RESEARCH ARTICLE
Natural Revegetation of a Boreal Gold Mine Tailings
Pond
Ian W. R. Young,1 Christian Naguit,1 Sara J. Halwas,1 Sylvie Renault,1 and John H. Markham1,2
Abstract
Understanding the natural revegetation of forests dis-
turbed by the dumping of mine wastes is vital for the
success of reclamation strategies. The Gunnar gold mine
tailings pond in southeast Manitoba has remained largely
unvegetated since the mine was closed in 1942, with lim-
ited vegetation developed on one side of the pond. We
examined the natural Picea mariana/Larix laricina for-
est that has developed on the pond to determine how the
plant community develops and what changes in the tailings
are associated with this development. Vegetation sampled
along transects showed a consistent pattern of succession
from Equisetum palustre to Salix spp., and Populus bal-
samifera, to Larix laricina and finally to P. mariana. Larix
laricina and P. mariana are moving into the site at the
rate of 1.5 m per year with L. laricina invading 4 years
Introduction
The main waste products of ore processing operations are
crushed rock material, referred to as tailings. The rate of
plant colonization of disturbed land is dependent on the
extent of the disturbance, the prevailing edaphic conditions,
and the presence of neighboring and residual vegetation
persisting since disturbance (Szarek-Łukaszewska 2009). In
the case of mine tailing dumps (hereafter referred to as ponds)
natural revegetation occurs at a very slow rate because of
their low nutrient status, poor aggregation, and low organic
carbon (Tordoff et al. 2000; Shu et al. 2005). Additionally,
highly acidic tailings ponds produced from sulfide-bearing ores
limit or entirely suppress revegetation because of the high
bioavailability of heavy metals (Winterhalder 1996).
Understanding the processes that occur during natural colo-
nization and succession on mine wastes is important for guid-
ing restoration programs (Alday et al. 2011a). Colonization
typically begins at the margins of tailings ponds from existing
vegetation (Shu et al. 2005). Wind-borne or animal-delivered
1 Department of Biological Sciences, University of Manitoba, Winnipeg, MB R3T
2N2, Canada
2 Address correspondence to J. H. Markham, email markhamj@cc.umanitoba.ca
© 2012 Society for Ecological Restoration
doi: 10.1111/j.1526-100X.2012.00913.x
Restoration Ecology
ahead of P. mariana. Both tree species show a similar pat-
tern of annual growth, showing positive correlations with
spring precipitation, a pattern also occurring on L. laricina
growing on a nearby site. The establishment of E. palus-
tre was accompanied by initially rapid decreases in com-
paction and conductivity of the tailings, and an increase in
inorganic nitrogen. Surface organic matter depth, coarse
organic matter mass, and soil organic carbon increased at
a constant rate, whereas subsurface coarse organic matter
had an initial rapid increase followed by a gradual increase.
As fern allies (and specifically members of the Equisetaceae
family) have a number of properties that facilitate succes-
sion on mine wastes, their use should be explored further.
Key words: Equisetum, mine tailings, revegetation, vege-
tation succession.
seeds and vegetative propagules from neighboring vegetation
provide pioneer plant species, establishing at pond margins and
favorable microsites (Shu et al. 2005). Therefore, the dispersal
mode of plants can strongly affect the trajectory of succession
(Alday et al. 2011b). Organic matter input to the tailings from
early successional plant turnover may help to stabilize the tail-
ings surface and allow for improved wind-borne sediment and
seed interception (Bradshaw 2000). The decomposition and
mineralization of deposited organic matter can fuel microbial
communities, and have a positive feedback on overall plant
health due to positive effects on soil structure and nutrient
cycling (Shrestha & Lal 2006). The early pioneer plant stage
will transition toward a woody community, accompanied gen-
erally by an increase in organic matter depth and the associated
nutrient pools (Smith et al. 1997).
Throughout the world there are hundreds of thousands of
mine sites where operators no longer exist (orphaned mines).
In these cases, the responsibility of site remediation of the
land, including tailings dumps, either falls to new operators,
governments, or of does not occur. The Province of Manitoba
(Canada) has approximately 150 abandoned or orphaned
mines, a few of which have mine tailings ponds. One of these,
the Gunnar gold mine in southeastern Manitoba, was in oper-
ation from 1936 to 1942 (Slivitzky 1996). Our previous work
(Renault et al. 2007) has shown that the chemical and physical
properties of the Gunnar tailings have prevented plant growth
on most of the pond. These properties included high bulk
1
Natural Revegetation of a Tailings Pond
density, low levels of nutrients (especially nitrogen), and low
organic matter content. The high level of elements (arsenic:
15 mg/kg1 , copper: 194 mg/kg, manganese: 1,185 mg/kg,
chromium: 111 mg/kg, aluminum: 5.4%, iron: 6.1%, vana-
dium: 162 mg/kg−1 , and zinc: 212 mg/kg−1 ) could also limit
plant growth and establishment as they exceed the Canadian
Interim Remediation Criteria for Soil (CIRCS) Agricultural
and/or Residential/Parkland guidelines (Slivitzky 1996). How-
ever, high acidity is not likely to be a concern for revegetation,
as it is on many tailings ponds, as the pH of the unvegetated
Gunnar tailings ranges from 6.4 to 8.5 (Renault et al. 2007).
The purpose of our study was to examine changes in both
tailings and vegetation associated with the natural revegetation
that has occurred on the exposed Gunnar mine tailings pond
since it was created over 70 years ago. Our hypothesis was
that natural succession will lead to more fertile conditions in
the tailings. This study provides insights into the changes that
need to be made in order to fully revegetate this mine site and
potentially other similar mine tailings sites.
Methods
Study Site and Experimental Design
The Gunnar mine tailings pond (50◦ 51.37 N, 95◦ 15.31 W)
cover 11 ha and contain 210,000 m3 of tailings. The elevation
of the pond varies by circa 1.8 m, being greatest at the west
end of the pond and grading to a wetland on the east, and to
a lesser extent toward the south. On the north and west side
of the tailings pond, vegetation from the surrounding forest
has invaded less than 10 m onto the tailings since the site was
abandoned in 1942, whereas on the southwest side of the pond
a Picea mariana forest has developed. The total forest zone
varies from 60 to 80 m in width. The remaining unvegetated
area of tailings is approximately 360 × 120 m.
Sampling
Between 2008 and 2009, nine transects were sampled along
the southwest side of the tailings pond running into the
natural vegetation. Transects were set up by selecting points
approximately every 30 m along the edge of the vegetation
(starting where less than 1% vegetation cover occurred)
running perpendicularly to the edge of the tailings pond.
The elevation along each transect and between transects was
measured every 2 m using a surveying transit.
The cover of vegetation was assessed along six of the
transects in 1 × 1 m plots every other meter the first 20 m of
the transect and every 3 m for the next 30 m. In every 1-m
section along the transect the closest P. mariana and Larix
laricina trees were selected within 5 m either side of the
transects. Selected trees were greater than 6 cm diameter at
breast height. Tree cores were collected 20 cm above ground
level. To compare the performance of trees on the site with the
surrounding area, an additional set of cores were taken from
a fen (wetland) dominated by L. laricina approximately 2 km
from the site. Trees were sampled over a 750 m transect with
2 Restoration Ecology
canopy dominant trees sampled circa every 10 m. In total, 63
L. laricina cores and 43 P. mariana cores were collected from
the tailings pond and 22 L. laricina cores were collected from
the nearby fen. Coarse organic matter (OM), that is, material
retained on a 2 mm sieve, was collected every 2 m along
the transects from an area of 0.0397 m2 in the litter layer
and separately to a depth of 20 cm from the tailings surface
(although there was a sharp separation between the tailings and
the litter layer there was no identifiable A layer). This material
was then dried to a constant mass at 65◦ C and weighed.
A hydraulic penetrometer (model HyPen1, Pike Agri-Lab
Supplies, Inc., Jay, ME, U.S.A.) was used to measure soil
compaction on all transects at a depth of 30 cm, to determine
the potential for roots to grow into the tailings. Measurements
were made at the same intervals as the vegetation and 3,
2 and 1 m before the start of the vegetated zone. Tailing
samples were also collected to 20 cm using a 2.5 cm diameter
corer for chemical properties. The electrical conductivity of
the air dried tailings was recorded with a Traceable Portable
conductivity meter (Fisher Scientific, Ottawa, Canada) using
a (1:1, V:V) water slurry method (Schofield and Taylor
1955). Inorganic nitrogen levels were measured using the
micro-diffusion method on 2M KCl extracts (Mulvaney 1996).
To determine the pH of the tailings a 0.01M CaCl2 solution
was added to the tailings (2:1, V:V). The organic matter of
the tailings was determined using dichromate oxidation (Kalra
& Maynard 1991).
Data Analysis
Tree cores were dried and sanded, and the width of rings was
measured to calculate growth increments. When determining
the year of recruitment, we assumed that trees were 1 year
older than the number of growth rings. Many of the cores
from the tailings pond had increments in the first 5 years of
growth with widths less than a standard deviation compared
to all other years. We interpreted this as suppression in the
understory. We therefore excluded these increments from the
analysis. For each core, the increment width was standard-
ized to standard deviation units and a mean of zero (Fritts
1976). Growth increments in years where less than 16 trees
of each species present were excluded from the analysis. To
account for any miscounted cores, a 3-year running average
(i.e. the previous, the present, and the following year) was
calculated for the mean increment values for each species
in each year. Variation in yearly growth increments between
L. laricina and P. mariana on the tailings pond, and between
L. laricina on and off the tailings pond were compared using
linear correlations. The mean growth increment and the year
of recruitment for each species were correlated with environ-
mental data from the nearest Environment Canada weather
station (Bissett, 51◦ 1.800 N, 95◦ 42.000 W, data available
from http://www.climate.weatheroffice.ec.gc.ca). We exam-
ined individual correlations between growth increment and
year of recruitment for the mean daily temperature from May
to September, mean daily temperature in April, mean daily
temperature in October, total precipitation from April to June,

total precipitation in July, and total precipitation in August.
As trees continued to recruit onto the site once other trees had
established, we used quantile regression (Cade & Guo 2000)
with the 90th percentile of the data to establish the relationship
between the distance along the transects and the maximum age
of trees. This relationship was used to estimate of the rate of
invasion onto unoccupied tailings. The changes in vegetation
were examined for the individual dominant species and for the
total cover of plant functional groups with plants classified as
mosses, ferns and allies, annuals, biennials, forbs, grasses and
sedges, and shrubs and trees. The diversity of plant species
was calculated using Simpson’s diversity index. Relationships
between the tailings variables and distances along the transect
were examined with least squares regression. Aikake Informa-
tion Criterion (AIC) was used to decide which model best fits
the data including null, linear, log transformed, and quadratic
models. Jump (version 8.01, SAS Institute) was used to con-
struct the models and perform the data analyses.
Results
All transects had a downward slope toward the edge of the
pond with a slope of 0.31 ± 0.14 cm/m (mean ± standard
deviation). However, the elevation at the start of each transect
varied by up to 69 cm and the average slope between adjacent
transects was 1.60 ± 1.53 cm/m. This suggests that consistent
changes along the transects are not solely due to the changes
in elevation along them as there is much more variation
between transects.
Tree age on the pond ranged from 7 to 49 years for Picea
mariana and from 9 to 48 years for Larix laricina (Fig. 1).
The distance along the transect had a significant effect on tree
age of both P. mariana and L. laricina. Although both species
continued to recruit into established vegetation, locations near
the start of the transects had only younger trees. Using the
upper 90th quantile of the data, the slopes of the regression
lines indicate that L. laricina and P. mariana have similar
rates of invasion, 1.45 m/yr and 1.56 m/yr, respectively. These
estimates are in agreement with the fact that the vegetation has
spread about 80 m in 66 years. The intercepts of the regression
suggest that L. laricina invades the site 5.7 years earlier
40
P. mariana
35
L. laricina
30
Tree age (yr)
25
20
15
10
5
0 from 17 families and 37 genera were observed along the
0 10 20 30 40
Distance (m)
Figure 1. Tree age along transects. Lines are quantile regressions on the
90th percentile for Picea mariana (dashed) and Larix laricina (solid).
Restoration Ecology
Natural Revegetation of a Tailings Pond
Figure 2. Standardized growth increments of Picea mariana (dashed
line) and Larix laricina (dotted line) growing on the Gunnar tailings
point and L. laricina growing in a nearby fen (solid line).
than P. mariana. We therefore assume that the vegetation is
colonizing the site at the rate of about 1.5 m/yr.
Both tree species showed the same trend in growth incre-
ment over time (Fig. 2). There was a significant (p = 0.017)
positive linear correlation in growth increments between
P. mariana and L. laricina (r = 0.390) on the tailings sites,
and a significant (p = 0.035) correlation between L. laricina
on the tailings pond and the nearby fen (r = 0.500). This sug-
gests that the same environmental conditions are required for
the growth of both species on the tailings pond and whether
or not the L. laricina are growing on the tailings or in nearby
wetlands. Both species on the tailings pond and L. laricina
off the tailings pond experienced reduced growth in the mid-
1990s. These were also years with low recruitment on the
tailings: no L. laricina recruitment occurred between 1991
and 1994 and no P. mariana recruitment occurred between
1989 and 1992. April and October mean daily temperatures
accounted for about half the variation in the growth increment
of trees both on and off the tailings (mean r = 0.454 ± 0.041
averaged across each combination of species, location, and
monthly temperature). Monthly precipitation was usually not a
significant predictor of growth; the only instance where there
was a significant correlation was between May precipitation
and P. mariana growth increment (r = 0.423, p = 0.029).
The cover of dominant species followed a consistent pattern
of distribution along the transects (Fig. 3). The cover of
Equisetum palustre rose rapidly within the first few meters
to a maximum cover of 52% and then gradually declined,
remaining present at less than 5% from 23 m onward. This
was followed by three Salix planifolia, which had an average
cover of 1% for the first 2 m and then reached a maximum
cover of 5%. This Salix species was absent by the 20 m point.
Populus balsamifera was absent for the first 4 m, reached peak
abundance at 8–12 m, then gradually declined and was absent
by 38 m. Larix laricina was absent for the first 8 m, reached
a peak abundance at 16 m, declining gradually until no trees
were present by 41 m. Finally, P. mariana was absent for the
first 4 m and continued to increase in abundance along the
transects up to a cover of 67%. A total of 45 plant species
50
transects (Appendix S1). The cover of plant functional groups
also followed a consistent trend across the transects. Only two
mosses were found, Pleurozium schreberi and Hylocomium
splendens with P. schreberi making up 98% of the total moss
3
Natural Revegetation of a Tailings Pond

Figure 3. Mean cover of dominant species and plant functional groups along transects.

cover. Mosses had less than 1% cover for the first 10 m and
then had an increase paralleling P. mariana. The ferns and
allies were represented by two Equisetum species, E. palustre
making up 95% of the cover. The two most abundant forb
species were Fragaria vesca and Pyrola asarifolia which
made up 45 and 39% of the forb cover, respectively. Total
forb cover rose to 26% at 18 m and then decreased to less than
1%. The most common shrubs were Salix species, making up
79% of the total shrub cover. Shrubs reached a maximum
cover of 21% at 8 m and then declined to less than 1%
by 35 m. Trees were absent from the transects for the first
2 m. The cover then rose to between 30 and 50%, being
made up of a mix of L. laricina and P. balsamifera. Past
38 m P. mariana became the dominant tree where it displaced
L. laricina and P. balsamifera and had a cover of over 60%.
Grass cover (not shown) reached a peak of 6% cover at 2
m, but otherwise ranged 1–3% across the transects. The only
annual and biennial found along the transects were Crepsis
tectorum and Sonchus arvensis, respectively, and they never
exceeded 1% cover. The total cover of legumes never exceeded
3% and they were absent from the first 10 and last 12 m of
the transects. The plant diversity increased along the transects
up to 30 m and then decreased (Fig. 4).
Tailings Physicochemical Parameters
The pH of the tailings was relatively homogeneous (7.3–7.7),
whereas the electrical conductivity and compaction decreased
significantly (p < 0.001 for both) along the transects, with
both showing a rapid initial drop followed by a more gradual

4 Restoration Ecology

Figure 4. Plant diversity along the transect.
decline (Fig. 5). The litter layer depth increased in a linear
fashion along the transects, although the depth also became
more variable with distance. In addition, there was a decrease
in litter layer depth in the zone dominated by P. balsamifera
(20–35 m). Coarse organic matter both in the litter layer and in
the tailings showed a significant increase along the transects.
The coarse organic matter in the litter layer increased linearly
and at a greater rate than the coarse organic matter in the
tailings that had a rate of increase that leveled off along the
transects. Soil carbon increased along the transects from 0.3
(a)
(c)
(e)
Figure 5. Variations in tailings properties along the transects. Tailings conductivity (a), compaction (b), surface litter layer (LL) depth (c), coarse organic
matter on the tailings surface and to a depth of 20 cm (d), soil carbon (e), and inorganic nitrogen (f). Only the best fitting model according to AIC is
shown.
Restoration Ecology
Natural Revegetation of a Tailings Pond
to 0.4%. Inorganic nitrogen levels along the transects could
not be described by any of the statistical models. However,
levels were always less than 6 mg/kg before the start of the
vegetation zone, the mean values at the 0, 2, and 4 m were
always greater than 20 mg/kg, after which the values varied
from 8.5 to 18.5 mg/kg for the remainder of the transect.
Discussion
The vegetation colonization at the Gunnar mine site follows
a pattern similar to other anthropogenically and naturally dis-
turbed forests (Smith et al. 1997), being colonized initially by
pioneer species (either woody or herbaceous) and eventually
succeeded by climax species. The transition from a Populus
balsamifera to a Picea mariana dominated forest is also typi-
cal of southern boreal forest succession on mesic sites (Frelich
& Reich 1995). However, most of the plant cover in this
study was made up of woody species recruiting on to the site
within the first few meters (or about 3 years) of the Equisetum
palustre colonization. In other mine successions, annuals make
up a much greater proportion of the early colonizing species
and dominate the site for longer, with woody species increas-
ing in cover at a much slower rate (Martinez-Ruiz & Marrs
2007; Alday et al. 2011b). In these other systems diversity
also peaked with the cover of annual species. On our study
(b)
(d)
(f)
5
Natural Revegetation of a Tailings Pond
site peak diversity was associated with the cover of peren-
nial forbs.
The initial colonization of the tailings by Equisetum spp.
resulted in the most rapid changes in soil properties and may
have facilitated plant succession. The presence of Equisetum
was associated with decreases in compaction and conductiv-
ity and an increase in soil inorganic nitrogen. Both decreased
bulk density and increased soil nitrogen have generally been
regarded as the processes associated with vegetation develop-
ment in newly exposed land (Crocker & Major 1955) and mine
waste sites (Wali 1999; Borden & Black 2005). Species in the
genus Equisetum have a number of properties that may allow
them to facilitate succession on mine tailings. Along with other
fern allies they have the ability to grow in metal rich substrates
(Cornara et al. 2007). Nitrogen fixation by associative dia-
zotrophs has also been found in their rhizomes (Uchino et al.
1994), which may result in increased soil nitrogen levels. Their
ability to move air through their rhizomes by convection could
provide a source of soil aeration (Armstrong & Armstrong
2009), which would be especially important in mine tailing
ponds given their poor physical structure. The ability to access
soil nutrients at great depths in the soil column may allow them
to act as nutrient pumps (Marsh et al. 2000). The high degree
of internal and external surface area and the low lignin con-
tent result in the rapid decomposition and mineralization of
Equisetum spp. litter (Marsh et al. 2000). The organic matter
from Equisetum shoot and root litter could also facilitate the
invasion of other plant species by chelating heavy metals (Shu
et al. 2005).
The development of vegetation on the site was accompa-
nied by a steady build up of organic matter, both on the
surface of and in the tailings. The newly formed litter layer can
provide several important modifications to the soil, through
surface structure protection, improved water retention, ther-
mal regulation, improved sediment and seed capture, and as
an energy source for saprophytic organisms (Bradshaw 2000).
Other studies suggest that increased organic matter can facil-
itate the recruitment of less heavy metal tolerant species on
mine tailings and other industrial wastes (Shu et al. 2005;
Szarek-Łukaszewska 2009; Wang et al. 2011). Soil organic
matter represents the major carbon and nitrogen reserves in
all terrestrial ecosystems (Smith et al. 1997) and accounts
for approximately 75% of the total terrestrial carbon pool
(Shrestha & Lal 2006). By returning unvegetated mine tail-
ings to their original undisturbed land use, large quantities of
atmospheric carbon dioxide can be sequestered and protected
(Shrestha & Lal 2006). Rates of carbon sequestration between
0.7 and 4 Mg C ha−1 yr−1 have commonly been observed in
forest ecosystems, and by initiating succession of these tail-
ings, atmospheric carbon can be effectively stored, thereby
reducing its environmental harm (Shrestha & Lal 2006). On
naturally and anthropogenically reclaimed mine spoils, rates
of carbon sequestration between 0.5 and 1.0 Mg C ha−1 yr−1
have been obtained (Shrestha & Lal 2006). Therefore, unveg-
etated mine wastes represent potential sinks for atmospheric
carbon emissions, should they be returned to their natural land
use. Given the deep litter layer that has developed on the older,
6 Restoration Ecology
naturally revegetated portion of the Gunnar mine tailings site,
the total amount of stored carbon needs to be assessed.
The other tailing property that modified was a reduction
in soluble salts (as measured by electrical conductivity). The
overall values of electrical conductivities were lower than
those recorded for the tailings of the nonvegetated zone
(287–360 mS/m2 , Renault et al. 2007). Mine wastes are
typically high in soluble salts owing to the dissolution and
weathering of carbonate and sulfide minerals (Borden &
Black 2005). Weathering and dissolution rates decrease with
time, and under the proper climatic conditions, leaching and
percolation allow for the removal of excess salts from the
soil solution. Another study examining potential mine soil
factors limiting early growth in Pinus strobus found that the
most influential chemical characteristic on tree growth was
soil soluble salt content (Torbert et al. 1988). This finding
agrees with the observation that weedy plant communities
tended to persist for a longer time period on naturalized
sites with high soil conductivity, while soils with lower salts
transitioned to a woody community more rapidly (Borden &
Black 2005).
The annual growth of trees on the mine site followed the
same pattern as trees growing off the site. Our results sug-
gest that the growth of selected trees is strongly dependent
on spring precipitation. Excessive precipitation in the spring
leaves the vegetated portion of the Gunnar tailings flooded
because of the elevation of the site and the inability of tailings
to promote drainage because of its compact nature. Flood-
ing creates anoxic conditions, which can reduce tree growth
because of decreased root respiration, organic matter decom-
position, and N mineralization. Growth studies of L. laricina
and P. mariana in peatlands have shown similar results where
poor drainage and high water tables can reduce tree growth,
while the opposite is true for peatlands where the water table
is lowered (Lieffers & MacDonald, 1989; MacDonald & Yin
1999). Combined with the rapid colonization of vegetation, the
tree growth data suggest that the site quickly starts to mimic
the growth of surrounding forests.
Regardless of the geographic setting of the anthropogeni-
cally disturbed substrate, biological and physical processes,
individually and interactively, contribute to improvements in
edaphic conditions that facilitate the progression of primary
succession (Bradshaw 2000). Techniques that draw on these
natural biological and physical processes have been suggested,
and incorporated into many technical restoration efforts world-
wide (Prach & Hobbs 2008). For example, natural invasion
can be accelerated and facilitated through active seeding pro-
grams where native seed is imported from adjacent areas to
account for the poor initial survival that is common to dis-
turbed substrates (Bradshaw 2000). Alternatively, or in addi-
tion to improving seed availability, substrate physical and
chemical properties may be amended to achieve the benefit
of years of vegetative growth and biomass turnover. Technical
restoration efforts may amend the disturbed substrate with low
cost organics or fertilizers in order to restore the substrate to
levels of fertility sufficient for plant establishment. Further-
more, mechanical treatment of compacted substrates through

various ploughing techniques can accelerate the naturalization
process, potentially bypassing the lengthy process of bulk den-
sity reduction normally accomplished through the action of
plant roots, soil biota, as well as abiotic processes such as
freeze-thaw and wet-dry cycles (Bradshaw 2000).
According to our data, it would take between 70 and 110
years for the site to be completely and naturally revege-
tated, assuming that the conditions of the nonvegetated tail-
ings are similar to the vegetated area, and that environmen-
tal conditions remain favorable for plant growth. The time
required to return to a community that approaches the undis-
turbed condition will vary tremendously between mine wastes
owing to their highly variable composition and toxicity. Smith
et al. (1997), while studying the natural afforestation of iron-
smelting slag, found that after 75 years the P. strobus forest
community that had developed was floristically similar to nat-
ural 60–80-year-old secondary growth forests of the area.
Several studies have shown that certain edaphic characteris-
tics of drastically disturbed forest ecosystems have the capacity
to return to near pre-disturbance, or reference site conditions,
given sufficient time (Anderson 1977; Smith et al. 1997).
However, it is important to note that while some edaphic
factors may return to natural levels quite rapidly (e.g. sur-
face organic carbon, microbial carbon, conductivity), others
can take hundreds of years to reestablish (e.g. organic car-
bon at depths and carbonate content), and some factors may
never have the potential to return to an undisturbed condition
(e.g. rockiness and depth to hardpan) (Schafer et al. 1980;
Insam & Domsch 1988). The rapid colonization found on this
site suggests that the facilitation effects of Equisetum species
may be of great benefit in establishing vegetation on mine
tailings.
Implications for practice
• Natural colonization of mine tailings can be achieved by
modification to their chemical and physical structure.
• Increases in surface litter allow for the establishment
of a more diverse plant community and accelerate
the progression toward the regional climax vegetative
community.
• The use of fern allies should be explored for use in the
remediation of compacted, nutrient poor, or metal rich
substrates.
Acknowledgments
This work was funded by the Manitoba Geological Survey
(MGS) and NSERC. Thanks to Manitoba Conservation for
the work permit (WPB 25041). We also would like to
thank R. Sheffield, C. Szczerski, S. Kunkel, J. Newediuk, J.
Christensen, K. Kostyra, and L. Smith for field assistance.
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Supporting Information
Additional Supporting Information may be found in the online version of this
article:
Appendix S1. Species present along transect and their functional group
classification.
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