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Restoring diverse grassland: What can be achieved where, and what will it do for us?
Summary
Elevated CO2 effects (a press perturbation) on plants communities may alter community
dynamics and as such the response to extreme climatic events (pulse perturbation). The
ongoing Giessen FACE experiment (GiFACE) in Germany in a high diversity permanent
grassland presents a unique opportunity to study the combined effects of a press
perturbation with a pulse perturbation. The experiment was set up in 1998, and in 2003
the area was subject to a severe drought that significantly reduced biomass production.
The objective of this study was to elucidate the role of species diversity in the recovery
of primary productivity after drought under elevated CO2. It can be shown that due to
the high diversity in this grassland, biomass production had recovered one year after
drought to at least pre-drought levels. However, the community effects of the drought
were much longer lasting, with increased effective diversity, as expressed through species
turnover and variation in species abundances, for three years after the drought. While the
recovery of primary productivity in 2004 was related to species richness, species richness
tended to be negatively associated with variation in biomass production in subsequent
years, and a variation in primary productivity was better explained by effective diversity.
In conclusion, while there were small effects of elevated CO2 on community structure
observed over the course of the experiment, these effects did not influence the recovery
of primary productivity in elevated CO2 plots after the drought.
Key words: FACE, stability, press perturbation, pulse perturbation, diversity effect
Introduction
The ability of ecosystems to withstand extreme climatic events is an important issue, with the
frequency of such events increasing with climate change (Smith, 2011). Furthermore, ecosystems
are under a number of anthropogenic pressures such as increased nitrogen deposition, elevated
CO2, and changes in land use. The change in the dynamics of ecosystems under the latter is often
subtle, potentially leading to surprise outcomes in the longer term (Ives & Carpenter, 2007). The
study of the combined effects of pulse perturbations (as in the occurrence of extreme climatic
events) and press perturbations will help identify similarities and differences in the mechanisms
that lead to stability in ecosystem functioning under multiple perturbations.
FACE (Free Air Carbondioxide Enrichment) experiments offer a unique opportunity to study
the combined effects of a constant anthropogenic change in the environment, a press perturbation
(sensu Ives & Carpenter, 2007), and extreme climatic events (pulse perturbations). Unfortunately,
not all FACE experiments also allow the study of diversity effects as such systems often host
low numbers of species. The Giessen FACE experiment in Germany, however, was set-up in
1998 in a diverse temperate grassland system without initial disturbance as the experimental
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area was not ploughed and had been under the same management regime for around 100 years
(Jger et al., 2003; Kammann et al., 2005). In 2003, 5 years after the initiation of the experiment,
Europe experienced an extreme climatic event (drought with heat-wave) that significantly reduced
primary productivity all over Europe (Ciais et al., 2005). This system thus presents us with the
unique opportunity of studying the interactions of a press perturbation with a pulse perturbation.
The overall objective of this paper was to investigate if and how species diversity contributed to
the recovery of primary productivity after drought and what the effect of the CO2 treatment (press
disturbance) had on this recovery. In particular, we asked the following questions:
1)
Did primary productivity decline in 2003 and how long did it take to recover?
2)
Does species diversity have an effect on the recovery of primary productivity after drought?
3)
Does elevated CO2 have an effect on species diversity and community dynamics?
4)
And thus, does elevated CO2 have an effect on the recovery of primary productivity after
drought?
It is hypothesised that the recovery after drought (pulse perturbation) is dependent on species
diversity effects due to species complementarity (Hector et al., 2010).
Methods
Study site
The Giessen FACE experiment (GiFACE) was set-up at a grassland research site near Giessen,
Germany (5032N, 841.3E, 172 m a.s.l.). The vegetation at this site is a diverse hay meadow
typical of extensively managed temperate grasslands. Monitoring of the site started in 1993 and
elevated CO2 treatment (20% above ambient concentration during the day) was initiated in three
replicates in 1998 and is ongoing since then. For a detailed description of the site and the treatment
consult Jger et al. (2003).
Sampling
Sampling of the vegetation occurs twice a year, once in spring (late May early June) and once in
late summer (late August early September) (Grters et al., 2006). The community composition
of the vegetation is assessed in two 4.5 m2 sampling quadrats in each plot (ring) by estimating
%-cover for each species using a semi-categorical system, whereby Braun-Blanquet cover
categories1 were used except when cover was in-between categories and directly noted. One week
after the vegetation survey the aboveground biomass is harvested in 14 small sampling units
scattered throughout the plot (ring).
Analyses
A comparison of within plot to between plot variation showed that there is spatial autocorrelation
for sampling units within plots. It was therefore decided that the results from sampling units
within plot should be averaged for further analyses. All analyses were conducted separately for
the two seasons of sampling (spring and summer), except for primary productivity, which was
compared on a yearly basis to answer the first question. Primary productivity was defined as the
oven-dried harvested biomass per season. The recovery of primary productivity after the drought
was defined as follows:
Degree of recovery = biomass2004 average(biomass98-02)
Species diversity was defined by the number of species per plot (species richness) and by
Simpsons dominance index (D) (Simpson, 1949). Furthermore, it was considered that not only
r=0.1%, +=0.5%, 2m=1%, 2a=10%, 2b=20%, 3=37.5%, 4=62.5% and 5=87.5% (% cover is the mid-point
of the cover class).
1
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species number, but also the effective diversity as expressed by the variation in abundance of
species from year to year was important in explaining diversity effects. Therefore, an index of
effective diversity (EFFDIV) was calculated, which provided a combined measure of species
turnover and changes in species abundance:
EFFDIV = i abs(Log10(abupi+1) Log10(abuci+1))
where abs is the absolute value, abup is the abundance of the species in the previous year and
abuc is the abundance of the species in the current year, i is an index ranging from 1 to the
total number of species. The log10 transformation of the abundances of species was undertaken
because of the multiplicative nature of cover data and because it improved the normality of the
data (Lepmilauer, 2003). Furthermore, years were classified according to the occurrence of the
drought in 2003. A pre-drought period was defined as 19982002, the drought year was treated
separately, and a post-drought period was defined (20042008). Based on preliminary analyses it
was found that while this classification of years into categories corresponds well to the dynamics
of primary productivity, the same classification could not be applied to effective diversity, which
continued to be high after the drought until 2006. Therefore, years were classified differently as
follows: 19982002: pre-drought period, 20032006: drought and recovery, and 20072008 postdrought
Stepwise, backward selection of explanatory variables was conducted using multiple linear
regression (proc glm) in SAS version 9.1. Only significant explanatory variables were retained.
For correlations between species richness or effective diversity and recovery Pearsons correlation
coefficient (r) was used and correlations expressed as percentages (r2 * 100).
Results
Effect of drought on primary productivity
Primary productivity in 2003 was significantly different from other years of the experiment (Fig.
1). There was no difference between ambient and elevated CO2 in terms of how much productivity
declined and primary productivity recovered to pre-drought levels in both treatments in 2004.
2003drought
Fig. 1. Harvested biomass by year (A = ambient CO2, E = elevated CO2). Error bars represent standard
deviations. Treatments are not significantly different. Differences in mean are observed between the
drought (2003) year and all other years.
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b) Summer releve
The variation of the recovery of biomass in 2004 was correlated to species richness with about
44 % of the variation in the spring biomass harvest explained by species richness (36% for the
late summer harvest). Interestingly, the variation in production from 2005 onwards was inversely
correlated to species richness and started to be positively correlated to effective diversity.
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