Dynamics of common reed (Phragmites australisTrin.) in Swiss fens with different management

Dynamics of common reed (Phragmites australisTrin.) in Central Europe have so far mostly beeninvestigated in connection with studies on reed`die-back' along lake shores. However, there hasrecently been increasing concern about reed expansionat terrestrial sites, such as fens and wet grasslands.In this paper we report on the results of fourseparate studies which monitored reed dynamics inSwiss fens with various mowing regimes over a periodof 4 to 15 years. The first study compared unmownplots with plots mown in winter in a triennialrotation; the second one included unmown plots, plotsmown in summer, and plots mown in winter; the thirdone compared plots mown in June and September withplots only mown in September; the fourth studyinvestigated only plots mown in September. Shootnumber and shoot size were recorded in permanentquadrats. In all studies the performance of P.australisfluctuated without trend or tended todecrease during the period investigated. Thedecreasing tendency concerned shoot size rather thanshoot number, and within a given study it was strongerfor plots with initially taller shoots. The variousmowing regimes did hardly influence these changes.Mowing in winter every three years reduced shoot sizein the year after mowing, but not on the long term.Mowing every year in late summer reduced the shootsize compared with unmown plots on the short term, butthis effect almost disappeared on the long term, aftermowing had become biennial. Mowing in June (inaddition to in September) caused no noticeableeffects. We conclude that other factors (e.g. weatherconditions, competition, or population processes) aremore important than management in determining theabundance of P. australisin the fen communitiesinvestigated here, although long-term effects ofmowing in summer still need more investigation. As apractical consequence it is suggested that at siteswhich are not strongly dominated by P.australis, as most of those investigated here,reducing the performance of this species should notconstitute a major target of nature conservationmanagement, nor can its dynamics be used as anindicator for management success before underlyingcauses are better understood.     

Differential and reversible responses of common fen meadow species to abandonment

Abstract. We studied the effects of abandonment on two common fen plant species. In mown and a chronosequence of abandoned fen meadows spanning 35 yr, we measured fitness traits of the sedge Carex davalliana and the forb Succisa pratensis. Cessation of mowing had little effect on fitness traits and seed production of C. davalliana, but seedling density decreased more than threefold. Population density of S. pratensis decreased with increasing community biomass, but was not affected by the cessation of mowing. However, flowering frequency increased threefold and seed production was 20% higher in fallow meadows. Consequently, seedling density of S. pratensis increased nearly threefold after abandonment. However, these changes were not dependent on the age of the fallow. In a common garden and germination experiment, we found no differences in either species between plants from fallows and mown fen meadows, except for the height of the flowering stalk of S. pratensis. The combined results from the common garden experiment and the field studies indicate that changes in fitness traits observed in fallows were mostly phenotypic and likely to be reversible. If other species react in similar ways, there is a high potential for re‐establishing traditional fen meadow communities from fallows by mowing.     

Shading by Phragmites australis: a threat for species-rich fen meadows?

Abstract. Common reed (Phragmites australis Trin.) has spread in fen meadows on the Swiss Plateau during the last decades. An increased dominance of this tall grass may reduce the plant species richness and displace rare or endangered species because of the additional shade. To investigate whether this has actually happened and whether shading by Phragmites was likely to be responsible for the reduction, the plant species composition was surveyed in 241 plots (4 m²) with differing above-ground biomass of Phragmites (ABP). Species richness and the occurrence of characteristic fen species were negatively related to site productivity (total above-ground biomass), but correlations with ABP were generally weaker. The main change associated with increasing ABP within species-rich fens (alliances Molinion and Caricion davallianae) was an increase of species characteristic of the more species-poor alliance Filipendulion. Thus, Phragmites did apparently not play a disproportionate role in the reduction of species richness in the plant communities studied. Selective clipping experiments and light measurements also indicated that shading by Phragmites does not have a strong effect upon the performance of other species in the community, at least in the short term. The reason appeared to lie in the later seasonal growth of Phragmites compared with the other species. These results suggest that the direct impact of shading by Phragmites on the species richness of fen meadows is probably lower than has been assumed. However, possible long-term or indirect effects still need to be investigated.     

How is Regeneration of Plants after Mowing Affected by Shoot Size in Two Species-Rich Meadows with Different Water Supply?

Mowing a meadow is an example of an equalizing process that reduces differences among species by removing aboveground biomass approximately 5 cm above ground. This regular disturbance that affects all plants prevents competitive exclusion of small species and thus allows coexistence of numerous species differing in shoot size. In this paper we search for the mechanism behind this by comparing the shoot biomass of 41 common species in dry and wet species-rich meadows in mown and recently abandoned plots in June (before mowing) and in October. We asked the following questions: i) Do the plants differ in proportion of biomass lost by mowing? ii) Are the mown plants able to compensate for biomass lost by mowing? iii) Is the compensatory ability of mown plants related to their size? iv) Is the compensatory ability of plants related to severity of disturbance (removed biomass)? v) Does water availability in meadows affect these features? Our results revealed that the earlier explanation of equalization of meadow plants after mowing due to the proportionally larger biomass loss in larger plants than small plants does not represent the entire mechanism. Even when larger plants in the wet meadow lost more biomass, the proportion of lost biomass was not dependent on plant size, and compensation ability (growth of mown in comparison with unmown plants) was not related to the lost biomass in this meadow type. On the contrary, the observed pattern could be explained by different compensation abilities of small versus tall plants. In addition, according to our expectations, the compensation for lost biomass in the wet meadow was higher than in the dry one.     

Effects of winter mowing on vegetation succession in a lakeshore fen

Abstract. To assess whether winter mowing in wetlands fulfils the aim of preventing succession towards drier communities, 34 permanent quadrats (15 m²) were surveyed annually from 1984–1985 to 2000 within large mown and unmown (control) areas (several ha) in a calcareous lake shore fen (W Switzerland). Three trends were noticed: decrease of aquatic species, spread of Cladium mariscus and establishment of woody species (especially Alnus glutinosa and Frangula alnus). None of these trends was prevented by mowing, but mowing did prevent the accumulation of C. mariscus litter and kept woody saplings small. Succession was generally slow and often occurred in the form of sudden, discrete changes. Plant species richness increased with mowing and remained constant without mowing. Soil disturbance by the mowing machine contributed more to the effects of management on species composition than the periodic removal of biomass. It is concluded that mowing every three years in winter is insufficient to preserve semi‐aquatic communities against succession but sufficient to maintain the plant species richness of a low productive, regularly flooded fen.     

The effects of mowing and fertilization on carbohydrate reserves and regrowth of grasses: do they promote plant coexistence in species-rich meadows?

After the cessation of regular management and after fertilization a single clonal species tends to dominate in many types of grasslands, whereas in regularly managed meadows these potential dominants usually attain a low cover. It has been hypothesized that plants reaching a high dominance in abandoned and fertilized meadows are selectively suppressed by mowing so that a balanced competition is maintained and competitive exclusion is postponed. We compared regeneration capacity and carbohydrate reserves accumulated by three species of clonal grasses, which markedly increase their dominance in irregularly mown, un-mown or fertilized meadows. Above-ground biomass and the amount of storage carbohydrates of the two largest species, (Molinia arundinacea, Calamagrostis epigejos) were reduced in a mown meadow. This effect was weaker in Bromus erectus, which produces smaller shoots. Shoots of Molinia were most impacted by mowing but their regeneration was efficient due to the large carbohydrate reserves in the shoot bases. Fertilization did not affect Bromus and Calamagrostis. In contrast, fertilized plants of Molinia produced larger storage organs and accumulated more carbohydrates. We conclude that plant size and growth form are important features promoting the ability of potential dominants to tolerate mowing and regenerate after it. Our results confirm that taller plants are selectively suppressed by this type of disturbance, thereby potentially promoting plant coexistence.     

Population structure of the threatened perennial Serratula tinctoria in relation to vegetation and management

Question: How does vegetation and management affect the population stage structure of Serratula tinctoria, a species indicative of highly endangered and species‐rich flood‐plain meadows? Are different management regimes suitable to support viable populations of S. tinctoria? Location: Flood‐plain of the northern Upper Rhine, Germany. Methods: We analysed the population structure of S. tinctoria at 24 meadow sites which differed in vegetation and management. In this comparative study the investigated meadows were either (1) late mown in September; (2) early mown in June or (3) mown in June and then grazed by sheep. Results: The structure of the surrounding vegetation had a clear effect on the population structure of S. tinctoria. The percentage of bare soil, as well as the cover of bryophytes, were positively related, whereas the Ellenberg N‐value of the established vegetation was negatively related to the density and proportion of seedlings. Generally, we found only slight differences between the population structure of S. tinctoria in early and late mown meadows. Both management regimes had high densities of seedlings. In contrast, the meadow pastures supported significantly lower densities of seedlings and generative adults but relatively high densities of juveniles and vegetative adults, indicating a greater importance of clonal propagation under grazing pressure. Conclusions: Our results clearly showed that viable populations of S. tinctoria may occur in all of the studied management regimes. Although S. tinctoria was considered to be highly sensitive to early mowing, our findings suggest a facilitation of the species at more nutrient‐rich sites by mowing in June, which is also a benefit for the integration of management in farming systems.     

Rotational fallows as overwintering habitat for grassland arthropods: the case of spiders in fen meadows

Regular mowing of grassland is often necessary for plant conservation, but uncut vegetation is needed by many arthropods for overwintering. This may lead to conflicting management strategies for plant and arthropod conservation. Rotational fallows are a possible solution. They provide a spatio-temporal mosaic of mown and unmown areas that may combine benefits to both plants and arthropods. We tested if rotational fallows enhance spider overwintering in fen meadows. Rotational fallows consisted of three adjoining strips 10 m wide and 35–50 m long. Each year, one of these strips was left unmown (fallow) in an alternating manner so that each strip was mown two out of three years. Spiders were sampled during spring with emergence traps in nine pairs of currently unmown fallow strips and completely mown reference plots. Fallows significantly enhanced orb-weavers (Araneidae), sac spiders (Clubionidae) and ground spiders (Gnaphosidae). However, only 4.7% of the total variation in community composition was attributable to fallows. Community variation was larger between landscapes (34.5%) and sites (38.2%). Also β diversity was much higher between landscapes (45 species) and sites (22 species) than between fallows and mown reference plots (10 species). We conclude that the first priority for spider conservation is to preserve as many fen meadows in different landscapes as possible. Locally, rotational fallows enhance overwintering of the above-mentioned spider families, which are sensitive to mowing in other grassland types as well. Thus, rotational fallows would probably foster spider conservation in a wide range of situations. However, stronger effects can be expected from larger and/or older fallow areas.     

Community structure and diversity of bryophytes and vascular plants in abandoned fen meadows

We examined effects of abandonment on species diversity and species composition by comparing 21 calcareous fen meadows in the pre-alpine zone of central and northeastern Switzerland. Meadows were divided into three classes of successional stages (mown: annually mown in late summer, young fallow: 4–15 years, and old fallow: >15 years of abandonment). In each fen, we measured litter mass in four 20 cm×20 cm plots, as well as (aboveground) biomass and species density (number of species per unit area) of bryophytes and vascular plants. Bryophyte biomass was reduced in abandoned fens, whereas litter mass and aboveground biomass of vascular plants increased. Species density of both taxonomic groups was lower in abandoned than in mown fens. Young and old successional stages were not different except for bryophytes, for which old successional stages had higher species density than young stages. We used litter mass and aboveground biomass of vascular plants as covariables in analyses of variance to reveal their effects on species density of both taxonomic groups. For bryophytes, litter mass was more important than vascular plant biomass in explaining variance of species density. This indicates severe effects of burying by litter on bryophyte species density. For species density of vascular plants, both vascular plant biomass and litter mass were of similar importance in explaining the decreased species density. Canonical correspondence analyses showed that abandonment also had an effect on species composition of both bryophytes and vascular plants. However, young and old successional stages were not different indicating fast initial changes after abandonment, but slow secondary succession afterwards. Furthermore, indicator species analysis showed that there was no establishment of new species after abandonment that might dramatically alter fen communities. Re-introduction of mowing as a nature conservation strategy may thus be very promising – even for old fallows.     

Is regular mowing the most appropriate and cost-effective management maintaining diversity and biomass of target forbs in mountain hay meadows?

Conservation of grassland biodiversity is a key issue in the EU agro-biodiversity policy. We assessed the effects of yearly mowing on target forb biomass in years with contrasting precipitation (2006–2007) in mountain fen and dry-mesophilous hay meadows in NE-Hungary. We hypothesised: (i) Species richness and biomass of target forbs is higher in mown than in abandoned stands. (ii) Mowing has more an effect on the biomass of target forbs, graminoids and litter than precipitation.     

Mowing management and density dependence in meadow passerine hatching success

Capsule Improving habitat quality through late or intermittent mowing may increase passerine population density without reducing reproductive success.

Aim To describe the relationship between passerine territory density (TD) and Whinchat Saxicola rubetra and Yellow Wagtail Motacilla flava hatching success, under different management regimes.

Methods TD was defined by the Territory Mapping method in nine sites selected in five French flood plains, in four intermittently mown meadows in Russia and in four additional French sites where mowing was partially postponed. Hatching success was assessed by the observation of prey carrying. Generalized Linear Mixed Models were used to analyse the influence on hatching rates of mowing management, early mowing and TD. Linear Mixed Models were used to study the effect of mowing management on TD.

Results In French early mown meadows, TD affected reproductive success. Higher hatching rates and TD were observed in Russia. Similar results in French study sites with postponed mowing confirmed the influence of mowing management on carrying capacity.

Conclusion In west-European grassland systems, late or intermittently mown patches within or at the edge of meadow areas, with potentially improved invertebrate-prey availability, should help to sustain bird demography and improve the efficiency of current conservation programmes.     

Reinforced Traditional Management is Needed to Save a Declining Meadow Species. A Demographic Analysis

The changes in agricultural practices during the last century have led to a drastic decrease in the number of traditionally managed hay meadows. Also, traditional management practices are often applied more cursorily in the remaining meadows. In combination with an increase in aerial anthropogenic nitrogen deposition, this has led to a loss of biodiversity. To investigate whether the current management is sufficient for maintaining viable populations of a typical meadow plant, Succisa pratensis, we experimentally reinforced the raking and mowing parts of the traditional management over four years in a two-by-two factorial experiment in three traditionally managed wooded hay meadows on the Baltic island of Gotland, Sweden. We found decreased litter and hay production in two of the three studied meadows as a result of our treatments. Plant sizes and asymptotic population growth rates (??) of S. pratensis increased, particularly in plots receiving the combined raking and mowing treatment. Stochastic long-term population growth rates (?? _s ) increased under the reinforced management: projected population sizes 50?years into the future showed a three-fold increase in raked plots and a 17-fold increase in plots that were both raked and mown. Because we found positive responses even in these seemingly well-managed meadows we conclude that it is essential that management is carried out more thoroughly to ensure viable population sizes. Our conclusion applies to most semi-natural grasslands receiving anthropogenic nitrogen, or where traditional management practices are less rigorously applied. We also suggest using biomass estimation instead of vegetation height as a measure of management strength.     

Permanent plot research in wet meadows and cutting experiment

Vegetation mowing is usually recommended for preserving wet meadows, in particular for the prevention of land warping and brushwood encroachment. The concept and the methods for the study of the influence of different cutting rhythms on the vegetation of two wet meadows, as well as some short term results are presented. Cutting experiments with permanent plots have been undertaken in vegetation communities such as Orchio-Schoenetum nigricantis and Caricetum elatae, abandoned for many years. The strategy of the study was to have diverse but complementary methodological approaches in order to assess changes in the vegetation and species abundance, especially concerning the botanical composition, the vegetation structure and the species vitality. The methods include counting of plants or part of plants, morphometry of Phragmites communis, biomass measurements, phenological records, structure relevés with the point-intercept technique as well as classical vegetation relevés with coverage estimation.On a short term base we can conclude, on the whole, that the yearly winter cut treatment induces a better vitality (number of individuals, flowering, biomass) of the studied species, while in the control plots the vitality is low, due notably to the important layer of accumulating litter and the lack of light on the ground, which hamper soil warming. The yearly summer cut treatment and the winter cut made every three years show in general intermediate characteristics. Furthermore, the vegetation structure is noticeably different according to whether the vegetation is cut (with hay removal) or not. Concerning the methodology, it is concluded that a diverse approach is a good way to assess changes in the vegetation with a minimum loss of information. If the rather subjective method of vegetation relevés with cover estimation may be suitable for long term survey on permanent plots, more quantitative and objective techniques have to be set if short term or precise changes in the botanical composition, the species vitality and the biomass, as well as structural changes, are sought.This paper was presented at the meeting of the working Group for Succession on Permanent Plots, held in Bern (Switzerland), 4th–7th September 1991.     

Recent changes in the plant composition of wetlands in the Jura Mountains

Aim

To assess vegetation changes in montane fens and wet meadows and their causes over 38 years.

Location

Wetlands, Jura Mountains (Switzerland and France).

Methods

Plots were inventoried in 1974 and re‐located in 2012 (quasi‐permanent plots) on the basis of sketches to assess changes in plant communities. The 110 plots belonged to five phytosociological alliances, two in oligotrophic fens (Caricion davallianae, Caricion fuscae) and three in wet meadows (Calthion, Molinion, Filipendulion). Changes between surveys were assessed with NMDS, and changes in species richness, Simpson diversity, species cover and frequency and the causes of these changes were evaluated by comparing ecological indicator values.

Results

Changes in species composition varied between alliances, with a general trend towards more nutrient‐rich flora with less light at ground level. Species diversity declined, with a marked decreasing trend for the typical species of each alliance. These species were partly replaced by species belonging to nitrophilous and mesophilous grasslands. However, no trend towards drier conditions was detected in these wetlands. The largest changes, with an important colonization by nitrophilous species, occurred in the Swiss sites, where grazing was banned 25 years ago. As a result of floral shifts, many plots previously belonging to fens or wet mesotrophic meadows shifted to an alliance of the wet meadows, generally Filipendulion. Moreover, communities showed a slight trend towards more thermophilous flora.

Conclusions

The investigated wetlands in the Jura Mountains have suffered mainly from eutrophication due to land‐use abandonment and N deposition, with a loss of typical species. Areas with constant land use (grazing or mowing) showed less marked changes in species composition. The most important action to conserve these wetlands is to maintain or reintroduce the traditional practices of extensive mowing and livestock grazing in the wetlands, especially in areas where they were abandoned 25 years ago. This previous land‐use change was intended to improve fen conservation, but it was obviously the wrong measure for conservation purposes.     

Langfristige Auswirkungen ehemaliger Bewirtschaftungsvarianten auf die Diversit?t von Blütenpflanzen, Moosen und Flechten eines Kalkmagerrasens

Jeschke M., Kiehl K., Pfadenhauer J. and Gigon A. 2008. Long-term effects of former management on the diversity of vascular plants, mosses and lichens in a calcareous grassland. Bot. Helv. 118: 95 – 109. Long-term effects of grassland management on the species richness of vascular plants, mosses and lichens were studied in a calcareous grassland near Schaffhausen (Switzerland). Experimental plots were treated for 22 years (1977 – 1999) with different mowing regimes, annual burning in late winter, or abandonment. From 2000 onwards, all plots were mown annually in autumn. Five years later (2004), we studied the after-effects of the previous treatments by recording the cover of vascular plants, mosses and lichens in nested plots with sizes from 0.01 m² to 16 m². Vascular plant species richness as well as the number of species characteristic for calcareous grasslands were significantly lower in formerly abandoned or burnt plots than in regularly mown plots, with larger differences on small subplots. There was no significant effect of the previous mowing time (July vs. October) or mowing frequency (annual or every two years) on vascular plants. Mosses and lichens had been eliminated by annual burning between 1977 and 1999, and only few moss species re-colonized the formerly burnt plots until 2004. The species richness of mosses did not differ between formerly abandoned and mown plots, but there were more mesophytic and fewer xerophytic species in the formerly abandoned plots. Mesophytic mosses were also more abundant in plots that used to be mown every second year than in those mown annually. Our data indicate that even after five years of similar management, i.e. annual mowing in autumn, the previous long-term management had strong after-effects on species richness and species composition of calcareous grasslands. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Eingereicht am 19. M?rz 2008, Angenommen am 30. August 2008, Redaktion: Sabine Güsewell     

Evaluating alternative mowing regimes for conservation management of Central European mesic hay meadows: A field experiment

Conservation status of hay meadows highly depends on their management. The main goal of this study was to assess the efficiency of different mowing regimes in maintenance of plant species richness and diversity of mesic hay meadows. The field experiment was carried out on a species rich, mesic hay meadow in Western Hungary. We evaluated the effects of four alternative types of management on the plant community after 7 years of continuous treatment: (1) mowing twice a year, typical traditional management, (2) mowing once a year in May, most practised currently by local farmers, (3) mowing once a year in September, often proposed for conservation management and (4) abandonment of mowing. Traditional mowing resulted in significantly higher number and higher diversity of vascular plant species than other mowing regimes. Mowing twice a year was the only efficient way to control the spread of the invasive Solidago gigantea, and mowing in September was more successful in it than mowing in May. We conclude that the traditional mowing regime is the most suitable to maintain botanical diversity of mesic hay meadows; however, other regimes should also be considered if certain priority species are targeted by conservation.     

Effects of light competition and litter on the performance of Viola palustris and on species composition and diversity of an abandoned fen meadow

Theeffects of light competition and litter on seedling recruitment and theperformance of established individuals were examined in Violapalustris. This polycarpic perennial plant was a common component ofspecies-rich fen meadows in northwestern Germany until the middle of thiscentury, but today is considered to be regionally endangered. From summer 1996until summer 1998 a bi-factorial field experiment combining three standing croptreatments (mowing, thinning, control) with two litter layer treatments (litterremoval, no litter removal) was carried out in an abandoned fen meadow toinvestigate the effects of these factors both on V.palustris and on aboveground species composition and diversity.MANOVAs (multivariate analysis of variance) revealed significant treatmenteffects for the performance of V. palustris related toexperimental manipulation. Mowing increased vital attributes including thenumber of rosette leaves, the percentage of individuals with chasmogamous andcleistogamous flowers and the number of rhizomes. In contrast, the length of theleafstalk, mean leaf area and the length of rhizomes were negatively associatedwith mowing. Litter removal significantly increased the number of V.palustris seedlings. The removal of the litter layer already resultedin an increase in aboveground species richness in the first year of theexperiment. After three years of experimental manipulation, both abovegroundspecies richness and diversity were positively associated with mowing.Redundancy Analysis showed that typical wet meadow species (Cardaminepratensis, Lychnis flos-cuculi, Lotuspedunculatus) and species of mesotrophic fens (Violapalustris, Agrostis canina, Potentillapalustris) were positively influenced by mowing. It can be concludedthat V. palustris recruitment and growth is promoted by aregular disturbance regime on fen meadows. The conservation of many typicalspecies of wet meadows and fens in northwestern Germany depends on moderategrazing or cutting. These management types prevent successional changes whichlead to an increase in standing crop and the development of a litter layer, bothof which can cause local extinction.     

Vegetation changes following sheep grazing in abandoned mountain meadows

Abstract. Sheep grazing was investigated as an alternative to traditional management of meadows in the Krkono?e Mts. Until the second World War these meadows were mown in mid‐summer and grazed by cattle for the rest of the season. Subsequent abandonment of the meadows has resulted in decreasing species richness. Degradation phases of the former communities have been replacing the original species‐rich vegetation. Significant changes were apparent six years after the introduction of sheep grazing. In grazed plots the proportion of dominant herbs (Polygonum bistorta and Hypericum maculatum) decreased and grasses (Deschampsia cespitosa, Festuca rubra, Agrostis capillaris, Anthoxanthum alpinum) increased. The increase in grasses was positively correlated with an increase in several herbs. The proportion of some herbs increased despite being selectively grazed (Adenostyles alliariae, Melandrium rubrum, Veratrum lobelianum). Any losses caused by grazing of mature plants were probably compensated by successful seedling establishment. Cessation of grazing resulted in significant changes in vegetation within three years. The cover of nitrophilous tall herbs and grasses (e.g. Rumex alpestris, Holcus mollis, Deschampsia cespitosa, Geranium sylvaticum) increased in the abandoned plots. In the plots grazed for nine years cover of species‐rich mountain meadow species increased (e.g. fine‐leaved grasses, Campanula bohemica, Potentilla aurea, Viola lutea, Silene vulgaris). The main conservation risk is the expansion of a competitive species with low palatability, Deschampsia cespitosa. This species can be suppressed by a combination of grazing and mowing. In order for grazing to be effective, the number of sheep should be proportional to meadow production. This may be difficult to maintain as production is variable and is impossible to predict at the beginning of a growing season. A large part of the biomass may thus remain intact in some years. Negative effects of grazing may be, at least partly, eliminated by a combination of cutting and grazing.