 

 

 

 

 

o.ooH mo.moq 05.05 mm.0~m H.ooH 0m.0~m ma.H0 mm.q0~ H0009
0.H 00.0 0~.0 -.o c.H oe.m mm.~ 00.0 000002 000000aumm
0.o 0H.m 00.0 0H.m ¢.N 00.5 0~.H Nq.0 0H005
0.0H 5N.m0 00.0 mm.5m 5.5H H0.50 m<.mH 0H.q¢ 000002 50003
o.~m 0H.~mm ~5.00 ~¢.50~ 0.05 00.5mm Hm.<q 05.~H~ 00>004
000005 meHHmM
m.ma mo.caH 0m.mHH m¢.¢5 o.ooH <¢.HmH 00.0HH 00.05 H0009
5.0m 05.00 00.nq 0~.5 N.m~ mm.wq cm.mm Hm.¢H 00000: 000000000:
m.o 00.0 I 00.0 c.m 00.0 0H.N Nm.¢ 0H=um
0.0 mm.oH 00.0 50.0 0.0 om.5H 55.HH m¢.0 000002 50003
0.05 Hm.0NH O0.50 H5.00 «.m0 Hm.mHH mm.w0 00.00 00>000
Hmwmz
o.ooH «0.500 0m.mON 0o.~0~ 0.00 mH.O5m Ho.0mH 0H.¢HN H0009
5.0 No.m~ 00.0H cm.m 0.0 Hq.om 0m.¢H no.0 00000: 0000000003
m.m 00.0H c~.m n0.m m.< 5m.mH 0H.HH m0.¢ 0H005
5.0H mm.oa oo.~m mm.0m N.oN mm.¢5 H¢.0m 00.0m 000002 50003
0.~5 N0.qmm 00.0NH «m.¢- m.m0 00.00N 00.0m 05.00H 00>0ma
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0.m 00.0w 0m.5H om.m m.~ 0m.o~ 0H.¢H 00.0 00000: 000000000:
N.5H Hm.¢m 00.H m~.ma m.o N0.0 NH.o 00.0 0H00m
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0.00 5m.5qm mm.05 00.H5N n.00 Hq.00m mm.5¢ wo.m0m 00>000
50HSm
H0009 53000 Aa\w Ama\wv H0009 5x000 Aa\wv 5~e\wv 500w000o
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.000H0.xmmuu 00maw0< 0:0 00 mmstmcH zomu H0000 000 20mm 00 00H00w000o .NH 0H009

48

.0000 we E\w 0H m0H=000 000000000 H00000H 000 HHmwaH 0000H0 mo 00H000H05000

 

 

 

 

 

 

 

 

 

 

 

00.0 m.mo mo.moq oo.o o.oo0 qm.o~m 0muo0

oo.q 0.00 00.0mm 5o.~ 0.05 oc.m¢~ 00mm amouom
mm.o 0.0 mm.Nm om.o «.0 ~m.om umae=m 00°00mx
mN.o 0.0 mo.m0 0m.o 5.0 on.m~ mc0uam

m0.o 0.0 ~w.o0 o0.o «.0 00.50 000:03

mm.o o.oo0 mo.om0 Nm.o a.ao «0.0m0 00000

mm.0 0.00 mo.n00 00.0 w.oo 00.500 0000

mad 0.0 00.00 o~.o o.o 5~.50 00.08% 00002
on.o o.- 05.00 00.0 m.a mo.w0 ws0uam

~0.o «.5 am.m0 «0.0 0.00 55.50 000:0:

«5.0 o.oo0 «4.500 No.0 o.oo0 m0.05m 0muo0

mm.m o.5e mm.o~m om.~ 0.50 mm.mq~ 00mm

5w.o 0.00 mq.m5 mm.o 0.50 00.00 umee=m ammuum cums
wo.o 0.00 50.50 om.o 0.00 00.50 0:0000

50.0 0.0 mm.m~ m~.o a.o wm.m~ 000:0:

om.0 o.oo0 00.5qm oo.0 m.m¢ 50.005 0muo0

om.¢ 0.05 «0.000 00.0 «.00 mm.c5m 0000

oq.o 0.5 m5.wm mq.o ~.m 00.5m “measm 000am
«0.0 o.m0 No.05 50.0 o.m~ 00.000 0:0000

m0.o 0.5 mm.o~ no.0 5.00 mm.m00 umua03

500\E\w N E\w 500\E\m N 0a\w 0000mm muHm

0500

 

.00000 30000 00m=ws< 0:0 00 00000 000 mH0000 Eomo H0000000 000H0500 .mH 0H009

DRY WT. IN g/m

49

 

 

 

Litter Types
D HERBS
400 — - FRUIT -
[:5 wooo 1 ,
LEAVES F;
523:5:
.
300 - "‘
200 r- \ "
100 £52 -*
232:;

 

 

 

 

 

 

 

 

 

 

 

Wi Sp Su F0
1976

 

Figure 6. Seasonal CPOM influx in g/m of streambank at the Smith site.

50

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Table 14. Leaf influx at the Smith site.

Yearly total in

g/m Of stream bank Cumulative Cum. %
Species 1976 1977 x total of total
Juglans nigra 87.82 97.92 92.52 92.52 25.9
Ulmus americana 68.58 60.29 64.44 156.96 43.9
£2£22§.SPP- 67.77 57.46 62.62 219.58 61.5
Populus tremuloides 47.53 47.70 47.62 267.20 74.8
Viburnum lentago 26.81 13.66 20.24 287.44 80.5
Prunus serotina 22.21 17.64 19.92 307.36 86.0
Spring fragmentsa 17.86 16.29 17.08 324.44 90.8
Acer saccharum 11.10 6.02 8.56 333.00 93.2
Quercus borealis 1.66 10.12 5.89 338.89 94.9
Salix nigra 5.10 5.89 5.50 344.39 96.4
Miscellaneous leaves 1.43 9.07 5.25 349.64 97.9
Corylus americana 5.89 3.82 4.86 354.50 99.2
Crataegus spp. 0.80 0.80 0.80
Populus grandidentata 0.61 0.76 0.68
Carpinus caroliniana 0.85 0.12 0.48
Carya ovata - 0.62 0.31
Quercus alba 0.14 0.36 0.25
Rosa palustris - 0.16 0.08
Acer saccharinum 0.14 - 0.07
Fraxinus nigra 0.14 - 0.07
Totals 366.44 348.00 357.24

 

aIncludes leaf particles, bud scales and reproductive parts which were
captured during the spring and early summer.

51

A comparison of leaf influx (Table 14) with the vegetation transects
(Table 5) at this location indicated that relatively small amounts of
leaf litter (as CPOM) which entered the stream originated more than 10 m
from the channel. The five dominant species (Table 14), which
contributed more than 80% by weight to the total, reached their greatest
basal area in the 0-10-m vegetational quadrats. 0f the upland species
at this site, only black cherry was well represented in the leaf input
totals. However, it was also of importance in the 0-10-m portion of the
riparian zone (Table 5).

CPOM influx to the 43rd Street site was 370 g/m in 1976 and 488
g/m in 1977, a 32% increase in the second year (Table 11). Of these
totals, 58% was collected in the infall traps giving a relatively high
lateral transport of 42% (Table 12). The four basic litter components
were remarkably consistent between the two years in respect to their
percentage of total inputs (Table 12). Therefore, the estimated
differences in CPOM inputs between 1976 and 1977 involved all litter
types, in contrast to the situation previously noted at the Smith site.
Leaves (71%) and twigs (19%) were the dominant categories of litter
(Figure 7).

Among the seasons, winter, spring and summer CPOM contributions
were consistently low and accounted for one-third of the total in both
years (Table 13). Inputs were lowest in the winter, averaging only 6.5%,
while spring and summer averaged 12% and 14% respectively, of the annual
total. Fall influxes were mostly leaves (83%) and twigs (9%), with
herbaceous materials and fruits of minor importance (Figure 7). Over
the rest of the year, leaves made up 50% of the litter, with twigs and

herbs comprising most of the remainder (Table A3). Direct infall was

52

 

 

 

 

 

 

 

 

 

Litter Types
DHERBS
40°“ 5553377 "
.LEAVES
§
E §
: 5
. \
§
\
W Sp Su F0
1976 1977

Figure 7. Seasonal CPOM influx in g/m of streambank at the 43rd
Street site.

53

the preponderant input in summer and fall, while lateral transport was
higher in the other seasons. Increased lateral transport was especially
evident in the early spring following snowmelt and thawing of the ground
(Table A6; 24 Mar 76, 23 Mar 77).

Although the total influx of leaves was almost 40% higher in 1977,
the percentages by processing types were quite similar in each year
(Tables 15 and 20). Leaves were commonly of the medium processing rate,
particularly black willow, black cherry and nannyberry (Viburnum

lentago). "Fast" leaves were mostly dogwoods, prickly ash (Zanthoxylum

 

americana) and black locust, while the most abundant "slow" leaves were
hawthorns.

Most CPOM reaching the stream probably originated within 20 m
of the channel, since only 5% of the basal area of woody plants was
found beyond that point (Table 6). The major leaf input was black
willow (Table 15), the dominant shrub at the channel margin. 0f the
other abundant leaf inputs, several reached their greatest basal
coverage in the 10-20-m quadrats (Table 6 - black cherry, hawthorn).
While the 0-10-m portion of the vegetation corridor was probably the
major litter contributor, this cannot be stated conclusively due to the
overlap in riparian species in the 0-10-m and 10-20—m quadrats.

The annual estimate of allochthonous CPOM at the Nagel site was
almost identical over the two years (Table 11). The amounts within the
different categories of litter were quite~ similar, with most of the
total made up of leaves (65%) and herbaceous material (26%) (Table 12).
A distinctive characteristic of this location was the high level of
lateral transport, accounting for over 60% of the annual sum.

Autumn was the time of greatest particulate organic movement into

54

Table 15. Leaf influx at the 43rd Street site.

 

Yearly total in

g/m 0f stream bank Cumulative Cum. %

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Species 1976 1977 2 total of total
Salix nigra 53.67 61.69 57.68 57.68 18.8
Prunus serotina 47.15 51.82 49.48 107.16 34.9
Crataegus spp. 38.83 52.86 45.84 153.00 49.9
Eggngg spp. 20.61 46.76 33.68 186.68 60.7
Viburnum lentago 28.79 22.55 25.67 212.35 69.2
Spring fragmentsa 24.65 20.51 22.58 234.93 76.6
Zanthoxylum americanum 10.98 17.96 14.47 249.40 81.3
Robinia psuedoacacia 8.76 19.42 14.09 263.49 85.9
Juglans nigra 10.28 16.71 13.50 276.99 90.9
Rosa palustris 5.83 10.56 8.20 285.19 93.0
Ulmus americana 3.20 11.72 7.46 292.65 95.4
Miscellaneous leaves 3.16 6.22 4.69 297.34 96.9
Physocarpus opulifolius 1.53 7.48 4.50 301.84 98.4
Malus coronaria 0.06 3.06 1.56

Fraxinus spp. 0.57 2.27 1.42

Quercus borealis 0.72 1.68 1.20

Corylus americana - 0.86 0.43

Hammamelis virginiana 0.04 0.48 0.26

Totals ’ 258.83 354.61 306.72

 

 

aIncludes leaf particles, bud scales and reproductive parts which were
captured during the spring and early summer.

55

the stream, averaging 64% of the whole. MBan inputs for the other
seasons ranged from 9 to 16% of annual totals, with summer the lowest,
winter intermediate and spring highest (Table 13; Figure 8). Though
leaves were almost 80% of the autumn litter, they accounted for only
27 to 45% during the rest of the year (Tables 12 and A4). Herbaceous
material averaged 40% of all CPOM inputs over the non-fall seasons.
Generally, lateral transport was higher than direct infall, even in the
autumn when litterfall was maximal.

Rapidly processed leaves were the most abundant type at the Nagel
location, averaging 60% of total leaf litter (Tables 16 and 20). These

were mostly basswood, with some dogwood and swamp rose (Rosa palustris).

 

"Medium" leaves, mainly black willow and nannyberry, averaged 28% of
the leaf sum. There were only two species contributing measureable

"slow" leaves, red oak (Quercus borealis) and quaking aspen (Populus

 

tremuloides).

 

Due to the vegetational pattern at this site, the transects (Table
7) do not readily identify the riparian area which contributed CPOM.
Leaf litter inputs were relatively meager at Nagel as a result of
reduced woody vegetation. Because of this, one clump of basswood
located next to trap number 4 contributed almost 50% of the leaf total.
Since these trees were located in the middle of the 30-m sampling
section they were not included in the vegetation transects. Except for
basswood, the dominant woody species in the 10-20-m and 20-30-m
quadrats (Table 7) are poorly represented in the litter data (Tables
16 and 20), Thus, it was likely that most leaf influxes came from the
0-10—m zone.

In the Kellogg Forest, the annual litter input was 326 g/m in 1976

400
g 300
“~5
0')
Z
...°
3 200
>-
a:
Q
100

Figure 8.

56

 

Litter Types

[:3 HERBS

" III FRLHT "
a: wooo

LEAVES

 

 

 

 

 

 

E30

Wi Sp Su
1976

 

3‘7///////////////////-

 

Seasonal CPOM influx in g/m of streambank at the Nagel site.

57

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Table 16. Leaf influx at the Nagel site.

Yearly total in

g/m Of stream bank _ Cumulative Cum. %
Species 1976 1977 x total of total
Tilia americana 73.28 44.99 59.14 59.14 47.7
Viburnum lentago 4.40 23.22 13.81 72.95 58.9
Salix nigra 14.65 10.95 12.80 85.75 69.2
Quercus borealis 5.38 16.51 10.94 96.69 78.0
Eggggg spp. 7.27 11.63 9.45 106.14 85.6
Rosa palustris 4.41 8.08 6.24 112.38 90.7
Spring fragmentsa 2.37 3.32 2.84 115.22 93.0
Populus tremuloides 1.96 2.39 2.17 117.39 94.7
Miscellaneous leaves 1.57 1.24 1.40 118.79 95.8
Acer rubrum 1.16 3.44 2.30 121.09 97.7
Physocarpus opulifolius 1.55 0.83 1.19 122.28 98.7
Fraxinus nigra 0.15 0.93 .54
Acer saccharum 0.54 0.12 .33
Ulmus americana 0.15 0.47 .31
Carpinus caroliniana 0.57 0.03 .30
Prunus serotina 0.19 0.15 .17
Totals 119.60 128.30 123.93

 

aIncludes leaf particles, bud scales and reproductive parts which were

captured during the spring and early summer.

58

and increased 24% to 405 g/m in 1977 (Table 11). Of these totals the
majority (81%) was direct infall. The yearly sums were mostly leaves
and woody material which averaged 82% and 17% of the whole (Table 12,
Figure 9).

0f the annual CPOM contributions, 80% entered in the fall. The
remainder of the year received fairly constant, but low amounts, which
were most reduced (4%) in the winter (Table 13). Lateral transport
exceeded direct infall in all seasons except autumn, especially in the
winter. Leaves were the dominant litter type in the fall period in
1976, and in 1977, throughout the entire year (Table A5; Figure 8).
Woody materials were the leading litter component in winter and spring
of 1976, probably as a consequence of severe weather conditions. Fruits
were especially abundant in the 1976 winter samples (Figure 7) and were
mostly ash samaras (Table A10).

Leaf litter collections in the Kellogg Forest were usually "fast"
types like ash and dogwood, which together accounted for 70% of the
total (Table 17). "Slow" leaves were almost nonexistent (30%), while
"medium" species, dominated by nannyberry, were moderately represented
(19%) (Table 20).

The vegetation analysis (Table 8) rather clearly indicates the
origin of the leaf litter. Green and black ash accounted for over
50% of the leaf input sum and made up almost 70% of the basal area in
the 0-10-m quadrats. The basal coverage of these two species decreased
by 60% the next 10 m and by over 90% beyond 20 m from the stream.
Basswood showed an opposite trend in ground cover and averaged only 5%
of the leaf total. Dogwoods, nannyberry and swamp rose reached their

greatest basal area at streamside and were important leaf contributors

59

 

 

0 . 0
_§§§§§

A

     

F0

 

 

 

 

 

0w.
0\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\80
a 5 $0.
mmmmm .
m Hmwm
mm-mm
... _ ... ._.

E\m z. .53 >05

 

1977

1976

l CPOM influx in g/m of streambank at the Kellogg

Seasona

59

 

 

0 - _
_§§§§§m

    

 

 

 

 

 

 

E\m z. .53 >05

66.7
7
.pw
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mm-mm
... ... ... ._.

onal CPOM influx in g/m of streambank at the Kellogg

Seas

9.

Figure

60

Table 17. Leaf influx of the Kellogg Forest site.

Yearly total in

g/m Of stream bank Cumulative Cum. %

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Species 1976 1977 E total of total
Fraxinus spp. 133.78 162.63 148.20 148.20 50.2
Corpus spp. 50.32 65.92 58.12 206.32 70.0
Viburnum lentago 15.41 21.80 18.60 224.92 76.3
Tilia americana 13.22 15.72 14.47 239.39 81.2
Rosa palustris 7.05 15.54 ' 11.30 250.69 85.0
Corylus americana 7.47 14.92 11.20 261.89 88.8
Spring fragmentsa 7.69 9.74 8.72 270.61 91.8
Carpinus caroliniana 7.99 7.58 7.78 278.39 94.4
Rhamnus catharticus 3.41 8.71 6.06 284.85 96.4
Ulmus americana 7.26 3.77 5.52 289.97 98.3
Miscellaneous leaves 1.13 2.36 1.74

Populus tremuloides 1.10 0.92 1.01

Quercus bicolor 0.36 1.71 1.03

Quercus borealis 0.93 0.67 0.80

Pinus strobus 0.36 0.08 0.22

Prunus serotina 0.20 - 0.10

Physocarpus opulifolius - 0.11 0.06

Totals 257.68 332.18 294.93

 

 

aIncludes leaf particles, bud scales and reproductive parts which were
captured during the spring and early summer.

61

(30% - Table 17). Species such as white pine (Pinus strobus) and red

 

maple, which were prominent beyond 20 m from the channel, were collected
in negligible amounts. In addition, the woody stem cover in the Kellogg
Forest was much higher in the 10-20-m and 20-30-m quadrats than at the
other locations (Table 9). If appreciable movement of leaves from areas
further than 10 m from the creek had taken place, measured totals should
have been higher than at the other sites. Therefore, it appeared that
most leaf litter contributed to the stream in the Kellogg Forest came

from the 0-10-m portion of the vegetative corridor.

CPOM Influxes Among the Sampling Sites

Over the two years of study, CPOM influxes were significantly
different (p<.05) among the locations (Table 18). Changes in riparian
vegetation appeared to play an important role in litterfall, since
CPOM totals (g/m or g/mz) were correlated (p<.05) with stand density
and shrub basal area (Table A9). Allochthonous litter was especially
reduced at the disturbed Nagel site (Figure 10), with significantly
lower (p<.05) amounts found at this location than a pooled value for
the others (Table 18). Between sites, CPOM collections were
significantly lower at Nagel than Smith (p<.05) in 1976 and 43rd Street
and Smith (p<.05) in 1977 (Table 18). Among the locations with no
disturbance, only one difference was significant; Kellogg Forest was
lower than Smith (p<.10) in 1976.

Removal of woody vegetation also affected the types of litterfall
at the Nagel site (Table 12; Figure 11). Herbaceous material accounted
for over 25% of CPOM influxes at the disturbed site, but only 6% at

the others. Wood influxes were much lower in total and relative

62

Table 18. Results of Kruskal-Wallis and Dunn's multiple comparison
tests for the combined trap totals.a

 

 

1976
Group ni Ri RiZ/ni 'RI Pooled'Rib
Smith 6 122 2480.67 20.3
43rd Street 6 78 1014.00 13.0 14.6
Kellogg Forest 6 63 661.50 10.5
Nagel ‘_§ '_31 228.17 6.2 6.2
24 300 4384.34

x2.01 (v=4-1=3) - 11.34 K912.69** (P<.01)

Group ._§mith __43rd Kel. For. __yagel _§901edb
Group 1 ‘81 j |Ri-20.3| |Ri—13.0| |Ri~10.5| 7Ri-6.27 |Ri-14.6|
Smith 20.3 0
43rd 13.0 7.3* 0
Kel. For. 10.5 9.8 2.5 0
Nagel 6.2 14.1* 6.8 4.3 o 8.4*

 

when ni=6, nj=6 ¢=.05=10.77 «=.10=9.77 when ni=18, nj=6 ¢=.05=6.53
Nagel<Smith (p<.05) Kel. For.< Smith (p<.10) Nagel<Pooled (p<.05)

 

 

 

1977
Group, ni Ri Rizlni ‘Ri. Pooled Rib
Smith 12 244 4181.33 18.7
43rd Street 6 116 2242.67 19.3 17.4
Kellogg Forest 6 85 1204.17 14.2
Nagel _§_ _49 266.67 6.7 6.7
30 465 7894.83
x2 05 (v=4-1-3) = 7.81 K=8.87* (p<.05)

.__ Group _§mith _43rd Kel. For. _§agel Fooledb
Group 1 R1 j IRi-18.7| lRi-19.3l lRi-14.2l lRi-6.7l lRi-17.4l
Smith 18.7 0
43rd 19.3 0.6 0
Kel. For. 14.2 4.5* 5.1* 0 *
Nagel 6.7 12.0 12.6 7.5 0 10.7

 

when ni=12, nj=6 «=.05=11.61 when ni=24, nj=6 ¢=.05=7.88
Nagel<Smith and 43rd Street (p<.05) Nagel<Pooled (p<.05)

 

aValues upon which rankings are based are in Table A8.

bPooled values represent combination of the Smith, 43rd Street and
Kellogg Forest sites.

*Significant at 5% level; **significant at 1% level.

63

 

 

. _ _ H _ _ _

_ _§§§ m
Iw.\\\\\\\\\\\. .N.
- 0000003 0
§§§ w.

 

 

 

       

 
 

 

a I E§ m
m mums - 0000000000.m0Em0000000000.§§m
_ _ _ _ 5 _ _ _ .

in z. .53 >05

_§n\\\\\\\\\\\\\\\\\\\\\\\\.s.

1977

19fl5

at the sites.

10. Annual CPOM totals by litter types

Figure

 

64

Litter Types

 

.. _ 0mmmm§\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\§ .N.
550.500n\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\§ a
0.50.000m.m...\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\x m

_§\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\ 0
m§.m.m.m.\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\§ .N.

 

 

 

m.03005030 5\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\x 0
00000000.000.0%:sugg_ususm w\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\ w.
_ _

nu :J nu
:J at

 

 

_
5
7]

._<._.O._. "—0 0x.

1976 1977
, with CPOM types expressed as a

cage of the total.

Figure 11. Annual CPOM influx
percen

65

amounts (6%) at Nagel. Leaf litter totals at Nagel were significantly
lower than Smith (p<.05) in 1976 and all other sites (p<.20) in 1977
(Table 19). In both years, leaf collections at Nagel were significantly
below (p<.05) the pooled estimate for the other locations.

Most allochthonous CPOM was transported to Augusta Creek by direct
infall, particularly at the undisturbed sites (72% - Table 11; Figure
12). While lateral transport exceeded direct infall at Nagel per meter
of stream bank (Table 11), total inputs on an areal basis (g/mz) were
dominated by direct infall (Table All). By either expression of the
data, however, lateral transport accounted for a higher percentage
of the CPOM total at the Nagel site than at the others. In contrast
to CPOM influxes from direct infall, lateral transport amounts were
similar among all sites (Tables 19 and A11).. These differences
suggested that the greatest impact from vegetational disturbance at
Nagel was a reduction in direct infall.

Autumn was the time of most (70%) litter influx to the stream
(Table 13; Figure 13), with this season's totals predominantly leaves
(range = 70-90%, Tables A2-A5). Due to leaf importance, the timing of
autumnal litterfall reflected the composition of riparian communities.
For example, CPOM influx at Nagel and Kellogg Forest was greatest in
September because the dominant ash and basswood leaves fell in that
month. In contrast, leaf-fall from nearstream woody species at the
other sites was more evenly divided between September and October.

During the rest of the year, CPOM influxes were low and quite
uniform (Table 13; Figure 13), although occassional severe storms
increased the variability. Intense low pressure fronts occur in this

region in winter and early spring (Eichenlaub, 1978), and associated

66

Table 19. Results of Kruskal-Wallis and Dunn's multiple cogparison
text for the combined trap totals (leaves only).

 

 

 

1976
Group, ni Ri RiZ/ni Ri Pooled Ri
Smith 6 114 2166.00 19.0
43rd Street 6 77 988.17 12.8 14.5
Kellogg Forest 6 70 816.67 11.7
Nagel _§_ ‘_39 253.50 6.5 6.5
24 300 4224.33
X2.05(v=4-1=3) = 7.81 Ks9.49* (p<.05)

___ Group l_§mith ‘_43rd Kel. For. _§agel Pooledb
Group 1 R1 1, IRi-19.0| |Ri-12.8| 781-11.7177Ri-6.57_7§I-14.57
Smith 19.0 0
43rd 12.8 6.2 0
Kel. For. 11.7 7.3 1.1 0 *
Nagel 6.5 12.5 6.2 5.2 0 8.0

 

when ni=6, nj=6 ¢=.05=10.77 when ni=18, nj=6 «=.05=6.53
Nagel<Smith (p<.05) Nageli<Pooled (p<.05)

 

 

 

 

1977
Group, ni Ri Rizni -Ri Pooled-RTb
Smith 12 210 3675.00 17.5
43rd Street 6 111 2053.50 18.5 17.6
Kellogg Forest 6 101 1700.17 16.8
Nagel ._§ 43 308.17 7.2 7.2
30 465 7736.83
x2 10(v=4-1=3)=6.25 Ks6.83 (p<.10)

___ Group Smith 43rd Kel. For. __§agel _Pooledb
Group 1 Ki 1 Iii-17.57 Iii-18.57 lRI-16.8| lRi-7.2 ”121-17.6,
Smith 17.5 0
43rd 18.5 1.0 O
Kel. For. 16.8 0.7* 1.7* 0*
Nagel 7.2 10.3 11.3 9.6 o 10.4*
when ni=12, nj=6 cc=.1o=1o.54 «=.15=9.64 s.zo=9.37

when ni=24, nj=6 ¢=.05=7.88
Nagel<43rd Street (p<.10) Nagel<Smith (p<.15)
Nagel<Kellogg Forest (p<.20) Nagel<Pooled (pf.05)

a
Values upon which rankings are based are in Table A8.

b
Pooled values represent combination of the Smith, 43rd Street and
Kellogg Forest sites.

*Significant at 5% level.

67

 

Trap Types
[:3 LATERAL TRANSPORT

80° " - INFALL

1“

C>

CD

CD
I

 

 

400 -

DRY WT. IN g/m

 

 

 

 

 

 

200 -

SM 43 NA KF SM 43 NA KF
1976 I977

   

1

 

     

Figure 12. Annual CPOM totals by sampler type at the sites.

 

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68

 

 

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68

 

   

 

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69

high winds, particularly with freezing rain, can cause high amounts of
litter to fall. Unusually large influxes of woody material occurred at
the Smith and Kellogg Forest sites after severe winds on November 10,
1975 and an ice storm on March 2, 1976 (Table A6). While the sampling
apparatus effectively retained much of this material, occassional trees
and large limbs which fell were not adequately sampled (cf. Christensen,
1975). Although litter movement to the stream was lowest in the winter,
due to the dormant state of the vegetation and snow cover, storm activity
served to increase allochthonous CPOM supply to the system.

Because of differences in nearstream plant associations, leaf
inputs were dissimilar. Most leaves of Smith and 43rd Street were of
the medium processing rate, while the "fast" leaves, ash, basswood and
dogwood, were dominant at Nagel and Kellogg Forest. (Tables 20 and A7;
Figure 14). Dogwoods were common at all four sites, particularly in
the O-lO-m portion of the riparian zone (Table 5-8). Ash and basswood
were most abundant along third-order reaches due to a wider floodplain
(Tables 7 and 8). The dominant "medium" leaf species were black cherry,
black willow, nannyberry, American elm and black walnut. Except for
black cherry, these are all floodplain species which had their greatest
coverage within 10 m of the channel (Tables 5-8). Slow leaves made up
a small portion of the leaf litter and were mostly upland species, red

oak, hawthorn and quaking aspen.

70

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I977

I976

sing category at the sites.

14. Annual leaf influx by proces

Figure

72

Literature Comparisons of Litterfall Estimates

 

Results from this study were expressed in g/m2 of channel surface
(Table 21 - Column 4), like most other investigations of particulate
organic influxes to running waters. With a continuous canopy, litterfall
levels over the stream and in the adjacent forest are similar (Fisher and
Likens, 1973; Dawson, 1976). Among Augusta Creek sites, only Nagel
lacked a complete canopy over the entire 30-m sampling section.
Therefore, the results from the direct infall samplers were extended
over the entire channel surface. CPOM inputs were a combination of
direct infall (Column 1) and lateral transport (Column 2) totals.

These particulate organic influxes are affected by channel width,
and generally decline as the channel widens (Cummins, 1975; Hynes, 1975).
This decrease in CPOM is a result of reduced canopy and lowered bank
length to stream bottom ratio (Meehan, et al., 1977). Highest litter
influxes (>500 g/mz) have been reported from small, first- or second-
order forested streams (Table 21). Results from the Smith site seem
representative of such systems. Among watercourses with 5-15 m widths
(Liston, 1972; Dawson, 1976; 'Post and DeLaCruz, 1977), CPOM inputs are
reduced, with 43rd Street and Kellogg Forest totals at the low end of
this range (Table 21). When available data (Table 21) on allochthonous
CPOM and mean channel width were ranked, a significant (p<.05)
negative correlation was found (Table A9).

While the importance of riparian vegetation to allochthonous CPOM
is generally recognized (Cummins, 1974), few investigations of litter
inputs to streams have included concurrent measurements of nearstream
woody cover. Dawson (1976) and McDowell and Fisher (1976) reported that

litter composition reflected the abundance of riparian species, but

73

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74

provided no numerical support for their observations. Liston (1972)
quantified the first 10 m of streamside vegetation and noted an apparent
correlation of leaf litter influx and basal area of woody species.

Along Augusta Creek, CPOM inputs showed a significant (p<.05) positive
correlation with stand density and mean shrub basal area (Table A9). In
addition, vegetation transects (Tables 5-8) indicated that the first

10 m of riparian cover contributed most CPOM to the aquatic system.

The significance of shrubs to stream CPOM inputs was denoted by
the afore mentioned correlations (shrubs made up most of the stand
density, Tables 5-8), and by the disproportionate share of leaf
contributions from shrubs (Tables 14-17) in respect to their basal area
in the stand (Table 9). In contrast to their role in providing bank
stabilization, shading and cover to small streams (Ware and Penfound,
1949; White and Brynildson, 1967; Karr and Schlosser, 1977), the
importance of shrubs to CPOM inputs has not been noted before.

Except where slope is severe, direct infall is usually the leading
source of allochthonous CPOM to headwater streams (MCDowell and Fisher,
1976; Winterbourn, 1976). This method of litter transport was dominant
at the Augusta Creek locations (Tables 21 and All). In most instances,
CPOM movement across the ground surface increases with greater slope
(Malmquist, et al., 1978). This observation was supported in this
study, since lateral transport totals were significantly correlated
(p<.05) with bank slope (Table A9). However, because slope was
moderate (<12Z, U.S.D.A., 1979), less than one-fourth of CPOM influxes
resulted from downslope movement (Table All). Only Sedell, et al.
(1974) have found higher litter contributions from lateral transport,

which seems to have resulted from extreme (>40Z) bank slope (McDowell

75

and Fisher, 1976).

Another aspect of allochthonous inputs involves seasonal changes.
Litterfall in temperate, deciduous forests is concentrated in the
autumn (Bray and Gorham, 1964; Gosz, et al., 1972; Grigal and
Gizzard, 1975). An average of 70% of annual CPOM inputs to Augusta
Creek were in this season (Table 13), which is similar to levels
reported for other streams in deciduous forests (Liston, 1972; Fisher
and Likens, 1973; Otto, 1975; Post and DeLaCruz, 1977). As generally
noted by these earlier studies, litter inputs were low, but rather
evenly distributed throughout the rest of the year (Table 13). The
rate of CPOM influx was lowest in the winter, which is characteristic
of north-temperate latitudes with pronounced snow cover (6032, et al.,
1972).

Among allochthonous CPOM types, leaf litter from riparian forest
species is usually the most abundant form. At the Augusta Creek
sampling areas, leaves averaged 70% of annual inputs (Table 12).

This patterns is consistent with the results of Fisher and Likens
(1973), Sedell, et a1. (1974), Otto (1975) and Winterbourn (1976),
whose values ranged from 50 to 90% 0f the yearly sum. Woody material,
which averaged 18% (Table 12), was the second largest category of CPOM.
Inputs of wood to forested headwaters have been shown to account for
about one-fifth (Fisher and Likens, 1973; Sedell, et al., 1974; Otto,
1975; Winterbourn, 1976) of yearly litter totals, although values up
to 70% have been reported (Anderson and Sedell, 1979). Other forms of
exogenous CPOM are usually of minor importance (Fisher and Likens,
1973), as was found in this study (Table 12).

Variations in leaf types entering headwaters, in response to

76

changes in riparian vegetation, have not received much prior attention.
Meehan, et a1. (1977) noted that along Oregon streams, "fast" leaves
tended to increase in proportion to "slow" leaves in a downstream
direction. A similar trend took place along Augusta Creek (Table 20).
Among the sampling locations, "slow" leaf totals and floodplain width
were negatively correlated (p<.05, Table A9). The general distribution
of forest types in this region supports this observation; uplands are
dominated by oak and hickory and floodplains by faster processed ash,
basswood and dogwood (Stearns and Kobriger, 1975). As floodplains
widen, a normal consequence is a great reduction in litter movement
from adjoining uplands to the stream (Table 20; Bell and Sipp, 1975).
Thus, leaf types may differ as a consequence of environmental change.

Allochthonous CPOM influxes to Augusta Creek were lower than most
comparable literature values (Table 21), possibly due to differences in
riparian vegetation. Forest stands along Augusta Creek had a high
presence of shrubs and low to moderate basal area (Table 10) while, as
judged by site descriptions, woody communities at the other locations
(Table 21) were more mature. Due to normal seral development (Bormann,
et al., 1970), CPOM inputs to Augusta Creek should increase as the
stands accrue biomass.

The effect of woody vegetation removal upon CPOM inputs is
demonstrated by the Nagel site results (Table 21), since litter totals
are much lower than all other locations. While such an impact from
alteration of nearstream vegetation has been suggested (Hynes, 1975),
it has not previously been quantified. Furthermore, the Nagel site has
experienced only selective cutting (Table 6). Extensive disturbances
like Clearcutting (cf. Likens, et al., 1977) should have a much more

severe effect on allochthonous CPOM.

77

Synthesis

Allochthonous CPOM Resources

 

Variations in allochthonous CPOM along headwater stream ecosystems
are important because of the significance of this material to the detrital
resource base. Just as lakes were once viewed as microcosms, lotic studies,
including most descriptions of allochthonous CPOM (Table 21), have often
focused on isolated reaches of running waters. Where CPOM influxes from an
entire catchment have been estimated, (Fisher and Likens, 1973), it was
assumed that CPOM was added at a constant rate along the channel. Results
from this study have demonstrated that litterfall will vary significantly
(p<.05) along a watercourse as a function of woody riparian cover (Table
A9), particularly in areas of vegetative disturbance (Table 21). CPOM
influxes also varied due to changes in channel width, narrower channels
had a more complete canopy and a larger bank-to-surface area ratio and
consequently, greater influx per channel area (Table 21).

Because of changes in channel width and vegetation, CPOM influxes
within the Augusta Creek catchment were unevenly distributed along the
stream (Table 22). If nearstream vegetation had been uniform, litter
influxes per unit of channel length would have been constant (Fisher
and Likens, 1973), and inversely proportional to channel width. While
the riparian zone was predominantly forested, two native associations were
present, shrub-carr and lowland forest, which differed in amount of litter
input (Table 22). Lowland forest along first- and second-order sections
made up 30% of the channel surface, yet accounted for one-half of the
CPOM total (Table 22),3 Including the shrub-carr community, first— and

3This result was not an artifact of forest distribution, since

lowland forest was as frequent along third-order reaches as along
smaller channels (50% of length, Table 4).

78

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79

second-order channels received 62% of the litterfall, while totalling

49% of the channel area.

Seasonal and annual allochthonous CPOM variations should also have
a critical influence on detrital levels within the stream system. In
both years of study, CPOM totals and types at a location were similar
(Tables 12 and 13). Even though most allochthonous CPOM entered the
stream in the autumn (70%), the remainder was quite evenly distributed
among the other seasons, affording a low, but continuous litter influx
at these times (Tables 13 and All). Such a litterfall pattern should
lend stability to the detritus base in Augusta Creek.

The major kinds of allochthonous CPOM play an important role in
detrital availability, since they range from rapidly degraded herbs and
fruits, to leaves of intermediate degradation rate, to refractory woody
material (Anderson and Cummins, 1979). Leaves and wood promote
continuity in the detrital resources of Augusta Creek, because they are
the dominant forms of external CPOM (70% and 23%, Table 23), and, because
they remain within the system for the longest time (Petersen and Cummins,
1974). Wood inputs were greatest along first- and second-order
tributaries, especially in sections of lowland forest, since 40% of the
catchment total came from these reaches (Table 23). In systems where it
is very abundant, wood may serve as an important source of dissolved
(DOM) and fine particulate organic material (FPOM) (Anderson and Sedell,
1979). However, the greatest significance of wood along Augusta Creek
may lie in its influence on the physical structure of the channel
(Likens, et al., 1978), and in.its retention of particulate organic
matter (POM), so that more complete biotic utilization of POM may

occur (Marzolf, 1978). The other litter forms, fruits and herbs, do

80

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Amaaa< .mN magma

81

not enter the stream in large amounts (Table 12). They are high quality
food resources (Meehan, et al., 1977), however, and may be important
nutritional sources for some detritivores.

Because of their differences in food quality (Hynes, 1975; Ward and
Cummins, 1979), changes in leaf influxes to Augusta Creek are noteworthy.
Variations in woody species occurrence (Tables 5-9) and life history
resulted in leaf litter differences. Diversity of leaf inputs was
greatest along first- and second-order tributaries, with species that
have medium rates of degradation accounting for 60% of the total and
"fast" and "slow" leaves 20% each (Table 23). In contrast, third-order
reaches received leaves of mostly "fast" (65%) and medium (26%) species.
Additionally, the autumn leaf-fall ended a month earlier in third-order
lowland forests. Since leaves are the most prevalent kind of
allochthonous CPOM, the variation in inputs to the stream should have
caused differences in food value of CPOM for detritivores.

The method and distance of CPOM transport to the stream are
critical in determining the area of undisturbed vegetation ("greenbelt")
which will insure normal litterfall to the aquatic system. The mode of
transport is of interest because it affects the condition of CPOM
entering the stream, since significant degradation can occur on the
floodplain if litter has a long residence time there (Merritt and Lawson,
1978). Direct infall was dominant at all sites throughout the year
(i = 78%, Table All), which indicated that CPOM entering the stream had
undergone little degredation in the floodplain. In conjunction with
litterfall estimates, streamside vegetation analyses showed that almost
all CPOM came from within 10 m of the channel. CPOM totals, types and

timing all reflected the composition of the woody riparian community.

82

Therefore, the quantity and quality of CPOM entering small running waters
is under control of the nearby terrestrial environment. These two areas
are functionally inseparable, and should be viewed and managed as a

single unit.

Other Estimates of Detrital Resources

 

One estimate of the elaboration of organic matter within the stream
is the FIR ratio, which involves an assessment of gross primary production
and community respiration (Odum, 1969). P/R values are from King
(1980),8 and disclose a great difference between the first-order section
(Smith) and the other sites (Table A12). In general, P/R increased with
stream order. Since CPOM influxes were reduced on an areal basis as the
channel widened and P/R increased, the community should show an
increased use of autochthonous production and a decreased dependence on
allochthonous sources with larger stream-order.

Another measure of detrital resources in Augusta Creek is the POM
standing crop. Average standing crop is similar between locations within
a habitat (pool or riffle) and does not change markedly over the year
(Table A14). These totals suggest an overall balance in storage,
processing, and import-export processes, which has been noted in other
headwater systems (Naiman and Sedell, 1979). A rather constant level of
benthic POM may act to buffer the community from the effects of seasonal
and annual differences in detrital inputs.

Seasonal and longitudinal changes in allochthonous CPOM may produce
changes in benthic POM size classes. CPOM standing crop was highest in
the winter and spring and lowest in the summer and fall (Table A14),

aInterpretation is difficult because the canopy was more open in

the area where P/R was estimated than in the litterfall sampling section
at 43rd Street and Kellogg Forest.

83

which corresponds with litterfall patterns. FPOM totals at the first—
order site are most reduced in the summer and fall, but show no seasonal
trend at the other sites. This difference could be a result of low
autochthonous production and FPOM export in the smallest tributaries. A
combination of pool and riffle totals shows an upstream increase in benthic
CPOM and a decrease in FPOM. Such a trend has been predicted in headwater

systems as a result of normal fluvial processes (Vannote, et al., 1980).

Biotic Responses to Changes in Allochthonous CPOM

 

Since allochthonous CPOM forms the base of the food web in forested,
headwater streams (Cummins, 1974; MEehan, et al., 1977), differences in
inputs should produce changes in detritivore populations and community
structure. Within communities, species tend to separate out along three
major niche dimensions, space, time and food (Pianka, 1974). As
riparian vegetation and channel width differ along Augusta Creek, so will
exogenous food resources available to the biota.

As judged by CPOM inputs (Table A11) and benthic CPOM (Table A14),
food resources for coarse particle feeders (shredders) appear to be
inversely correlated with stream width. Increased CPOM should result in
a greater density of shredders. While numbers of the dominant species
should increase, higher diversity may also occur, since each species
should use less of the total range of food (Ricklefs, 1979). More
diversity (specialization) could increase the efficiency of resource
utilization (Pianka, 1974), so the most efficient biotic processing of
CPOM may occur in the smallest tributaries. Shredder numbers are
highest at the Smith site (Table A13; Cummins, unpub. data), with a rich
array of functionally obligate forms like the cranefly, Tipula, and the

caddisfly, Lepidostoma. The seasonal aspect of CPOM influxes should also

 

84

be an important factor in shredder density differences along the creek.
Much higher litterfall occurred along first-order sections in winter and
spring (Table All). An extended period of CPOM availability may
encourage greater shredder diversity in these reaches by allowing more
separation of life cycles over time.

The greatest effect of CPOM (food supply) changes on shredder
populations should be found in areas where natural inputs have been
altered. The Nagel site not only had the lowest litter inputs (Table
21), it also had fewer log jams or other debris-retaining structures.
This last condition apparently resulted from riparian tree removal, since
the channel has not been disturbed. Debris dams are a major site of
shredder activity (Anderson and Sedell, 1979), so a paucity of these
structures should lower shredder populations. Less food will cause
emigration (drift-Walton, 1978), reduced growth and/or survivorship.

At the four locations, shredder numbers (spring and fall) and leaf
processing rates were lowest at the Nagel site (Cummins, unpub. data).
Reduced rates of leaf degredation, as a consequence of small shredder
numbers, have been reported in several other studies (Anderson and
Sedell, 1979).

Reduced CPOM at the Nagel site should particularly affect specialist
(obligate shredder) species, since faculative representatives would
utilize available CPOM, but switch to other foods when necessary (Cummins
and Klug, 1979). An increase in a generalist species in response to
reduced food (CPOM) levels has been noted in two streams with severe
vegetative disturbance in their catchments (Webster and Patten, 1979).

As Ricklefs (1979) has pointed out, one of the most consistent effects

of disturbances on community structure is to increase the dominance of a

85

few species.

The frequency of vegetationally altered sections and their
distribution will influence CPOM inputs and community structure. The
processing efficiency of shredders in such sections should decline,
because generalists sacrifice efficiency in the conversion of food to
growth for the ability to use a greater variety of resources (Pianka,
1974; Cummins and Klug, 1979). POM is sequentially reduced in size as
it passes downstream (Vannote, et al., 1980), however, the Nagel section
could be a break in this processing chain. In addition, reduced CPOM
inputs to these reaches lowers the supply of POM (mostly as FPOM) to
downstream communities. While increased autochthonous production in
disturbed areas may replace some of the lost allochthonous resources
(Gelroth and Marzolf, 1978), endogenous production supports different
functional groups (Cummins, 1974).

Shredder populations should reflect food quality differences
(dissimilar leaf influxes) which occurred along Augusta Creek. Food
quality may directly affect shredder growth rates, which should influence
life cycle length, size at maturity and fecundity (Anderson and Cummins,
1979). Due to early-dropped, mostly "fast" leaves (Tables 20 and A2-A5),
shredder growth in third-order sections should begin earlier, but be
compressed into a shorter time period. In contrast, the leaf input
pattern to first— and second-order channels should extend the period of
similar shredder diets (well-conditioned leaves). This should allow
increased temporal segregation, which is a common means of coexistence
among closely related stream invertebrates (Vannote and Sweeney, 1980).
The greater variety of leaf inputs to the smaller tributaries may also

increase species diversity, because it permits smaller niche breadth,

86

i.e., more specialization (Pianka, 1974). As mentioned earlier,
shredders are more numerous and exhibit greater species richness in
first- and second-order sections of the system.

One other influence on contemporary leaf influxes, particularly to
third-order channels, is Dutch elm disease. Before this fungal
introduction, American elm was a leading component of local lowland
forests, especially along streams with extensive floodplains (Curtis,
1959). In the Augusta Creek catchment, the species was most common
where the channel (and floodplain) were widest. Ash and basswood,
previous co—dominants, have replaced elms in the community (Thompson,
1972). Consequently, the diversity of allochthonous leaf litter has
declined with respect to stream detritivores, since a "medium" species
has been replaced by "fast" leaves. More extreme changes in food
quality, resulting from the conversion of deciduous forests to conifers,
have reduced detritivore diversity (Huet, cited in Hynes, 1970:231;
Wallace, et al., 1970). Because the range of food resources is
important to temporal separation of invertebrates within a functional
group (Vannote and Sweeney, 1980), shredder diversity in downstream
areas may have declined since the loss of elms.

CPOM influxes to Augusta Creek are reduced to FPOM by physical
abrasion and microbial and invertebrate processing. Fine particle
feeders (collectors) in small, woodland streams are dependent on such
processes for much of their food supply (Cummins, 1974). Because CPOM
inputs were reduced as the stream widened (Table 21), FPOM generation
and export from the smallest tributaries should be very important to
collectors in larger orders.

Some of the highest quality food resources available to collectors

87

are well-conditioned leaf fragments and invertebrate feces (Cummins and
Klug, 1979). Such high quality types of food may increase collector
growth rate (King, 1978), which could shorten the life cycle period (Ward
and Cummins, 1979). A number of the common species of collectors in
Augusta Creek have two generations per year, autumnal and vernal
(Cummins, 1974). While it is possible that increased levels of leaf
fragments and shredder feces (high inputs and shredder numbers) may

cause better growth in vernal generations in first- and second-order
sections, downstream transport of FPOM and greater autochthonous FPOM
may have a compensatory effect for third-order reaches.

The use of smaller-sized POM by filtering collectors as streams get
larger (Wallace, et al., 1977) may be related to allochthonous input
patterns and in-stream CPOM degradation. CPOM influxes and benthic
CPOM totals declined as Augusta Creek widened, while the benthic FPOM
sum increased and became smaller in size (Tables All and A14). POM
transport data from the stream indicates a downstream decrease in mean
particle size (Sedell, et al., 1979). Wallace, et al. (1977) and Alstad
(1980) have demonstrated mesh size increases in an upstream direction
in the most common family of net-spinning caddisflies, Hydropsychidae.
While they contend this is largely due to effects of the current, changes
in food particle size would offer an additional hypothesis.

Predators are another important functional group in headwaters.
Animals provide the highest quality food in streams (Anderson and Cummins,
1979), with production among lower trophic levels usually reflected in
predator production (warren, et al., 1964; Hynes, 1970). Levels of
non-predaceous invertebrates (Table A13) indicate that animal food

resources are greatest in the smallest orders. The higher number of

88

predators in an upstream direction may be a response to these conditions.
Since food availability is important to ecological relationships
among aquatic invertebrates (Cummins, 1973), variations in allochthonous
CPOM should influence community structure and function. CPOM influxes
(Table 21) and P/R values (King, 1980) from Augusta Creek suggest that
allochthonous detrital resources should decline and autochthonous
resources increase with larger stream-order due to increased channel
width. Results from this study also demonstrated that CPOM influxes
change within. stream-order (Tables 22 and 23). From my observations
on other streams, such heterogeneous allochthonous influxes should occur
along most small streams, since riparian communities normally are quite
variable. Some of the implications of these resources changes to the

biota, especially shredders, have been discussed.

In order for the factors which control community relationships to
be better understood, some quantitative measurements within streams seem
necessary. First of all, the arrayanuilevel of food resources should be
carefully estimated. While microbial conditioning rates and nutrient
levels have been determined for general types of detritus, these values
have not been closely related to detrital totals in the aquatic system.
Secondly, our understanding of how species use food resources is poor.
The functional groups concept (Cummins, 1973) has been useful in
identifying general roles of the biota in organic matter degredation,
however, many species are not confined to a single functional group
(Anderson and Sedell, 1979). Even within a functional group, our
information on resource partioning among species is limited to a few
organisms (e.g. Hydropsychidae - Wallace, et al., 1977; Alstad, 1980;

Limnephildae - Cummins, 1964; Mackay and Kalff, 1973). Until the

89

resource needs of species (or even genera - Wiggins and MacKay, 1978)
are identified, throughout their life cycles, our understanding of how
environmental factors affect the community species complex will be
severely limited. Headwater streams typically exhibit rich and diverse
invertebrate communities (Patrick, 1975). The natural variation in
allochthonous CPOM influxes may be an important agent in maintaining

this richness, both within and among streamrorders.

Allochthonous CPOM and the Riparian Zone

 

The stability of headwater stream ecosystems, which has been
defined as their "ability to withstand and recover from perturbation"
(Webster and Patten, 1979), is controlled by the terrestrial environment.
Although small streams do not have a high standing crop of living biomass,
they do have a large detritus standing crop which maintains organizational
integrity (Odum, 1969). Since endogenous primary production is low and
exogenous CPOM is high in forested streams, CPOM from nearstream
vegetation accounts for much of the detrital base (Vannote, et al.,
1980). Results from the Nagel site demonstrated that alteration of
riparian vegetation can significantly reduce CPOM inputs. Where large-
scale vegetation removal has occurred, in-stream detritus reservoirs have
greatly decreased (Webster and Patten, 1979). Recovery of such systems
depends on re-establishment of nearstream vegetation (Likens, et al.,
1977; Gurtz, et al., 1980). Thus, the presence of natural riparian
vegetation is critical if small streams are to exhibit the attributes of
stability, as defined above.

Nearstream disturbances are infrequent along Augusta Creek, so their

effect on allochthonous CPOM, available light and nutrient levels is

90

small. Consequently, biological features typical of forested head-
waters, such as low levels of primary producers and abundant CPOMEfeeding
invertebrates (Cummins, 1974), are found along most of the stream's
length.

Results from this study suggest that small watercourses in
multiple-land—use catchments should exhibit characteristic attributes
of woodland stream structure and function if natural riparian vegetation
is maintained. Since most drainage basins are under multiple-uses, the
management implications of this conclusion are important. While "the
valley rules the stream" (Hynes, 1975), the most significant portion of
the valley is the area next to the channel. Although a ZO-m corridor
along Augusta Creek was critical for CPOM influxes, this is not the
entire region required for protection of the aquatic environment.
Because of hydrologic (Maddock, 1978) and biotic interactions (Merrit
and Lawson, 1978), the entire floodplain should not be disturbed hi
order to insure system integrity. Where relief is higher than in this
basin, further consideration must be given to the amount of slope
(Trimble and Sartz, 1958; Karr and Schlosser, 1977). Within these
constraints, a "stream corridor" concept provides a prudent and

effective way to maintain and enhance our running water resources.

91

SUMMARY AND CONCLUSIONS

Significant differences in allochthonous CPOM influxes were found
along Augusta Creek. Litterfall variations in this study were correlated
with changes in nearstream.woody vegetation and channel width. Influxes
were greatest to a first-order reach in dense forest (647 g/m2) and
lowest in a section with selectively cut riparian vegetation (108 g/mz).
As channel width increased, litterfall declined on a surface area
basis, due to decreased bank to channel area ratio and reduced
vegetative canopy. While CPOM influx variations in multiple-land-use
catchments have received little previous attention, results from Augusta
Creek indicate that such differences characterize inputs to most small
lotic systems.

In respect to the drainage basin, CPOM influxes were unevenly
distributed among stream—orders. While first- and second—order sections
made up 49% of the stream's surface, they accounted for over 62% of the
litter entering Augusta Creek. This result suggests that the greatest
interface between aquatic and terrestrial ecosystems occurs in smallest
channels, with food resources for coarse particle feeders inversely
correlated with stream width.

Most CPOM was transported to the stream by direct infall (78%).

Low relief afforded relatively little lateral litter movement, however,
bank slope and lateral transport were correlated. The transport data

indicated that CPOM entering the aquatic system had undergone little

92

degredation in the riparian zone.

At each site, CPOM influxes were similar in totals and types in
both years of study. While autumn was the season of greatest litter
influx (70%), a low, but constant amount of litter entered the creek
during the rest of the year. Leaves were the dominant type of litterfall
(70%), with wood accounting for most of the remainder. This combination
of constant annual influxes, continual litterfall throughout the year,
and a variety of leaf types, should lend stability to the detrital
resource base of the ecosystem.

Vegetational changes along the stream resulted in influxes of
leaf litter with differential decomposition rates. In first- and
second-order tributaries, leaf inputs were mostly of species with medium
processing rates (60%), while "fast" leaves were dominant (65%) along
third-order channels. Although never abundant (<20%), "slow" leaves
decreased with greater stream size and were inversely correlated with
floodplain width. Furthermore, leaf-fall to the creek was finished
almost a month earlier along third-order sections. These data suggest
that allochthonous CPOM resources are more diverse, and may have an
extended period of availability to detritivores in smaller tributaries.

A comparison of vegetative transects and leaf influxes demonstrated
that almost all of the allochthonous CPOM originated from a 20-m
corridor of vegetation along the channel. Within this zone, nearstream
shrubs were particularly important litter sources, as indicated by the
significant correlations (p<.05) of shrub basal area and stand density
(mostly shrubs) to CPOM inputs. Protection of this corridor appears
critical if normal levels of allochthonous CPOM are to be maintained.

Even though the basin has been subjected to many different land

93

uses, especially agricultural, most of the riparian zone (>90%) appears
undisturbed. This natural "greenbelt" probably is one of the most
important factors in the maintenance of typical headwater stream

structure and function in Augusta Creek.

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APPENDIX

109

'Table A1. Woody vegetation along Augusta Creek.a

.Acer rubrum L. - red maple - Tb

Acer saccharinum L. - silver maple - T

Aoer saccharum Marsh - sugar maple - T

Betula lutea Michx. f. - yellow birch - T
Carpinus caroliniana Walt - musclewood - S

Carya ovata (Mill.) K. Koch. - shagbark hickory - T
Cornus amomum Mill. - silky dogwood - S

Cornus racemosa Lam. - gray dogwood - S

Cornus stolonifera Michx. - red osier dogwood - S
Corylus americana Walt. - hazel - S

Crataegus spp. - hawthorn - S

Fraxinus nigra Marsh. - black ash - T

Fraxinus pennsylvanica Marsh. - red ash - T
Hammamelis virginiana L. - witch hazel - S

Ilex verticillata (L.) Gray. - black alder - S
Juglans nigra L. - black walnut - T

Larix laricina (Du Roi) K. Koch. - tamarack - T
Lonicera xylosteum L. - European fly honeysuckle - S
Morus rubra L. - red mulberry - S

0st a virginiana (Mill.) K. Koch. - ironwood - T
Physocarpus opfiIifolius (L.) Maxim. - ninebark - S
Pinus strobus L. - white pine - T

Populus grandidentata Michx. - large-toothed aspen - T
Populus tremuloides’Michx. - quaking aspen - T
Prunus avrum L. - sweet cherry - T

Prunus serotina Ehrh. - black cherry - T

Prunus virginiana L. - choke-cherry - S

Pyrus coronaria L. - wild crab-apple - S

Pyrus malus L. - ap le - S

Quercus alBa L. - w ite oak - T

Quercus bicolor Wild - swamp white oak --T
Quercus borealis Michx. f. - northern red oak - T
Quercus veIutina Lam. - black oak - T

Rhamnus catharticus L. - buckthorn - S

'Rhus typhina L. - staghorn-sumac - S

Rhus vernix L. - poison-sumac - S

'RIBES americanum Mill. - wild black currant - S
Rohinia psuedoacacia L. - black locust - T

Rosa alustris Marsh. - swam -rose - S

SEIIX aIEa E. - white willowP- T

Salix discolor Mhhl. - pussy willow - S

Salix nigra L. - black willow - S

Sambucus canadensis L. - common elder - S

Tilia americana L. - basswood - T

Ulmus americana L. - American Elm - T

Ulmus rubra Mufil. - slippery elm - T

Viburnum lenta o L. - nann berr - S

VIEurnum o qus L. - high-gush Zranberry - S
Vitis aestIvaIIs Michx. - summer grape - S
zanthoxylum americanum Mill. - prickly ash - S

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

aAll species observed in the transects and/or the input traps -
scientific nomenclature of Gleason and Cronquist (1963).

bT-tree, S-shrub - by the criteria of Braun (1961), Gleason and
Cronquist (1963) and Harlow (1957).

110

 

 

Table A2. Total CPOM input by category and season at the Smith site.
1976 1977
a a
Season I L T I L T %
Lb 269.08 97.33 366.41 51.3 271.65 76.32 347.97 63.6
Annual W 272.53 47.75 320.28 44 8 56.96 27.08 84.04 15.4
Totals F 6.40 0.12 6.52 0.9 93.23 1.08 94.31 17.2
H 6.80 14.16 20.96 2 9 3.30 17.56 20.86 3.8
Total 554.81 159.36 714.17 99 9 425.14 122.04 547.18 100.0
L 0.63 4.62 5.25 4.4 4.14 6.44 10.58 50.5
Winter W 89.15 14.18 103.33 86.4 2.74 2 40 5.14 24 6
F 2.07 0.12 2.19 1.8 - - - -
H 5.14 3.64 8.78 7.3 1.73 3.48 5.21 24.9
Total 96.99 22.56 119.55 99.9 8.61 12.32 20.93 100.0
L 9.59 7.27 16.86 9.2 10.04 16.60 26.64 37.5
Spring W 144.88 16.45 161.34 88.4 24.04 12.04 36.08 50 8
F - _ _ - .. .. -
H 1.18 3.20 4.38 0.8 0.66 7.64 8.30 11.7
Total 155.64 26.92 182.58 100.0 34.74 36.28 71.02 100.0
L 18.39 3.49 21.88 58.4 22.46 1.60 24.06 62.0
Summer W 4.91 5 59 10.50 28.0 6.28 3.40 9.68 25.0
F 3.73 - 3.73 10.0 2.32 0.40 2.72 7.0
H - 1.35 1.35 3.6 0.01 2.32 2.33 6.0
Total 27.03 10.43 37.46 100.0 31.07 7.72 38.79 100.0
L 240.47 81.94 322.41 86.1 235.01 51.68 286.69 68.8
Fall W 33.59 11.53 45.12 12.0 23.90 9.24 33.14 8.0
F 0.60 - 0.60 0.2 90.91 0.68 91.59 22.0
H 0.48 5.97 6.45 1.7 0.90 4.12 5.02 1.2
Total 275.14 99.44 374.58 100.0 350.72 65.72 416.44 100.0

 

 

8I a direct infall in g/mz; L = lateral transport in g/m; T = total

g/m of bank.

bL-Leaves, W-Wood, F-Fruit, H-Herbaceous Material.

in

111

 

 

 

 

Table A3. Total CPOM input by category and season at the 43rd Street
site.
1976 1977
Season I L T8 % I L Ta %
Lb 164.78 94.08 258.86 69.9 229.94 124.68 354.62 72.8
Annual W 38.54 36.41 74.95 20.2 38.93 52.00 90.93 18.7
Totals F 4.83 11.14 15.97 4.3 9.65 9.20 18.85 3.9
H 6.03 14.38 20.41 5.5 3.54 19.48 23.02 4.7
Total 214.18 156.01 370.19 99.9 282.06 205.36 487.42 100.1
L 3.10 5.29 8.39 32.8 3.92 11.20 15.12 50.6
Winter W 3.41 6.71 10.12 39.6 6.74 1.56 8.30 27.8
F 1.48 0.25 1.73 6.8 - 1.00 1.00 3.3
H 2.28 3.06 5.34 20.9 1.47 4.00 5.47 18.3
Total 10.27 15.31 25.58 100.1 12.13 17.76 28.89 100.0
L 10.34 5.56 15.90 33.6 7.29 17.84 25.13 43.7
Spring W 14.44 12.65 27.09 57.2 8 45 14.96 23.41 40 7
F 0.51 0.26 0.77 1.6 - - - -
H 1.36 2.22 3.58 7.6 0.95 8.08 9.03 15.7
Total 26.65 20.69 47.34 100.0 16.69 40.88 57.57 100.0
L 22.95 6.96 29.91 61.2 31.20 9.56 40.76 55.5
Summer W 5.86 7.71 13.57 27.8 5.26 15.56 20.82 28.4
F 1.76 1.63 3.39 6.9 4.77 2.64 7.41 10.1
H 0.22 1.80 2.02 4.1 0.08 4.36 4.44 6.0
Total 30.79 18.10 48.89 100.0 41.31 32.12 73.43 100.0
L 128.39 76.27 204.66 82 4 187.53 86.08 273.61 83.8
Fall W 14.83 9.34 24.17 9. 7 18.48 19.92 38.40 11.8
F 1.08 9.00 10.08 4.1 4.88 5.56 10.44 3.2
H 2.17 7.30 9.47 3. 8 1.04 3.04 4.08 1.2
Total 146.47 101.91 248.38 100. 0 211.93 114.60 326.53 100.0
8I = direct infall in g/mz; L = lateral transport in g/m; T total in

g/m of bank.

bL-Leaves, W-Wood, F-Fruits, H-Herbaceous Material.

 

112

 

 

Table A4. Total CPOM input by category and season at the Nagel site.
1976 1977
Season I L Ta % I L Ta %
Lb 50.98 68.53 119.51 62 a 60.71 67.60 128.31 67.5
Annual W 5.43 11.77 17.20 9.0 5.87 4.48 10.35 5.4
Totals F 4.32 2.16 6.48 3 4 0.65 - 0.65 0.3
H 14.31 33.94 48.25 25.2 7.26 43.48 50.74 26.7
Total 75.04 116.40 191.44 100.0 74.49 115.56 190.05 99.9
L 2.21 3.53 5.74 21 1 1.60 3.08 4.68 33.5
Winter W 1.32 0.89 2.21 8 1 0 29 0.96 1.25 8 9
F 3.77 1.30 5.07 18 6 - - - -
H 4.16 10.05 14.21 52.2 1.82 6.24 8.06 57.6
Total 11.46 15.77 27.23 100 0 3.71 10.28 13.99 100.0
L 1.64 4.83 6.47 34.1 2.30 20.56 22.86 54.8
Spring W 2.51 2.25 4.76 25.1 1 90 1.36 3.26 7.8
F 0.06 0.86 0.92 4.8 - - - -
H 2.45 4.39 6.84 36.0 1.41 14.20 15.61 37.4
Total 6.66 12.33 18.99 100.0 5.61 36.12 41.73 100.0
L 2.43 3.56 5.99 34 7 6.78 1.36 8.14 43.6
Summer W 0.60 5.31 5.91 34.2 2.98 1.96 4.94 26.4
F 0.49 - 0.49 2.8 0.28 - 0.28 1.5
H 0.64 4.24 4.88 28 3 0.56 4.76 5.32 28.5
Total 4.16 13.11 17.27 100.0 10.60 8.08 18.68 100.0
L 44.70 56.61 101.31 79.2 50.03 42.60 92.63 80.1
Fall W 1.00 3.32 4.32 3.4 0.70 0.20 0.90 0.8
F - - - 0.37 - 0.37 0.3
H 7.06 15.26 22.32 17.4 3.47 18.28 21.75 18.8
Total 52.76 75.19 127.95 100.0 54.57 61.08 115.65 100.0

 

 

a

g/m of bank.

bL-Leaves, W-Wood, F-Fruits, H-Herbaceous Material.

I = direct infall in g/m2; L = lateral transport in g/m; T = total in

113

 

 

Table A5. Total CPOM input by category and season at the Kellogg
Forest site.
1976 1977
a a a
Season I L T % I L T A
Lb 212.74 44.91 257.65 79.0 267.42 64.72 332.14 82.0
Annual W 44.18 13.43 57.61 17.7 57.59 5.68 63.27 15.6
F 6.42 1.26 7.68 2.4 3.10 0.08 3.18 0.8
H 1.05 2.35 3.40 1.0 0.22 6.28 6.50 1.6
Total 264.39 61.95 326.34 100.1 328.33 76.76 405.09 100.0
L 1.85 1.76 3.61 20.3 5.62 7.00 12.62 75.
Winter W 4.10 2.28 6.38 35.8 1.94 1 04 2.98 17.7
F 5.77 1.26 7.03 39.5 - - - -
H 0.39 0.39 0.78 4.4 0.10 1.12 1.22 7.3
Total 12.11 5.69 17.80 100.0 7.66 9.16 16.82 100.0
L 2.34 0.68 3.02 10.6 5.89 5.12 11.01 56.0
Spring W 22.24 2.79 25.03 88.3 5 06 2 16 7.22 36 7
F _ - _ - _ _ _ _
H 0.17 0.14 0.31 1.1 0.10 1.32 1.42 7.2
Total 24.75 3.61 28.36 100.0 11.05 8.60 19.65 99.9
L 19.09 0.75 19.84 65.0 13.84 1.28 15.12 45.8
Summer W 4.44 4.53 8.97 29.4 14.35 0.28 14.63 44.3
F 0.36 - 0.36 1.2 1.14 0.08 1.22 3.7
H - 1.35 1.35 4.4 0.02 2.00 2.02 6.1
Total 23.89 6.63 30.52 100.0 29.35 3.64 32.99 99.9
L 189.46 41.72 231.18 92.6 242.07 51.32 293.39 87.4
Fall W 13.40 3.83 17.23 6.9 36.24 2.20 38.44 11.5
F 0.29 - 0.29 0.1 1.96 - 1.96 0.6
H 0.49 0.47 0.96 0.4 - 1.84 1.84 0.5
Total 203.64 46.02 249.66 100.0 280.27 55.36 335.63 100.0

 

 

aI = direct infall in g/mz; L = lateral tranSport in g/m; T = total in
g/m of bank.

bL-Leaves, W-Wood, F-Fruits, H-Herbaceous Material.

114

 

 

 

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115

Table A7. Leaf species collected in the litter traps arrayed by
processing types.a

 

Fast

Acer saccharum - sugar maple - S,Nb

Cornus spp. - dogwood - S,43,N,KF
Fraxinus spp. - ash - 43,N,KF

Robinia psuedoacacia - black locust - 43
Rosa palustris - swamp rose - 43,N,KF
Tilia americana - basswood - N,KF
Zanthoxylum americanum - 43

 

 

 

 

 

Medium

Acer rubrum - red maple - N

Acer saccharinum - silver maple

Carpinus caroliniana - musclewood - KF
Carya ovata - shagbark hickory
Hammamelis virginiana - witch hazel
Juglans nigra - black walnut - 8,43
Prunus serotina - black cherry - 3,43
Salix spp. - willow - S,43,N

Ulmus americana - American elm - S,43,KF
Viburnum lentago - nannyberry - S,43,N,KF
Miscellaneous Leaves - S,43,N,KF

Spring Fragments - S,43,N,KF

Physocarpus opulifolius - 43,N

Corylus americana - hazel - S,KF

 

 

 

 

 

 

 

 

 

 

 

Slow

Crataegus spp. - hawthorn - 43

Pinus strobus - white pine

Populus grandidentata - large-toothed aspen
Populus tremuloides - quaking aspen - S,N,KF
Quereus bicolor - swamp white oak - KF
Quereus borealis - northern red oak - S,43,N
Rhamnus catharticus - buckthorn - KF

Pyrus coronaria - wild crab-apple - 43

 

 

 

 

 

 

 

 

 

aProcessing types after Petersen and Cummins (1974) or K.W. Cummins
(pers. com.).

bStudy sites where more than 1 g/m of this species was collected.
S-Smith, 43-43rd Street, N-Nagel, KF-Kellogg Forest.

116

 

 

 

 

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117

 

 

 

 

 

Table A9. variables used in rank correlations and the correlation
results.
Smith 43rd Nagel Kel. For.

Variables 1976 1977 1976 1977 1976 1977 1976 1977
Total inputs (g/mzof bank) 714 547 370 487 191 190 326 405
Total inputs (g/m ) 733 561 258 340 108 108 279 348
Lateral transport (g/m) 159 122 156 205 116 115 62 77
Total leaves (g/m of bank) 366 348 259 355 120 128 258 332
"Slow" leaves (%) 13.8 17.2 15.3 16.2 6.1 14.7 .1.1 1.0
Bank slope (%) 6.0 7.0 2.5 1.5
Mean channel width (m) 1.9 7.1 7.0 8.3
Mean floodplain width (m) 12 25 75 100
Mean basal area (m2/ha) 17.2 11.7 6.5 30.2
Mean stand density 7617 5182 1467 4329
Mean shrub b. area (m2/ha) 4.4 3.4 1.5 2.3

Correlation Resultsa

Variables n N
Total inputs (g/m) vs. mean stand basal area 8 20
Total inputs g/m2) vs mean stand basal area 8 26
Total inputs (g/m) vs mean shrub basal area 8 44**
Total inputs (g/m ) vs mean shrub basal area 8 38*
Total inputs (g/mg vs mean stand density 8 44**
Total inputs (g/m ) vs mean stand density 8 38*
Total leaves vs mean stand basal area 8 16
Total leaves vs mean shrub basal area 8 40*
Total leaves vs mean stand density 8 36*
Lateral transport vs. bank slope 8 42*
"Slow" leaves vs mean floodplain width 8 -36*
Total inputs (g/mz) vs. mean channel width 10 -62**
from Table 19
Critical values of N n m=.05 a=.01
N=4 3 01-6 (n-l) 8 36 44
=variable pairs 10 46 58

 

aKendall's rank correlation test as described in Sokal and Rohlf (1969).

The Nagel site was excluded due to vegetation disturbance.

The results

of Post and delaCruz (1977) were not included because mean channel

width could not be determined.

*Significant at 5% level; **significant at 1% level.

 

118

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Table A10. Total fruit collections at the sites by species and trap
type.
1976 1977
Site Species I L Ta I L Ta
Cornus spp. 3.28 - 3.28 1.49 - 1.49
Viburnum lentago 1.04 - 1.04 0.18 - 0.18
Smith Rosa palustris 1.66 - 1.66 - - -
Juglans nigra 0.38 - 0.38 91.42 1.08 1.08
Miscellaneous 0.06 0.12 0.18 0.14 - 0.14
Total 6.40 0.12 6.52 93.23 1.08 94.31
Crataegus spp. 0.88 10.52 11.40 - 7.30 7.30
Viburnum lentago - - - 1.28 - 1.28
43rd Cornus spp. 0.05 - 0.05 1.46 - 1.46
Prunus serotina 1.96 0.20 2.16 4.91 1.90 6.81
Rosa palustris 1.57 0.42 1.99 2.00 - 2.00
Miscellaneous 0.37 - 0.37 - - -
Total 4.83 11.14 15.97 9.65 9.20 18.85
Rosa palustris 0.57 - - - - -
Nagel Tilia americana 3.10 2.16 5.26 0 31 - 0.31
Cornus spp. 0.65 - - - - -
Viburnum lentago - - - 0.34 - 0.34
Total 4.32 2.16 6.48 0.65 - 0.65
Fraxinus spp. 4.20 1.26 5.46 1.66 O 08 1.74
K. For. Tilia americana 0.64 - 0.64 - - -
Cornus spp. 0.38 - 0.38 0.53 - 0.53
Viburnum lentago 1.20 - 1.20 0.91 - 0.91
Total 6.42 1.26 7.68 3.10 0.08 3.18

 

 

a

g/m of bank.

I = direct infall in g/mz. L = lateral transport in g/m. T = total in

119

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Table A12. Seasonal production/respiration ratios (P/R) from selected
riffle sections of Augusta Creek, 1974-1975 (King, 1980).

 

Season Smith 43rd Street Nagel Kellogg Forest
Winter 0.23 i .013 0.97 i .12 0.96b 1.62b
Spring 0.39 i .35 1.76 i .46 1.70 i .46 2.29 + .82
Summer 0.22 i .05 1.23 i .22 1.16 i .19 1.25b
Fall 0.23 i .09 1.04 i .25 2.00 i .32 2.02 + .34

 

a
Values are means of 3 replicates :_1 standard deviation.

bTwo replicates.

Table A13. Riffle invertebrate community functional group analysis at
the Augusta Creek sites from the River Continuum Project,
Summer, 1976 (Vannote, Cummins, Minshall, Sedell).

 

Functional
Group Smith 43rd Street Kellogg Forest
# Shredders/m2 1600 43 86
Z 6.7 0.4 1.1
# Collectors/m2 21306 9207 7740
% 88.7 95.1 96.0
# Scrapers/m2 986 367 151
X 4.1 3.8 1.9
# Predators/m2 132 65 86
X 0.6 0.7 1.1
# Detritivores/mz 22906 9250 7826
X 95.4 95.6 97.1

Total 24024 9682 8063

 

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