Papers by Angela Breeuwer
Unie zijn deze zogenaamde 'agri-environment schemes' een belangrijk speerpunt van het landbouwbel... more Unie zijn deze zogenaamde 'agri-environment schemes' een belangrijk speerpunt van het landbouwbeleid geworden.
Evaluatie agrarisch natuurbeheer: effecten op weidevogeldichtheden
Evaluatie Agrarisch Natuurbeheer: effecten op weidevogeldichtheden Frank Willems1, Angela Breeuwe... more Evaluatie Agrarisch Natuurbeheer: effecten op weidevogeldichtheden Frank Willems1, Angela Breeuwer2, Ruud Foppen1 ,Wolf Teunissen1, Hans Schekkerman3, Paul Goedhart4, David Kleijn2 & Frank Berendse2 ... Voor Kievit en Scholekster bestonden geen verschillen. ...

Plant Ecology, 2009
To predict the role of ombrotrophic bogs as carbon sinks in the future, it is crucial to understa... more To predict the role of ombrotrophic bogs as carbon sinks in the future, it is crucial to understand how Sphagnum vegetation in bogs will respond to global change. We performed a greenhouse experiment to study the effects of two temperature treatments (17.5 and 21.7°C) and two N addition treatments (0 and 4 g N m -2 year -1 ) on the growth of four Sphagnum species from three geographically interspersed regions: S. fuscum, S. balticum (northern and central Sweden), S. magellanicum and S. cuspidatum (southern Sweden). We studied the growth and cover change in four combinations of these Sphagnum species during two growing seasons. Sphagnum height increment and production were affected negatively by high temperature and high N addition. However, the northern species were more affected by temperature, while the southern species were more affected by N addition. High temperature depressed the cover of the 'wet' species, S. balticum and S. cuspidatum. Nitrogen concentrations increased with high N addition. N:P and N:K ratios indicated P-limited growth in all treatments and co-limitation of P and K in the high N treatments. In the second year of the experiment, several containers suffered from a severe fungal infection, particularly affecting the 'wet' species and the high N treatment. Our findings suggest that global change can have negative consequences for the production of Sphagnum species in bogs, with important implications for the carbon sequestration in these ecosystems.

Plant Ecology, 2007
To evaluate the effects of changes in water level and temperatures on performance of four Sphagnu... more To evaluate the effects of changes in water level and temperatures on performance of four Sphagnum mosses, S. magellanicum, S. rubellum, S. imbricatum and S. fuscum were grown at two water levels, -5 cm and -15 cm, and at two temperatures, 15°C and 20°C. These species differ in their position along the microtopographical gradient and in their geographical distribution. Height increment, subcapitulum bulk density, biomass production, capitulum water content and cumulative evaporation were measured. Height increment and biomass production of S. magellanicum was lower at low water table than at high water table, whereas height increment and biomass production of S. rubellum, S. imbricatum and S. fuscum were unaffected. Height increment of S. magellanicum, S. rubellum and S. imbricatum was higher at high temperature than at low temperature. Biomass production of only S. magellanicum and S. rubellum was higher at high temperature than at low temperature, corresponding with their more southern distribution. Cumulative evaporation of S. magellanicum and S. rubellum was lower at low water table and could be explained by hampered water transport towards the capitula. We conclude that changes in water table and temperature may alter the Sphagnum composition on raised bogs, which may result in changes to important ecosystem processes. Therefore, it is important that species composition and changes therein are taken into account when evaluating global change effects on raised bog ecosystems.

Oikos, 2008
Despite their low primary production, ombrotrophic peatlands have a considerable potential to sto... more Despite their low primary production, ombrotrophic peatlands have a considerable potential to store atmospheric carbon as a result of their extremely low litter decomposition rates. Projected changes in temperature and nitrogen (N) deposition may increase decomposition rates by their positive effects on microbial activity and litter quality, which can be expected to result in enhanced mass loss and N release from Sphagnum and vascular plant litter. This is the first study that examines the combined effects of increased temperature and N deposition on decomposition in bogs. We investigated mass loss and N release at four bog sites along a gradient from north Sweden to northeast Germany in which both temperature and N deposition increased from north to south. We performed two litterbag experiments: one reciprocal experiment with Eriophorum vaginatum litter and one experiment using recalcitrant (Sphagnum fuscum) and more degradable (Sphagnum balticum) Sphagnum litter collected from the most northern site. We measured mass loss and N release during two (Sphagnum) and three (E. vaginatum) years. The N concentration and decomposability of the E. vaginatum litter did not differ between the sites. Mass loss from E. vaginatum litter increased over the gradient from north to south, but there was no such effect on Sphagnum litter. N loss of all litter types was affected by collection site, incubation site and time and all interactions between these factors. N release in Sphagnum was positively related to N concentration. We conclude that decomposition of vascular plants and Sphagnum litter is influenced by different environmental drivers, with enhanced temperatures stimulating mass loss of vascular plant litter, but not of Sphagnum. Enhanced N deposition increases Sphagnum litter N loss. As long-term consequences of climate change will presumably entail a higher vascular plant production, overall litter decomposition rates are likely to increase, especially in combination with increased temperature.

Oecologia, 2009
Increased N deposition in Europe has aVected mire ecosystems. However, knowledge on the physiolog... more Increased N deposition in Europe has aVected mire ecosystems. However, knowledge on the physiological responses is poor. We measured photosynthetic responses to increasing N deposition in two peatmoss species (Sphagnum balticum and Sphagnum fuscum) from a 3-year, north-south transplant experiment in northern Europe, covering a latitudinal N deposition gradient ranging from 0.28 g N m ¡2 year ¡1 in the north, to 1.49 g N m ¡2 year ¡1 in the south. The maximum photosynthetic rate (NP max ) increased southwards, and was mainly explained by tissue N concentration, secondly by allocation of N to the photosynthesis, and to a lesser degree by modi-Wed photosystem II activity (variable Xuorescence/maximum Xuorescence yield). Although climatic factors may have contributed, these results were most likely attributable to an increase in N deposition southwards. For S. fuscum, photosynthetic rate continued to increase up to a deposition level of 1.49 g N m ¡2 year ¡1 , but for S. balticum it seemed to level out at 1.14 g N m ¡2 year ¡1 . The results for S. balti-cum suggested that transplants from diVerent origin (with low or intermediate N deposition) respond diVerently to high N deposition. This indicates that Sphagnum species may be able to adapt or physiologically adjust to high N deposition. Our results also suggest that S. balticum might be more sensitive to N deposition than S. fuscum. Surprisingly, NP max was not (S. balticum), or only weakly (S. fuscum) correlated with biomass production, indicating that production is to a great extent is governed by factors other than the photosynthetic capacity.

The effect of temperature on growth and competition between Sphagnum species
Oecologia, 2008
Peat bogs play a large role in the global sequestration of C, and are often dominated by differen... more Peat bogs play a large role in the global sequestration of C, and are often dominated by different Sphagnum species. Therefore, it is crucial to understand how Sphagnum vegetation in peat bogs will respond to global warming. We performed a greenhouse experiment to study the effect of four temperature treatments (11.2, 14.7, 18.0 and 21.4 degrees C) on the growth of four Sphagnum species: S. fuscum and S. balticum from a site in northern Sweden and S. magellanicum and S. cuspidatum from a site in southern Sweden. In addition, three combinations of these species were made to study the effect of temperature on competition. We found that all species increased their height increment and biomass production with an increase in temperature, while bulk densities were lower at higher temperatures. The hollow species S. cuspidatum was the least responsive species, whereas the hummock species S. fuscum increased biomass production 13-fold from the lowest to the highest temperature treatment in monocultures. Nutrient concentrations were higher at higher temperatures, especially N concentrations of S. fuscum and S. balticum increased compared to field values. Competition between S. cuspidatum and S. magellanicum was not influenced by temperature. The mixtures of S. balticum with S. fuscum and S. balticum with S. magellanicum showed that S. balticum was the stronger competitor, but it lost competitive advantage in the highest temperature treatment. These findings suggest that species abundances will shift in response to global warming, particularly at northern sites where hollow species will lose competitive strength relative to hummock species and southern species.

Ecosystems, 2010
A large proportion of northern peatlands consists of Sphagnum-dominated ombrotrophic bogs. In the... more A large proportion of northern peatlands consists of Sphagnum-dominated ombrotrophic bogs. In these bogs, peat mosses (Sphagnum) and vascular plants occur in an apparent stable equilibrium, thereby sustaining the carbon sink function of the bog ecosystem. How global warming and increased nitrogen (N) deposition will affect the species composition in bog vegetation is still unclear. We performed a transplantation experiment in which mesocosms with intact vegetation were transplanted southward from north Sweden to northeast Germany along a transect of four bog sites, in which both temperature and N deposition increased. In addition, we monitored undisturbed vegetation in control plots at the four sites of the latitudinal gradient. Four growing seasons after transplantation, ericaceous dwarf shrubs had become much more abundant when transplanted to the warmest site which also had highest N deposition. As a result ericoid aboveground biomass in the transplanted mesocosms increased most at the southernmost site, this site also had highest ericoid biomass in the undisturbed vegetation. The two dominant Sphagnum species showed opposing responses when transplanted southward; Sphagnum balticum height increment decreased, whereas S. fuscum height increment increased when transplanted southward. Sphagnum production did not differ significantly among the transplanted mesocosms, but was lowest in the southernmost control plots. The dwarf shrub expansion and increased N concentrations in plant tissues we observed, point in the direction of a positive feedback toward vascular plant-dominance suppressing peat-forming Sphagnum in the long term. However, our data also indicate that precipitation and phosphorus availability influence the competitive balance between Sphagnum, dwarf shrubs and graminoids.

Basic and Applied Ecology, 2009
Climate change can be expected to increase the frequency of summer droughts and associated low wa... more Climate change can be expected to increase the frequency of summer droughts and associated low water tables in ombrotrophic peatlands. We studied the effects of periodic water table drawdown in a mesocosm experiment. Mesocosms were collected in Southern Sweden, and subsequently brought to an experimental field in the Netherlands. Two water table treatments were applied: one with constant water tables at 5 cm below the moss surface, and one in which the water table was allowed to drop, resulting in water tables fluctuating between 5 and 21 cm below the moss surface. Sphagnum growth, as well as Sphagnum and vascular plant abundance, were assessed for 2 1 2 years. Our results show that the abundance of graminoid species increased most in the constant water table treatment. In contrast, ericoid species cover increased when water tables were allowed to fluctuate. Furthermore, Sphagnum cuspidatum production decreased with fluctuating summer water tables, while Sphagnum magellanicum responded oppositely. From these results we conclude that increased occurrence of periods with low water tables may bring about a shift in dominant Sphagnum species as well as a shift from graminoid to ericoid vascular plant cover, resembling the shift from hollow to lawn or hummock vegetation. The difference in response within functional groups (vascular plants, Sphagnum) may add to the resilience of the ecosystem.
Basic and Applied Ecology, 2009
In European peatlands which have been drained and cut-over in the past, re-vegetation often stagn... more In European peatlands which have been drained and cut-over in the past, re-vegetation often stagnates after the return of a species-poor Sphagnum community. Re-introduction of currently absent species may be a useful tool to restore a typical, and more diverse, Sphagnum vegetation and may ultimately improve the functioning of peatland ecosystems, regarding atmospheric carbon sequestration. Yet, the factors controlling the success of re-introduction are unclear. In Ireland and Estonia, we transplanted small and large aggregates of three Sphagnum species into existing vegetation. We recorded changes in cover over a 3-year period, at two water levels (À5 and À20 cm).

Biological Conservation, 2009
Since 1992 the European Union helps member states to reverse the loss of biodiversity in agricult... more Since 1992 the European Union helps member states to reverse the loss of biodiversity in agricultural landscapes by the financial support of agri-environment schemes. Long-term studies investigating the effects of these schemes are an essential prerequisite for the development of an effective policy to restore biodiversity on farmland. In Dutch meadow landscapes almost all agri-environment schemes focus on the restoration of meadow bird populations by postponement of the mowing date. Between 1990 and 2002 we measured long-term changes in meadow bird densities in areas with and without agri-environment schemes in the Netherlands, both before and after the start of the contract. During these years bird territories were surveyed during five field visits between 15 March and 15 June. Densities of black-tailed godwit (Limosa limosa), and redshank (Tringa totanus) were higher in the areas with management agreements, but these differences were already present before the start of the contracts. After the start of the management contracts densities of black-tailed godwit and oystercatcher (Haematopus ostralegus) did not increase, while those of lapwing (Vanellus vanellus) and redshank even declined relative to the control areas. It is concluded that the current agri-environment schemes are not sufficient to restore meadow bird populations in Dutch agricultural landscapes. In addition to the prescribed postponement of the mowing date, it is probably necessary to raise groundwater levels and to reduce fertilization to allow for the development of an open vegetation structure that will increase chick survival to sufficiently high levels.
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Papers by Angela Breeuwer