Vegetation re‐establishment on land previously subject to control of Pteridium aquilinum by herbicide

Abstract. Vegetation development after conventional bracken control with herbicide in the uplands is often slow and rarely results in the development of the target community. Two experiments were established on contrasting sites to investigate cost‐effective methods for the re‐establishment of vegetation dominated by Calluna vulgaris (heather), which is the usual target community seen as the desired replacement for Pteridium aquilinum in the uplands by many land managers. At one site, where grazing intensity was low and shelter was sufficient to prevent rapid Pteridium litter loss, the necessary treatments to establish Calluna were to disturb the litter and add a source of Calluna seeds. A difference between continued Pteridium control or a relaxation of control was not observed over the time‐scale of the experiment. At the second site, where grazing intensity was high and shelter minimal, vegetation dominated by Rumex acetosella had developed on a thin but dense litter layer. To enhance Calluna regeneration it was necessary to prevent sheep grazing, disturb the remaining litter layer and add a source of Calluna propagules. A nurse crop hindered Calluna establishment due to its vigorous growth. At neither site was an initial application of fertilizer an aid to Calluna establishment. It is possible to devise methods for the re‐establishment of Calluna at sites where previous attempts at Pteridium control alone have not had the desired effect on vegetation development. However, the precise choice of methods used is dependent on the grazing management at the site. High levels of livestock grazing prevented Calluna establishment, but some trampling may aid litter breakdown. The ability to adjust stocking rates and access to the site by machinery are critical factors to take into account in producing an effective strategy for long‐term Pteridium control and its replacement by target vegetation.     

Transition from heathland to grassland initiated by the heather beetle

This paper discusses the transition from heather-dominated vegetation to one dominated by grass. The dominating species involved are Calluna vulgaris and Deschampsia flexuosa. The study covered three years, 1980–82, and was done at the Hoorneboeg heathland in the Netherlands. Several factors account for this transition: damage caused by the chrysomelid heather beetle Lochmaea suturalis Thomson, soil type, and interaction between beetle damage and Deschampsia when already present underneath or in the immediate vicinity of a Calluna canopy. There was a time-lag between beetle damage and grass establishment or expansion. In fact, this transition concerns a type of vegetation succession and can be treated as such. This succession therefore involves several causative factors external to the system Calluna-Deschampsia. The successional process is conceived as a stochastic one defined by transition probabilities which depend on a time component, a space component and a logistic response component. All these components are discussed in this paper, but the discussion of yet another component, the crowding of Deschampsia itself, will be treated separately.     

Competition between heather and grasses on Scottish moorlands: Interacting effects of nutrient enrichment and grazing regime

Abstract. Considerable losses and degradation of heathlands (in moorlands and lowlands) have been reported across Europe, with Calluna vulgaris (heather) being replaced by other species, often grasses. Increasing atmospheric nitrogen deposition and overgrazing have been suggested as the driving factors behind this change. This possibility was investigated in a study of the interacting effects of nutrient inputs and grazing on heather and three grass species (Nardus stricta, Deschampsia cespitosa and D. flexuosa) in the field, on a moorland in northeastern Scotland. In addition, the interacting effects of increasing nutrients and Calluna canopy height on N. stricta and D. cespitosa were studied using turves in an outdoor experimental area. In the field, fencing had a larger effect than fertilizer on the growth of all species, except for N. stricta, the species most unpalatable to herbivores. Fencing led to an increase in the height of the Calluna canopy, which may reduce light availability for the grasses. In the turf experiment, the height of the Calluna canopy affected the diameter of the grass tussocks and percentage of green matter (i.e. live leaf material), with plants under the more closed Calluna canopies being smaller. This study suggests that the slow‐growing, evergreen Calluna is a more effective competitor than the faster growing grasses when it has a tall, intact canopy, even at increased levels of nutrient supply. However, overgrazing promotes gap formation in the Calluna canopy, providing an opportunity for grasses to take advantage of increased nutrients. Thus the conservation of heather moorlands requires an understanding of the grazing level which allows Calluna to maintain sufficient canopy structure to outcompete grasses for light.     

Effects of prescribed burning on plant available nutrients in dry heathland ecosystems

Heathland management is an important tool with which to modify ecosystem impacts caused by atmospheric nutrient deposition. Since changes in nutrient availability as a result of management measures affect the outcomes of heathland succession and species competition, studies on this issue are important from both a nature conservation and management point of view. This study reports the effects of prescribed burning on nutrient availability in dry heathland soils and the nutrient content of the two competing heathland species Calluna vulgaris and Deschampsia flexuosa, with particular reference to N and P. We hypothesise that winter prescribed burning leads to additional N availability, which enhances the importance of P in the context of nutrient limitation in heathland ecosystems. In the nature reserve “Lueneburg Heath” (NW Germany) we examined the availability of nutrients in the humus horizons and in the leachate as well as the relevant C:element ratios in Calluna and Deschampsia before and after a burning experiment. Our results show that prescribed burning resulted in drastically increased NH₄⁺ availability in the O-horizon. We observed only short-term effects (for NO₃⁻, PO₄³⁻, Mg) and insignificant effects on the availability of other nutrients (K, Ca). As a consequence of an increased nutrient availability in the humus horizons and a limited nutrient uptake by plants after burning, leaching increased significantly for N, Ca, K, and Mg after burning treatment. No significant changes were found in the foliar C:N ratios for either species after prescribed burning, although Deschampsia showed an increased deficiency for all the other nutrients, particularly for P, as expressed by increased foliar C:P and N:P ratios. By contrast, the nutrient content of Calluna did not change significantly, suggesting that prescribed burning favours the competitive capacity of Calluna as against Deschampsia. We assume that water shortage as a result of changes in the microclimate was mainly responsible for the deterioration of the nutrient content of Deschampsia. This gives Calluna a competitive advantage, enabling it to out-compete Deschampsia on burned heathlands, with respect to the key factor P-limitation.     

Root turnover as determinant of the cycling of C,N, and P in a dry heathland ecosystem

Root production and turnover were studied using sequential core sampling and observations in permanent minirhizotrons in the field in three dry heathland stands dominated by the evergreen dwarfshrub Calluna vulgaris and the grasses Deschampsia flexuosa and Molinia caerulea, respectively. Root biomass production, estimated by core sampling, amounted to 160 (Calluna), 180 (Deschampsia) and 1380 (Molinia) g m^-2 yr^-1, respectively. Root biomass turnover rate in Calluna (0.64 yr^-1) was lower compared with the grasses (Deschampsia: 0.96 yr^-1; Molinia 1.68yr^-1)). Root length turnover rate was 0.75–0.77 yr^-1 (Deschampsia) and 1.17–1.49 yr^-1 (Molinia), respectively. No resorption of N and P from senescing roots was observed in either species. Input of organic N into the soil due to root turnover, estimated using the core sampling data, amounted to 1.8 g N m^-2 yr^-1(^Calluna), 1.7 g N m^-2 yr^-1 (Deschampsia) and 19.7 g N m^-2 yr^-1 (Molinia), respectively. The organic P input was 0.05, 0.07 and 0.55 g P M^-2 yr^-1, respectively. Using the minirhizotron turnover estimates these values were20–22% (Deschampsia) and 11–30% (Molinia) lower.When the biomass turnover data were used, it appeared that in the Molinia stand root turnover contributed 67% to total litter production, 87% to total litter nitrogen loss and 84% to total litter phosphorus loss. For Calluna and Deschampsia these percentages were about three and two times lower, respectively.This study shows that (1) Root turnover is a key factor in ecosystem C, N, and P cycling; and that (2) The relative importance of root turnover differs between species.     

The effects of grazing and nutrient inputs on grass-heather competition

Summary

A combination of field and pot experiments was used to investigate the relative importance of grazing pressure and resource availability in grass-heather competition. In pots where nutrients were added, Nardus acquired the additional nutrients more rapidly than Calluna and increased its above-ground biomass to a greater extent. However, in the field, when fencing and fertilizer treatments were applied to a fine-scale mosaic of Calluna and grasses, including Nardus, nutrient addition only benefited Nardus in unfenced areas. In areas where sheep and deer were excluded, nitrogen addition increased heather cover; Nardus cover declined despite the added nutrients, possibly because the absence of grazing allowed heather to close its canopy above the Nardus, a shade-intolerant species. Effects were site dependent, with fencing increasing heather cover more rapidly on some plots than others; sites where grass cover was declining were the ones where grasses showed the smallest uptake of fertilizer. The results demonstrate that nitrogen additions well above the suggested critical load for heather moorland do not lead to a significant loss of heather cover if herbivory is absent, at least not within three years. It is suggested that the loss of heather moorlands to grass-dominated vegetation in Scotland may principally be driven by grazing, rather than nutrients.     

Competition between two grass species with and without grazing over a productivity gradient

Soil nutrient-level and herbivory are predicted to have opposing effects on the allocation pattern of the competitive dominant plant species. Lower stem and higher leaf allocation are favoured when plants are grazed, whereas a higher stem allocation is favoured at high nutrient levels. Grazing by hares and geese can prevent invasion of the tall Elymus athericus, into short vegetation of Festuca rubra, at unproductive stages of salt-marsh succession but not at more productive stages. We hypothesise that the negative effect of herbivory on Elymus decreases due to increasing soil nitrogen levels and shifts the competitive balance towards this species. We tested how simulated grazing and nitrogen availability affected the competitive balance between adult plants of both grass species in a greenhouse experiment. Elymus had a higher above-ground biomass production, invested relatively more in stem and root tissue and had a larger shoot length than Festuca. The above-ground relative yield of Elymus in mixtures of both species increased with increasing nitrogen levels. This indicates that Elymus was the superior competitor at high soil fertility. Although clipping removed relatively more biomass from Elymus than from Festuca and exceeded the observed biomass removal in field conditions, it did not change the competitive balance between both species. Decreasing effects of herbivory due to increasing nitrogen levels are not a likely explanation for the invasion of Elymus in productive marshes. The results suggest that once Elymus has established it can easily invade vegetation dominated by Festuca irrespective of grazing by herbivores such as hares and geese. Herbivory by small herbivores may mainly retard the invasion of this plant by influencing establishment itself.     

Herbivory of clonal populations: cattle browsing affects reproduction and population structure of Yucca elata

The hypothesis that cattle browsing on inflorescences of the soaptree Yucca elata reduces reproductive success was investigated by comparing recruitment and population structure in six populations protected from grazing, six grazed during the flowering season and five grazed outside the flowering season. Cattle consumed 98% of inflorescences, which were found to be highly nutritious. Reduced recruitment in flower-grazed populations could not be attributed to reduced flower survival, as recruitment in non-flower grazed populations was also reduced. Changes in population structure were due to cattle browsing small caudices, including both genets and ramets. An alternate hypothesis of limited germination in soils compacted by cattle was not supported. Cattle browsing of inflorescences reduced reproductive effort, which may be due to the inability of the plants to resorb nutrients after flowering. Browsing also increased branching, probably through lack of apical dominance, whereas physical trampling increased procumbency. Cattle browsing implies a lack of genetic recruitment, possible local extinction of the yucca moth Tegeticula yuccasella, the exclusive pollinator of Y. elata, and local reduction in insect and bird biodiversity.     

The influence of different grazing regimes on Phragmites‐ and shrub vegetation in the well‐drained zone of a eutrophic wetland

Abstract. The effects of grazing by cattle and horses on vegetation development were studied in the well‐drained border zone of the Oostvaardersplassen nature reserve, a managed eutrophic wetland in the young Zuidelijk Flevoland polder in The Netherlands. At the start of the study period, 12 yr after the area was enclosed by dykes, the vegetation was dominated by Phragmites australis and tall herbs, particularly Cirsium arvense. Over the next 8 yr, different plant communities developed under different grazing regimes. In all communities, C. arvense was gradually replaced by Urtica dioica, and stands dominated by these two species expanded at the expense of P. australis. The shrub Sambucus nigra invaded the stands of C. arvense and U. dioica. Grazing affected the rate of these developments and the degree to which the grass Poa trivialis became dominant. When cattle were enclosed at a relatively high stocking rate in an area of Phragmites australis and tall herbaceous vegetation during summer, P. trivialis became dominant within 4 yr. The introduction of herbivores led to a faster spread of S. nigra, which contains cyanogenic glycosides which only ruminants can detoxify. Horses, as hind‐gut fermenters, did not feed on S. nigra. Grazing pressure and herbivore species, therefore, are two important variables that can be used to manage the development of Phragmites‐ and shrub vegetation: the greater the grazing pressure by cattle or horses the greater the area dominated by grasses, and a relatively high grazing pressure by cattle will retard S. nigra expansion.     

Nest trampling and ground nesting birds: Quantifying temporal and spatial overlap between cattle activity and breeding redshank

Conservation grazing for breeding birds needs to balance the positive effects on vegetation structure and negative effects of nest trampling. In the UK, populations of Common redshank Tringa totanus breeding on saltmarshes declined by >50% between 1985 and 2011. These declines have been linked to changes in grazing management. The highest breeding densities of redshank on saltmarshes are found in lightly grazed areas. Conservation initiatives have encouraged low‐intensity grazing at <1 cattle/ha, but even these levels of grazing can result in high levels of nest trampling. If livestock distribution is not spatially or temporally homogenous but concentrated where and when redshank breed, rates of nest trampling may be much higher than expected based on livestock density alone. By GPS tracking cattle on saltmarshes and monitoring trampling of dummy nests, this study quantified (i) the spatial and temporal distribution of cattle in relation to the distribution of redshank nesting habitats and (ii) trampling rates of dummy nests. The distribution of livestock was highly variable depending on both time in the season and the saltmarsh under study, with cattle using between 3% and 42% of the saltmarsh extent and spending most their time on higher elevation habitat within 500 m of the sea wall, but moving further onto the saltmarsh as the season progressed. Breeding redshank also nest on these higher elevation zones, and this breeding coincides with the early period of grazing. Probability of nest trampling was correlated to livestock density and was up to six times higher in the areas where redshank breed. This overlap in both space and time of the habitat use of cattle and redshank means that the trampling probability of a nest can be much higher than would be expected based on standard measures of cattle density. Synthesis and applications: Because saltmarsh grazing is required to maintain a favorable vegetation structure for redshank breeding, grazing management should aim to keep livestock away from redshank nesting habitat between mid‐April and mid‐July when nests are active, through delaying the onset of grazing or introducing a rotational grazing system.     

Plant nutrient mobilization in temperate heathland responds to elevated CO2, temperature and drought

Temperate terrestrial ecosystems are currently exposed to increased atmospheric CO₂ and progressive climatic changes with increased temperature and periodical drought. We here present results from a field experiment, where the effects of these three main climate change related factors are investigated solely and in all combinations at a temperate heathland. Significant responses were found in the top soils below the two dominant species (Calluna vulgaris and Deschampsia flexuosa). During winter incubation, microbial immobilization of N and ammonification rate decreased in response to warming in Deschampsia soil, and microbial immobilization of N and P decreased in warmed Calluna soil. Warming tended to increase microbial N and P in Calluna but not in Deschampsia soil in fall, and more microbial C was accumulated under drought in Calluna soil. The effects of warming were often counteracted or erased when combined with CO₂ and drought. Below Deschampsia, the net nitrification rate decreased in response to drought and, while phosphorus availability and microbial P immobilization decreased, but nitrification increased in response to elevated CO₂. Furthermore, leaf litter decomposition of both species decreased in response to drought. These complex changes in availability and release of nutrients from soil organic matter turnover and mineralization in response to elevated CO₂ and climate change may influence the future plant carbon sequestration and species composition at temperate heathlands.     

Low vegetation recovery after short-term cattle exclusion on Robinson Crusoe Island

Livestock browsing and grazing are considered serious threats to the conservation of the Juan Fernández Islands’ flora, Chile. Nowadays the elimination of grazing by cattle is very difficult. In order to prevent the␣entrance of cattle into the native forests, an 8.3 km-long fence was established on the main island (Robinson Crusoe). The response of the vegetation was evaluated during 27 months of exclusion. The changes in abundances and composition of species were monitored in 12 permanent plots, each of 25 m², located in ungrazed and grazed areas. We expected an increase in plant height and total ground cover in the ungrazed area, and also a directional compositional change towards forest species. We found five different patterns of variation or non-variation for the 22 species observed: (i) one species (Acaena argentea) diminished in abundance in the ungrazed area; (ii) another one (Conium maculatum) increased in the ungrazed area; (iii) one species (Vulpia bromoides) increased in the grazing area; and some species displayed no net variation, but they showed (iv) a nearly constant occurrence, or (v) they appeared and disappeared sporadically. Contrary to our hypothesis, the vegetation showed no net differences in cover on both sides of the fence. As expected, plant height increased in the ungrazed area. In this area, we detected no noteworthy changes in floristic composition towards forest species. On the contrary, four new pastureland species appeared outside of the exclusion area during the last year of evaluation. Other ungrazed sectors of the island showed qualitative differences from grazed sectors, such as major height and density of plants, and lower intensity of browsing, grazing, and trampling, attributable to a reduction of herbivorous pressure. The modest responses reported in this study could be related to the short lapse of time since the exclusion, soil compaction, overgrowth by a single species (A. argentea), great seasonal variations in different species’ abundances, the reduction in the number of cattle grazing the area previous to the fence’s installation, and persistent herbivory by rabbits (Oryctolagus cuniculus L.). We conclude that the effect of grazing by cattle is␣less than expected, and that there are other factors that delay the recovery of the Juan Fernández pasturelands.     

Grass competition is more important than seed ingestion by livestock for Acacia recruitment in South Africa

Although grazing livestock may have direct negative effects on woody species through herbivory and trampling, (heavy-)grazing is often associated with woody plant encroachment. Two main mechanisms can explain the positive effects of grazers on woody cover: (1) Grazers reduce the interspecific competition with trees and may reduce fuel load, and (2) gut passage through livestock increases seedling establishment by scarification (increased germination) and fertilization by dung (higher survival and growth of seedlings). We tested the effects of fire, grass, cattle ingestion (transit) and dung on germination, seedling height and survival as well as on recruitment of Acacia sieberiana in a sub-humid grassland of South Africa. About 8,000 seeds were planted in the field in a randomized block design. The removal of grass by grazing and/or fire had the most important effect on Acacia recruitment in savanna. Our findings highlight the hierarchy of the main factors affecting Acacia recruitment, which provides new insights to the understanding of woody plant encroachment.     

Consequences of grazing on the vertical transmission of a fungal Neotyphodium symbiont in an annual grass population

Symbiosis between cool‐season grasses and vertically transmitted fungal endophytes are common and significantly impact on ecosystem function. This makes the understanding of the underlying mechanisms to symbiotic individuals frequency in local populations much more interesting. Most studies have been focused on the differential fitness between symbiotic and non‐symbiotic counterparts (relative fitness), barely considering other mechanisms. We performed a microcosms experiment to evaluate whether grazing alters the dynamics of the endophyte Neotyphodium occultans in the annual grass Lolium multiflorum by simultaneously modifying the relative fitness and the endophyte efficiency to be transmitted from host plants to seeds. Grazing was simulated by means of clipping and trampling on symbiotic and non‐symbiotic plants growing separately, in soils obtained from paddocks, differing in their agronomic management history (natural grassland vs. ryegrass promotion). Seed production showed a complex pattern as it depended on the symbiotic status of the plants, the level of grazing and the agro‐ecological context. Grazed plants produced three times fewer seeds than ungrazed plants only in microcosms with endophyte‐symbiotic plants in soils from ryegrass promotion. Endophyte benefits on seed production were exclusively observed in ungrazed plants in the same soil. Symbiotic plants produced symbiotic and non‐symbiotic seeds in all the treatments. While the production of non‐symbiotic seeds by these plants was not affected by grazing and the soil, grazing reduced the production of symbiotic seeds in both contexts. Grazing negative effect on the density of fully infected spikes determined a significant increment in the transmission failures which were not modified by agro‐ecological contexts. Therefore, grazing can modulate symbiosis dynamics through reducing seed production and endophyte transmission efficiency. Transmission has been disregarded, but it is a context‐dependent process that could lead to a gradual reduction in the symbiotic plants frequency in a population if the mutualism effectiveness does not outweigh transmission failures.     

The impact of red deer on liverwort-rich oceanic heath vegetation

Background: There is concern about increasing numbers of large herbivores including red deer (Cervus elaphus), but little is known about their impact on bryophytes.

Aims: This study set out to determine the effect of different localised densities of red deer on the internationally important Northern Atlantic hepatic mat, characteristic of oceanic heath vegetation, at four locations in the Scottish Highlands where sheep have been absent for decades.

Methods: Thirty 7 m × 7 m plots were randomly located in each study area. The standing crop dung pellet group count method was used to estimate red deer density. Species richness, diversity and cover of hepatic mat liverworts were obtained from 1 m × 1 m quadrats placed at random within the sample plots. Calluna vulgaris cover, ericoid height, rock cover, gradient and altitude were also recorded.

Results: Model simplification in analysis of covariance revealed a consistent pattern of decreasing cover of hepatic mat and Calluna with increasing red deer density at all four study areas. Northern Atlantic hepatic mat cover, diversity and species richness were positively correlated with Calluna cover.

Conclusions: The data suggest that Calluna cover is reduced (through trampling and browsing) at high local densities of red deer which has had cascading effects on the Northern Atlantic hepatic mat. Alternative explanations are discussed.     

Competition between grass‐cutting Atta vollenweideri ants (Hymenoptera: Formicidae) and domestic cattle (Artiodactyla: Bovidae) in Argentine rangelands

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Light grazing of saltmarshes is a direct and indirect cause of nest failure in Common Redshank Tringa totanus

The Common Redshank Tringa totanus breeding population on British saltmarshes has declined by over 50% since 1985, with declines linked to changes in grazing management. Conservation initiatives have encouraged low‐intensity grazing of less than one cattle per hectare, but Redshank have continued to decline, even in regions where light grazing was predominant. This study quantified effects of grazing intensity on Redshank nest survival over six lightly grazed saltmarshes with livestock densities between 0 and 0.82 cattle per hectare, in the Ribble Estuary, northwest England. We assessed whether grazing resulted in nest mortality directly through cattle trampling and/or indirectly through grazer modification of habitat that accelerates predation risks. Cattle density was recorded both during the Redshank breeding season and for 1 year prior to the study, to account for both short‐term trampling effects and the longer term effects on vegetation. Results showed that risk of nest loss to trampling increased from 16% at 0.15 cattle per hectare to 98% at 0.82 cattle per hectare in the breeding season. The risk of a nest being predated increased from 28% with no grazing to 95% at 0.55 cattle per hectare based on all year grazing data. These results suggest that even light conservation grazing at less than one cattle per hectare can reduce Redshank nest survival rates to near zero. It may therefore be appropriate to reduce saltmarsh grazing intensities, or change the timing of saltmarsh grazing to reduce the number of livestock present during the Redshank breeding season.     

A reappraisal of cyclical processes in Calluna heath

Small-scale, short-term cycles constitute an important aspect of vegetation dynamics. It has been widely accepted that such cycles are characteristic of unmanaged heath communities where the dominant, Calluna vulgaris, undergoes an age-related series of changes in its growth and morphology. This is thought to give rise to a repetitive sequence of changes in the occupancy of gaps which form in the Calluna canopy and are first colonized by other species, with Calluna re-establishing later. Recently, doubts have been cast as to whether the changes are in fact often repetitive. A new study has shown much greater variability than was previously suspected in the transitions occurring in gaps. There is evidence that, in some instances though by no means all, Calluna can re-establish and thereby initiate a genuine cycle. It seems, however, that such re-entry is seldom by means of seedling establishment, but more frequently by development of adventitious shoots and roots on stems which have come to lie across the gap and have been covered by moss, moist litter or humus.     

Managed livestock grazing for conservation outcomes in a Queensland fragmented landscape

Protecting native biodiversity is a difficult prospect in extremely modified landscapes, especially where high‐impact exotic species are widespread. Using new data and a review of the literature, this paper comments on the use of livestock grazing to manage the invasive and highly combustible pasture grass species, Buffel Grass (Cenchrus ciliaris) and thereby help conserve fire‐sensitive Brigalow (Acacia harpophylla) vegetation in reserves in Queensland, Australia. We cite evidence that shows that grazing is a potentially useful management tool in such cases and its use can be compatible with the protection of both fire‐sensitive vegetation and other native plant species within the understorey. However, there are limitations in implementing grazing within conservation reserves including the lack of a clear understanding of the influence of grazing on biodiversity and resource condition. Importantly, we highlight secondary invasion by the exotic grass Indian Couch (Bothriochloa pertusa) as a key emerging threat that may undermine the biodiversity benefits gained by grazing in reserves. Grazing can be a useful tool for conservation management in particular scenarios, but the associated risks demand accompanying monitoring and reporting of positive and negative impacts to ensure the fundamental aim of biodiversity protection is being achieved.