Nutrient limitation and vegetation changes in a coastal dune slack

Abstract. Basiphilous pioneer species are among the most endangered plant species in The Netherlands. They find most of their refuges in young coastal dune slacks, especially on the Wadden Sea islands. For the purpose of nature management it is important to know which processes control the presence of basiphilous pioneer communities, and to learn about the nature of slacks harbouring the concerning successional sequences. In a large dune slack on the Island of Terschelling, we assessed soil nutrient status and tested for nutrient limitation in four chronosequential stages: 2, 6, 37 and ca. 80 yr of age. Stage 2 harboured a basiphilous pioneer vegetation; in the stages 3 and 4 a dense vegetation of dwarf shrubs and grasses occurred. Soil organic matter and nutrient concentrations in each stage were measured in 1991. In 1992 and 1993 fertilizers were applied to all stages to detect nutrient limitation. Rates of accumulation of organic matter, nutrients and above-ground biomass were estimated. When interpreted as successional stages, the different stages represent a sequence as expected on the basis of general successional theory. There was a peak in yearly nutrient accumulation between the 6- and 37-yr old stage and a steady state after ca. 80 yr. Between the first two and the latter two stages a shift occurred from allogenic to autogenic succession which correlated with a shift in emphasis from available nutrients to light availability as limiting resources. Basiphilous pioneer species suffered only deficiency of nitrogen, probably because of their low phosphorus requirements. It is concluded that in dune slack habitats, in addition to a low nutrient availability in general, a very low phosphorus availability favours basiphilous pioneer species to species showing co-limitation of nitrogen and phosphorus as found in some grasses and dwarf shrubs. A comparison between the effects of lime addition and the effects of nitrogen and phosphorus additions suggests that, in the early stages, soil buffering increases the availability of nitrogen and inhibits the availability of phosphorus. Sod cutting is an effective technique for restoring basiphilous pioneer vegetation, when slacks are acidified only superficially and buffering-mechanisms can be reactivated. Yearly mowing and removing of standing crop may prolong the life-span of basiphilous pioneer vegetation, when soil acidification has not yet dropped below pH 6.     

Is succession in wet calcareous dune slacks affected by free sulfide?

Abstract. Consequences of sulfide toxicity on succession in wet calcareous dune slacks were investigated. Sulfide may exert an inhibitory effect on dune slack plants, but several pioneer species exhibit ROL (Radial Oxygen Loss) and thereby protect themselves against free sulfide. Under oxic conditions free sulfide will be oxiginated to harmless sulfate. However, successive species when not capable of ROL may be sensitive to free sulfide and cannot invade the area. Therefore, the occurrence of free sulfide may have a stabilizing effect on the pioneer vegetation. Data on the vertical distribution of oxygen, redox and sulfide were collected in mesocosms with Littorella uniflora or Carex nigra, with and without microbial mats and compared to control mesocosms. Also, in situ data were collected in a dune slack on the Frisian Island of Texel. In the mesocosms, free sulfide was detected only at nighttime in C. nigra populated mesocosms and in unvegetated units, but not in L. uniflora vegetated mesocosms. In the field, sulfide and redox profiles showed distinct differences between the groundwater exfiltration and infiltration site of the dune slack. At the exfiltration site, sulfide was only occasionally found; in contrast, measurable amounts of free sulfide were regularly found at the infiltration site of the slack. Since Phragmites australis dominates in the infiltration site of the slack, the results suggest that free sulfide accelerate the succession, rather than slowing it down by the exclusion of some plant species.     

Effects of microbial mats on germination and seedling survival of typical dune slack species in the Netherlands

The role of microbial mats in wet dune slack succession is often discussed. We tested if presence of microbial mats may retard dune slack succession by lowering the germination and seeding survival of successor species. This hypothesis was tested on a set of typical dune slack species of the Frisian Islands in two climate chamber experiments. The species were separated into early-, intermediate- and late successional species. There were large differences in germination rates between species (2% – >200% compared to the reference), but within a species the high germination rates were mostly found on sand without a microbial mat. Only the germination of Agrostis stolonifera appeared to be stimulated by the presence of a well-developed microbial mat, they were even higher than in the reference. Seedling survival also did not show different responses between successional stages. Seedlings placed on top of a microbial mat showed for most species lower survival rates compared to seedlings that were planted or placed on top of the sand. Growth was the only measured variable that differed between successional groups. Species of the early- and intermediate successional stages grew significantly better if a microbial mat was present whereas late successional species were not stimulated. Early and intermediate successional species seem to be favored by the presence of a microbial mat. An explanation for this may be that they can profit from the enhanced nitrogen availability caused by N₂-fixation by cyanobacteria in the microbial mat. This revised version was published online in June 2006 with corrections to the Cover Date.     

Vegetation development in dune slacks: the role of persistent seed banks

Abstract. The soil seed bank composition was determined at four sites in the dune slack ‘Koegelwieck’ on the Dutch Wadden Sea island of Terschelling. At three different sites in the slack, where sod-cutting experiments down to the mineral sand had been carried out, the established vegetation and seed bank were assessed after 5, 9 and 39 yr of undisturbed development, respectively. In addition, a fourth site in the slack was investigated, where vegetation development had proceeded for 80 yr since plant colonization of bare soil and where nowadays a vegetation dominated by Calamagrostis epigejos and Salix repens occurs. Together these four sites can be regarded as a chronosequence of dune slack formation. Clear time sequences were detected in the seed bank data. Many late successional species showed a significant increase in the number of seeds during the succession. Some of the early successional basiphilous pioneer species such as Anagallis minima, Centaurium littorale, Littorella uniflora, Radiola linoides and Samolus valerandi, showed either a decrease during the time of succession or an optimum in the vegetation while remaining present in the seed bank in low but detectable numbers. They could, therefore, play a role in re-establishment of the vegetation after sod-cutting. One of the target species, Schoenus nigricans, established within a few years after removal of the sod. However, no seeds of this species have been detected in the soil below either of the successional stages. Based on the species disappearance from the established vegetation and based on the independent data of Thompson et al. (1997) an estimation of seed longevity could be made for several Red List species of wet dune slacks.     

Fen‐meadow succession in relation to spatial and temporal differences in hydrological and soil conditions

Question: In fen meadows with Junco‐Molinion plant communities, falling groundwater levels may not lead to a boosted above‐ground biomass production if limitation of nutrients persists. Instead, depending on drainage intensity and micro‐topography, acidification may trigger a shift into drier and more nutrient‐poor plant communities. Location: Nature reserve, central Netherlands, 5 m a.s.l. Methods: Long‐term study (1988‐1997) in a fen meadow along a gradient in drainage intensity at different scales. Results: Above‐ground biomass increased only slightly over ten years, despite a lower summer groundwater table. The accountable factors were probably a limited availability of nutrients (K in the higher well‐drained plots, P in the intermediate plots and N in the lower hardly drained plots), plus removal of hay. Junco‐Molinion species increased in dry sites and Parvo‐caricetea species increased in wet sites, presumably primarily because of soil acidification occurring when rainwater becomes more influential than base‐rich groundwater. The extent of the shift in species composition depends primarily on the drainage intensity and secondarily on microtopography. Local hydrological measures have largely failed to restore wetter and more basic‐rich conditions. Conclusions: Acidification and nutrient removal, leaching and immobilization resulted in the succession towards Junco‐Molinion at the cost of Calthion palustris elements. Lower in the gradient this change was reduced by the presence of buffered groundwater in slightly drained sites. To conserve the typical plant communities of the Junco‐Molinion to Calthion gradient in the long term, further acidification must be prevented, for example by inundation with base‐rich surface water.     

Succession and fluctuation in a wet dune slack in relation to hydrological changes

Vegetation changes in a wet dune slack complex have been studied over a period of 23 yr. The vegetation was recorded in 40 permanent plots: all plots in 1964, 1977, 1983 and 1987 and some also in five other years. There were large fluctuations in annual precipitation during this period, including some extremely dry (1976) and wet (1985) years. Many species of wet calcareous and pioneer habitats declined in cover between 1964 and 1977, when there were prolonged dry summer periods, though few species were lost from the sampled quadrats. Practically all calciphilous and pioneer species disappeared between 1977 and 1983, when there were relatively many wet summers. It is suggested that extremely dry conditions temporarily obscure the impacts of hydrological changes in local hydrological systems, by retarding the vegetation succession. After a period of excessive precipitation the (acidifying) effects of ground-water withdrawal rapidly became evident in the species composition of the dune slack.     

Wind dispersal in freshwater wetlands: Knowledge for conservation and restoration

Questions: For wetland plants, dispersal by wind is often overlooked because dispersal by water is generally assumed to be the key dispersal process. This literature review addresses the role of seed dispersal by wind in wetlands. Why is wind dispersal relevant in wetlands? Which seeds are dispersed by wind and how far? And how can our understanding of wind dispersal be applied to wetland conservation and restoration? Methods: Literature review. Results and conclusions: Wind is a widely available seed dispersal vector in wetlands and can transport many seeds over long distances. Unlike water, wind can transport seeds in all directions and is therefore important for dispersal to upstream wetlands and to wetlands not connected by surface water flows. Wind dispersal transports seeds to a wider range of sites than water, and therefore reaches more sites but with lower seed densities. Many wetland plant species have adaptations to facilitate wind dispersal. Dispersal distances increase with decreasing falling velocity of seeds, increasing seed release height and selective release mechanisms. Depending on the adaptations, seeds may be dispersed by wind over many km or only a few m. The frequency of long‐distance wind dispersal events depends on these adaptations, the number of produced seeds, the structure of the surrounding vegetation, and the frequency of occurrence of suitable weather conditions. Humans reduce the frequency of successful long‐distance wind dispersal events in wetlands through wetland loss and fragmentation (which reduce the number and quality of seeds) and eutrophication (which changes the structure of the vegetation so that seed release into the wind flow becomes more difficult). This is yet another reason to focus on wetland conservation and restoration measures at increased population sizes, prevention of eutrophication, and the restoration of sites at short distances from seed sources.     

Restoration of the Cirsio dissecti‐Molinietum in The Netherlands: Can we rely on soil seed banks?

Abstract. Vegetation and soil seed banks of a threatened Atlantic fen meadow community were studied using recent phytosociological records and seedling emergence from soil samples. Similarly managed but differently degraded stands that suffered different levels of species impoverishment were compared. The actual vegetation was related to a set of phytosociological references representing the subassociations of the community. DCA positions of reference relevés from the different subassociations were overlapping, suggesting that in all references many common species occur. Recent records were positioned in‐between the seed bank samples and the references. The soil seed banks of all stands were dominated by ordinary species. Most character species had at most sparse seed banks and no seedlings of locally extinct character species, mentioned in historic floristic records, were detected. In contrast species of pioneer and small‐sedge communities as well as those of heathlands were abundant in the seed banks. Based on the vertical distribution of seeds in the soil layers most fen meadow species were classified into transient or short‐term persistent seed bank types. We concluded that complete restoration of the Cirsio dissecti‐Molinietum without reintroduc‐tion is only likely in stands that were degraded only a few years ago. On the other hand, the presence of viable seeds of Nanocyperion and Parvocaricetea species is promising for the restoration of these communities even after decades. Recreation of pioneer habitats by sod cutting will preserve these species.     

Nutrient supply in undrained and drained Calthion meadows

Abstract. Plant species-rich Calthion meadows on mesotrophic fen peat soil extensively cut for hay are among the endangered semi-natural vegetation types in northwestern Europe. They are often badly affected by lowering the groundwater table (drainage) and fertilization. In a comparative study of an undrained site with a Calthion meadow and an adjacent drained site, availability of N, P and K was biologically assessed under field conditions (for two years) as well as in a greenhouse (for 18 weeks) by measuring shoot responsiveness. Also, experimental wetting of intact turf samples taken from both sites was applied in order to study the interaction between nutrient supply and anaerobic soil conditions. It was concluded that the above-ground phytomass yield in the undrained site was restricted by a major shortage of N-supply and a moderate shortage of K-supply by the fen peat soil. The above-ground phytomass yield of the drained site was only reduced by a strongly limited supply of K by the soil. The extent of K-deficiency was larger for the drained site. No P-deficiency was observed in any of the drained or undrained sites. Rewetting turf samples, taken from the drained site, did not change above-ground phytomass yields, suggesting that nutrient supplies were not affected by rewetting. Leaching has likely resulted in a strong reduction of K-supply in the drained site. It is assumed that a shortage in K-supply from the peat soil may have become an important environmental constraint for characteristic plant species of Calthion meadows. This may hamper the development of this meadow type on drained peat soils after rewetting by groundwater discharge.     

Nutrient limitation in species-rich lowland fens

Abstract. Nitrogen, phosphorus and potassium were supplied to some Belgian fens of varying nutrient status and productivity. Plant growth in the lowest productive fen with a species-rich Caricion davallianae vegetation was strongly P-limited. N was ineffective when applied alone, but increased the effect of P-addition when applied together. Summer biomass and plant nutrient concentrations were monitored for four years, and showed partial recovery of nutrient limitation. In a more productive fen dominated by Carex lasiocarpa and in a fen meadow, nutrient limitation was less strong. N limited growth in the productive fen, and N and K were co-limiting in the fen meadow. The P-concentration in the productive fen vegetation showed a marked increase after P-fertilization, but it did not result in higher standing crop. The significance of P-limitation for the conservation of species rich low productive fens is discussed. P-limitation may be an essential feature in the conservation of low productive rich fens: because it is less mobile in the landscape than N and/or because it is an intrinsic property of this vegetation type. Plant nutrient concentrations and N:P-ratios may be used as an indication for the presence and type of nutrient limitation in the vegetation. We found N:P-ratios of 23 to 31 for a P-limited site and 8 to 15 in N-limited sites. This was in agreement with critical values from the literature: N:P > ca. 20 for P-limitation and N:P < 14 for N-limitation. Thus, this technique appears valid in the vegetation types that were studied here.     

The effects of groundwater discharge,mowing and eutrophication on fen vegetation evaluated over half a century

Questions: Were continued groundwater discharge and mowing regimes sufficient for vegetation preservation from 1944 to 1993? Which has a stronger effect on vegetation development; groundwater discharge or mowing? What is the role of surface water eutrophication as driver of vegetation change? Location: Het Hol, The Netherlands (ca. 92 ha, 52°13′N, 5°05′E). Methods: Hydrology was simulated for the late 1940s, early 1960s and 1987. Vegetation maps (1944, 1960, 1975 and 1993) were compared for biotope cover. Vegetation recordings in 1944 and 1987 were compared. Surface water quality was compared between 1950 and 1987. Which sites were mown was reconstructed from an interview. Effects of periodic mowing and groundwater discharge on vegetation development were tested for correlation. Results: Biotope diversity reduced significantly through decrease of semi‐aquatic and tall‐herb biotopes, and expansion of forest. The quagfen terrestrialization sere nearly disappeared from 1987 recordings, while the reed sere did well concerning abundance and species richness. Several typical (rich) fen species disappeared from recordings, while new species were mostly field margin species. Periodic mowing and discharge combined are correlated with increasing species numbers. The P‐concentration in surface water increased while N‐concentration decreased. Conclusions: Preservation of the reed sere was successful, whereas preservation of the quagfen sere was not. Periodic mowing and discharge stimulate species richness, discharge more so than periodic mowing. But slight eutrophication likely induced a shift from P‐limitation to N‐limitation, which stimulated the reed sere at the expense of the quagfen sere.     

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.     

Hydrology of Dutch Cirsio‐Molinietum meadows: Prospects for restoration

Abstract. Fen meadows (Cirsio dissecti‐Molinietum) are seriously threatened by desiccation, acidification and eutro‐phication. In The Netherlands several projects were launched to restore damaged fen meadows. This review describes how successes and failures of these restoration projects depend on hydrological systems. Six hydrological systems have been distinguished, which all provide the site conditions required by this community. Nowadays, the best developed fen meadows are found in the higher Pleistocene landscape of The Netherlands, where they depend on base‐rich groundwater discharging from local or large groundwater systems. Fen meadows of the lower Holocene landscape usually occur in man‐made surface water systems. Almost all stands have been severely deteriorated. Restoration of fen meadows in the Pleistocene landscape is promising when the hydrology is only slightly disturbed or when hydrological measures are taken in combination with sod cutting. Restoration prospects of fen meadows in the Holocene landscape are low. Until now a complete regeneration of Cirsio‐Molinietum meadows has not been realized. Restoration measures failed to restore high pH values in the top soil. It is hypothesized that viable seeds of many target species lack in the soil seed bank. In addition, the dispersal capacities of these species seem to be limited.     

Forest species in an agricultural landscape in the Netherlands: Effects of habitat fragmentation

Abstract. For 312 forest patches on sandy soils in the Netherlands, effects of fragmentation are studied of forest habitat in the past on the present occurrence of forest plant species. Using regression techniques, the numbers of forest edge, interior, zoochorous and anemochorous species, as well as occurrence of 24 individual species were related to patch area and connectivity measures. Connectivity was defined as the amount of forest habitat around patches within three zones up to 1000 m. Plant categories were distinguished by habitat type and dispersal mechanism. The results showed that number of total species and number of species of all habitat and dispersal categories increased with area. The occurrence of ten individually studied species were also positively related to area. Most of them were interior species. The number of zoochorous species increased with increasing connectivity. Also occurrence of ten individually studied species were affected by connectivity. Interior zoochorous species showed the highest percentage of affected species. The relationship of interior, animal-dispersed plants to connectivity can be explained by the limited distances covered by their dispersal agents (forest birds and ants) in a non-forest habitat. Also, some anemochorous plants appeared to be affected by connectivity, especially those with heavy seeds and potentially short distance dispersal. As not all species within a certain dispersal or habitat category react similar to area or isolation, it is suggested that differences in underlying processes of fragmentation such as local extinction and colonization need more focus.     

Vegetation response after restoring the connectivity between a river channel and its floodplain

Questions: How is restored inundation affecting the ground‐water composition in a floodplain? To what extent has the floodplain plant community composition changed? What are the effects of flooding frequency and intensity and water quality on the floodplain vegetation changes? Location: Demer river, Flanders, Belgium. Methods: 75 10 m x 10 m plots were surveyed in 1997 and in 2003. Vegetation changes were quantified using a Detrended Correspondence Analysis on the combined 1997–2003 data‐set. Groundwater quality in the floodplain was analysed on macro‐ions. Inundation intensity and frequency were related to vegetation changes. Results: The inundation water was rich in nutrients, although only for NH₄ and ortho‐PO₄, and a significant increase was recorded in the groundwater. Plant species characteristic of very nutrient‐rich herb vegetation disappeared or decreased while the cover and species richness of species from wet, moderately productive habitats increased. Very few new species colonized the floodplain. Conclusion: Nutrient‐rich flooding water does not necessarily result in increased abundance of species from nutrient‐rich habitats, very likely because N becomes less available by deni‐trification and is transformed into biomass by tall herbs. Either micro‐site limitation or dispersal limitation could explain the colonization problems of new species. The data do not allow distinction between these two mechanisms.     

Vegetation succession in a floating mire in relation to management and hydrology

Abstract. The vegetation succession in a floating mire was studied in relation to management and hydrological conditions in a former peat cutting area in the northern part of the Netherlands. An existing map showed that in 1956 the vegetation consisted mainly of meadows, reedbeds and rich fen vegetation while a recent survey revealed that this picture had completely changed in 1989. The area had mainly become woodland but part had remained open, due to the continuation of a mowing regime. The timing of the mowing appears to be critical, especially during the earlier phases of the succession. Winter-mowing favours the development of eutrophic reedbeds, while summer-mowing promotes mesotrophic sedge communities. In the course of time these succession lines converge increasingly towards acidic vegetation types. The development from open water to embryonic bog differed greatly between sites with a similar management regime. Mostly this took only a few decades but in a few sites the vegetation had hardly changed since 1956. Whereas the former sites were acid throughout the profile, the latter showed a high pH from top to bottom. It is obvious that acidification does not occur as long as alkaline surface water can move freely underneath the floating mat. As soon as a given site becomes disconnected from the surface water the depletion of bases by infiltrating rainwater is no longer compensated. A rough estimate of the acidification rate shows that in an infiltration area three decades are sufficient for a floating mat of 40 cm to become completely decalcified. This corresponds well to the observed succession from rich fens to embryonic bogs in the same period. It is suggested that the best way to preserve rich fen vegetation is to start the succession anew by digging turf ponds.     

Occurrence of pollen and spores in relation to present-day vegetation in a Dutch heathland area

Abstract. Pollen, spores and fungal remains in moss cushions along a transect, traversing a Betula-Quercus forest and a surrounding heathland, were analysed in order to study the relation between present-day vegetation and recent pollen deposition. Pollen and spores are divided into local types from plant species encountered along the transect and regional types, not present along the transect. Relative percentages and absolute concentration of the palynomorphs were estimated. Radiocaesium activity in the moss was measured to assess the minimal duration of palynomorph accumulation (i.e. the age of the moss samples). The absence of simultaneous trends in the regional pollen types indicates that the samples do not have major differences in age. The moss represents the later stages in the heath mosaic cycle. This is supported by the radiocaesium content of the moss. Thus, in this case the concentration values of regional pollen approach deposition values. The values of the local pollen types calculated on such a rather constant regional pollen flux is considered to represent real differences in the local deposition. The local pollen types can be arranged topographically into five groups characteristic of the local vegetation types and their ecotones. The arrangement of the curves of types from phyco-, zoo-, and myco-coenoses suggests their possible origin, either locally from the heath area or from the Betula-Quercus forest, or regionally from elsewhere. Analyses and topographical arrangement of the curves of unknown types contribute in this way to their identification and their possible source of origin.     

Higher acidification rates in fens with phosphorus enrichment

Question: Why is bryophyte succession in eutrophicated fens faster than in natural fens? Location: Mineral‐rich fens in The Netherlands and NW Europe. Methods: Literature review on the ecology of four bryophyte species in various successional types as observed in Dutch fens. Results: Bryophyte succession in eutrophicated fens from the brown moss Calliergonella cuspidata to Sphagnum squarrosum is much faster than in natural fens with species shifts from Scorpidium scorpioides to Sphagnum subnitens. Under P‐poor conditions, the brown moss stage is stabilized as long as mineral‐rich water is supplied. This is because S. scorpioides is tolerant of rainwater, is a strong competitor and can counteract acidification to some extent while S. subnitens is intolerant to groundwater and has low growth rates and low acidification capacity. In contrast, the Sphagnum stage is stable after rapid succession from rich‐fen mosses under P‐rich conditions. Calliergonella cuspidata has suboptimal growth in rainwater, possibly due to ammonium toxicity, while the high growth rates of S. squarrosum in nutrient‐rich and highly acidic groundwater allow early establishment and rapid expansion. Conclusions: If measures to improve fen base status occur in environments of increased nutrient (P) availability, the management may not lead to the desired restoration of brown moss stages, but instead to rapid acidification by S. squarrosum.     

Relationships between vegetation zonation and altitude in a salt-marsh system in northwest Spain

Abstract. Vegetation zonation in salt marshes has traditionally been attributed largely to altitudinal differences, since altitude determines the temporal pattern of tidal flooding and is thus closely related to proximate determinants of the distribution of species and plant communities. We investigated the distribution of vascular plants and plant communities along a series of altitudinal transects in two salt marshes in the northwestern Iberian Peninsula. Our results indicate that altitudinal range varies significantly both among species and among communities, and confirm that salt-marsh vegetation characteristics (species cover and composition) can be predicted on the basis of altitude, particularly at the lower levels of the profile.