CH 13

C&S in Pastures Chap 13 2/11/01 8:56 am Page 261
13 Competition and Succession in

Re-created Botanically Diverse Grassland
Communities
Ross Chapman*
CSIRO Plant Industry, Centre for Mediterranean Agricultural Research, Wembley,
Australia
Introduction Despite their acknowledged environmental val-

ues, agricultural intensification over the 20th cen-
Since clearing of native forests began in prehistoric tury has led to the widespread loss of these habitats
times, species-rich grasslands have typically consti- from many areas. This loss of habitat has been par-
tuted a major habitat within the landscape of tem- ticularly acute in lowland regions and has endan-
perate regions of Europe (Green, 1990; Rychnovská gered many of the once widespread constituent
et al., 1994). The maintenance of this distinct suc- flora and fauna (Perring and Farrel, 1983; Shirt,
cessional stage or sere depends upon continuous 1987; Batten et al., 1990). In recent years, there has
human intervention to arrest or deflect otherwise been growing concern about the implications of the
natural successional processes. This intervention loss of such diversity. In 1992, both the European
usually occurs in the form of either grazing with Community (Council Directive 92/43/EEC) and
domestic livestock or haymaking. Because their the United Nations (UNCED, 1993) indepen-
evolution and preservation are directly linked to dently took steps to protect and restore biodiversity
human activities, these grasslands are described as in threatened habitats. Within Great Britain, recog-
plagioclimax or semi-natural communities. nition of both the environmental value and the
Such grasslands have traditionally supported endangered status of traditional species-rich grass-
the production of a number of livestock-based land led to the establishment of specific schemes to
agricultural commodities, including meat, milk protect and re-create these communities within the
and wool. In addition, these communities provide agricultural landscape (MAFF, 1992). However, the
a range of important environmental functions for process of agricultural intensification has changed
the wider ecosystem (Fig. 13.1). They provide a many characteristics of both the soil and the land-
habitat not only for a wide range of constituent scape. These modified conditions alter the competi-
flora (Rodwell, 1991), but also for a diverse selective and successional processes that occur within
tion of vertebrate and invertebrate fauna (Curry, grassland communities, and this creates substantial
1987; Green, 1990). Furthermore, these grasslands problems for the successful restoration of such
enhance the aesthetic qualities of the rural land- botanically diverse grassland communities.
scape and so increase the recreational and amenity Engineering works, such as those associated
values of the countryside (Green, 1990; Hopkins with mineral extraction and the construction of
and Hopkins, 1994; Fig. 13.2). road and rail networks, are also responsible for a
*Present address: c/o Orchard House, Carlton Scroop, Grantham, UK

© CAB International 2001. Competition and Succession in Pastures
(eds P.G. Tow and A. Lazenby) 261
C&S in Pastures Chap 13 22/10/01 1:39 pm Page 262
262 R. Chapman
substantial environmental disturbance and habitat operations. These problems arise from not only the
loss. Within Great Britain, the Department of the internal conditions of the modified ecosystems, but
Environment has recognized the potential for also the nature of the linkages between that ecosys-
nature conservation within strategies for the recla- tem and the surrounding landscape. The environ-
mation of land disturbed by such operations mental and ecophysiological origins of these
(DOE, 1989a,b, 1991). As a response, many constraints are discussed and some potential solu-
restoration plans now include a component for tions are offered.
nature conservation, and this frequently constitutes
the re-creation of species-rich grasslands. However,
soils and landscapes of reclaimed sites are typically The Impact of Habitat
heavily disturbed. This creates a number of prob-
lems for both the successful establishment and the
Productivity and Grazing
subsequent maintenance of these grasslands that are Livestock on the Composition and
very different from those experienced within inten- Diversity of a Community
sive agricultural systems.
This chapter reviews the problems imposed by Habitat productivity
competition and succession in botanically diverse
grasslands re-created in temperate north-western Habitat productivity may have a profound effect
Europe on both extensified agricultural land and upon both the botanical composition and the
sites restored after disturbance during engineering diversity of a grassland community.
Milk
Meat Fibre
Feed
production
Agricultural
values
Botanically
diverse
grassland
communities
Environmental
values
Habitat Aesthetic
for flora values
Recreational
Habitat values
for
fauna
Amenity
values
Invertebrates Vertebrates
Fig. 13.1. Schematic representation of the agricultural and environmental values of botanically diverse
grassland communities.
Re-created Botanically Diverse Grassland Communities 263
gated through nutrient competition (Grime, 1979).

The physiological basis by which ‘stress tolera-
tors’ come to dominate under conditions of low
productivity is still to be established. Some have
argued that these species have a superior capacity to
compete for nutrients when under very low levels of
supply (Tilman, 1982; Goldberg and Novoplansky,
1997). Others, in contrast, have maintained that
their dominance is simply due to the adoption of a
growth strategy that creates a reduced demand for
these resources while minimizing losses (Grime,
1979; Vázquez de Aldana and Berendse, 1997).
Fig. 13.2. A visually attractive species-rich grassland
community in the Yorkshire Dales, northern England.
Botanical diversity
Botanical composition
Following observation from numerous habitats, Al-
The type of species present within any grassland Mufti et al. (1977) proposed that a humpback rela-
community is greatly influenced by the fertility and tionship existed between the productivity and the
productivity of that particular habitat (Grime, botanical diversity of a vegetative community (Fig.
1979). Under conditions of high productivity, as 13.3). According to this model, at very high produc-
typically occur on fertile soils with high levels of tivity (greater than 1000 g dry matter (standing crop
nutrient supply, communities are typically domi- plus litter) m−2), aggressive competition excludes all
nated by fast-growing species, which rapidly estab- but the most competitive species and this imparts a
lish a tall canopy, such as Dactylis glomerata and very low level of diversity to the community. At
Lolium perenne. These species have been described reduced levels of productivity, stresses create gaps
as possessing a ‘competitive’ vegetative strategy within the matrix of competitor species which can be
(Grime, 1979). In contrast, habitats of very low exploited by less competitive species. At extremely
productivity, such as occur on very infertile soils, low levels of productivity (less than 200 g dry matter
tend to be dominated by very slow-growing, ‘stress- m−2), only the most stress-tolerant species can sur-
tolerating’ species, such as Molinia caerulea and vive, which again induces very low levels of diversity
Festuca ovina (Grime, 1979). Between these two in the community. The greatest diversity occurs at
extremes lies a continuum of productivity niches. low to intermediate levels of productivity (350–750 g
Although the mechanisms involved in the dis- dry matter m−2), where the greatest overlap of ‘com-
persal of species along this productivity gradient are petitor’ and ‘stress tolerator’ niches occurs.
still debated (Wilson and Tilman, 1993; Goldberg Various workers have subsequently attempted to
and Novoplansky, 1997; Vázquez de Aldana and verify this relationship in grassland communities.
Berendse, 1997), it is clear that some degree of com- Smith (1994) combined data from three contrast-
petition is involved. Willems (1983) asserted that ing sites and found no link between botanical
fast- and tall-growing species quickly come to domi- diversity and productivity or fertility. Oomes
nate the canopy under productive conditions associ- (1992), in contrast, compared botanical diversity
ated with high fertility. This leads to the rapid and productivity in 27 grassland sites, and found a
extinction of photosynthetically active radiation as it humpback response very similar to that proposed
passes through the upper layers of the canopy. As a by Al-Mufti et al. (1977). This relationship is fur-
consequence, less radiation is available for species of ther substantiated by the findings of Vermeer and
shorter stature, which exist lower in the canopy, and Berendse (1983) and Willems (1983).
the community becomes dominated by just a few When considering the relationship between
tall and fast-growing ‘competitor’ species. botanical diversity and productivity, Marrs (1993)
Furthermore, it has been alleged that these competi- and Pegtel et al., (1996) both noted that, while the
tor species may be quick to establish an extensive humpback model of Al-Mufti et al. (1977) may
root system, which, even under conditions of high generally prevail, the exact relationship may be site-
fertility, will efficiently deplete all available nutri- or vegetation type-specific and that the form of the
ents, thus causing less aggressive species to be subju- response may well be modified by such influences
264 R. Chapman
Species density (number of species 0.25 m–2) 30
20
10
0
0 200 400 600 800 1000 1200 1400 1600 1800 2000 2600
Maximum standing crop + litter (g m–2)
Fig. 13.3. The relationship between maximum standing crop plus litter and species density of herbs at 14
sites in northern England. grasslands; woodlands; tall herbs. (Reproduced from Al-Mufti et al., 1977.)
as soil pH levels and the intensity and frequency of community. That animals are selective in their graz-
disturbances. ing habit is well established (e.g. Gibb et al., 1989).
This behaviour creates spatial heterogeneity in the
canopy architecture of the community, with the
The impact of the grazing animal mean canopy height being shorter in the more fre-
quently grazed patches than in the less grazed
The composition and diversity of the community patches. This difference in canopy architecture cre-
may also be profoundly influenced by grazing live- ates niches for plants of contrasting growth habit
stock. In particular, animals may significantly influ- (Putman et al., 1991) and leads to the establishment
ence the physical structure of the community of a mosaic of subhabitats within the pasture com-
through the partial or complete destruction of the munities. Furthermore, excretion of dung and urine
canopy of competitive and dominant species. This by grazing animals leads to a localized accumulation
may arise either directly, through the effects of of nutrients. The elevated fertility of the affected
defoliation, or indirectly, through the effects of areas will favour more competitive species, such as L.
treading, urine scorch and burrowing or scraping perenne and Trifolium repens. The patchy distribution
by livestock (Grime, 1979). The effect of this dis- of excreta will create further subhabitats, which add
turbance is to create a niche which opportunistic to the spatial diversity of the pasture community.
‘ruderal’ species can exploit free from competition
(Grime, 1979; Smith and Rushton, 1994). These
ruderal species are typically fast-growing and yet Conclusions
relatively uncompetitive annuals or short-lived
perennials that are capable of producing high seed It has been argued that the peak in botanical diver-
yields, such as Bromus mollis or Medicago lupulina. sity occurs when the physical structure of the sward
Maintenance of maximum diversity in a grassland allows the niches of competitor, stress tolerator and
community therefore requires regular disturbance ruderal species to overlap. This occurs under condi-
by grazing animals in order for ruderal strategists to tions of low to moderate productivity coupled with
be retained alongside the competitor and stress tol- regular livestock-induced disturbances. Under these
erator species (Smith and Rushton, 1994). conditions, competitor species are able to form an
Grazing animals can also influence the spatial extensive but spatially incomplete matrix across the
diversity and botanical composition of a pasture community. Interstices are maintained within this
matrix by a combination of stresses, such as nutri- intensification. Fuller (1987) has accurately detailed
ent deficiencies, and disturbances, such as poaching this loss of habitat from lowland England and
by grazing animals. It is these gaps that provide the Wales. In 1932 there existed approximately 7.2 mil-
niches which stress-tolerating and ruderal species lion ha of unimproved semi-natural grassland habi-
exploit. Maintenance of the highest levels of diver- tat within this region. By 1984 there were only
sity therefore requires that fertility and disturbance 0.6 million ha of unimproved lowland grasslands
levels are finely balanced to ensure that species dis- (Fig. 13.4), which represents a habitat loss of some
playing all three vegetative strategies can coexist. 92%. The process of intensification has affected the
The restoration and subsequent maintenance of grassland habitats in several ways.
sustainable botanically diverse grasslands will thus
require careful management of both fertility and Fertilizer applications
grazing to achieve the correct level of productivity
and vegetation disturbance. Traditional management of grasslands excluded the
use of mineral fertilizers (Green, 1982; Archer,
1985; Fuller, 1987). Instead, the avoidance of
nutrient depletion depended essentially on biologi-
The Re-creation of Botanically cal nitrogen- (N2-) fixation, the recycling of dung
Diverse Grasslands on Formerly and urine from stock grazing on pastures and, in
Intensive Agricultural Land some instances, on the return to meadows of farm-
yard manure collected from stock housed over win-
ter (Smith and Jones, 1991; Younger and Smith,
The impact of agricultural intensification on 1994). This practice tended to maintain habitat fer-
botanically diverse grassland communities tility at the levels appropriate for the maintenance
of high levels of botanical diversity (Wilkins and
The 20th century has witnessed a catastrophic loss Harvey, 1994).
of botanically diverse semi-natural grasslands from Over recent decades, the adoption of mineral fer-
temperate regions of north-western Europe. This tilizers as a tool to manage grasslands has become
has largely been as a consequence of agricultural more widespread and it has been estimated that by
8
Area of unimproved grassland (millions of hectares)
0
1932 1984
Fig. 13.4. The decline in unimproved semi-natural grassland in England and Wales between 1932 and 1984
(data extracted from Fuller, 1987).
266 R. Chapman
the mid-1980s over 85% of all grasslands from this sexual reproduction to sustain their populations
lowland England and Wales had been affected by (Smith and Jones, 1991). Modernization of grass-
mineral fertilizer use (Elsmere, 1986). The increased land management has seen silage replacing hay as
use of mineral fertilizers clearly benefited the agricul- the more common method for conserving winter
tural productivity of these grasslands. However, feed (Hopkins and Hopkins, 1994). Silage produc-
applications of even small quantities of mineral fer- tion characteristically involves defoliating the sward
tilizers to these botanically diverse grassland commu- very much earlier. This practice of early and more
nities encourages the dominance of fast-growing, frequent defoliation prevents many species from
aggressive and competitive species. In particular, the successfully setting seed and leads to their gradual
abundance of tall-growing grasses, such as extinction, thereby reducing the community’s
Brachypodium pinnatum, Arrhenatherum elatius, D. botanical diversity (Smith et al., 1996).
glomerata and L. perenne, is increased (Kirkham et
al., 1996; Smith et al., 1996; Willems and van
Extensification of grassland management
Nieuwstadt, 1996). Aggressive growth by these com-
petitor species quickly excludes slower-growing, less
The adoption of more intensive systems of grass-
aggressive species, thus reducing the botanical diver-
land production, such as the application of mineral
sity of the community (Van Hecke et al., 1981;
fertilizers, the installation of field drainage systems
Berendse et al., 1992; Mountford et al., 1993; Smith,
and the adoption of silage making have seriously
1994; Kirkham et al., 1996; Smith et al., 1996).
degraded the botanical diversity of grassland com-
munities. The adoption of more extensive manage-
Drainage ment practices will therefore be fundamental to any
attempt to re-create these communities. However,
A fundamental step in the intensification and
experiments investigating the impact of extensifica-
‘improvement’ of the agricultural landscape has fre-
tion, such as the cessation of all applications of fer-
quently been the installation of a field drainage sys-
tilizers or manures, brought little immediate
tem (Fuller, 1987). An effective drainage system
improvement in the diversity of the communities
will lower the water-table beneath the grassland
studied (Olff and Bakker, 1991; Mountford et al.,
community. A direct consequence of this will be
1994). These failures were caused by: (i) residual
the loss of any species associated with wet or water-
fertility maintaining productivity and competition
logged conditions, such as Caltha palustris and
at undesirable levels; and (ii) slow rates of succes-
Glyceria fluitans. Furthermore, the lower water-
sional change, due to limitations in the rate of
table will result in increased levels of soil aeration.
recruitment of additional species into the extensi-
This will enhance the rate of soil microbial activity,
fied community. These studies suggest that a suc-
organic matter breakdown and the cycling of nutri-
cessful restoration plan will require the inclusion of
ents in general and N in particular (Berryman
practices that actively reduce productivity, through
1975; Oomes et al., 1997). The consequent
manipulation of nutrient cycles and soil fertility,
increase in nutrient supply raises the productivity
and the enhancement of the recruitment and suc-
of the community. This will lead to the displace-
cessional processes. The problems and potential
ment of slow-growing, stress-tolerating species by
solutions associated with productivity adjustment
more competitive species, thus suppressing the
will be discussed individually in the next section.
diversity of the habitat.
The impact this has on successional development
and community composition will be discussed
Defoliation practice immediately afterwards.
Traditional grassland management practices in tem-
perate Europe typically included the production of Manipulation of nutrient cycles and soil
a summer hay crop, which was conserved and used fertility
as winter feed. This practice provided a defoliation-
free period, lasting from late spring through to Excess fertility may be reduced by instigating a net
summer. This allowed many of the constituent flow of nutrients out of the restored ecosystem.
species to flower and set seed, a process vital for the This may be achieved effectively through the
maintenance of many species that depend upon removal of either vegetation or nutrient-rich top-
soil. Alternatively, a similar effect may be achieved yielded approximately 12 t dry matter ha−1 year−1
by slowing the rate at which nutrients are cycled at the start of the experiment. After 10 years of
within the ecosystem. Various management options unfertilized cut-and-clear management, productiv-
are available to meet these objectives. ity on the sandy soil had declined to approximately
5 t dry matter ha−1 year−1, while on a clay-on-peat
Vegetation management soil productivity remained between 6 and 8 t dry
matter ha−1 year−1. The successional responses that
Supply of nutrients to any plant community may followed these changes in production will be dis-
be considered as a dynamic process, influenced by cussed later.
the rates of nutrient input from external sources, Other investigations have examined the impact
internal nutrient cycling through constituent of cut-and-clear practices on the particular nutrient
plant, animal and microbial components of the cycling processes operating within contrasting com-
system, and nutrient exports. Perhaps the simplest munities in more detail (Table 13.1). These results
way to reduce the chemical fertility and excess pro- collectively show that the cutting and clearing of
ductivity of a grassland habitat would be to insti- vegetation from unfertilized but formerly inten-
gate a net export of nutrients out of the system by sively managed grasslands led to a fertility decline
both stopping all further inputs, while simultane- as a result of nutrient export. The general trend was
ously cutting and removing all herbage produced for yield-limiting deficiencies of potassium (K) and
at that site. Several experiments have investigated N to be induced within 2–9 years after the intro-
the efficiency with which such practices reduce the duction of these practices, but the development of
productivity of grassland communities. Bakker yield-limiting phosphorus (P) deficiencies took
(1989) successfully demonstrated that the cutting about 20 years. These contrasting responses may be
and removal of a hay crop from a formerly inten- attributed to the differing methods by which nutri-
sively managed grassland led gradually, over a ents are cycled within grassland systems and are dis-
number of years, to a decline in productivity. Olff cussed below.
and Bakker (1991) confirmed these observations
and reported that practices that involved taking POTASSIUM CYCLING. The cation exchange sites of
two cuts of hay per year gave a greater rate of pro- a soil may readily adsorb K ions from the soil solu-
ductivity decline than those based around the pro- tion and retain them in a form that is immobile
duction of a single hay crop. Similar findings were and yet still available for plant uptake (Russel,
again reported by Berendse et al. (1992). The latter 1973). However, soils with a low number of cation
authors, however, found that the rate of decline in exchange sites, such as those with a low clay con-
productivity induced by a cut-and-clear manage- tent, have a limited ability to retain K in this form;
ment practice varied according to soil type. In any excess K ions remaining in the soil solution are
their experiment, annual herbage productivity was therefore prone to leaching from the soil profile
compared on two contrasting sites, one on a sandy (Olff and Pegtel, 1994).
soil and one on a clay-on-peat soil. Both sites Vegetation management also has a profound
Table 13.1. Soil impoverishment and the induction of production-limiting nutrient deficiencies by cut-and-
clear practices.
Authors’ soil Apparent induction of Nutrients remaining

Study description nutrient deficiencies unlimited
Oomes (1991) Sand K after 9 years P and N

Olff and Pegtel (1994) Sand K and N after 2–6 years;
P after 19 years
Oomes et al. (1996) Peat K after 10 years P and Na
Pegtel et al. (1996) Gley podzol N after 3 years; K (moderate)
after 3 years; P after 20 years
aN remained unlimited in this study; this is possibly due to the installation of a field drainage system
artificially enhancing the N mineralization rate in peaty soils (Oomes, 1991).
268 R. Chapman
impact on the cycling of K. The uptake of K by additional 20 kg mineral N ha−1 year−1 in some
plants is regulated principally by supply rather than industrialized regions of Europe (Oomes et al.,
demand; thus, when this nutrient is available in 1997). These processes will affect the time it takes
sufficient quantities, the vegetation will readily for the cut-and-clear process to deplete the soil N
absorb more than is required for growth (Robson et supply.
al., 1989), a phenomenon known as ‘luxury The processes by which K and N are cycled
uptake’. Under many management practices, such within grassland ecosystems are therefore quite dif-
as grazing (Holmes, 1989), mulching (Oomes, ferent. The relative speed with which either nutri-
1991; Oomes et al., 1996) and manure recycling ent reaches a state of production-limiting deficiency
(Lecomte, 1980; MAFF, 1982), this plant-absorbed under a cut-and-clear management will depend
K is returned to the soil and retained within the upon the initial size of the nutrient pool at the start
grassland ecosystem. However, if the nutrients con- of the process and the ability of the site to buffer N
tained within the vegetation are exported from the losses. These phenomena are likely to be site-
system entirely, the phenomenon of luxury uptake specific and may explain why some studies found
will lead to the rapid depletion of plant-available K, cut-and-clear processes to induce K deficiencies
giving the observed rapid establishment of produc- before N (e.g. Oomes, 1990), while others found
tion-limiting deficiencies. the opposite to be the case (e.g. Pegtel et al.,
1996).
NITROGEN CYCLING. The cycling of N within
grassland systems is a complex process (Whitehead, PHOSPHORUS CYCLING. Mineral P applied in fer-
1995). A large proportion of soil N may be tilizers to grassland ecosystems is rapidly but
retained in the immobile and unavailable organic reversibly converted into several forms that are
fraction of the soil. Organic N is gradually unavailable for plant uptake (Fig. 13.5). Some P
degraded by microbial activity to release ammo- may be incorporated into inorganic complexes,
nium (NH4+). Like K, NH4+ is freely adsorbed by some may be adsorbed on to colloidal surfaces and
soil cation exchange sites, where it may be pro- some may become incorporated into the soil
tected from leaching and remain available for plant organic matter through plant and microbial
uptake. This NH4+ may, however, be oxidized to processes (Gough and Marrs, 1990). These
form nitrate (NO3−). This form of N is poorly processes are largely reversible and the equilibrium
adsorbed by soil particles and is very prone to point depends critically upon the concentration of
leaching losses. P in the soil solution. Intensive agricultural man-
Plants are able to absorb N only from the soil’s agement, therefore, tends to establish a substantial
pool of mineral N (NH4+ and NO3−). The fast grow- pool of unavailable P. Following the cessation of
ing and highly productive species that are common fertilizer applications and the commencement of a
in intensified grassland communities are able to cut-and-clear management, the vegetation will
accumulate N in large quantities within leaf and begin to deplete the soil’s pool of plant-available P.
stem tissues. Because of this, cut-and-clear practices As this occurs, additional P will tend to be
are therefore able to rapidly remove large quantities returned to solution from the various unavailable
of mineral N from recently extensified grassland pools within the soil which effectively buffers the
communities that are dominated by more competi- decline in the pool of plant-available P (see Fig.
tive species (Olff and Pegtel, 1994). 13.5; Marrs, 1993). Because prolonged intensive
A combination of high plant uptake of mineral agricultural management establishes considerable
N and leaching of NO3− may therefore lead to a reserves of unavailable P within the soil, a cut-and-
rapid depletion of the soil mineral N pool. clear process may take many years before it is able
However, the main determinant of soil N fertility is to induce P productivity limitations within exten-
the size and mineralizability of the soil organic N sified grasslands (e.g. Marrs, 1993; Pegtel et al.,
pool, which may return as much as 200 kg N ha−1 1996).
year−1 to the mineral N pool (Berendse et al.,
1992). Reducing the soil N supply is therefore a
Topsoil removal
longer-term process, which depends upon the
depletion of the soil organic N pool. Furthermore, Plant nutrients are typically concentrated in the
pollutants contaminating rainwater may return an surface soil; concomitantly, the microbial popula-
(a) tions, which are critical to nutrient cycling, are also

concentrated in the same upper layers of soil
Fertilizer
P (Leeper, 1973; Russel, 1973). Several workers have
explored the inclusion of topsoil removal in restora-
tion schemes as a tool for lowering soil fertility and
community productivity. Berendse et al., (1992)
found that stripping the topsoil from a previously
Available/ intensive agricultural grassland did indeed lower
Colloids soluble Complexes
the productivity of the habitat.
P
A similar study reported by Oomes et al. (1996)
examined the impact of topsoil removal on nutrient
cycling. They reported that supplies of both N and
P were substantially reduced (Table 13.2) and that,
unlike cut-and-clear processes, this reduction in
Organic
P chemical fertility was almost immediate. This was
probably achieved through both the removal of a
significant component of the soil nutrient reserve
and a reduction in the rates of cycling through
(b)
Removal modified levels of microbial activity. Furthermore,
of this reduction in nutrient supply was accompanied
vegetation
by a lower level of biomass production (see Table
13.2).
While topsoil removal may be an effective way
to rapidly reduce the productivity of a site, the
Available/ operation will require substantial use of plant and
Colloids soluble Complexes equipment. Because of the costs incurred, this
P process is unlikely to be a practical method of
manipulating fertility, unless either the restoration
value of a particular habitat is extremely high or the
topsoil may be sold off-site to reduce costs.
Organic
P Manipulation of the water-table
As discussed previously, the installation of a field
Fig. 13.5. Schematic representation of the fluxes of drainage system lowers the water-table, increases
P between the soluble, complexed, colloidal and microbial respiration and enhances nutrient
organic pools within grassland soils following (a) cycling. The potential therefore exists to slow the
fertilizer application and (b) removal of vegetation. rate of nutrient cycling, reduce the productivity
and lessen the level of competition in extensified
Table 13.2. Dry-matter production and soil P grasslands by raising the water-table. This may also
uptake from a hay crop taken in June from a soil increase N losses from the system by increasing
with the upper 5 cm removed (top soil removed) rates of denitrification. Various experiments have
and a soil with an intact soil profile (intact soil) investigated this possibility. An experiment by
(data extracted from Oomes et al., 1996). Oomes (1991) artificially raised the water-table
under a grassland on a peaty soil. This was achieved
Topsoil removed Intact soil
by extracting groundwater from wells and pumping
Dry-matter yield 2.7 4.6 it into ditches surrounding experimental plots. The
(t ha−1) water then penetrated the soil within the plots by
Uptake of P 7.29 13.5 flowing along irrigation pipes spaced at 5 m inter-
(kg ha−1) vals and buried at 50 cm depth. This allowed the
Uptake of N 47.25 75.9
water-table to be raised by approximately 20 cm in
(kg ha−1)
the summer and by 25–30 cm in winter. The
270 R. Chapman
elevated water-table successfully reduced the supply Successional responses to reduced

of K and, to a lesser extent, P to the vegetation over competition in extensified agricultural
a 3 year period. This indicated that changing the grasslands
site hydrological condition effectively modified the
nutrient cycles. Lowering the productivity of extensified agricul-
Similar reports by Berendse et al. (1994) and tural grassland communities will create gaps within
Oomes et al. (1996) discussed the effect of raising the matrix of competitor species within the grass-
the water-table by 10–40 cm on a humic clay-over- land community. This will generate niches suitable
peat soil. In that instance, the elevated water-table for the re-establishment of less competitive species,
reduced both annual N mineralization and above- which, if colonized successfully, should lead to the
ground N accumulation by approximately 20%, natural regeneration of a botanically diverse grass-
indicating that the treatment had successfully land community. Indeed, Bakker (1989), Olff and
manipulated both the soil’s hydrological condition Bakker (1991), Olff and Pegtel (1994) and Willems
and the N cycle. Extractable P and K in the top and van Nieuwstadt (1996) all found that practices
0–10 cm soil layer also declined markedly as a which successfully suppressed community produc-
result of raising the water-table. Oomes et al. tivity were also associated with simultaneous
(1996) speculated that the decline in P may have increases in botanical diversity. Thus, as new niches
been due to an enhanced rate of fixation into were re-established in the extensified community,
insoluble forms by the calcium-rich groundwater. they were rapidly colonized by previously absent
The change in K availability in the upper soil lay- species.
ers is likely to have been caused by root distri- This, however, has not always been the result.
bution patterns. The elevated water-table A restoration attempt by Oomes et al. (1996) suc-
concentrated a greater proportion of the roots in cessfully lowered the productivity within a grass-
the upper, drier layers of soil. Uptake of K would land community, but this was not associated with
therefore have largely been limited to a shallow any change in botanical diversity. In this study,
depth of soil, causing an appreciable decline in therefore, the community was clearly limited in its
availability over the course of this investigation. ability to recruit new species into the re-created
Furthermore, Oomes et al. (1996) observed that a niches. Berendse et al. (1992) and Pegtel et al.
reduction in productivity was associated with a sig- (1996) similarly reported reductions in productiv-
nificant change in community structure. ity that failed to yield the anticipated increase in
Gradually, taller ‘competitor’ species were replaced botanical diversity.
by less aggressive species of shorter stature that These apparently contradictory results may be
were better adapted to conditions of low nutrient explained by considering the seed-bank dynamics
availability. and propagule dispersal patterns of grassland
species, along with the proximity of the experimen-
tal site to external sources for colonizing popula-
Conclusions
tions. Many grassland species may be described as
The re-creation of botanically diverse communities generally possessing either a transient or a short-
in extensified agricultural ecosystems is likely to be lived seed bank (Thompson and Grime, 1979;
severely constrained by high levels of competition Thompson, 1987; Grime et al., 1988). As discussed
from relatively few species induced by excess soil above, intensification of grassland management
fertility. However, either instigating a net flow of excludes all but the most competitive species from
nutrients out from the extensified community or the vegetation. Under such management, any less
slowing the rate of nutrient cycling within the com- competitive species will be unable to regularly sup-
munity presents potential opportunities to reduce plement their seed bank and will be quickly elimi-
this fertility. This will lead to the establishment of nated from the system as their seed banks are
levels of productivity and competitive intensity exhausted (Hutchings and Booth, 1996; Bekker et
appropriate for the re-establishment of botanically al., 1997; Kirkham and Kent, 1997).
diverse grassland communities. The subsequent re- The recruitment of species into niches re-cre-
creation of such a community will depend upon a ated in extensified grassland communities will thus
successful successional response to the habitat’s depend upon the dispersal of propagules from alter-
modified fertility status. native sources in nearby vegetation. Seeds of many
grassland species are generally not well adapted for species atypical of undisturbed grasslands, but also
dispersal, e.g. Agrostis capillaris, Poa pratensis and demonstrated a high degree of shrub encroach-
Phleum pratense (Thompson, 1987), and will typi- ment. These results clearly indicate that alternative
cally be dispersed within a radius close to the seral pathways exist once a grassland is re-created.
maternal plant (Carey and Watkinson, 1993; The seral succession necessary for the re-establish-
Hutchings and Booth, 1996). Migration of grass- ment of the distinctive plagioclimax grassland com-
land species along vegetative corridors therefore munity will only occur under appropriate grazing
tends to be extremely slow; van Dorp (1993) esti- pressure and livestock management. The actions of
mated that dispersal of populations of grassland the grazing animal are therefore essential for the
species along one-dimensional corridors linking dif- successional re-development of species-rich pasture
ferent grassland habitats in The Netherlands was communities.
limited to a maximum of 4 m year−1.
It may therefore be anticipated that the estab-
Conclusion
lishment of natural successional processes in exten-
sified grasslands will depend critically upon the The successional development of botanically
spatial connections between the restored site and diverse grassland communities therefore requires
the surrounding landscape. If the restored site is effective species recruitment and appropriate graz-
close to suitable undisturbed vegetation (as in the ing management. Natural successional processes
experiments of Bakker, 1989; Olff and Bakker, require the establishment of an open ecosystem
1991; Olff and Pegtel, 1994; Willems and van with strong linkages with appropriate external plant
Nieuwstadt, 1996), there will be an effective rain of communities. If these conditions are not estab-
propagules on to the site and natural successional lished, the restored ecosystem will remain isolated
processes may proceed rapidly. In contrast, the and effectively closed. Succession will then be lim-
experiments of Berendse et al. (1992), Oomes et al. ited by the extremely poor rate of migration from
(1996) and Pegtel et al. (1996) were all conducted external populations. Under such circumstances,
on sites isolated from other botanically diverse the restoration practitioner must either accept the
grassland communities. The geographical isolation re-establishment of incomplete communities by
of these communities would prevent an influx of purely natural means (van Dorp, 1993) or must aid
propagules into the restored site, despite the cre- the process of recruitment by artificially assisting
ation of appropriate niches for colonization. the process of succession (Hutchings and Booth,
1996; Watt et al., 1996; Stevenson et al., 1997).
Once recruitment limitations have been overcome,
The role of the grazing animal
it is vital that an appropriate livestock management
Once the level of productivity has been adjusted regime is adopted to ensure that successional devel-
and recruitment commenced, correct successional opment proceeds along the correct trajectory.
development depends upon the adoption of an
appropriate defoliation regime. This is well illus-
trated in an experiment conducted by Gibson and The Re-creation of Botanically
Brown (1992), which investigated the effect of
grazing intensity on the re-creation of a calcicolous
Diverse Grasslands on Land
species-rich pasture on a former arable field. In par- Disturbed by Engineering
ticular, these workers were investigating whether Operations
the role of the grazing animal in creating a plagio-
climax pasture community is: (i) to arrest a linear The re-creation of botanically diverse grasslands on
successional process at a specific seral stage; or (ii) land restored following engineering operations poses
to divert the successional process along an entirely a number of problems. As with sites affected by
separate seral trajectory. In this experiment, only intensive agricultural practices, these will include
vegetation under the heaviest grazing treatment problems with plant competition, recruitment and
began to develop a species composition similar to succession. However, the environmental context of
that of the undisturbed pasture ‘target’ community. disturbed sites will typically be very different from
Vegetation that was grazed less intensively not only agricultural sites. As a consequence, the exact nature
became dominated by competitor and ruderal of the problems faced may differ substantially.
272 R. Chapman
The environmental conditions of efficiency with which it is cycled through that sys-
disturbed sites tem. A simplified representation of this N cycle is
presented in Fig. 13.6. Engineering operations are
Soil nutrient availabilities likely to disturb this N cycle by disrupting the
microbial processes and invertebrate populations.
Many engineering processes, such as opencast min-
ing and road verge construction, conclude with the
reconstruction of the soil profile using original top- SOIL MICROBIAL ACTIVITY. Mechanical handling of
and subsoils. This soil will, however, have been soils during the stripping and replacement processes
severely disturbed. This disturbance will certainly leads to a severe loss of soil macropores (King,
include damage induced by the mechanical strip- 1988). Such poor soil structure will compromise
ping and replacement of the medium, and may also the mineralizability of any organic N that remains
entail prolonged storage in stockpiles. While these in the substrate after restoration or is subsequently
processes may not adversely reduce the availability incorporated (Hassink, 1992; Hassink et al., 1993;
of soil P and K (Scullion, 1994b; Chapman and Killham et al., 1993). Because of the low soil
Younger, 1995), they may change many of the soil organic matter content, skeletal substrates are also
physical qualities, including bulk density, structure likely to display poor structural stability and pore
and drainage, (Abdul-Kareem and McRae, 1984; size distribution and this will further inhibit N
Baker et al., 1988; King, 1988). However, perhaps
mineralization (Bradshaw, 1997).
the greatest damage will be inflicted on the soil N
Fungi are essential for not only the efficient
cycle. A study by Davies et al. (1995) indicated that
breakdown of the soil organic matter, but also the
as much as 0.25 g N m−2 may be lost from a soil
recovery and maintenance of the soil structural sta-
reinstated from a stockpile. A detailed analysis of the
bility. Stockpiling soils significantly suppresses the
N dynamics during the soil handling process
revealed that these losses were due to transforma- soil fungal populations (Harris et al., 1993), and
tions that occurred during the storage and soon this will further impede the cycling of N in restored
after replacement. N supply rates in restored ecosys- soils.
tems have been shown to be closely correlated with
total soil N content (Skeffington and Bradshaw, SOIL INVERTEBRATE POPULATIONS. Detailed studies
1981); thus such a major loss of total soil N will have indicated that populations of earthworms and
severely compromise the soil’s overall fertility. other decomposer invertebrates (e.g. Isopoda and
Many industrial processes generate substantial Diplopoda) are unusually low on restored sites
quantities of inert skeletal wastes, such as colliery (Scullion et al., 1988a,b; Scullion, 1994a; Wheater
spoil, Leblanc waste, fly ash and chalk marl
and Cullen, 1997). This slows the rate of decompo-
(Bradshaw and Chadwick, 1980; Ash et al., 1994;
sition of plant litter and other organic matter
Mitchley et al., 1996). Natural weathering
(Majer, 1997) and so directly reduces the efficiency
processes release some P and K from these sub-
stances and availabilities of these nutrients may vary with which N is cycled within the restored ecosys-
from deficient to adequate (Bradshaw, 1997). In tem. Furthermore, low populations of earthworms
contrast, N is virtually absent from these substrates; inhibit soil structural development and this in turn
the major limitation to community productivity suppresses the ability of soil microorganisms to
therefore typically arises from extreme N deficien- mineralize organic N.
cies (Bradshaw, 1997).
Thus deficiencies of macronutrients, and espe- CONCLUSION. Low levels of activity from both
cially of N, severely limit plant growth and vegeta- microbial and invertebrate decomposers are likely
tion establishment on both restored soils and to inhibit the breakdown of organic matter in
skeletal substrates remaining following industrial restored ecosystems. This will impede the efficient
disturbance. cycling of N through the restored ecosystem (see
Fig. 13.6) and cause a greater proportion of the N
pool to accumulate in the macro-organic and
Nitrogen cycling in restored soils
humic N pools. This will significantly reduce the
The N availability of any system depends not only proportion of the ecosystem’s already depleted N
on the total amount of N present, but also on the pool becoming mineralized and plant uptake will
(a) Plant
N
Uptake Senescence
Soil Macro-organic
mineral N matter (litter
plus dead
roots)
Mineralization
Mineralization Decay
Soil
humic N
(b) Plant
N
Reduced
pool
Uptake Senescence
Soil Reduced Enhanced Macro-organic

mineral N pool pool matter (litter
plus dead
roots)
Reduced
mineralization
Reduced Reduced
mineralization Enhanced decay
pool
Soil
humic N
Fig. 13.6. A simplified representation of the N cycling pathways in the soil and plant components of an
ecosystem in (a) an undisturbed and (b) a restored environment, showing reduced rates of decay and
mineralization leading to a reduced level of plant uptake and an accumulation of N within the macro-
organic fraction.
274 R. Chapman
be severely reduced. The implications that this has extremely diverse community. This process of nat-
for ecosystem productivity will be discussed later. ural succession, however, occurred over a period of
several centuries. This time-scale will be prohibi-
tively long for most practitioners. Bradshaw (1997)
Landscape conditions
recently reviewed the natural successional processes
Many interactive processes that link ecosystems that operate on disturbed land. Examination of the
operate over relatively short distances; these include successional dynamics has demonstrated that
the dispersal of seeds from grassland species and the ecosystem establishment is constrained principally
migration of populations of wingless invertebrates. by the processes of both plant recruitment and fer-
Industrial disturbances may occupy great areas of tility development.
landscape; for example, opencast coal pits in Successional development of a restored ecosys-
England frequently involve drastic disturbance over tem commences with the establishment of a vegeta-
hundreds of hectares (Fig. 13.7; British Coal tive cover. Skeletal soil media are naturally void of
Opencast Executive, 1991). Disturbances of this any form of seed bank. In other instances, such as
scale will effectively prevent the establishment of following mineral extraction or road building, the
many linkages between the restored and undis- substrate may be composed of heavily disturbed soil.
turbed ecosystems. An ecosystem developing within The combined effects of earlier land-use practices
a disturbed site will therefore often be geographi- and soil handling processes during the engineering
cally isolated, with few opportunities for the operations will typically eliminate species character-
recruitment of absent species. This restricts the rate istic of botanically diverse grasslands from the seed
of colonization of appropriate plant species and so bank. Natural colonization will therefore depend
compromises natural successional processes. entirely upon migration of propagules from other,
Furthermore, the recruitment of invertebrates vital undisturbed sites. However, disturbed sites are often
for efficient N cycling is likely to be inhibited, thus in locations far removed from appropriate undis-
restricting ecosystem fertility (Scullion, 1994a). turbed vegetation, and this typically restricts the rate
of immigration. An examination of the flora of sev-
eral disturbed habitats by Ash et al. (1994) demon-
Natural successional processes in disturbed strated that, even after 100 years of natural
ecosystems colonization, ecosystem development was still prin-
cipally restricted by low rates of species recruitment.
Natural successional processes can be extremely The first plants to naturally colonize disturbed
successful in the re-creation of botanically diverse habitats are typically, although not exclusively, slow-
grasslands on disturbed sites. An example of this is growing stress-tolerating species, which are well
Millers Dale in Derbyshire, UK. This dale was adapted to nutrient deficiencies but are not neces-
abandoned after extensive disturbance during min- sarily typical of botanically diverse grassland com-
eral extraction in ancient times. Since then, natural munities (Ash et al., 1994). Further natural
processes have led to the establishment of an development of botanically diverse grassland com-
munities requires the development of ecosystem fer-
tility, followed by the recruitment of a more
appropriate complement of species. Because of the
low growth rate of the primary colonizing species,
soil organic matter accumulation will occur slowly.
In addition, the litter and organic residues produced
by these species are highly resistant to microbial
degradation (Palmer and Chadwick, 1985; Berendse
et al., 1989; Robles and Burke, 1997). The develop-
ment of ecosystem fertility and productivity will
therefore only occur slowly. Furthermore, as habitat
productivity approaches the critical low to moderate
Fig. 13.7. An aerial view of an opencast coal pit, level, the recruitment of further species more typical
showing massive and widespread disturbance to the of botanically diverse grasslands will be constrained
landscape. by poor linkages with external and undisturbed
grassland communities. Thus natural recruitment of and reducing the overall diversity. However, the
species to disturbed sites of enhanced fertility will be extreme N deficiency creates a niche where
a slow process (Chapman and Younger, 1995). legumes, with their independent N supply, can gain
Natural succession will therefore be too slow for a significant competitive advantage. Under such
most restoration schemes and some level of artificial conditions, legumes may quickly come to dominate
assistance will be necessary for the re-creation of species less well adapted to N deficiencies and so
botanically diverse grassland communities on dis- may severely threaten the diversity of the re-created
turbed sites. communities (Figs 13.8 and 13.9; Hodgson, 1989;
Chapman and Younger, 1995). Clearly, therefore,
the successful re-creation of botanically diverse
Assisted succession and competitive grasslands on disturbed soil will require measures to
interactions in disturbed ecosystems curb the competitive advantage of legumes.
Artificial introduction of propagules Manipulating the N supply of the restored

ecosystem
As the establishment of an initial vegetative cover
on a disturbed site is recruitment-limited, a funda- The strong competitive ability of legumes in
mental step in an assisted successional development restored ecosystems may be moderated by manipu-
will be the sowing and establishment of an artifi- lating the N cycle. Legume N2-fixation will cause
cially prepared seed mixture. An approach for the level of soil N to gradually build up over time.
devising a suitable seed mixture for re-creating As ecosystem fertility increases, non-legumes will
botanically diverse grassland communities in dis- compete more effectively with legumes, and com-
turbed ecosystems has been reported by Hodgson munity diversity and sustainability should increase.
(1989). This method allows the preparation of mix- However, given that initial N levels may be as
tures of species that are both typical of species-rich much as 2.5 t ha−1 less than equilibrium values
grasslands and suitably adapted for conditions of (Davies et al., 1995), this may take an unacceptable
low nutrient availability. length of time. Alternatively, soil N may be supple-
mented by the addition of organic amendments,
e.g. sludge (Bradshaw, 1997). Sludge, however, may
Competitive relationships on nutrient-deficient
be rich in heavy metals and the impact of these on
soils
competition within botanically diverse communi-
The successful re-creation of a botanically diverse ties is currently unknown. Alternative sources of N
grassland community in disturbed ecosystems will may come from applications of farmyard manure or
demand medium- and long-term sustainability, slurry. Application of slurry direct to the sward runs
especially in the face of severe N deficiencies. This the risk of suppressing broad-leaved species through
N deficiency severely reduces the productivity of re- scorch damage to the leaves (Anon., 1985;
created botanically diverse grasslands on these sites Chapman, 1988); this may be overcome by inject-
(Table 13.3). In contrast to extensified agricultural ing the slurry directly into the soil. Farmyard
land, reduced productivity prevents the more com- manure has long been applied to many traditionally
petitive grasses from dominating the community managed botanically diverse communities (Younger
Table 13.3. Hay yields from a traditionally managed botanically diverse grassland on an undisturbed site
in Teesdale and two similar communities re-created on land reclaimed following opencast coal mining,
all in northern England.
Butterwell (restored following Acklington (restored following

Teesdale (undisturbed)a opencast coal mining)b opencast coal mining)c
5.4 t ha−1 2.25 t ha−1 2.2 t ha−1

aFrom Younger and Smith, 1994.
bR. Chapman, unpublished data.
cFrom Chapman and Younger, 1995.
276 R. Chapman
Herbage legume content

Simpson’s diversity index
50 15
14
Legume content of herbage (%)
40
13
Simpson’s diversity index

30
12
11
20
10
10
9
0 8
1 2 3
Year since re-establishing a botanically diverse grassland community
Fig. 13.8. The relationship between herbage legume content and Simpsons’s diversity index over the first
3 years following the establishment of a botanically diverse grassland on land restored after opencast coal
mining (data extracted from Chapman and Younger, 1995).
petitive species, such as L. perenne and Rumex cris-

pus, while reducing the abundance of T. repens.
Overall community diversity, however, was unaf-
fected. The small response to the applied N may be
due to the timing and/or the concentration of the
application.
TIMING OF APPLICATION. Grassland species display

different phenological abilities to respond to pulses
of N (McKane et al., 1990; Bilbrough and
Caldwell, 1997). It is possible that at the time of
application, only a minority of species were in a
Fig. 13.9. A species-rich grassland community re- condition to utilize the enhanced N supply. The
created on a former opencast coal site, remaining species will, at best, have been unaf-
demonstrating aggressive growth by legumes, fected by the N applications or even suppressed by
induced by low soil N content, which is suppressing the increased vigour from the positively responding
overall community diversity.
species.
and Smith, 1994) and so may allow an increase in CONCENTRATION OF APPLICATION. Grassland
the N capital of the ecosystem with little risk to the species have been shown to differ in their ability to
community’s broad-leaved components. utilize contrasting pulses of N (Campbell and
An experiment by Chapman et al. (1996) inves- Grime, 1989; Bilbrough and Caldwell, 1997). In
tigated the possibility of controlling the aggressive particular, competitor species are best able to
legume component by increasing the total N sup- respond to the high-concentration N pulses that are
ply with mineral fertilizers. Applications of N associated with the style of mineral fertilizer appli-
increased the abundance of some of the more com- cation used in this experiment. This will have given
the observed positive response from a minority of Conclusions

competitive species with no actual enhancement of
overall community diversity. The fundamental objective that underlies all ecolog-
Both phenological and N pulse problems may ical restoration projects is the re-creation of stable
be overcome by spreading the N application over and sustainable habitats. However, the successful re-
time. This will ensure that increased N supply ben- establishment of botanically diverse grassland com-
efits species from a wider range of phenological munities on both agricultural and disturbed sites is
groups. Furthermore, individual N applications will severely constrained by issues of competition and
be of lower concentration than those used in this succession. Both the origin of these problems and
experiment. This may enable the less competitive their potential solutions relate to a combination of
species to utilize the applied N with increased effi- the conditions within the restored ecosystem and
ciency, thereby obtaining a positive response from a the nature of the linkages between that ecosystem
wider range of constituent species and improving and the external environment.
the community’s overall diversity. Competition problems arise from inappropriate
soil fertility, and so relate directly to the internal con-
Defoliation management ditions of the restored ecosystem. Typically, agricul-
tural habitats will be characterized by excess nutrient
A study by Chapman et al. (1996) investigated the availability, while disturbed habitats will tend to be
potential for strategic grazing by sheep to suppress associated with nutrient deficiencies. Nutrient avail-
the aggressive legume growth in restored ecosystems. abilities may be manipulated by managing the
Measurements made included herbage production cycling of nutrients within the ecosystem. Enhancing
and Simpson’s equitability index, which measures the rate of nutrient turnover may increase the pro-
the evenness with which constituent species are rep- portion of the total nutrient pool available for plant
resented in the community. The grazing treatment uptake, and so effectively enhance the productivity
effectively reduced legume biomass and this was of an otherwise infertile environment. This may be
associated with an increase in the equitability index. achieved on disturbed sites by increasing the popula-
Defoliation by sheep, therefore, reduced the produc- tions of decomposer invertebrates and enhancing the
tivity of the legumes and allowed other species to activity of the soil microorganisms. In contrast, slow-
compete more successfully in the re-created commu- ing the rate of nutrient cycling might retain more
nity. That initial study, however, failed to identify nutrients within fractions unavailable for plant
the mechanism underlying this change in competi- uptake, thus reducing the fertility of the environ-
tive relationships. A subsequent investigation, which ment. Raising the water-table presents one opportu-
focused on the most abundant legumes in the com- nity for achieving this in intensified agricultural
munity, demonstrated that the impact of strategic environments. Ecosystem fertility may also be effec-
grazing arose from the selective defoliation of T. tively manipulated by importing or exporting nutri-
repens (Chapman and Younger, 1997). This signifi- ents from or to external sources. Nutrient export
cantly reduced the subsequent growth of that may be effectively achieved through the removal of
species and simultaneously released companion vegetation or soil from the restored ecosystem. The
grasses from competitive suppression and so depletion of nutrients through the cutting and clear-
enhanced the diversity of the overall community. ing of vegetation is easy to implement, but may take
It is possible that herbivory by invertebrates many years to achieve the desired outcome. Topsoil
may provide similar benefits. Wardle and Barker removal, in contrast, may require greater initial cost
(1997) reported that invertebrate herbivores selec- and effort from the restoration practitioner, but will
tively impeded the growth of T. repens in botani- yield an immediate benefit in reduced site fertility.
cally diverse grasslands in New Zealand. If applied Nutrients may be readily imported through the
to disturbed ecosystems, this could be expected to application of either mineral fertilizers or organic
reduce the competitiveness of the legume compo- manures. While the pool of N in disturbed sites may
nent and stabilize the diversity of re-created grass- be increased following biological N2-fixation by
land communities. The opportunity to stabilize legumes, this may take an unacceptable length of
these communities through the managed reintro- time. Furthermore, aggressive legume growth may
ductions of herbivorous invertebrate populations is severely endanger the stability of botanically diverse
clearly an area that requires further research. grasslands re-created in nutrient-poor disturbed
278 R. Chapman
environments. However, research has shown that If all the restrictions on propagule recruitment
strategic grazing may successfully check the growth are overcome, there exist a number of alternative
of the legumes and maintain the stability of the re- successional trajectories which the restored ecosys-
created community. tem might follow. The adoption of the appropriate
The successional development of botanically trajectory is a vital prerequisite for the successful re-
diverse grassland communities on restored ecosys- creation of a species-rich grassland. The application
tems depends upon both the recruitment of appro- of the correct grazing and defoliation management
priate propagules and the establishment of an therefore plays an essential role in achieving this
appropriate successional trajectory. Seeds of appro- objective.
priate species will typically be absent from the seed The successful re-creation of a stable and sus-
bank. Because of this, recruitment will depend tainable botanically diverse grassland community
upon the immigration of propagules from sources therefore requires consideration of competitive and
outside the restored ecosystem. This process will be successional processes within the restored ecosys-
greatly influenced by the landscape context of the tem. These are related to not only those ecological
restored ecosystem. Location of the restored ecosys- processes that are internal to the restored ecosys-
tem adjacent to an appropriate and undisturbed tem, but also those processes that link that system
botanically diverse grassland community will allow to the external environment. Managing these
the effective exchange of propagules. However, processes may require manipulation of nutrient
many grassland species are able to disperse seeds pools and nutrient cycles, careful attention to graz-
over only a very short range. Increasing the distance
ing and defoliation practices and controlling the
between the restored ecosystem and an undisturbed
colonization of the restored habitat by appropriate
botanically diverse grassland community will
plant species.
quickly weaken and sever these linkages.
Successional development then becomes con-
strained by the slow rate of natural recruitment.
Under these circumstances, the successful re-cre- Acknowledgements
ation of a botanically diverse grassland community
may require the enhancement of succession I would like to thank Dr Amanda Ellery and Dr
through the artificial introduction of appropriate David Strong for providing useful comments on
propagules using specially prepared seed mixtures. the original manuscript.
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C&S in Pastures Chap 13 2/11/01 8:56 am Page 261

13 Competition and Succession in

Introduction Despite their acknowledged environmental val-

*Present address: c/o Orchard House, Carlton Scroop, Grantham, UK

Re-created Botanically Diverse Grassland Communities 263

gated through nutrient competition (Grime, 1979).

Species density (number of species 0.25 m–2) 30

Re-created Botanically Diverse Grassland Communities 265

Re-created Botanically Diverse Grassland Communities 267

Authors’ soil Apparent induction of Nutrients remaining

Oomes (1991) Sand K after 9 years P and N

Re-created Botanically Diverse Grassland Communities 269

(a) tions, which are critical to nutrient cycling, are also

elevated water-table successfully reduced the supply Successional responses to reduced

Re-created Botanically Diverse Grassland Communities 271

Re-created Botanically Diverse Grassland Communities 273

Soil Reduced Enhanced Macro-organic

Re-created Botanically Diverse Grassland Communities 275

Artificial introduction of propagules Manipulating the N supply of the restored

Butterwell (restored following Acklington (restored following

5.4 t ha−1 2.25 t ha−1 2.2 t ha−1

Herbage legume content

Simpson’s diversity index

petitive species, such as L. perenne and Rumex cris-

TIMING OF APPLICATION. Grassland species display

Re-created Botanically Diverse Grassland Communities 277

the observed positive response from a minority of Conclusions

Re-created Botanically Diverse Grassland Communities 279

Re-created Botanically Diverse Grassland Communities 281

Whitehead, D.C. (1995) Grassland Nitrogen. CAB International, Wallingford, UK.

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