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Dynamics of Typha latifolia L. populations in treatment


wetlands in Estonia

Martin Maddison a,∗ , Tõnu Mauring a,b , Kalle Remm a , Merje Lesta a , Ülo Mander a
a Department of Geography, Institute of Ecology and Earth Sciences, University of Tartu, 46 Vanemuise Street, Tartu 51014, Estonia
b Institute of Technology, University of Tartu, 1 Nooruse Street, Tartu 50411, Estonia

a r t i c l e i n f o a b s t r a c t

Article history: The aim of this paper is to evaluate and compare broadleaved cattail (Typha latifolia L.)
Received 10 December 2007 biomass production and the nitrogen (N) and phosphorus (P) content in phytomass in three
Received in revised form treatment wetland systems and to propose suitable areas for treatment wetlands in Esto-
10 June 2008 nia for raw material production. The biomass samples (roots/rhizomes, shoots with leaves
Accepted 10 June 2008 and spadixes) and litter were collected from 1 m × 1 m plots—15 plots in the Tänassilma
semi-natural wetland, 15 plots in the Põltsamaa free water surface (FWS) constructed wet-
land (CW), and 10 plots in the Häädemeeste FWS CW. The average aboveground biomass of T.
Keywords: latifolia varied from 0.37 to 1.76 kg DW m−2 in autumn and from 0.33 to 1.38 kg DW m−2 in win-
Assimilation ter. The greatest average nitrogen (22,950 mg N kg−1 ) concentration was found in spadixes in
Biomass production 2002, and the phosphorus (6500 mg P kg−1 ) concentration was measured in roots–rhizomes
Broadleaved cattail in 2003. Average standing stock of nitrogen and phosphorus was higher in aboveground than
Nitrogen belowground phytomass. In FWS CWs with high hydraulic and nutrient loadings, however,
Phosphorus the harvesting of aboveground biomass is not an effective means for the removal of nutri-
Wastewater treatment ents. Cattail biomass is a valuable insulation material, whereas the fibre from spadixes
mixed with clay gives elasticity to clay plasters. According to our estimates, about 5412 km2
could be used for Typha cultivation in Estonia.
© 2008 Elsevier B.V. All rights reserved.

1. Introduction agricultural irrigation systems (Ray and Inouye, 2007) and agri-
cultural drainage (Borin and Tocchetto, 2007), wastewater from
Constructed wetlands are widely used for wastewater treat- a tannery (Calheiros et al., 2007), farms (Hunt et al., 2003),
ment throughout the world. These systems can also be used dairy (Newman and Clausen, 1997; Gottschall et al., 2007), fish
for raw material production. farms (Maltais-Landry et al., 2007), landfill (Maehlum, 1995;
Cattails are very often a part of natural and constructed Bulc, 2006), woodwaste (Masbough et al., 2005), contaminated
treatment wetland ecosystems (Kadlec and Knight, 1996; groundwater from industrial areas (Machate et al., 1999) and
Vymazal, 2007). Cattail has been planted in constructed wet- mining (Mitsch and Wise, 1998; Lee and Bukaveckas, 2002), as
lands for primary and/or secondary treatment of domestic well as urban runoff (Scholz and Xu, 2002).
and industrial sewage (Ennabili et al., 1998; Shannon et al., Free water surface (FWS) wetlands covered by cattails can
2000; Mander et al., 2001a; Lim et al., 2003; Solano et al., be considered to be valuable biotopes supporting biodiver-
2004; Toet et al., 2005; Álvarez and Bécares, 2006), industrial sity (Lacki et al., 1991; Kadlec and Knight, 1996; Wild et al.,
wastewater (Maine et al., 2006; Ye et al., 2001), runoff from 2001).


Corresponding author. Tel.: +372 7375 816; fax: +372 7375 825.
E-mail address: martin.maddison@ut.ee (M. Maddison).
0925-8574/$ – see front matter © 2008 Elsevier B.V. All rights reserved.
doi:10.1016/j.ecoleng.2008.06.003
e c o l o g i c a l e n g i n e e r i n g 3 5 ( 2 0 0 9 ) 258–264 259

Although cattail is a persistent plant, spreads aggres- inhabitants) and from the food processing industry. It was con-
sively, and has a tremendous reproductive potential, the structed in 1997 (Mander et al., 2001a). First, about 1200 cattail
long-term results of biomass production in wastewater treat- plants were planted in the soil at the bottom of the second
ment wetlands are confusing. The production of vegetation and third ponds, and 500 young reed plants were planted in
in constructed wetlands can vary from 700 to 11,000 g m−2 , the fourth pond in summer 1998. Within a few years, the reed
standing stock of nitrogen (N) from 12.5 to 585 g m−2 and phos- disappeared and the cattail colonised all of the ponds. The
phorus (P) from 1.8 to 112.5 g m−2 (Kadlec and Knight, 1996; cattail rhizomes–roots became woven together and accumu-
Vymazal, 2004). There is some experience in Estonia in the lated decaying leaf litter to form floating mats on the wetland
use of cattail for construction materials. Cattail chips mixed surface.
with clay are used in the production of safe and cost-efficient The Häädemeeste wastewater treatment system consists
building blocks. The material is light and has good thermal of a conventional treatment plant, five infiltration ponds (total
insulation properties. Fibre material from cattail spadixes is area 0.23 ha) planted with common reed, and a cattail (T. lat-
used as clay plaster reinforcement. The fibre is an ideal mate- ifolia L.) free water wetland (0.72 ha). The system is located
rial to avoid cracks in clay plaster. Ready-made dry fibre and half a kilometre from the Baltic Sea coast and treats the
clay mixtures and cattail chips and clay blocks are produced municipal water (max Q = 160 m3 day−1 ) of the settlement of
and sold on the market (Mauring, 2003). Häädemeeste. The system was built in 1999. The primary pur-
The amount of reproduced biomass and content quality pose of the wetland is the removal of N and P (Mauring, 2002).
of macrophytes must be determined for macrophyte-based After 5 years (2004), the cattail basin has been colonised by
material producing wastewater treatment wetlands. another local native species—common reed (Phragmites aus-
We evaluated the annual biomass production and deter- tralis).
mined standing stock of N and P in the phytomass of The average annual wastewater and nutrient loadings of
broad-leaved cattail (Typha latifolia L.) in two FWS constructed the studied areas are presented in Table 1 (Ministry of the
wetlands (CW) and in one semi-natural treatment wetland Environment, 2004).
in Estonia. Based on the GIS analysis of various cartographic
sources, we proposed suitable areas for treatment wetlands 2.2. Sampling and analysis of plant fractions
for raw material production.
The cattail and reed phytomass measurements were per-
formed in autumn after the vegetation period, i.e. at the end
2. Materials and methods of August and the beginning of September and also the fol-
lowing winter, i.e. at the end of January and the beginning of
2.1. Site description February, in the years 2002–2006. The samples were collected
from 1 m2 plots. The information on phytomass sampling and
Common cattail and reed biomass production and cattail nutrient analysis in the plant fractions is given in Table 2.
nutrient standing stock were studied in three wetlands: in the The belowground root–rhizome samples were collected to
subsurface flow semi-natural wetland in Tänassilma (58◦ 22 W a depth of 50 cm in Tänassilma and a depth of 20 cm in Hääde-
25◦ 31 N) and in two FWS CWs in Põltsamaa (58◦ 38 W 25◦ 58 N) meeste and Põltsamaa using an auger (Ø108.6 mm). It was
and Häädemeeste (58◦ 5 W 24◦ 29 N) (Maddison et al., 2005). not possible to bore deeper in Häädemeeste and Põltsamaa
The Tänassilma semi-natural wetland (total area 228 ha) because of the thickness of the clay soil. Root samples were
is located in a primeval valley at the head of the Tänassilma washed clean of soil. Cattails were divided into four fractions:
River. The wetland is adapted to a high pollutant load of 15,000 roots–rhizomes, shoots with leaves, spadixes and litter. All
population equivalents and has received precipitation run-off samples were dried to a constant weight at 70 ◦ C.
and untreated municipal wastewater from the town of Vil-
jandi from 1948 to 2005. The upper reach of the wetland (69 ha) 2.3. GIS analysis
used to be species-rich grassland and former swamp, but after
wastewater had been discharged into the wetland, vegetation Different GIS data layers were used for selection of areas suit-
downstream was replaced by dense stands of broadleaved cat- able for cattail cultivation. First, eutrophic peat soils and gleic
tail. This area has almost no surface overflow and acts as a root soils were selected from a 1:100,000 soil map. Then the follow-
system and peat filter (Nõges and Järvet, 2002). The new efflu- ing categories were excluded: forest according to CORINE 2000
ent of treated wastewater from Viljandi treatment plant flows land cover, settlements according to 1:50,000 base map, pro-
in a narrow ditch in the centre of the wetland. The average tected areas, planned protected areas, valuable habitats, areas
water level decreased in 2004, when the wastewater effluent designated for Natura 2000 network from Estonian Nature
was changed. Infosystem (EELIS) and 200-m wide buffer zones at coastlines
The Põltsamaa CW is a cascade of four serpentine ponds of the sea, lakes and rivers.
with a total area of 1.2 ha. The mean depth of the first
pond is 100 cm, and of the other three approximately 70 cm. 2.4. Statistical analysis
This system is located in the flood plain of the Põltsamaa
river. The system is designed for the secondary treatment of The statistical analysis was carried out using the STATISTICA
wastewater from a conventional treatment plant, which con- 7.0 (StatSoft Inc.) program. The normality of the variables
sists of mechanical filters, sedimentation tanks and aeration was verified using the Lilliefors’ and Shapiro–Wilk W-tests.
tanks. It treats wastewater from the town of Põltsamaa (∼5000 Biomass production and nutrient content variables were nor-
260 e c o l o g i c a l e n g i n e e r i n g 3 5 ( 2 0 0 9 ) 258–264

Table 1 – The average annual (2002–2003) hydraulic (L m−2 year−1 ) and nutrient (g m−2 year−1 ) loadings of Tänassilma
semi-natural wetland and Põltsamaa and Häädemeeste CWs
Loading Wetlands

Tänassilma (69 ha) Põltsamaa (1.21 ha) Häädemeeste (0.95 ha)

Hydraulic (L m−2 year−1 ) 278 26436 10421


N (g m−2 year−1 ) 13.8 849.7 58.7
P (g m−2 year−1 ) 2.6 170.3 22.2

Table 2 – The pattern of phytomass sampling and analysis in the Tänassilma semi-natural wetland and Põltsamaa and
Häädemeeste CWs
Sampling and analysis Fractions Wetlands

Tänassilma Põltsamaa Häädemeeste

Autumn Winter Autumn Winter Autumn

Roots–rhizomes 15 – 15 – 10
Number of collected Litter 15 15
samples Shoots
Spadixes

Roots–rhizomes 2002–2006 – 2002–2006 – 2002–2006


Year of collected Litter 2003–2006 2003–2006
samples Shoots
Spadixes

Roots–rhizomes 15 – 9 – 10
Number of analysed Litter 4 6
samples Shoots
Spadixes

Roots–rhizomes 2002–2003 – 2002–2003 – 2002–2003


Year of analysed Litter 2005 2005
samples Shoots
Spadixes

mally distributed. The 95% confidence intervals were used and 0.07–0.35 kg m−2 and 0.09–0.48 kg m−2 correspondingly in
to compare mean values of results. The level of significance Tänassilma and Põltsamaa. The average biomass of spadixes
˛ = 0.05 was accepted in all cases. in Põltsamaa varied from 0.03 to 0.10 kg DW m−2 . The corre-
sponding results in Häädemeeste were 0.20–0.31 kg DW m−2 ,
and in Tänassilma 0.03–0.20 kg DW m−2 . The average phy-
3. Results and discussion tomass of spadixes in winter was 0.15 kg DW m−2 in
Häädemeeste, and the same results varied in Tänassilma
3.1. Biomass and Põltsamaa from 0.02 to 0.16 kg DW m−2 and from 0
to 0.35 kg DW m−2 , respectively. The average aboveground
The average aboveground biomass of T. latifolia varied biomass of reed did not vary significantly from cattail, but
from 0.37 to 1.76 kg DW m−2 in autumn and from 0.33 to there were more roots–rhizomes of reed than cattail.
1.38 kg DW m−2 in winter for the three wetlands throughout Up to 1 kg less aboveground phytomass of cattail was left
the study period. The corresponding results for P. aus- in winter than at the end of the vegetation period in autumn.
tralis were from 0.61 to 1.32 kg DW m−2 and from 0.61 to Phytomass loss was less when there were more spadixes in
1.02 kg DW m−2 kg, respectively (Fig. 1). autumn. This is because fruiting shoots are more weather-
Biomass measurement results showed that 1 ha of CW can resistant (Linde et al., 1976). There was half a kilogram less
annually produce enough raw cattail materials for the insula- aboveground reed phytomass in winter than in autumn. Cat-
tion of up to three houses and fibre material up to 25 houses. tail biomass has a large temporal and spatial variation in
The belowground biomass of the cattail varied from 0.61 to productivity. This makes it difficult to use them as a raw mate-
1.31 kg DW m−2 , and reed from 1.60 to 1.69 kg DW m−2 . rial for building and fibre production.
The differences in aboveground phytomass values between It is known that plants do not always bear fruit in stable
the studied wetlands in autumn were only significant quantities every year, but rather alternate between a year of
in Tänassilma in 2004 and Põltsamaa in 2006. The esti- high production and year of low production. It is possible that
mated biomass of cattail shoots varied from 0.56 to heavy fruiting of cattail may also produce an imbalance within
1.07 kg m−2 in Tänassilma, 0.27–1.58 kg m−2 in Põltsamaa, and the plant which affects the next year’s production (Linde et
0.47–1.21 kg m−2 in Häädemeeste. The phytomass of cattail al., 1976). There were more spadixes in Häädemeeste than in
litter was between 0.20 and 0.57 kg m−2 in Häädemeeste, Põltsamaa and Tänassilma in 2002 and 2003, but only a few
e c o l o g i c a l e n g i n e e r i n g 3 5 ( 2 0 0 9 ) 258–264 261

Fig. 1 – Aboveground phytomasses (kg m−2 ) of cattail (T. latifolia) and reed (P. australis) in Tänassilma semi-natural wetland,
and Põltsamaa and Häädemeeste CWs in autumn (September; a) and winter (January; w) from 2002 to 2006.

cattail shoots survived, and common reed colonised the free Average nitrogen and phosphorus standing stock was
water wetland surface in 2004. There were more spadixes in higher in aboveground than belowground phytomass (Table 3).
Tänassilma in 2003 than in 2002, but there were only a few The average standing stock of nutrients in Tänassilma, Põlt-
spadixes and low production of aboveground phytomass in samaa and Häädemeeste belowground phytomass varied
2004. Cattail population on floating mats creates less spadixes from 11.6 to 19.4 g N m−2 and from 1.6 to 4.6 g P m−2 and above-
because fruiting shoots are heavy, and they cannot stand up ground from 17.0 to 32.3 g N m−2 and from 2.6 to 6.0 g P m−2 .
if they are not fixed to the soil. It also caused a large loss of The corresponding results in winter were 4.4–7.5 g N m−2 and
aboveground phytomass in Põltsamaa in winter. 0.6–1.0 g P m−2 . The standing stock of nutrients depends on
The average aboveground cattail biomass values both the elements’ concentrations in the plant tissue as well
(0.3–1.8 kg DW m−2 ) in Tänassilma semi-natural wetland as on the amount of plant biomass (Vymazal, 2004).
and Põltsamaa and Häädemeeste CWs were lower than We found that N and P were stored in reserve organs
reported by Toet et al. (2005): 2.09 kg m−2 , Ennabili et al. (1998): after the fruiting stage. Stored nutrients are available to
2.16 kg m−2 and Fernandez and de Miguel (2005): 2.23 kg m−2 , help the plant develop new shoots the following spring. For
but were similar to those found in Germany (1.3–1.45 kg m−2 ; instance, Vymazal (2004) reported that P. australis translo-
Wild et al., 2002). The cattail roots and rhizome biomass cates reserve products very late in the season, and the
values (0.6–1.3 kg DW m−2 ) in the studied systems were sim- harvesting of this plant during the growing season may lead
ilar to those recorded by Romero et al. (1999), varying from to serious damage to the stand. Therefore, we recommend
0.7 to 1.6 kg m−2 , but were lower than those measured by that the harvesting of cattail be undertaken after nutrient
Ennabili et al. (1998): 3.5 kg m−2 . The average aboveground translocation.
standing stock values of reeds in the studied areas were The nutrient balance of treatment wetlands depends on
once again lower than those reported by Lesage et al. (2006): many factors and can vary from year to year. In warm and
1.5 kg m−2 , Vymazal (2004): 2.09 kg m−2 , Ennabili et al. (1998): sunny summers the accumulation of N and P in both below-
2.3 kg m−2 , Bragato et al. (2005): 2.5 kg m−2 and Toet et al. ground and aboveground phytomass can significantly increase
(2005): 2.85 kg m−2 . (Mander et al., in press). In subsurface flow CWs with low
hydraulic and nutrient loadings the nutrient removal can be
3.2. Nutrients significant, although we should also consider that the pool of
nutrients in filter material and sediments plays a more impor-
The greatest average nitrogen concentration (22.95 g N kg−1 ) tant role in phytomass production than direct wastewater
was found in spadixes in 2002, and phosphorus (6.5 g P kg−1 ) inflow. For instance, the Tänassilma system received signif-
concentration was measured in roots–rhizomes in 2003. Phos- icantly higher loadings from the 1950–2000 (Nõges and Järvet,
phorus concentration in litter, shoots and roots–rhizomes 2002), which is nowadays lowered to 13.8 g N m−2 year−1 and
was higher in 2003 than in 2002, but was lower in spadixes. 2.6 g P m−2 year−1 (Table 1). Therefore, the high nutrient accu-
For spadixes the maximum average P concentration was mulation in Tänassilma is largely thanks to the accumulated
measured at 4.76 g kg−1 , while in the case of litter the great- N and P in sediments. Thus the calculation of removal effi-
est P concentration was 2.7 g kg−1 . N and P concentrations ciency based only on initial wastewater loadings and annual
in cattail shoots and litter were lower in winter than in nutrient uptake does not provide adequate results. A simi-
autumn, but were higher in spadixes and roots–rhizomes lar situation can be seen in the Häädemeeste FWS CW with
(Table 3). low initial loadings. In Põltsamaa, on the other hand, both
262 e c o l o g i c a l e n g i n e e r i n g 3 5 ( 2 0 0 9 ) 258–264

Table 3 – The range (min–max) of N and P concentrations (in both soil and plant fractions; g kg−1 ) and standing stock
(g m−2 ) in cattail plant fractions in the Tänassilma semi-natural wetland and Põltsamaa and Häädemeeste CWs in
autumn and winter
Elements Fractions Wetlands

Tänassilma Põltsamaa Häädemeeste

Autumn Winter Autumn Winter Autumn

Soil 23.6–28.0 26.6–31.3 1.6–28.2 0.8–5.4 0.4–1.45


Roots–rhizomes 15.6–25.2 19.0–27.0 15.7–33.3 16.0–34.0 11.0–21.0
N (g kg−1 ) Litter 8.2–19.2 8.0–12.0 9.5–19.9 7.0–14.0 7.3–18.1
Shoots 12.7–24.1 14.7–27.4 12.8–23.1
Spadixes 16.3–23.8 13.0–21.0 11.6–27.1 10.0–21.0 10.9–21.8

Roots–rhizomes 7.0–26.3 – 6.9–23.0 – 5.2–30.6


−2
Litter 1.9–6.4 3.0–7.0 0.5–9.3 0.6–6.3 0.5–11.9
N (g m )
Shoots 10.4–32.2 6.4–61.6 2.5–46.2
Spadixes 0.5–6.3 1.2–4.5 1.3–4.4 0.5–0.6 0.3–11.3

Soil 1.2–4.4 1.9–5.6 0.9–6.3 0.5–1.5 0.2–1.1


Roots–rhizomes 1.2–4.6 2.8–4.3 2.5–8.6 3.7–5.2 0.9–5.9
P (g kg−1 ) Litter 1.6–3.7 0.6–1.3 1.3–3.8 0.2–1.9 0.9–2.9
Shoots 1.6–4.3 2.3–4.4 1.7–3.9
Spadixes 2.5–4.7 2.7–4.1 3.0–5.6 2.2–3.3 2.4–4.6

Roots–rhizomes 0.6–3.3 – 1.1–7.2 – 0.5–7.2


Litter 0.4–1.5 0.2–0.8 0.1–2.0 0.01–0.7 0.1–1.9
P (g m−2 )
Shoots 1.5–6.0 1.1–10.2 0.4–6.4
Spadixes 0.1–1.3 0.3–0.9 0.2–0.9 0.1–0.1 0.1–2.4

hydraulic and nutrient loadings are very high (Table 1), which 3.3. Suitable areas for energy and treatment wetlands
would result in low removal efficiency if the plants were har- in Estonia
vested: 0.5% of N and 0.3% of P of initial wastewater loadings.
Thus removed nutrients via the harvesting of aboveground The percentage of wet and moist land in Estonia is rel-
biomass from heavily loaded CWs in temperate and cold cli- atively high (Mander et al., 2001b). Peat soils cover 21%
mates yield a very small portion of the inflow load, usually as (9497 km2 ) of Estonian territory, and gleysols cover even
little as <1%, and harvesting does not usually increase removal more—33% (15,138 km2 ). Not all of this territory is accept-
efficiency (Vymazal, 2004). able for Typha plantation. After exclusion of not suitable
Harvesting may be feasible if there is an application for largely overlapping areal categories (forest: 21 198 km2 , Natura
macrophytes, e.g. construction (Mauring, 2003) or energy pro- 2000 network: 21,1053 ha, protected areas: 5919 km2 , special
duction (Mander et al., 2001b; Ciria et al., 2005). However, the areas of conservation: 7500 km2 , valuable habitats: 231 km2 ,
stability of cattail re-growth after harvesting has not been planned protected areas: 995 km2 settlements: 1365 km2 , and
thoroughly researched (Hellsten et al., 1999). 200 m wide coastal buffer zone at water bodies: 7684 km2 )

Fig. 2 – Potentially suitable areas for treatment wetlands for raw material production in Estonia (marked in grey).
e c o l o g i c a l e n g i n e e r i n g 3 5 ( 2 0 0 9 ) 258–264 263

5412 km2 of potential territory for energy and treatment wet- Bulc, T., 2006. Long term performance of a constructed wetland
lands remains. About 464,000 ha of the potential polygons are for landfill leachate treatment. Ecol. Eng. 26 (4), 365–374.
larger than 10 km2 (Fig. 2). About 464,000 ha of these are larger Calheiros, D.S.C., Rangel, A.O.S.S., Castro, P.M.L., 2007.
Constructed wetland systems vegetated with different plants
than 10 km2 . Presumably, cattail and reed plantation are not
applied to the treatment of tannery wastewater. Water Res. 41
the only possible land use for these potential areas. In existing (8), 1790–1798.
peat-mining areas, restoration should be undertaken after the Ciria, M.P., Solano, M.L., Soriano, P., 2005. Role of macrophyte
excavation of peat deposits. Thus, these areas can be consid- Typha latifolia in a constructed wetland for wastewater
ered as potential biomass production wetlands. treatment and assessment its potential as a biomass fuel.
Biosystems Eng. 92 (4), 535–544.
Ennabili, A., Ater, M., Radoux, M., 1998. Biomass production and
NPK retention in macrophytes from wetlands of the Tingitan
4. Conclusions
Peninsula. Aquat. Bot. 62 (1), 45–56.
Fernandez, J., de Miguel, E., 2005. Results of the LIFE-Environment
Average aboveground biomass of T. latifolia varied from 0.3 to project “Macrophytes”, executed in Lorca (Murcia, España). In:
1.8 kg DW m−2 , and the same results for P. australis were from Fernandez, J., Cirujano, S., de Miguel, E. (Eds.),
0.6 to 1.3 kg DW m−2 . Phytodepuration: Proc. Intern. Meeting. Llorca, Spain, pp.
The results showed that cattails in CW have a large tem- 151–157.
poral and spatial variation in productivity, which makes it Gottschall, N., Boutin, C., Crolla, A., Kinsley, C., Champagne, P.,
2007. The role of plants in the removal of nutients at a
difficult to use them as a potential source for building mate-
constructed wetland treating agricultural (dairy) wastewater,
rial production. Thus, further investigations in this field are
Ontario, Canada. Ecol. Eng. 29 (2), 154–163.
required because of the increasing need for Typha wool as a Hellsten, S., Dieme, C., Mbengue, M., Janauer, G.A., den Hollander,
valuable ingredient in clay plasters. N., Pieterse, A.H., 1999. Typha control efficiency of a
The nutrient uptake capacity of cattail was consider- weed-cutting boat in the Lac de Guiers in Senegal: a
ably high. The average N and P concentration was higher preliminary study of mowing speed and re-growth capacity.
in belowground than in aboveground biomass, whereas the Hydrobiologia 415, 249–255.
Hunt, P.G., Matheny, T.A., Szögi, A.A., 2003. Denitrification in
belowground stock of both nutrients was lower than the
constructed wetlands used for treatment of swine
aboveground stock. wastewater. J. Environ. Qual. 32 (2), 727–735.
The harvesting of aboveground biomass would have Kadlec, R.H., Knight, R.L., 1996. Treatment Wetlands. Lewis
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mined lands in east-central Ohio. Ohio J. Sci. 91 (4), 154–158.
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Lee, A.A., Bukaveckas, P.A., 2002. Surface water nutrient
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