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THE ROLE OF EMERGENT AQUATIC VEGETATION IN THE
NUTRIENT BUDGETS OF MANAGED MARSHES

BY

Karl Eugene Ulrich

A DISSERTATION

Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of

DOCTOR OF PHILOSOPHY

Department of Zoology

198k

ABSTRACT

THE ROLE OF EMERGENT AQUATIC VEGETATION IN THE NUTRIENT
BUDGETS OF MANAGED MARSHES

BY

Karl Eugene Ulrich

Wetlands have been used as sites for the disposal of
nutrient-enriched waters. The emergent vegetation is
important in the nutrient budgets and hydrology of these
systems. However, little is known about how most emergent
plant species respond to variations in the absolute
quantities and proportions of substrate macronutrients. More
infdrmation is also needed on the establishment and
harvesting of the vegetation.

Nutrient uptake experiments were conducted on Phragmites

 

australis, Zizania aquatica, Typha latifolia, Typha

angustifolia, and Sparganium eurycarpum. Plants were grown

 

outdoors in a sand substrate in 30 l polyethylene tubs during
the 1980-82 growing seasons. Nitrate, phosphate and
potassium fertilizers were supplied using a factorial
treatment arrangement. Nitrate-N levels were 0, 7.8, 23.h,
46.8 or 93.6 g m-Z; phosphate-P was 0, 2.1, 6.3, 12.5 or 25.0
g m-Z; K was 0, 7.8, 23A or h1.6 g m—Z.
Nitrate and phosphate treatments and their interaction
strongly affected plant growth. Potassium treatment had
little or no effect. Nitrate strongly controlled
distribution of dry weight. The ratio of belowground to

aboveground dry weight within a species varied by a factor of

about 3 with highest ratios associated with low nitrate.
Tissue content of N or P was increased 2 to 3 times over that
necessary to obtain maximum biomass production. N:P ratios
less than about A were found with severely N limited growth:
ratios were greater than about 12 with severely P limited

growth. '§L eurycarpum was least able to tolerate extremely

 

high nitrate levels.

Attempts to establish the above 5 species plus Spartina

 

pectinata in a clay-lined man-made marsh had mixed success.

 

s; eurycarpum rapidly established itself but was subject to

 

decline and invasion by other species. Typha spp.
established stands more slowly but maintained better stand

density and purity. P; australis sent out mainly floating

 

tillers from propagule clumps rather than rhizomes and

vertical shoots. §4 pectinata and g; aguatica failed to

 

establish.

Multiple harvests of gypha spp. resulted in biomass
removal of 130% of single harvest controls and N and P
removal of about 150% of the controls.

Nutrient budgets develOped for three man—made
marsh-ponds showed a net removal of fixed N from flow-through

waters but a variable situation with total P.

ACKNOWLEDGEMENT S

My appreciation is extended to those persons and
organizations whose assistance and guidance have made this
dissertation possible.

This research was supported by a grant to Dr. Thomas
Burton from the the Office of Water Research and Technology
(Project # B-O55-MICH) of the U.S. Department of the
Interior. Additional funds were provided by a grant to Dr.
Burton from the Michigan Sea Grant Program (Project R/Cw-S)
of NOAA, U.S. Department of Commerce. I especially wish to
thank Dr. Burton for his support and guidance on this
project. I also wish to thank the other members of my
guidance committee for their time and efforts in guiding this
project. These persons are Dr. Stephen Stephenson, Dr.
William Cooper and Dr. C.D. McNabb.

My thanks are also extended to Cathy Sleight who
provided extensive technical assistance and to Judy Ulrich

who provided extra help when it was most needed.

ii

TABLE OF CONTENT S

 

 

3352

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

MATERIALS AND METHODS . . . . . . . . . . . . . . . 8
NUTRIENT UPTAKE EXPERIMENTS . . . . . . . . . . 8
ESTABLISHMENT AND HARVEST EXPERIMENTS . . . . . 1“
MARSH NUTRIENT BUDGETS . . . . . . . . . . . . 18

RESULTS AND DISCUSSION . . . . . . . . . . . . . . . 19

 

 

 

 

NUTRIENT UPTAKE EXPERIMENTS . . . . . . . . . . 19
Phragmites australis . . . . . . . . . . . 19
ZiEanié aguatiEa . . . . . . . . . . . . . 32
TYBha spp. O O O O O O. O O O O O I O O O O 32
Sparganium eurycarpum . . . . . . . . . . 50
General Discussion . . . . . . . . . . . . 55

ESTABLISHMENT EXPERIMENTS . . . . . . . . . . 63

HARVEST EXPERIMENTS . . . . . . . . . . . . . 68

MARSH NUTRIENT BUDGETS . . . . . . . . . . . . 72

CONCLUSIONS . . . . . . . . . . . . . . . . . 82

LIST OF REFERENCES . . . . . . . . . . . . . . . . . 89

 

APPENDIX 0 o o o o o o o o o o o o o o o o o o o o o 100

iii

LIST OF TABLES

Table 1. Analysis of Variance on the effects of
treatments and their interactions on estimates of
P; australis cumulative shoot dry mass after 108
days growth (n=246).

 

Table 2. Results of analysis of variance on the
effects of treatments on the ratio of 3; australis
root + rhizome dry weight to shoot dry weight.
Based on weights of materials harvested at end of
experiments and arcsin transformed prior to
statistical analysis.

 

Table 3. Results of analyses of variance on the
effects of treatments on %N, % , and %K of E;
australis shoots and roots plus rhizomes. Based on
weights of materials harvested at end of
experiments.

Table A. Results of analyses of variance on the
effects of treatments on harvest weight, %N, %P,
and %K of g; aquatica shoots (n=66,30,29,30

respectively).

 

Table 5. Effects of variable nitrate, phosphate,
and potassium treatments on Z. aquatica shoot dry
weight. root dry weight and fISsue content of N, P
and K. Weight values are in g per plant;
percentges are as per cent of 60 C dry weight.

 

Table 6a. Analysis of variance on the effects of
treatments and their interactions on.TL
angustifolia cumulative above round dry mass at
the end of—Ehe growing season %n=17u).

Table 6b. Analysis of variance on the effects of
treatments and their interactions on.TL
angustifolia cumulative below round dry mass at
the end of the growing season %n=174). .

Table 7a. Analysis of variance on the effects of
treatments and their interactions on T; latifolia
cumulative above ground dry mass at the end of the
growing season (n=173).

 

I iv

Page

20

25

26

34

35

37

37

38

Page
Table 7b. Analysis of variance on the effects of

treatments and their interactions on 2; latifolia
cumulative below ground dry mass at the end of the
growing season (n=173). 38

 

Table 8a. Results of analysis of variance on the

effects of treatments on the ratio of T;

angustifolia root + rhizome dry weight to shoot dry
weight. Based on weights of materials harvested at

end of 1981 experiments and arcsin transformed

prior to statistical analysis. “0

 

Table 8b. Results of analysis of variance on the

effects of treatments on the ratio of 2; latifolia

root + rhizome dry weight to shoot dry weight.

Based on weights of materials harvested at end of

1981 experiments and arcsin transformed prior to
statistical analysis. 40

 

Table 9a. Analysis of variance on the effects of
treatments and their interactions on S; eurycarpum
cumulative above ground dry mass at the end of the

growing season (n=1hh). 53

 

Table 9b. Analysis of variance on the effects of
treatments and their interactions on S; eurycarpum
cumulative below ground dry mass at the end of the

growing season (n=6A). 53

 

Table 10. 1982 3; australis aboveground and

belowground dry weights, %N and %P with various
combinations of nitrate-N and phosphate-P

treatments. Standard errors are in parentheses. 64

 

Table 11. 1982 TL_latifolia aboveground and

belowground dry weights, %N and %P with various
combinations of nitrate-N and phosphate-P

treatments. Standard errors are in parentheses. 65

Table 12. 1982 3; angustifolia aboveground and
belowground dry weights. %N and %P with various
combinations of nitrate-N and phosphate-P

treatments. Standard errors are in parentheses. 66

Table 13. Effects of harvest treatments on shoot

weight and nutrient content of Typha spp. All

units g m-2. * = material removed from plot; other

values are shoot standing crOps. 69

Table 1A. Results of harvest experiments on

vegetation established in M2. Dry weights are in

kg ha-l. * = material removed from plot; other

values are standing crops. 71

Table 15. Input/output balance of water and

water-borne chemical constituents of M1 during

1980-82. Water is reported as 10 6 1; all other

units are kg. 73

Table 16. Input/output balance of water and

water-borne chemical constituents of M2 during

1980-82. Water is reported as 10 6 1; all other

units are kg. 76

Table 17. Input/output balance of water and

water-borne chemical constituents of M3 during

1980-82. Water is reported as 10 6 1; all other

units are kg. 79

Table A1. 1980 P. australis aboveground dry weight
(g m- -2) estimates from regressions. Number of
replicates and standard errors are in parentheses. 100

Table A2. 1980 Z. aquatica aboveground dry weight

(g per plant) estimates from regressions. Number

of replicates and standard errors are in

parentheses. 10A

 

Table A3.1981 T. angustifolia aboveground dry

weight (g m-2) estimates from regressions. Numbers

in parentheses represent number of replicates;

standard error of the mean; average number of

shoots per m2; and standard error of the mean for

the number of shoots, respectively. 106

Table AA.1981 T. latifolia aboveground dry weight

(g m- -2) estimates from regressions. Numbers in
parentheses represent number of replicates;

standard error of the mean; average number of

shoots per m2; and standard error of the mean for

the number of shoots. respectively. 108

Table A5. 1982 P. australis aboveground dry weight

(g m- -2) estimates from regressions. Numbers in
parentheses represent number of replicates;

standard error of the mean; average number of

shoots per m2; and standard error of the mean for

the number of shoots, respectively. 110

vi

Table A6. 1982.3; angustifolia aboveground dry

weight (g m-2) estimates from regressions. Numbers

in parentheses represent number of replicates;

standard error of the mean; average number of

shoots per m2; and standard error of the mean for

the number of shoots. respectively. 111

 

Table A7. 1982 $1 latifolia aboveground dry weight

(g m-2) estimates from regressions. Numbers in
parentheses represent number of replicates;

standard error of the mean; average number of

shoots per m2; and standard error of the mean for

the number of shoots, respectively. 112

 

Table A8, 1982 S; eurycarpum aboveground dry

weight (g m-2) estimates from regressions. Numbers

in parentheses represent number of replicates;

standard error of the mean; average number of

shoots per m2; and standard error of the mean for

the number of shoots. respectively. 113

 

vii

LIST OF FIGURES

Figure 1. Average areal dry weight (60 C) of P.
australis shoots after 106 days growth during 1980
with variable combinations of nitrate and phosphate
fertilization. Vertical lines are standard errors
of the means.

Figure 2. Total areal seasonal belowground
production of P. australis as a function of nitrate
or phosphate treatment level during 1980. Vertical
lines are standard errors of the means.

 

Figure 3. Relationship at end of 1980 growing
season between nitrate or phosphate treatment level
and distribution of production aboveground and
belowground by P. australis. Vertical lines are
standard errors of the means.

 

Figure 4. P. australis shoot concentration of N as
per cent of— dry weight (60 C) at end of 1980
growing season for different combinations of
nitrate and phosphate fertilization. Vertical
lines are standard errors of the means.

 

Figure 5. P. australis shoot concentration of P as
per cent of_ dry weight (60 C) at end of 1980
growing season for different combinations of
nitrate and phosphate fertilization. Vertical
lines are standard errors of the means.

Figure 6. Seasonal areal uptake of N by L.
australis shoots under variable combinations of
nitrate and phosphate fertilization during 1980.

 

Figure 7. Seasonal areal uptake of P by L.
australis shoots under variable combinations of
nitrate and phosphate fertilization during 1980.

Figure 8. Comparison of the average 1980 N, P, and
K content of P;_australis shoots with that of roots
plus rhizomes. Vertical lines are standard errors
of the means.

 

viii

21

23

24

27

28

30

31

33

Figure 9. Average areal dry weight (60 C) of T.
latifolia and T. angustifolia aboveground and
belowground tissues in early September, 1981 with
variable combinations of nitrate and phosphate
fertilization.

 

 

Figure 10. Average aboveground to belowground
tissue dry weight ratios in early September, 1981
for T. latifolia and L. angustifoia at various
levels of nitrate treatment.

 

Figure 11. Average aboveground and belowground
tissue dry weights in early September, 1981 for 2;
latifolia and 24_angustifolia at various levels of
fiitrate treatment.

 

Figure 12. Seasonal accumulation of aboveground
dry weight by L. latifolia and L. angustifolia
during 1981 tub experiments.

Figure 13. Aboveground and belowground N content
as a function of dry weight for T. latifolia and L.
angustifolia tissues harvested during early
September, 1981 from tub experiments.

 

Figure 14. Aboveground and belowground P content
as a percentage of dry weight for L. latifolia and
L. angustifolia tissues harvested during early
September, 198i from tub experiments.

 

Figure 15. Areal content of N and P of L.
latifolia and L. angustifolia aboveground tissues
during early September, I981 with variable nitrate
and phosphate treatments.

Figure 16. Areal content of N and P of L.
latifolia and L. angustifolia during early
September, 1981 with variable nitrate and phosphate
treatments.

 

Figure 17. Average areal dry weight (60 C) of S.
eurycarpum aboveground and belowground tissues 1n
early September, 1982 with variable nitrate and
phosphate treatments.

Figure 18. Aboveground and belowground N content

as a percentage of dry weight for S. eurycarpum
tissues harvested during early September, 1982.

ix

39

41

43

44

45

46

47

48

52

54

Page
Figure 19. Aboveground and belowground P content

as a percentage of dry weight for SL eurygarpum
tissues harvested during early September, 1982. 56

Figure 20. Early September, 1982 areal content of

N in L. eurycarpum aboveground and belowground

tissues w1th variable nitrate and phosphate

treatments. 57

 

Figure 21. Early September, 1982 areal content of

P in L. eurycarpum aboveground and belowground

tissues with variable nitrate and phosphate

treatments. 58

Figure 22. Variation in the ratio of %N to %P in

§L eurycarpum aboveground and belowground tissues

durifig early September, 1982 as a function of

nitrate and phosphate treatment. 59

 

INTRODUCTION

Emergent aquatic vegetation has potential for nutrient
and other contaminant removal from sewage-enriched
substrates (Wile et al..1981). In addition, wetland
systems can have other beneficial effects on water quality
and hydrology. They may serve as retention structures that
detain water during flood periods and reduce peak flows of
streams. During storage, substantial improvements in water
quality may occur. Certain nutrients such as phosphorus
may be sorbed to or precipitated into the sediments. Other
nutrients such as nitrogen. sulfur, and carbon may be lost
to the atmosphere as gases due to microbial transformations
which also reduce the oxygen demand of the water. A large
prOportion of the sediment load may be dumped into the
marsh as the water velocity declines. Tchobanoglous and
Culp (1980) found that natural wetlands used for treatment
of wastewater removed 60% to 90% of the suspended solids.
This sediment load may include adsorbed P, heavy metals,
and organic toxicants. Wetlands may also act to reduce
pathogens (Seidel. 1976a, 1976b; de Jong. 1976; Wellings.
1976; Wile ggugi, 1981) Because Of these properties. both
natural and artificial marshes have been used for a variety
of purposes such as treatment of urban stormwater runoff
(e.g. Hickock, 1978; Tolman. 1978), treatment of municipal
and industrial wastewater (e.g. Grant and Patrick, 1970;
Lee g£‘§;., 1976; Steward and Ornes. 1975; Ewel. 1976; de
'Jong, 1976; Farnham and Boelter, 1976; Richardson gglg;..

2
1976; Small, 1976; Seidel. 1976a. 1976b; Spengler 23 al.

1976a. 1976b; Whigham and Simpson, 1976a, 1976b; Turner

In In

g;., 1976; Cederquist. 1977; Fetter 33Ԥ;., 1978; Sloey
g;.. 1978; Wile gg‘gi, 1981), pretreatment of drinking
water (Czerwenka and Seidel. 1976), and for management of
dredge slurries (Lee g§.§;., 1976).

Since emergent plants represent a harvestable and
potentially valuable biomass resource (Pratt and Andrews,
1981 Farnham. 1978) as well as a potential means of
removing nutrients from a system. information is needed on
their growth and nutrient uptake over wide ranges of
nutrient supply. The nutrient uptake characteristics are
most critical for nutrients such as phosphorus and
potassium which have no volatile forms and therefore cannot
be removed from a system in the gaseous state. Although
80-90% of the phosphorus detained by a marsh is usually
trapped in the sediments or stored in belowground plant
tissues and not easily removeable by harvest (Steward,
1976: Turner 33 §;., 1976; Sloey 23 §;.. 1976.I1978),
harvest may nevertheless be desireable if uses exist for
the biomass. Marshes have natural rates of primary
production that are among the highest of all ecosystems
(Westlake. 1963; Whittaker and Likens. 1973). Some uses
for the biomass as a source of fiber or energy have already
been established (Bjork. 1967; Pratt and Andrews. 1981).
However. there is a need for more information concerning
the Optimal means of establishment. nutrient application.

and best harvesting. It may be valuable to study*

3

interspecific differences in nutrient utilization since it
has been shown by Garten (1978) that certain species of
emergent macrophytes are quite distinct in their leaf
mineral compositions. These differences may be of
practical importance in designing artificial wetland
systems or in planning the utilization of natural systems
for the improvement of water quality.

The focus of this dissertation research was to gather
the necessary data for the culture of emergent marsh
vegetation with the primary goal of removing unwanted
macronutrients from artificially enriched systems. Major
hypotheses that were tested included the hypothesis that
nutrient treatments affect both net primary production and
distribution of this production. Also. the hypothesis that
nutrients would be taken up in excess of needs for maximum
growth was examined. To accomplish these goals, a number
of common. highly productive emergent plant species that
showed promise as species for cultivation in highly
enriched waters were selected. The species most thoroughly
studied prior to this investigation included several
species of bulrush (Scirpus). Many species of Scirpus have
the advantages of high productivity and nutrient uptake.
They also are receptive to frequent harvesting with rapid
shoot regrowth from the cut stubs so that stands can be
maintained with young nutrient-rich tissues (Hanseter.
1975). S; lacustris has an extensive literature and
history of culture in EurOpe (e.g. Seidel, 1976; de Jong.

1976). In Wisconsin, g; validus, §L acutus. and §;

L1

fluviatilis have been examined in some detail (e.g.
Spengler 333;... 1976a, 1976b; Sloey_e_t_a_l.. 1978). _S_._
robustus has been investigated in California (Cederquist,
1977). The present study was directed towards other
promising taxa, most of which have had more limited
research into their nutrient removal capabilities.

Phragmites australis (Cav.) Trin. ex Steudel, appeared
to be an ideal species for this purpose because of its
cosmOpolitan distribution, and its capacity for high rates
of biomass production (Bjork, 1967). Primary production
and nutrient uptake in natural 3; australis (= E; communis)
stands (Mason and Bryant. 1975; Dykyjova, 1978) and the
effects of these stands on sediment and water chemistry
(Bayly and O'Neill. 1972; Banoub, 1975) have been
extensively examined. Changes in nutrient content with
physiological age and differences between tissue types have
also been studied (Kvet. 1973). Dykyjova 22 g; (1971;
1972) grew Ea australis in outdoor hydroponic culture using
a standard culture nutrient solution with a variety of
dilutions. However. they did not vary the ratios of any of
the supplied nutrients. Stands apparently recover well
after harvesting and may even show an increase in the size
and density of new shoots (Dykyjova and Husak, 1973).
However, 2;_australis does not respond well to multiple
harvests within a single growing season.

Wild rice (Zizania aquatica L.) was selected because
it was one of the few marsh grasses with established

economic uses as a crOp species and also because of its

5
value as a wildlife food (Fasset. 1957; Correll and
Correll. 1972). Because it is an annual, most of its
production remains aboveground where harvest should be
relatively easy. Peak biomass can reach over 2000 g m-2
and averages about 1200 g dry weight In"2 (Whigham and
Simpson., 1977). Whigham and Simpson (1976a) conducted
some experiments on the ability of this species to accept
municipal wastewater irrigation. Lee and Stewart (1981.
1983) have looked for correlations between sediment and
plant tissue nutrients in natural;L aquatica stands.
There has also been considerable research done on g;
aguatica at the University of Minnesota (e.g. Grava and
Raisanen, 1978). However, their research has focused on
maximizing grain production and control of disease rather
than whole plant production and nutrient uptake. There
.exists limited data on nutrient uptake variability and
growth patterns over widely variable nutrient application
rates. One major difficulty with the use of wild rice is
its high susceptibility to disease. especially
figlminthosporium blight (Kernkamp gg‘gi, 1976).

Species of cattail (Typgg) have received some
attention for use in wastewater treatment. Spangler gg‘gl
(1976b) investigated 2; latifolia L. growth on artificial
substrates but rejected its use primarily on the basis of
its regrowth characteristics. Cut shoots (mostly leaves)
showed very limited regrowth because they generally lacked
active intercalary meristems at the time of cutting. Most

' new growth came from shoots develOping from underground

6
buds which surfaced several weeks after the harvests. They
were unable to sustain a high biomass yield with biweekly
harvests. Hanseter (1975) also found that (in gypgg spp.
stands) shoot density and size decreased after harvest.
However, even with their limited capacity for rapid
regrowth after harvest, gypgg spp. warrant more attention
because of their large peak standing crops and ability to
maintain nearly pure stands. Two temperate circumboreal
species of cattail (g; latifolia and g; angustifolia L.)
have overlapping ranges which includes the northeastern
one-third of the U.S. Ecological differences between 2;
latifolia L. and 2; angustifolia L. have been documented
(Grace and Wetzel, 1981,1982; McNaughton, 1975). It is of
interest to know if these differences extend to factors of
importance in the culture of these species for wastewater
renovation. That is, what are the differences between
these two species in nutrient uptake, tissue production,
distribution of production, ease of establishment, and
regrowth after harvest. Even if Eypgg dominated marshes
are not appropriate for multiple harvest systems to
maximize P removal, they may be valuable for their N
stripping abilities. For example, a Eypgg dominated
Wisconsin marsh (Spangler g§.g;, 1976b) reduced ammonium-N

1 in the influent to below levels of

values from 8 mg N 1-
detection in the effluent and reduced nitrate-N values from
0.2 to 2.1 mg 1'1 in the influent to 0.1 to 0.2 mg 1'1 in
the effluent (Sloey 33 §;.,1978).

The genus Sparganium is taxonomically related to the

7

cattails and is sometimes included in the family Typhaceae
rather than being placed in the monogeneric Sparganiaceae
(Voss, 1972). .§a eurycarpum is the largest and most robust
of the North American burreeds (Fasset, 1957). It is
potentially worthy of cultivation due to its often high
primary productivity (van der Valk and Davis, 1978) and
resultant potential for removal of nutrients from
artifically enriched substrates. It resembles Eypgg spp.
in its poor regrowth characteristics when using multiple
harvests (Hanseter, 1975).

§a eurycarpum often forms large, nearly monospecific
stands in waters up to 1 m deep. In Pentwater Marsh
(Oceana County, Michigan), a Lake Michigan riverine coastal
marsh, it is the dominant cover type in the deepest water
of the emergent vegetation zone. This marsh is receiving
extensive study of its vegetation and the effects of the
vegetation on the nutrient budget of the marsh as part of
the Michigan Sea Grant Program (Burton and Kelley, 1983).
A second reason for studying this plant was therefore to
provide information on the growth and nutrient uptake of
this plant species under controlled conditions so that this
information might be used to help predict the response of
Pentwater marsh to additional influxes of nutrients.

Prairie cordgrass (Spartina pectinata Link) is a
productive wetland grass species chosen for study in
comparison with the other two grass species. It is a
perennial C4 grass and it was desired to examine how this

species would differ from the C3 species studied in

8
prOperties important to its growth on sewage-enriched

substrates.

MATERIALS AND METHODS
NUTRIENT UPTAKE EXPERIMENTS

The response of 5 species of emergent aquatic plants
to varying rates of nitrate, phosphate, and potassium
fertilization was examined using a series of experiments
with each experiment in an individual tub. These
experiments were conducted outdoors at the Water Quality
Management Facility (WQMF) of the Institute of Water
Research at Michigan State University. Plants were grown
in 30 1 polyethylene tubs (.125 m opening area) using 20 l

of sand as a growth medium. The sand contained L.44 g m-2

of available P,41.1 g m"2 of available N, and42.3 g m-2
of available K as determined by the Michigan State
University Soil Testing Laboratory.

During 1980, experiments were conducted on.§;
australis and;L aquatica. 2; australis propagules
consisted of 5 cm long size-matched rhizome sections with
20 cm of shoot attached. Rhizomes were collected locally
from a single clump growing in water 20 cm deep. It is
unknown if all of the prOpagules were of the same clone.

Average dry weight was 1.2 g (s=0.4g). Average total N, P,
and K fractions of the propagules were 3.12%, 0.51%, and

9
2.47% respectively. Rhizomes were planted immediately

after collection with 15 cm of vertical shoot extending
above the sand surface.

Five §;_gguatica seeds were introduced to each tub
used for this species. Once established, wild rice plants
in each tub were thinned to one seedling. Seeds of the
Netum variety were obtained from the University of
Minnesota. All tubs were partially submerged in the
water-level controlled marshes of the WQMF in order to
provide a uniform and more natural temperature regime.

Tubs were arranged in double rows extending through the
shallow terrace in areas where all vegetation had been
removed. Assignment of species and chemical treatments was
completely random. The sand medium in the tubs was
completely isolated from the surrounding marsh water.

Treatments consisted of single applications of nitrate
(CaNOB), phosphate (NaZHPOu) and potassium (KCl) solutions
mixed into the sand prior to the rhizome or seed planting.
A 4x4x4 factorial arrangement of the three nutrient
treatments was used. The amount of nitrate-N added to each

-2

tub was 0, 7.8, 23.4, or 46.8 g m of sand surface

(equivalent to 0, 48.8, 147, or 293 g m-3 of sand); the
amount of phosphate-P added was 0. 2.1. 6.3. or 12.5 g m‘2
(equivalent to 13.1. 38.8, or 78.1 g m'3); the amount of
potassium, added was 0, 7.8, 23.4, or 41.5 g m'2
(equivalent to 0, 48.8, 147, or 260 g m'3). There were
five replicates of each treatment combination. Other

necessary plant nutrients were added from a single solution

10

to all tubs in the following amounts (g m'2)1MgSOu (38.5);
Fe(SOu)3 (1.13); H2B03 (2.48); MnSOu.H20 (0.088);
CuSOh°5H20 (0.096); ZnSOLL (0.112); 00012.6H20; (0.112);
(NH4)2M°O4 (0.080) (Modified from Dykyjova & Veber, 1978).
Deionized water was added to bring the tub solution level 2
cm above the sand. During the growing season, water level
in the tubs was maintained above the sand surface using
deionized water to supplement the rainwater inputs.

Experiments °n.21 angustifolia and‘gg latifolia were
conducted during 1981. The set-up of these experiments was
similar to that used for 2; australis and g; aquatica.
Eypgg prOpagules cOnsisted of 5 cm long size-matched
rhizome sections with 20 cm of shoot attached. All
rhizomes were collected locally from monospecific clumps
growing in water 10-30 cm deep. It is unknown if
propagules were all of the same clone. Average propagule
dry weight was 7.0 g (s=2.2g) for E;_latifolia and 1.1 g
(s=0.3g) for 24_angustifolia. Average total N, P, and K
fractions of the propagules were 2.65%, 0.52%, and 2.42%
respectivley for 2; latifolia and 2.33%. 0.51%, and 2.35%
for 2a angustifolia. Experimental treatments were the same
as those used for 2a australis and g; aquatica. However,
instead of being placed in marshes, 1981 tubs were
partially buried in a pit filled with water-saturated sand.
Tubs were again arranged in double rows with chemical
treatments and species assigned at random. This relocation
was necessary to avoid fluctuating marsh water levels,

damage to experimental plants by muskrats, and

11
contamination by blackbird droppings experienced during
1980.

During 1982, initial nutrient uptake experiments were
conducted on §a eurycarpum. In addition, further
experiments were conducted on g; latifolia, g;
angustifolia, and on‘zL australis at higher levels of
nitrate and phosphate treatments than were previously used.
.§1 eurycarpum propagules also consisted of 5 cm long
size-matched rhizomes with 20 cm of shoot attached.
Rhizomes were collected locally from a clump started two
years earlier from a single rhizome collected from
Pentwater Marsh and grown in one of the WQMF marsh-ponds.
Thus, all propagules were genetically identical. Average
prOpagule dry weight was 2.0 g (s=0.8 g). Average N and P
fractions of the propagules were 1.81% and 0.22%
respectively. Propagules were planted and tubs positioned
as described for the 23253 spp.

Experimental treatments for theԤL eurycarpum tubs
consisted of single applications of nitrate (CaN03) and
phosphate (NaZHPOh) solutions mixed into the sand prior to
the rhizome planting. A 5x5 factorial treatment
arrangement was used. The amount of nitrate-N added to
each tub was 0, 7.8, 23.4, 46.8 or 93.6 g m"2 of sand
surface (equivalent to 0, 48.8, 147, 293, or 156 g m-B’of
sand); the amount of phosphate—P added was 0, 2.1, 6.3,
12.5 or 25.0 g m"2 (equivalent to 0, 13.1, 38.8, 78.1 or
156 g m'3). There were seven replicates of each treatment

combination for a total Of 175 tubs planted with g;

12
eurycarpum. Other necessary nutrients were added from a
single solution to all tubs as with the previous
experiments except that 46.8 g m-2 of K as KCl was also
added to each tub. Other aspects of the experimental setup
were identical with that used during 1981. Only the two
highest treatment levels of nitrate and phosphate were used
on 21 australis, 1; angustifolia, and‘gg latifolia during
1982.

Shoot growth rates for each species in all three years
were estimated from meter stick measurements of shoot
height above sand level (sand surface to tip of longest
leaf) and caliper measurements of maximum shoot diameter at
tub rim level. Measurements were made at intervals of
about three weeks. These measurements were used in
conjunction with weights of shoots from the parent stands
to develop multiple regression expressions to estimate
shoot dry weight accumulation. Equation 1 is for 2;
australis, equation 2 for g; aquatica, equation 3 for g;
latifolia, equation 4 for 2; angustifolia, and equation 5
for_S_L eurycarpum. 95% confidence intervals on slopes and-

intercepts are shown below each equation.

In W = 1.83 In H + 0.70 In D - 8.17 (r=.98;p§.99) (1)
(1.7211-94) (0.5710-83) (- 8.471- 7.87)

ln W =- O. 86 In H + 1. 72 ln D + 1. 74 (r=.93;p;.99) (2)
(- 2. 061-o3“) (0. 9212- 52) (- 2. 6716 1)

1n W- -1.55 ln H + 0.88 In D - 7. 80 (r=.98;p;.99) (3)
(1. 28;1.82) (0.64;1.12) (- 7. 13;-8. 46)

In W = 1.11 ln H + 1.35 ln D - 7.14 (r=.98;p;.99) (4)
(0.93;1.29) (1.19;1.52) (- 6. 701-7- 58 )

13

In W = 1.27 ln H + 1.36 In D - 9.11 (r=.98;p1.99) (5)

W in each equation is equal to shoot dry weight (g), H is
equal to shoot height (cm) and D is equal to shoot diameter
(mm). Equation 1 was developed using 98 g; australis
shoots harvested during September, 1980; Equation 2 from 20
Z.aquatica shoots harvested in August, 1980; equations 3
and 4 used 70 E; latifolia and 138 2; angustifolia shoots
respectively that were collected in August, 1981; equation
5 used 30 §1 eurycarpum shoots cut during August, 1982.
Shoots for 2;,australis,‘gL angustifolia, T; latifolia and
.§; eurycarpum regressions were from parent stands;.§L
aquatica shoots were grown from the same seed used in the
nutrient uptake experiments. Both flowering and
non-flowering shoots were used. Estimates were corrected
for bias introduced by the logarithmic transformations
using the method of Baskerville (1973). In order that
extrapolation might be possible to weights at other drying
temperatures, 10 samples of each species were dried at 90 C
and 105 C in addition to 60 C. It was found that the
weights at the three temperatures varied by less than 1%
for all species. Shoot losses over the growing season were
presumed small and no attempt was made to estimate them.
During late August and early September, shoot samples
of each species were obtained by clipping them at sand
level while root and rhizome samples were water-sifted from

the sand. All samples were dried to constant weight at 60

14

C. Dried samples were coarse ground in a Wiley Mill,
thoroughly mixed, and subsampled for fine grinding through
a .1 mm screen. A 0.2 g sample of the finely ground tissue
was digested using a modified microkjeldahl digestion
(Nelson and Sommers, 1973). The digestion included a
reducing step so that any nitrates or nitrites present were
included in the N determinations. Lithium sulfate was
substituted for potassium sulfate in the digestion so that
the same digestate could be used for K determinations as
well as for N and P. K was determined by flame emission
spectrOSCOpy. N was determined using method 351.1
(Colorimetric, automated phenate, U.S. EPA, 1974) while P
was determined using method 365.1 (colorimetric, automated
ascorbic acid, U.S. EPA, 1974). As a check on the
procedure, U.S. National Bureau of Standards standard plant
materials (pine needles, orchard leaves and tomato leaves)
of known elemental composition were carried through the
same analysis procedures. The procedures used in this
study resulted in average recoveries of 105% (s=5.7%), 103%
(s=6.3%). and 97% (s=6.6%) of the standard N, P, and K

respectively.
ESTABLISHMENT AND HARVEST EXPERIMENTS

Experiments on the establishment and harvest of g;
australis, g; aquatica, 1; latifolia, 1; angustifolia, §a
eurycarpum, and.§L pectinata were conducted on three

Man-made marshes (M1, M2, M3) on the WQMF. The marshes

15

were constructed in 1973. All three were 0.4 ha in surface
area with a terrace design. The outer, shallow terrace
occupied 25% of the total marsh surface area and was
designed to Operate at a depth of 15 cm but can be filled
to 45 cm. The second terrace occupied 50% of the total
area with a designed operation depth of 60 cm while the
middle terrace was designed to operate at a depth of 90 cm.
All marshes were sealed with low organic matter clays
during construction.

One Of the three marshes (M2) had little emergent
vegetation established in it as of the spring of 1980.
This resulted from a 1974 drawdown to repair a control
structure that had killed most of the originally planted
vegetation. All three of the marshes were originally
stocked with}:L latifolia by transplanting approximately
600 rhizomes per marsh into the shallow terrace during
October and November, 1973 (McNabb et al, 1975). At the
start of this project, the shallow terrace of M2 contained
only scattered patches of g; latifolia, g; angustifolia, g;
australis, and other minor emergent components. Because of
the availability of unoccupied space, M2 was chosen for the
establishment experiments. Immediately prior to the start
of experiments, the other two marshes (M1 and M3) had
nearly continuous and dense stands of 2;,latifolia,‘2;
anggstifolia, and shoots of intermediate morphology which
may have been the hybrid QLx glauca growing on the shallow
terraces and extending out over part of the middle

terraces. These two marshes were utilized for harvest

16
experiments on established stands of 2yp§_.

The shallow terrace of M2 was prepared for plant
establishment by first removing all existing vegetation
from areas where experiments were to be conducted. With
the water drained, the sediments of the shallow terrace

were fertilized with urea at the rate of 45 g N m'2

by
raking granulated urea evenly into the sediments. Other
nutrients were assumed to be already present in excess
since the marsh had previously been repeatedly flushed with
secondary municipal wastewater during 1973-1975.

Wastewater inputs to the marshes are normally low in N
because the water flows through two sewage lagoons with
high N stripping properties before entering the marshes
(King and Burton, 1979).

The shallow terrace of M2 was divided into plots 2.5 m
long around the circumference of the marsh. The plots
extended about 5 m from land to the edge of the shallow
terrace. Nine plots were prepared for each of the six
previously mentioned species. Plots were assigned to
species in blocks of 3 contiguous plots. During the early
summer of 1980, an attempt was made to establish the five
perennial species selected for study (2; australis, Ea
latifolia, Tg’angustifolia, §1 eurycarpum, and g;
pectinata). 2; australis, E; latifolia, and 1;
angustifolia propagules were collected locally from shallow
water populations by digging up belowground sections from
monospecific stands. §;,eurycagpum was collected from

Pentwater Marsh and §;_pectinata was collected from a ditch

17
along a local railroad right-of-way. Clumps of roots and
rhizomes about 20 cm in diameter were transplanted into the
prepared plots. Each plot had 10 evenly spaced clumps
imbedded into the clay/organic surface layer.

During April, 1980 g; aquatica plots were planted with
about 500 g per plot of Netum variety seeds. The seed was
spread evenly over the surface and then raked into the
clay/Organic surface sediments.

Also during 1980, harvest experiments were begun on
plots of zypgg spp. growing on the shallow terraces of M1
and M3. Nine 5 m long plots with dense stands of Eypgg
spp. were selected in each marsh for two different
harvesting regimes and controls with three replicates in
each marsh. Plant harvesting began during early June,
1980. Three plots were harvested four times each at three
week intervals while three other plots were harvested twice
each at six week intervals with the remaining three plots
in each marsh serving as controls. Biomass removed and N,
P, and K content of this biomass was determined for all
plots. Control plots were also sampled for standing crOp
and nutrient content.

During 1981, the plots in M1 and M3 were monitored for
recovery from the harvest treatments. Samples were taken
during both early and late summer to measure standing crop
and nutrient content.

Similar harvest experiments were conducted on M2 plots
during 1981 on the newly established vegetation. Only

plots of 2; latifolia, T; angustifolia, and_S_L eurycarpum

18

were dense enough to conduct reasonable harvest
experiments. Recovery of M2 plots was monitored during

1982 by measurement of standing crop and nutrient content.

MARSH NUTRIENT BUDGETS

Nutrient budgets for M1, M2, and M3 were constructed
for 1980, 1981 and 1982 by measuring controlled inputs and
outputs of nutrients and water. All three marshes were
flushed periodically except during the winter months using
water from Lake 2 of the four lake WQMF facility. This was
done on a rotating basis with each marsh receiving inputs
for one week at a time while the other two marshes received
no inputs. Inputs were measured at the control structure
of Lake 2 and outputs were measured at the output channels
of the three marshes using water level recorders and
V-notched weirs. Water samples were collected daily when
water was flowing from these input and output structures.
Laboratory analyses of water samples included
determinations of pH, specific conductivity, nitrate-N,
nitrite-N, ammonium-N, total Kjeldahl nitrogen, total P,
soluble molybdate reactive P, chloride, potassium,
alkalinity, and hardness. These data were used to

construct input/output budgets for the marshes.

19
RESULTS AND DISCUSSION

NUTRIENT UPTAKE EXPERIMENTS
Phragmites australis

The original emergent shoot generally died but new
shoots typically appeared from the rhizome in about two
weeks. The number of shoots per tub in September varied
from 1 or 2 to over 50. There was no significant growth
response to K+ fertilization (Table 1). Total K uptake by
2; australis in pots not fertilized with K exceeded the
1.31 g m"2 supply of exchangeable and soluble K of the
sand. Thus, more K was available to the plants than was
indicated by the sand nutrient analysis, possibly due to
mineral weathering or input from precipitation. Since it
was found that K+ treatment did not significantly affect
growth, growth values for the nitrate and phosphate
treatments were averaged over all levels of K treatment
(Figure 1). There were strong and significant (Table 1)
growth responses to the nitrate and phosphate treatments
and to the interaction of nitrate and phosphate treatments
(Table 1). 2; australis growth responded to nitrate and
phosphate fertilization in much the manner predicted by the
Mitscherlich (1954) "rule" for many crap plants. That is,
growth increase per unit increase in nitrate and phosphate
fertilization declines at higher treatment levels.

Root and rhizome production of §;_australis increased

20

 

mm. ”mm." mm assassaeesH Bessemeaa a-a-z

 

.m.: :mm.o m sowpomuech escapeehe xnm
.m.: :mm.o a :owpomhoacH escapeene Ma:
mm. o:m.~m m :oapomumusH p:ch«oha mnz
so. mNL.H m amazemeae a
mo. mmo.am m escapeeua m
mm. “cm.ww~ m pcosamoua z
m we swam m mm soapmwhm> Ho condom

 

.Acdmucv gpzohm name moH poems mama ape poonm
o>fiamassso mwamppmsm 4M.mo moamsaamc no mzowpompcuca hams» use
mpzospMOAp Ho maommao on» so oocmwhm> mo mamhamc< .fi manna

 

21

 

 

 

 

 

 

 

 

 

 

 

 

 

10°C
'NE 2'84 \‘N
0.... RV = 6:8 w§
'2 A
*' R
1- R
1 \
o % §
:3 *2 N
= 2 N
.. A a
2 2 a
z / S
. a N
A N
. in / k

 

TREATMENT (g m'a)

Figure 1. Average areal dry weight (60 C) of 2; australis
shoots after 106 days growth during 1980 with varia e
combinations of nitrate and phosphate fertilization.
Vertical lines are standard errors of the means.

22
with nitrate-N content of the substrate but only through
the intermediate level of 14.6 g m-2 (Figure 2). Root and
rhizome production as a fraction of total production
declined with increasing nitrate treatment (Figure 3).
Root and rhizome production increased with phosphate supply
only up to the 1.31 g m-2 treatment level (Figure 2). The
fraction of the total production in the roots and rhizomes
showed no significant (Table 2) relationship to phosphate
treatment (Figure 3). Fiala (1976) found the lowest root
and rhizome to shoot biomass ratios associated with the
most productive habitats in natural 2; australis stands in
Czechoslovakia. N content of the top 20 cm of sediment was
found to be the best predictor of root and rhizome biomass
of all parameters examined by Fiala. N0 simple pattern
could be discerned by Fiala with respect to the effects of
a number of habitat variables on the growth of underground
organs of‘g; australis. Thus, the importance of substrate
nitrogen in determining the distribution of production in
§L_australis was suggested both by the present study and by
Fiala's (1976) results.

Shoot N and P concentrations were not significantly
affected by the level of K treatment (Table 3). Therefore,
shoot levels of N and P were averaged over all levels of K
treatment (Figures 4 and 5). Lowest tissue N
concentrations occurred with the lowest level nitrate
treatment only when no phosphate was added. Otherwise,
tissue N concentrations were lowest at intermediate rates

of nitrate addition. This patterm of lowest N at

23

 

__ N TM '1 ' .1:
--- PTRT

 

 

 

 

 

 

 

s"

E

23

1.. --

:1: .

52

m -~~~~

3 z’ ‘

111 ,’

2 . "

C) 1’

E

a I __

a:

+

g...

<3

8
0 l r T I
P 0 2.1 6.3 12.5
N 0 7.8 23.4 46.8

N on P LEVEL(9 m’z;

Figure 2. Total areal seasonal belowground production ofgL
australis as a function of nitrate or phosphate treatment
level during 1980. Vertical lines are standard errors of the
means.

 

24

2.5

 

__ NTRT
'“ PTRT

 

 

BELOW TO ABOVE GROUND RATIO

 

 

 

 

.J

6.3 12.5
23'“ “608

N on P LEVEL (9 m'z)

25'010
00
\]N
O
mH

Figure 3. Relationship at end of 1980 growing season between
nitrate or phosphate treatment level and distribution of
production aboveground and belowground by 2; australis.
Vertical lines are standard errors of the means.

25

 

.m.a ao.fi m an leases neesm\.m+.m psesamena a

 

.m.s me.o m an efipmm neesm\.m+.m escapades a
me. Hm.m m an edema neesm\.m+.m acesamsaa z
a as .mam a as : sesame aespmmsm> me measem

 

.mwmhamcm HMOH»MH»M»M op

poaum cospomm:MMp canons use masoeHmomxo mo use as ecumo>nmz MHmHMOpMEIMo magmHos
so oommm .pst03 hue poozm op unmaoz and oEoNdsm + pooh maaMhpmsm .m we owamm
can so mpsospmogp mo mpoowmo on» so consaam> we mwmzamsm Ho measmom .N manna

 

Table 3.

26

Results of analyses of variance on the effects of

treatments on %N, %P, and %K of P. australis shoots and
Based on wEIghts of materials

harvested at end of experiments.

roots plus rhizomes.

 

Source of Variation

'2’.

NN’d‘d

NW’U‘GZZNH‘U‘UZZ

Treatment
Treatment
Treatment
Treatment
Treatment
Treatment
Treatment
Treatment
Treatment
Treatment
Treatment
Treatment
Treatment
Treatment
Treatment
Treatment
Treatment

Treatment

 

Effect n _ng F Sig. of F
Shoot %N 41 3 7.41 .99
R.+R. %N 34 3 8.96 .99
Shoot %N 41 3 2.18 .89
R.+R. %N 34 3 1.59 .89
Shoot %N 41 3 0.41 n.s.
R.+R. %N 34 3 0.15 n.e.
Shoot %P 41 3 0.62 n.s.
R.+R. %P 34 3 4.40 .99
Shoot fiP . 41 3 1.75 .83
R.+R. %P w 34 3 6.67 .99
Shoot %P 41 3 0.76 n.s.
R.+R. %P 34 3 0.19 n.e.
Shoot %K 41 3 4.13 .99
R.+R. %K 34 3 0.96 n.e.
Shoot %K 41 3 1.72 .82
R.+R. %K 34 3 0.57 n.e.
Shoot %K 41 3 5.43 .99
R.+R. %K 34 3 5.31 .99

 

27

 

      
   

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

2.5
-N-O
13:3581;
ii N'4618
\‘ ..
s N A .. A
o - § \ F \
\ H- N N
:5 1) a 7% A
A a 7A
R A - A
A 7% 7%
O‘J'J" .47 ii l§ _J ’\\
p-o p-2.1 P-6.3 P-12.5

2

TREATMENT (9 Ii )

Figure 4. 2;.australis shoot concentration of N as per cent
of dry weight (50 C) at end of 1980 growing season for
different combinations of nitrate and phosphate
fertilization. Vertical lines are standard errors of the
means.

28

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

0.15
g 8‘: P"=12?5 1%
" \
:2 2 W Lu
2 / " / a
" / A 6 R
g; // x/ /’ //R§
// // v/ F //N\
I / / / '1 / §
'0 / / / M / L
/ "h / 7 / \
/ / t / / R
/ / / / \
¢\ 2 / / S
2 / / 7%
/ « V 8 g R
C 11‘ —--1 Q — )- LN
N-O N=-7.8 "-53.4 N-46.8
TREfifllMENTXgIfi')

Figure 5. ‘2; australis shoot concentration of P as per cent
of dry weight (60 C) at end of 1980 growing season for
different combinations of nitrate and phosphate

fertilization. Vertical lines are standard errors of the
means.

29

intermediate rates of N fertilization often occurs under
conditions of severe soil nutrient deficiencies with crop
plants when whole plants are sampled for nutrients (Smith,
1962). N is normally transported from mature tissues to
active growth areas resulting in a ”dilution” of N during
growth (Smith, 1962). Since plants with poor N supply grow
very slowly, the transport of N from mature tissue to
actively growing areas ofgL australis was depressed, .
resulting in higher per cent N at the lowest treatment
level compared to intermediate treatment levels. In
contrast, total N uptake was directly correlated with level
of nitrate added (Figure 6) because of the overriding
effect of the quantity of tissue produced (Figure 1).

Except with a nitrate-N level of 46.8 g m'z, minimum
shoot P concentrations were found to be associated with the
lowest level of phosphate at a given level of nitrate
treatment (Figure 5). Association of the lowest shoot P
concentrations with the lowest phosphate treatment level
may reflect the relative N as compared to P shortage
existing in the untreated sand. Otherwise, tissue P levels
might be expected to exhibit a pattern similiar to that of
the tissue N levels since both nutrients are mobilized from
older tissues. The total quantity of N uptake was strongly
dependent on the phosphate treatment (Figure 6). This was
especially true at the higher nitrate treatment levels.
The same type of relationship exists for P uptake (Figure
7). That is, the amount of P taken up is dependent

strongly on both the nitrate and the phosphate treatment

3O

 

7

 

. w
Q\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\ 5

villi/WM 2

P3603
NT

\.
TREATMENT (g

 

P- 201

-II.

-0

III
p

 

 

m¥<hn= 2 #001m

 

under variable combinations of nitrate and phosphate

Seasonal areal uptake of N by P. australis shoots
fertilization during 1980.

Figure 6.

31

 

 

 

 

 

 

 

 

 

 

    

 

1.0
IIIP"0
':-§.1
97‘ N P' 1:25
E
2
III
a:
,5
. 7
= 7
°‘ 2
g /
. a
3‘, /
K
o- , L . . n ,/
TREATMENT(9

 

Figure 7. Seasonal areal uptake of P by §;_australis shoots
under variable combinations of nitrate and phosphate
fertilization during 1980.

32
level.
Shoot N concentrations were found to be significantly
higher than root plus rhizome N concentrations (Figure 8).
P and K concentrations showed no significant between tissue

differences.
Zizania aquatica

Z. aquatica differed from all the other species
studied in that it is an annual established from seed. Its
biomass production response to individual treatment
combinations was also markedly lower, probably due to the
much smaller propagule size. Since 2; aquatica did not
approach its maximum areal biomass production, production
values are given on a per plant basis (Table 4).

Extremely low production values and high mortality
with certain treatments of §;_aguatica did not allow for
reliable nutrient analysis on samples from every treatment.
However, nutrient content patterns were similar to those
found with the species started from rhizomes. That is,
tissue P was highest with high phosphate-low nitrate
treatment while tissue N was greatest with low P-high N
treatment. However, P showed a much greater range than did

N (Table 5).

Typha spp.

For bOth‘EL angustifolia and 1; latifolia, N, P and K

33

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

1.5

gaggamzoue (WWW , 1 .
*_ ‘TTW'W'WI
I
(D
E
1 r11
39

M't'" F41 1....
C U
N

P
TISSUE - NUTRIENT

Figure 8. Comparison of the average 1980 N, P, and K content

of P. australis shoots with that of roots plus rhizomes.
VerficaI Iines are standard errors of the means.

34

 

 

em. sm.m mm mm aeesm asesemena
.m.: om.o mm mm seesm usesnmeae
He. ee.a am an neeam passeseas
em. aga.~ mm nausea eeeam aseseesaa

a as .mfim a an. usages seaemfiae> as seasem

 

 

soaumsdm 4m.mo MR use

.Ahao>dpoommop .om.mm.om.mcu:v meoonm
.mR .zx .anmwos pmo>hms :o masos»Moup

mo mpoommo on» so oosmaum> mo momzamsm mo mpasmom .3 manna

Weight values are in g per plant; percentges

 

Effects of variable nitrate, phosphate, and potassium
treatments on Z. aquatica shoot dry weight, root dry weight and tissue

content of N, P and K.

Table 5.

are as per cent of 60 C

1ght.

dry we

 

{é

Root

%N

%K Dry Wt.

Shoot
%P

%N

Dry Wt.

Treatment
(2 m-Z)

35

N
(\1\ {\OO ON ON

|| 0 0 0 0 0| | 0 0| | 0 0
ON «mm-o '41-. NH
ON NOON \OO O
«401 v-ow-I—o v-ON HN

|| 0 0 0 0 0| | 0 0| | 0 0
OO OOO OO 00
N O\ L\

' O\L\ OOO {\41' 'Cv-I
| O 0 0 0 0| | 0 0| . .
OH rev-0O Ov-I woe-4

v-‘N #OOWOd’O \Ol-fi

:-
Nv-IONO\00\O\OHNL\OC'\#O

OOv-IOOOOOOOv-IHOOON

 

['mm O\V\\-fi \OOVN O\€'\\O
0 0 0 0 0| 0 0 0| 0 0 0
e-IN NNN NNN v-cNN
ON v-Iv-H-fi O\OH 5:“
,'m: Hum can NNN
| 0 0 0 0 0| 0 0 0| 0 0 0
OO OOO OOO OOO
'\O:I' \O‘fid’ 'OMW'BNO
| 0 0 0 0 0 0 0 0 0 0 0
u—«N e-Iv-Iv-I e-Ie-Iv-I NNN
cos-mambo mo L\L\
Ov-Ov-tv-dOb-O-d'Omev-dNON
OOOOOONNOOJ’WOONO

-:NC'\ «NM HNn v-INC"

ONONONONONONONON

OOOOd’d’d’d’OOOO

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36
treatment factors showed significant effects on end of
experiment shoot weight and on root plus rhizome weight
(Tables 6 and 7). All first and second order interactions
were also significant (Tables 6 and 7). This shows that
growth response to N, P or K fertilizers was dependent upon
levels of the other two fertilizers. The magnitude of the
production response to nitrate treatment was far greater
than the response to phosphate treatment which was in turn
far greater than the response to K treatment. Because of
the small increase in plant growth as a function of added
K, production responses to nitrate and phosphate treatments
were averaged over all levels of K treatment (Figure 9).
Both species showed similar patterns of response. However,
significant differences did occur. For example,‘gL
angustifolia distributed a significantly greater (Table 8)
proportion of its net primary production to belowground
tissues (Figure 9). The;L angustifolia belowground to
aboveground dry weight ratio averaged 2.58 compared to 1.97
for 2;_latifolia (Figure 10). The difference in the two
ratios largely resulted from the higher aboveground
production of‘g; latifolia. Over all treatments, the
average aboveground dry weight of 2L latifolia of 440 g m"2
(Sx=28) was significantly greater (Table 8) than the 360 g
m-2 (si=25) mean aboveground dry weight of T; angustifolia.
There was no significant difference in the belowground
ground production of_'I_'_L angustifolia and g; latifolia
(Table 8).

Averaged over all levels of other factors, belowground

37

Table 6a. Analysis of variance on the effects of
treatments and their interactions on T. an stifolia
cumulative above ground dry mass at t5? enE c? {Re growing
season (n=174).

 

 

Source of Variatign DF F §i5;,0£_§
N Treatment 3 1186.4 .99
P Treatment 3 42.4 .99
K Treatment 3 6.9 .99
N-P Treatment Interaction 9 14.2 .99
N-K Treatment Interaction 9 4.8 . .99
P—K Treatment Interaction 9 2.6 .95
N-P-K Treatment Interaction 22 2.8 .95

 

 

Table 6b. Analysis of variance on the effects of
treatments and their interactions on T. a stifolia
cumulative below ground dry mass at tfig’eng of tfie growing
season (n=174).

 

 

§ggrce of Variation D? F Sig. of F
N Treatment 3 1110.3 .99
P Treatment 3 28.6 .99
K Treatment 3 6.8 .99
N-P Treatment Interaction 9 7.0 .99
N-K Treatment Interaction 9 3.0 .99
P-K Treatment Interaction 9 1.9 .95

N—P-K Treatment Interaction 27 1.8 .98

38

Table 7a. Analysis of variance on the effects of
treatments and their interactions on T. latifolia
cumulative above ground dry mass at t5? en 0

 

 

 

season (n=173). 8 growing
Source of Variggion DF F 4§;g:_q£_§
N Treatment ' 3 1142.7 .99
P Treatment 3 31.9 .99
K Treatment 3 5.4 .99
N-P Treatment Interaction 9 15.7 .99
N-K Treatment Interaction 9 3.7 .99
P-K Treatment Interaction 9 3.1 .98
N-P-K Treatment Interaction 27, 4.3 .99

 

 

 

Table 7b. Analysis of variance on the effects of
treatments and their interactions on T. latifolia
cumulative below ground dry mass at tRE'end OT Efie growing
season (n=173).

 

 

Source of Variation _DF P Sig. of F
N Treatment 3 189.2 .99
P Treatment 3 20.8 .99
K Treatment 3 4.0 .99
N-P Treatment Interaction 9 9.6 .99
N-K Treatment Interaction 9 4.9 .99
P-K Treatment Interaction 9 1.9 .95

N-P:§LTreatment Interaction 27 53.3, .99

39

 
   

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and belowground tissues in early September, 19
phosphate fertilization.

Figure 9.

40

Table 8a. Results of analysis of variance on the effects
of treatments on the ratio of‘z; anggstifolia root +
rhizome dry weight to shoot dry we ght. Based on weights
of materials harvested at end of 1981 experiments and
arcsin transformed prior to statistical analysis.

 

 

Source of Variation Effect n 'gf F Sig. of F
N Treatment R.+R./Shoot Ratio 110 3 28.9 .99
P Treatment R.+R./snoot Ratio 110 3 5.8 .99
K Treatment R.+R./Shoot Ratio 110 3 3.9 .99

 

 

Table 8b. Results of analysis of variance on the effects
of treatments on the ratio of T;_latifolia root + rhizome
dry weight to shoot dry weight. Based on weights of
materials harvested at end of 1981 experiments and arcsin
transformed prior to statistical analysis.

 

 

Source of Variation Effect n df F Sig. of F
N Treatment R.+R./Shoot Ratio 111 3 6.07 .99
P Treatment R.+R./Shoot Ratio 111 3 0.62 .99

K Treatment R.+R./Shoot Ratio 111 3 0.64 .99

 

41

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42
production in both species increased only up to the 23.4 g
m’2 treatment level of nitrate (Figure 11). Shoot
production continued to increase up to the highest
treatment level of nitrate. Nitrate was the only factor
that significantly (Table 8) affected the proportion of
biomass distributed aboveground as compared to belowground.
This effect of nitrate appears to be at least partially
independent of the increased production brought about by
the nitrate fertilization.

Aboveground shoot weight from the May planting to the
September harvest resembled a logistic type of increase
following an initial lag period (Figure 12). The shape of
this response curve was very similiar for both species.
Total shoot dry weight produced was slightly greater for T;
latifolia with T; angustifolia producing more numerous but
lighter shoots.

Although shoot dry weight production averaged over all

2 N,

levels of phosphate and K leveled off at 23.4 g m-
total N content of the tissues continued to increase up to
the 46.8 g m'2 treatment level in both 2; latifolia and T;
angustifolia (Figures 13 and 1h). Reaction to phosphate
treatment was similar. Dry weight increases leveled off
after the 2.1 g m"2 treatment level while total P content
increased up to the highest treatment level of 12.5 g m'2
of phosphate (Figures 13 and 14).

There was no significant difference (Table 8) between
'2; latifolia and T; angustifolia in aboveground or

belowground %N (Figure 15) or %P (Figure 16) when these

’43

 

 

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L19
were averaged over all treatments. Early September T;_
latifolia tissues averaged 0.75% N and 0.078% P aboveground
and 0.63% N and 0.136% P belowground. ‘1; angustifolia
averaged 0.86 %N and 0.086 %P aboveground and 0.73 %N and
0.157 %P belowground. Differences between aboveground and
belowground %N and %P were significant for both species
(Table 8).

The ratio of %P in belowground tissues to %P in
aboveground tissues varied markedly between treatments for
both species (Table 8). ‘Lowest ratio values of close to 1
occurred with high nitrate-low phosphate treatments. High
ratio values of over 3 were associated with high
phosphate-low nitrate treatments. Shoot production values
(Figure 9) indicated that nitrate limited growth and that
phosphate was present in great excess of needs for growth
in the high phosphate-low nitrate treatments resulting in
the highest belowground to aboveground %P ratio.
Apparently. phosphate in excess of that needed for growth
was taken up and stored primarily in the beIOWground
tissues.

The ratio of %N in belowground to %N in aboveground
tissues showed a relative constancy compared with %P. It
appears that the excess N taken up is distributed at a near
constant ratio between aboveground and belowground tissues
in both T;_latifolia and T; angustifolia.

Maximum values of %N and %P within a species and
tissue type with various combinations of nitrate and

phosphate treatments were 2 to h times the minimum values

50
(Figure 15). Highest %N values occurred with low phosphate
treatment and high nitrate treatment. Similarly, highest
%P values occured with low nitrate and high phosphate
treatment (Figure 16).

Tissue content of N and P expressed on an areal basis
(Figures 13 and 1h) showed much greater variability between
treatments than did N or P expressed as a percentage of
tissue dry weight (Figures 15 and 16). This areal
variability was due to the large variability in tissue
production between treatments. Typhg spp. aboveground
areal N or P content can be more than doubled by increasing
the respective fertilizer application beyond that needed
for maximum growth (compare Figure 9 with Figure 13). For
example. 2; angustifolia aboveground dry weight production
was approximately equal with treatments of either 12.3 g

2 2

m" of phosphate and 23.h or h6.8 g m' of nitrate.

However. the N content with the lower nitrate treatment was

only 5.6 g In"2 2

compared with 13.6 g m' for the higher
nitrate treatment. Belowground N or P tissue content can
also be increased substantially without increasing dry
weight by addition of fertilizer beyond that needed for
maximum growth (compare Figure 9 with Figure 16). This
trend for belowground P is especially significant due to
the differential concentration of excess P in belowground

tissues.

Sparganium eurycarpum

51

Early September mean shoot dry weight for S;

2 while belowground

eurycarpum ranged from 20 to 1035 g m-
dry weight ranged from 55 to 1280 g m'2 (Figure 17). Both
nitrate and phosphate treatments and their interactions
significantly affected production levels (Table 9). The
addition of nitrate resulted in a production response range
from near zero growth with zero added nitrate to maximum
belowground production at 23.4 g m'2 and maximum

2 to a high mortality

2

aboveground production at 46.8 g m“
rate or stunted growth at the 93.6 g m" treatment level.
Phosphate at the highest level did not result in any
increased mortality. Only relatively small increases in
aboveground or belowground dry weight occurred with
phosphate treatment levels greater than 6.3 g m-z.

The ratio of early September dry weight belowground to
dry weight aboveground was strongly and significantly
(Table 9) affected by nitrate treatment but not by
phosphate treatment (Figure 17). This response was
especially evident when comparing dry weight production
responses as the nitrate treatment level increased from
23.4 to 46.8 g m"2 (Figure 17). Above the 2.1 g In"2
treatment level of phosphate there was both an increase in
the aboveground dry weight and a decrease in the
belowground dry weight.

Tissue concentrations of N as per cent of dry weight
reached their maximum with the higher treatment levels of

nitrate combined with the lowest treatment levels of

phosphate (Figure 18). Similarly. tissue concentrations of

52

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53

Table 9a. Analysis of variance on the effects of treatments
and their interactions on S; eurycarpum cumulative above
ground dry mass at the end of the growing season (n=144).

 

 

Source of Variation DF F Sig. of F
N Treatment 4 132.4 .99
P Treatment 4 95.5 .99
N—P Treatment Interaction 16 179.0 .99

 

 

Table 9b. Analysis of variance on the effects of treatments
and their interactions °n.§; eurycarpum cumulative below
ground dry mass at the end of he growing season (n=64).

 

 

Source of Variation DF F Sig. of F
N Treatment 4 135.9 .99
P Treatment 4 29.1 .99

N-P Treatment Interaction 16 49.0 .99

54

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55
P peaked with high phosphate treatment levels combined with
low nitrate treatment levels (Figure 19). These maxima
corresponded with very low dry weight values (Figure 17).
This association explains the contrasting situation found
when areal uptake of N (Figure 20) or of P (Figure 21) was
compared with %N (Figure 18) or %P (Figure 19). Figures 20
and 21 are very similar and indicate the close
interdependence of nitrate and phosphate on the uptake of
one another. High uptake of one of the nutrients is
dependent upon high uptake of the other. %N was
consistently higher aboveground than belowground. %P was
not consistently higher either aboveground or belowground.
There was no tendency for excess phosphorus to be
differentially concentrated in belowground tissues as was

found with Typha spp.

General Discussion

Since N or P is believed to be the nutrient limiting
growth in most wetlands (Sculthorpe, 1967). the ratio of %N
to %P should resemble the ratio found under controlled
conditions of similiar nutrient supply in emergent aquatics
such as §; eurycarpum (Figure 22). This ratio may be a
better indicator of nutrient limitation than are the
"critical concentrations" of N or P used by others to
determine nutrient deficiencies in submerged species
(Gerloff and Krombholz. 1966). N:P ratios show less
variability between tissue types of different physiological

56

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57

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60
age than do absolute concentrations of N and P. "Critical
concentrations" of N and P also increase with increasing
concentrations of other nutrients (Bates. 1970). There is
also more temporal variability in amount of structural
tissues in emergent than in submergent species which results
in more variability in nutrient concentrations. The shoot
NzP ratio for §¢ eurycarpum plants with severe growth
limitation due to low P ranged from 13 to 19.5.
’Belowground. the range for the same treatments was 9.5 to
21.1. For severely N limited treatments the same ratio
ranged from 1.0 to 4.2 for shoots and 1.3 to 2.6 for
combined roots and rhizomes. Intermediate values of the
tissue N:P ratios were found with treatments where tissue
dry weight production was less strongly inhibited by low N
or P supply.

If these N:P ratios are to be used in comparisons with
natural stands to determine N or P deficiencies. tissues
analyzed from natural stands should ideally be of the same
developmental stage as the plants in this study unless it
can be demonstrated that the ratio is constant over the
growing season. For most emergent aquatic plants. shoot %N
during the early part of the growing season has been as much
as 3 times that found at the end of the growing season
(Dykyjova, 1978). This seasonality for N has been found in
some instances to be coupled with a relatively constant %P
which has led to a declining ratio with tissue maturity.
However, once the tissue has reached maturity and before

senescence. the ratio has usully remained relatively

61

constant. Dykyjova's (1978) data for several emergent
plant species showed little change in the NxP ratio from
early September through October. Since §L eurycarpum
tissues in this study were harvested at maturity in
September for nutrient analyses. comparisons of N:P ratios
of natural stands with data from this study should be based
on tissues collected during the latter part of the growing
season.

Shoot %N and shoot %P declined at approximately the
same rate over the course of the growing season for S;
eurycarpum stands in Pentwater Marsh. Michigan (Burton and
Kelley. 1983). This constant ratio allowed comparison
throughout the growing season with the tub-grown‘g;
eurycarpum. From May to September. 1981 the NsP ratio in
shoots ranged from 5.3 to 6.6. The N:P ratios for
Pentwater Marsh of 5.3 to 6.6 for the 1981 growing season
indicated that N was more limiting than was P. However.
the ratio was above that found with severe N limitation
with the tub grown §; eurycarpum. Therefore. it is likely
that the Pentwater S; eurycarpum was only moderately
deficient in N. Belowground Pentwater g; eurycarpum N:P
ratios were less appropriately compared with the tub-grown
data since the belowground tissues in the natural stand
were the result of more than one years growth. Belowground
samples from the natural stands would therefore include
much senescent material which may have lost P at a rate
greater than it lost N. Pentwater Marsh belowground NxP
ratio showed much greater variability both temporally and)

62

between samples from the same date than did the aboveground
ratio. The range during 1981 was from 5 to 11 with an
average value of 8. Using these values in comparison with
the tub-grown g; eurycarpum belowground tissues would point
more towards a phosphorus deficiency which. on the basis of
shoot nutrient comparisons, would be an incorrect
diagnosis.

N:P ratios similar to those found with §; eurycarpum
shoots were also found with the other species studied. The
above discussion on §; eurycarpum also, therefore. applies
to these other species.

Bernatowicz (1969) examined nutrient content of water
plants of Lake Warniak using the "critical concentration”
criteria of Gerloff and Krombholz (1966). Twelve of the
sixteen species examined were below the "critical
concentration" of .13% P while only the emergent species 2;
latifolia and §;_australis were below the critical
concentration of 1.3% N. The average NsP ratio was 10.5
for the Lake Warniak species. This analysis pointed out
the inappropriateness of using the same "critical
concentration" values for emergent and submergent plant
species. It also showed that the NxP ratio is less
dependent on the amount of structural tissues present than
are nutrient concentrations.

Differences existed in the abilities of the four
species of perennial plants studied to thrive at very high
levels of nitrate fertilization. This was determined

during the 1982 growing season when all four species were

63
grown under identical conditions using higher levels of
nitrate and phosphate fertilizers than had been used in
either of the two preceding years. Dry weight production
results are shown in Figure 17 for g; eurycarpum and in
Table 10-12 for the other three species. None of the .
species exhibited a significantly greater production of
aboveground or belowground tissues as treatment nitrate-N

increased from 46.8 to 93.6 g m-2

from 12.5 to 25.0 g m'z. ‘S; eurycarpum showed the most

or phosphate-P increased

severe response to the highest nitrate level with very high
mortality and stunted growth among the survivors. None of
the other species produced as much total dry weight at the
highest treatment level of nitrate as the same species did
at the second highest level. 2; australis experienced a
slight decline at the highest nitrate level while the m
spp. showed a more pronounced decrease. However, the
deleterious effects were not nearly as strong as those

found with S; eurycarpum.
ESTABLISHMENT EXPERIMENTS

Attempts to establish vegetation in Marsh M2 met with
mixed success. Some species grew rapidly and established
dense stands while others showed little growth and high
mortality of propagules. Successful species included 2;_
latifolia, 1;.angustifolia. and S; eurycarpum. S;
eurycarpum showed the most rapid and complete

establishment. Its plots were uniformly covered with

64-

 

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65

 

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66

 

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67
nearly pure stands of S; eurycarpum before the end of the
second month after planting. By this time. there was
little evidence of the original planting pattern since all
areas in each plot were homogeneously covered with
vegetation.

With.T;_latifolia and T;_angustifolia there was also
considerable growth and spread from the originally planted
clumps. However, these species did not completely fill in
their plots by the end of the first growing season. There
was still visible evidence of the original planting pattern
with gaps in certain areas by the beginning of the second
growing season. This was especially true in areas away
from the shoreline where the water depth was greater.

“g; aquatica seeds showed low germination and high
seedling mortality so that plots sown with this seed became
only sparsely covered with a fewfig; aquatica plants. Part
of the problem with the establishment of this species may
have been the presence of large amounts of submerged
macrophytes which were piled onto the plots from the deeper
terraces during times of strong winds resulting in shading
and uprooting of §;_aquatica seedlings.

‘2; australis also did very poorly in M2. Many of the
initial clumps died while most of the survivors did very
little spreading within their plots. Some of the surviving
clumps sent out numerous floating tillers over the water
surface. Some of these exceeded 10 m in length. However.
these same clumps showed very little local spreading within
their plots. This poor establishment may have been a

68

reflection of less than ideal substrate composition for the
spread of the plants. Most of the plots had only a very
thin ( 2 cm) soft organic layer over the heavy clay seal.
This substrate may not have been conducive for the spread
of P;_australis rhizome systems.

S; pectinata was the least successful of all of the
perennial species. Almost all of the initial clumps died
during the first summer. The few that survived showed
virtually no spread from the original clumps. Again. the

nature of the substrate was suspect.
HARVEST EXPERIMENTS

A summary of the harvest experiments onqupha spp.
plots in M1 and M3 (Table 13) showed the limited capability
of cattails to sustain a high yield with multiple harvests
over the course of a single growing season. After the
original late June, 1981 harvest. subsequent harvests
yielded an average of only 10% of the original harvest
biomass with each of 4 subsequent harvests at 3 week
intervals and an average of 19% with each of 2 subsequent
harvests at 6 week intervals. If the harvested stands had
been left standing and subjected to a single harvest in
late July. an examination of the control data indicates
that about 11% more biomass could have been harvested than
was harvested in late June. Therefore. the total amount of
biomass removal by the multiple harvests using either the 3

or 6 week harvest intervals was less than 130% of the

69

 

 

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7O
amount removable with a single harvest.

Increases in nutrient removal using multiple harvests
were somewhat greater than increases in biomass removal.
The maximum aboveground harvestable standing crop of N. P,
and K occurred in the control plots on June 25. Removal of
N amounted to 151% of this control mean using 3 week
harvest intervals and 150% of this control mean using 6
week harvests. P removal amounted to 154% of the control
mean with 3 week harvests and 150% of the control mean with
6 week intervals. K removal showed the smallest increase
with multiple harvests. K removal averaged 135% of the
control mean with 3 week harvest intervals and 130% of the
control mean with 6 week harvest intervals.

Harvesting had a marked effect on the vegetation
during the subsequent 1981 growing season (Table 13).
Effects were especially pronounced at the start of the 1981
growing season with harvested plots showing much lower
standing crops of nutrients and biomass. Plots that had
been harvested every three weeks had, on June 9. 1981. only
about 50% of the standing crop of nutrients and biomass of
the plots that had been harvested every six weeks and only
about 30% of that found in the unharvested plots. However.
differences rapidly diminished over the course of the 1981
growing season. By mid-August there was very little
difference between plots receiving the various treatments.

Results of harvest experiments on newly established
stands of 2;.latifolia. T;_angustifolia. and S; eurycarpum

in M2 (Table 14) show some differences between species in

71

 

 

 

 

 

 

 

 

 

 

 

 

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72
their response to harvesting at two different harvesting
dates. .2; latifolia and T; angustifolia showed greater
standing crOps of biomass and of N, P. and K at the
mid-August harvest date compared with the early July
harvest date. The Opposite pattern was true for the‘SL
eurycarpum plots. These patterns appear to be related to
the establishment course of the three species in M2. .2;
latifolia and T; angustifolia were still expanding from
their original plantings at the beginning of the 1981
growing season. .S; eurycarpum. on the other hand. was
experiencing a visually evident decline from the 1980
growing season. There was a noticeable deterioration in
the health of the S; eurycarpum stands with some shoots
appearing discolored and lacking the vigor of the previous
years growth. Invasion of the S; eurycarpum stands by
Scirpus spp. and Sagittaria sp. was also noticeable. It
was believed that propagules for these other species were
brought along with the S; eurycarpum from Pentwater Marsh
but they did not initially constitute an important

component of the newly established stands.

MARSH NUTRIENT BUDGETS

Input/output data for the three marshes M1. M2 and M3
(Tables 15-17) indicated that the marshes were all net
importers of water carried N. N may be either stored in
accumulating sediments, and/or lost as gases (NH3. N2, N20)

to the atmosphere. The three marshes reacted somewhat

73

 

 

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74

 

 

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76

 

 

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77

 

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79

 

 

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80

 

 

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000 00.0 00.0 00.0 00.0 00.0 00.0 00.0 000000 m0\0
-00 0 00000 0-000 2-200 z-mmz z-moz z-moz 000 3000 0 0000

 

.A.U.DCO0V m0 wands

82

differently for phosphorus. The import/export balance of
total P varied between net input and net output between
marshes and between years. Unbalanced chloride budgets for
some years probably indicate some unmonitored losses of
water and water carried nutrients since chloride is not
generally detained to any great extent by any physical.
chemical, or biological processes occurring in marshes.
These losses could represent seepage through muskrat

tunnels. around weirs. etc.

CONCLUSIONS

This research has revealed both similarities and
differences between the species in their abilities to
establish themselves from rhizome sections, incorporate
nutrients into their tissues, and recover from the stress
of multiple harvests. Some of these differences should
receive consideration when the role or potential role of
these species in natural and artificial systems is
discussed. For example. when establishing stands for the
purpose of maximizing biomass or nutrient uptake, it is
necessary to consider the substrate texture, operational
water depth of the system. and nutrient supply. All of the
perennial species studied may be easily propagated using
rhizome cuttings. However, they differ in other
prOperties.

In this study, all species spread about equally

 

 

83
rapidly through the sand substrate used in the tub
experiments. However. there were great differences between
the species in their abilities to colonize a heavy clay
substrate. .§; eurycarpum spread very rapidly in such a
situation and may be the best species of those studied for
use in a situation where a clay substrate cannot be avoided
and rapid establishment is necessary. Both species of
cattail are also suitable for such a substrate if rapid
establishment is not essential. Plantings of 2; latifolia
and 2;_angustifolia at the WQMF eventually resulted in very
dense. nearly monospecific stands. 2; australis appears to
be a very poor colonizer of clay substrates. When placed
in such a substrate the plants sent out floating horizontal
tillers rather than roots, rhizomes and vertical shoots.
At the WQMF E; australis is much more successful in the
looser soils immediately adjacent to the marshes than in
the heavy clay sediments of the marshes.

Differences in success on different substrates for the
species used in this study have been reported in the
literature. Around Danish lakes it was found that 2;
angustifolia did better in softer sediments that did 3;
australis (Boye-Peterson. 1917). In Finland. the capacity
of‘gL australis to invade coarser sediments has been used
as an explanation of its greater development along Open
shores than is found with Schoenoplectis lacustris
(Maristo, 1941). Somewhat contradictory is the finding of
Brand (1896) that Scirpus lacustris has harder rhizomes

that Phragmites australis and can therefore grow in

84
gravelly areas where E; australis cannot. The natural
substrates of most common occurrence for g; aquatica are
sand and organic (Fasset, 1957). It has been reported that
T; angustifolia grows on more organic substrates than 2;
latifolia. Most of the literature reports on substrate
preference are descriptive rather than experimental and
often do not account for such covariates as degree of
disturbance and substrate chemistry. Substrate texture
should therefore be examined more carefully and under more

controlled conditions before recommendations are made for

 

artificial plantings in a prepared substrate.

While all of the species overlapped in their water
depth utilization. §; eurycarpum appeared to be the most
successful in the deepest waters of the WQMF marshes while
2; australis was most successful at the land-water
interface. There was no clear depth separation of the
cattails at the WQMF but it has been reported from other
studies that 1; angustifolia has a greater depth tolerance
that E;_latifolia (Grace and Wetzel, 1982) At the WQMF the
two Typhg species did not grow in as deep of water as was
found with §; euraycarpum or extend onto dry land as did 2;
australis. Penetration into the sod of the area
surrounding the WQMF marshes was accomplished only by‘g;
australis. It has been reported in other situations that
‘3; australis is capable of invading stands of other species
(Buttery and Lambert, 1965). Thus, either a tolerance for
low water level or the penetrating abilities of the 2;

australis rhizomes may be responsible for its appearance in

85
the sod. In many areas of Europe 2; australis has been
reported to be the dominant species in deeper water areas
(Bjork. 1967). This contrasting situation is probably due
to the greater resistance of P; australis to wind and wave
stress due to its more linear form when compared to Typha
spp. (Hutchinson, 1975).

The ranges of chemical environments of occurrence for
some of the species of this study have been roughly mapped
out (Hutchinson. 1975; Moyle, 19h5). In the north central
U.S., §L_australis is typical of low alkalinity to
moderately hard waters. The two species of Typhg occupy
the above type of waters plus harder waters and waters very
high in sulfate. There is no sharp distinction between T;
latifolia and T; angustifolia in chemical environment of
occurrence (Hutchinson, 1975) although they have been
reported to have somewhat different pH (Hotchkiss and
Dozier. 19h9) and salinity tolerances (McMillan. 1959). Ԥ;
eurycarpum and 2;.aguatica are typical species in areas of
low sulfate and low alkalinity and hardness. The natural
range of §; eurycarpum in low nutrient water relates to the
present findings of the tub experiments where.§L eurycarpum
did the most poorly of the species studied at high nutrient
concentrations.

Most wetlands have anaerobic substrates with large
amounts of organic matter present. Under these conditions.
nitrate is rapidly denitrified and ammonia is the dominant
form of available nitrogen (Patrick and Mahapatra, 1968).
Our tubs differed from this situation in that there was

 

86

initially almost no organic matter present in the substrate
and most of the added nitrate was therefore not
denitrified. It is therefore of interest what the response
of the plants would have been had ammonia been used as the
nitrogen source. Some differences would have occurred
because of differences in toxicity and uptake paths of the
two nitrogen forms. Ammonia is normally the more toxic of
the two forms (Mengel and Kirkby, 1979) and might be
tolerated only at lower maximum levels. However. since
most emergent aquatics are normally found associated with
high ammonia levels, they might be more tolerant than
typical land plants of high ammonia levels. Ammonium ions
are positively charged while nitrate ions are negatively
charged. This difference leads to different
antagonist/protagonist relationships (to plant uptake) with
other ions in solution. Because of the ionic charges,
nitrate would be expected to stimulate potassium uptake and
depress phosphate uptake while the opposite would be true
for ammonium. This could result in different N:P ratios in
plants in tubs with the same NzP fertilizer ratios but
using ammonia instead of nitrate as the nitrogen source.
The spacing of fertilizer applications over the course
of the growing season could also make a difference in the
total nutrient uptake. Some of the tub experiment plants
took up over 80% of the applied nitrogen which indicates
that little was available for the plants by the end of the
growing season. Emergent aquatics in natural stands

normally take up most of their nutrients in the spring.

87
However, the tub grown plants represented expanding
colonies which sent up new shoots later in the season than
most natural stands. The tub grown plants would also,
therefore. have been expected to take up more nutrients
later in the season.

As has been found in other studies, these new data
indicate that very large stands of vegetation would be
needed to remove the amount of phosphorus loading resulting
from the domestic sewage of a large population. Spangler
et al (1976c) estimate that the phosphorus removable by
Scirpus validus shoot harvests would accommodate only 25
persons ha"1 if all of the input P was to be removed in the
shoots. This assumes shoot removal of 3.5 g m"2 and a per
capita P input of 1.3 kg per annum. Others have achieved
removal of up to 5 g P 111'2 yr'1 with emergent marsh species
(Sloey'gtual, 1978). Several harvests per season were
needed to achieve even this removal. Maximum shoot P
uptake in the tub experiments were in the range of 1 to 2 g
In.”2 for each of the species studied. This study therefore
reinforces the concept that marshes and other wetlands are
not suitable for the long term treatment of large
quantities of high P wastewaters.

It is likely that different plants of the same species
studied but collected from different sites could vary
markedly in their growth responses to various levels of
fertilization. .2; australis is known to be exceptionally
diverse genetically (Bjork, 1967). Polyploidy is

widespread with the basic number of chromosomes considered

 

88
to be either 6 (Love and Love. 1961) or 12 (Bjork, 1967).

The tetraploid (48 chromosomes) is apparently the most
common worldwide but other variations including triploidy,
hexaploidy, heptaploidy. and octaploidy (based on n=12)
have been reported. Substantial genetically determined
morphologic differences between shoots have also been found
within a ploidy level as well as between levels (Bjork.
1967). Ecotypic differences have also been found to exist
forqupha spp. (McNaughton, 1966) and §;_aquatica (Elliot,
1977) and probably also exist for g; eurycarpum. Care must

therefore be exercised in extending these results to other

 

m1. .-.v.‘lv .r'u_-
.

populations.

'§;,australis. T;_angustifolia, T;_latifolia, and‘g;
eurycarpum remain as species with good promise for
cultivation in artificial marshes. Their utilization in
such systems should consider the interspecific differences
found in this study. In addition. attempts should be made
to develop new varities or find naturally occurring
ecotypes with greater nutrient uptake capabilities and/or

aboveground production potential.

89
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49

 

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1980 P. australis aboveground dry weight (g m-2) estimates from re

Number of replic

Table A1.

Table A1 (cont'd.).

 

“9 63

Days After Planting
35

K 1“

Treatment (g m-2)
N P

 

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106

Days After Planting
K M m

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