Vegetation change in Southeast Greenland? Tasiilaq revisited after 40 years

Questions: Are there changes in species composition of the oceanic, Low‐Arctic tundra vegetation after 40 years? Can possible changes be attributed to climate change? Location: Ammassalik Island near Tasiilaq, Southeast Greenland. Methods: Species composition and cover of 11 key vegetation types were recorded in 110 vegetation survey plots in 1968–1969 and in 11 permanent plots in 1981. Recording was repeated in 2007. Temporal changes in species composition and cover between the surveys were tested using permutation tests linked with constrained ordinations for vegetation types, and Mann–Whitney tests for individual species. Changes in vegetation were related to climate change. Results: Although climate became warmer over the studied period, most of the vegetation types showed minor changes. The changes were most conspicuous in mire and snowbed vegetation, such as the Carex rariflora mire and Hylocomium splendens snowbed. In the C. rariflora mire, species number and cover of vascular plants and cover of bryophytes increased, whereas in the H. splendens snowbed species numbers of vascular plants, bryophytes, and also lichens increased. Lichen richness increased in the Carex bigelowii snowbed and cover of bryophytes in the Salix herbacea snowbed. No such changes occurred in the Alchemilla glomerulans meadow, Alchemilla alpina snowbed and Phyllodoce coerulea heath. There was no change of species composition within the Salix glauca scrub, A. alpina snowbed, lichen grassland and the Empetrum nigrum and Phyllodoce coerulea heaths. Most changes resulted from increasing frequency or cover of some species; there were very few decreasing species. Most of the increasing species indicate drier substrate conditions. Conclusions: Only minor changes in species composition and cover were detected in the vegetation types studied. These changes were probably caused by milder winters and warmer summers during the years before the 2007 sampling. Climate warming may have reduced the duration of snow cover and soil moisture, particularly in snowbed and mire habitats, where species composition change was most pronounced. However, its magnitude was insufficient to cause a major change in species composition. Thus, on the level of plant community types, tundra vegetation near Tasiilaq was rather stable over the last 40 years.     

Ecology of Racomitrium lanuginosum in British blanket mire - evidence from the palaeoecological record

Abstract

The moss Racomitrium lanuginosum (Hedw.) Brid. is widely distributed in the cool-oceanic north and west of the British Isles, where it is an important element in the vegetation of blanket mire. It has been described as occurring on the present-day surface of British blanket mire in two situations (i.e. where the local mire water table may be lowered, despite high atmospheric humidity), on the tops of tall hummocks and adjacent to areas of peat erosion, where it may be dominant in the vegetation. Accordingly, the occurrence of R. lanuginosum is widely perceived as indicative of drier mire conditions and/or mire degradation. In contrast, recent palaeoecological studies have documented the recurrence of R. lanuginosum in the absence of either hummock upgrowth or peat erosion. Such studies suggest that R. lanuginosum may also occur on blanket mire as a component of climatically-mediated mire development. This paper presents critical new data to document the decomposition of R. lanuginosum and describes the results of recent palaeoecological studies that have negated the exclusive role of R. lanuginosum in mire drying/degradation. We present evidence for the active role of R. lanuginosum in persistent blanket mire development, suggesting the moss may occur during periods of climate change towards increased wetness, after a sustained period of drier conditions. The results are of wider relevance in evidencing the sensitivity of oceanic blanket mire to past climate change.     

Impact of rewetting on the vegetation of a cut‐away peatland

Abstract. We tested whether rewetting improved environmental conditions during peatland restoration and promoted colonization and development of mire vegetation. Vegetation change was monitored in a cut‐away peatland one year before, and four years after, rewetting. Colonizers before rewetting were perennials, mostly typical of hummocks or bare peat surfaces. The main variation in vegetation was related to variation in the amounts of major nutrients and water table level. The wettest site with the highest nutrient level had the highest total vegetation cover and diversity, as well as some species typical of wet minerotrophic mires. Raising the water table level above, or close to, the soil surface promoted development of wet minerotrophic vegetation. Diversity initially decreased because of the disappearance of hummock vegetation but started to recover in the third year. Eriophorum vaginatum and Carex rostrata were both favoured, and bryophytes typical of wet habitats colonized the site. Moderate rewetting promoted the development of Eriophorum vaginatum seedlings and an increase in the cover of tussocks. Bryophytes typical of disturbed peat surfaces spread. In the control site development continued slowly towards closed hummock vegetation. The study showed that raising the water level to, or above, soil surface promotes conditions wet enough for a rapid succession towards closed mire vegetation.     

Compositional differentiation,vegetation‐environment relationships and classification of willow‐characterised vegetation in the western Eurasian Arctic

Question: How does willow‐characterised tundra vegetation of western Eurasia vary, and what are the main vegetation types? What are the ecological gradients and climatic regimes underlying vegetation differentiation? Location: The dataset was collected across a wide spectrum of tundra habitats at 12 sites in subarctic and arctic areas spanning from NW Fennoscandia to West Siberia. Methods: The dataset, including 758 vegetation sample plots (relevés), was analysed using a TWINSPAN classification and NMDS ordination that also included analyses of vegetation‐environment correlations. Results: Based on the TWINSPAN classification, eight vegetation types characterised by willow (cover of upright willows >10%) were discerned: (1) Salix glauca‐Carex aquatilis type, (2) Aulacomnium‐Tomentypnum type, (3) Salix‐Betula‐Hylocomium type, (4) Salix lanata‐Brachythecium mildeanum type, (5) Salix‐Pachypleurum type, (6) S. lanata‐Myosotis nemorosa type, (7) Salix‐Trollius‐Geranium type and (8) Salix‐Comarum palustre‐Filipendula ulmaria type. Willow‐characterised vegetation types were compositionally differentiated from other tundra vegetation and were confined to relatively moist valley and sloping tundra sites, from mire to mineral soils. These vegetation types were encountered across a broad latitudinal zone in which July mean temperature ranged from 6 to 10°C. Conclusions: Willow‐characterised tundra vegetation forms a broad category of ecologically and geographically differentiated vegetation types that are linked to dwarf shrub tundra, shrub tundra or mire. Because of complex ecological gradients underlying compositional differentiation, predicting the responses of willow‐characterised tundra vegetation to a warming climate may be complicated.     

Vegetation and soils of the Meikle Kilrannoch ultramafic sites

Summary

The vegetation of two ultramafic sites (MK1 and MK2) at Meikle Kilrannoch are described. MK1 is dome shaped and has much weathering bedrock whilst MK2 is flatter, lacks weathering bedrock and has probably been entirely peat covered. Six vegetation Groups were recognized: I, high-level blanket bog; II and III, grass heath (with II more sedge-rich); IV, dwarf shrub heath; V and VI, debris (an open vegetation on stony skeletal soils). Vegetation maps were produced which had five mapping units: Group I, Groups II and IH combined, Group IV, Groups V and VI combined, and non-vegetated eroding peat. Three main soil types occur: peat, which underlies Group I; a complex of freely draining magnesian brown soils and imperfectly drained magnesian gleys which bear Groups II–IV; and skeletal soils, derived either from weathering bedrock (MK1 only) or ultramafic drift, which bear vegetation in Groups V or VI. The rare plants are commonest in, but not restricted to, debris vegetation on skeletal soils. The likely plant toxicity of soil magnesium at the sites is reaffirmed but it is suggested that the low plant cover in the debris is more likely to result from low nutrients or intensive frost action or both. Floristic differences between MK1 and MK2 are discussed and quantified for Lychnis alpina (which had 68000individuals on the former site and 46 on the latter) but the causes of the differences remain unexplained.     

Response of the moss Racomitrium lanuginosum to changes in sheep grazing and snow‐lie due to a snow‐fence

Question: What are the responses of Racomitrium lanuginosum moss to altered snow‐lie and sheep use? Location: ACarexbigelowii‐Racomitrium lanuginosum heath on a Scottish montane plateau affected since 1986 by a fenced ski corridor. Methods: Permanent quadrats were set up along transects 45 m long perpendicular to the snow‐fence. Cover was assessed over a 12‐year period from 1990. Pellet‐group clearance counts provided data on sheep usage between 1990 and 1996. Snow‐lie was mapped in the springs of 1991–1996. Results: The snow‐fence created a gradient in sheep use and altered the duration of snow‐lie. At the start of monitoring Racomitrium cover was lower immediately adjacent to the fence, and after 12 years its cover was significantly reduced within 10 m of the fence. Further away from the fence Racomitrium cover was relatively stable. The loss of Racomitrium was correlated both with increased snow‐lie and heavier sheep usage. Grass cover increased near the fence and was related to sheep use. Dicranum fuscescens responded differently to Racomitrium, increasing significantly near the fence. Conclusions: We found that changes in snow‐lie and grazing pressure quickly brought about vegetation change in this montane ecosystem. Racomitrium was the most sensitive species to the changes in grazing and snow‐lie caused by the fence, having the biggest initial changes. Loss of Racomitrium permitted increases of species more resistant to grazing including Dicranum fuscescens and grasses.     

An integrated assessment of ecosystem carbon pools and fluxes across an oceanic alpine toposequence

Alpine ecosystems are predicted to be severely affected by climate change. Cold, wet oceanic-alpine environments may also accumulate large total ecosystem carbon (C) pools, but have rarely been investigated. We assessed C pools and fluxes on a toposequence of oceanic-alpine habitats from blanket mire and boreal Calluna heath to Racomitrium heath, Nardus snowbed and alpine Calluna heath. We quantified C pools in vegetation and soils for each habitat and compared these with C inputs from net primary production (NPP) and outputs via decomposition, measured in a 3-year litter bag experiment. We also investigated principle drivers (temperature, moisture, community composition) of C pool and flux differences between habitats. Total ecosystem C pools were large; 11–26 kg C m^?2 in alpine habitats and 50 kg C m^?2 in blanket mire. Within the alpine zone C storage was greatest in the snowbed. Litter decomposition was slow in all habitats (k?=?0.09–0.29 y^?1) while NPP was within the range reported for continental alpine systems. C pool sizes and C fluxes did not vary consistently with altitude but reflected topographic gradients of temperature and moisture within the alpine zone. Oceanic-alpine ecosystems contain large stores of C which may be vulnerable to the effects of climate change.     

Nutrient limitation and nutrient‐driven shifts in plant species composition in a species‐rich fen meadow

Question: We studied the development and persistence of the effects of nutrient pulses on biomass production and species composition in a fen meadow. Location: Nature reserve, central Netherlands, 5 m a.s.l. Methods: Single pulse fertilization with N and P in a factorial design on an undrained central and a drained margin site in a species‐rich fen meadow (Cirsio dissecti‐Molinietum). Biomass production and species composition were monitored during four years. Results: At the central site, N addition boosted biomass production, but only during one year. The species composition was not changed. P fertilization increased the biomass production and changed the species composition from a vegetation dominated by Carex panicea to a grassland community with abundant Holcus lanatus, but not before the second year. At the margin site, P fertilization changed the species composition in a similar way, but biomass production was not increased. N fertilization had no effect. At both sites the P induced shift in species composition persisted for four years although the P effect declined during the experiment. Conclusions: The biomass responses show that N was limiting in the central site. Another nutrient, besides N and P (probably K) must have been limiting in the marginal site. The fast decline of the N effect on biomass is ascribed to increased denitrification and biomass removal. The delay in the P effect on biomass and species composition and the persistence of the P effect on species composition are ascribed to fast immobilisation and subsequent slow release of fertilizer P in the peat soil. Recurrence of the P pulses is expected to cause permanent changes in species composition.     

Post-drainage changes in vegetation composition and carbon balance in Lakkasuo mire,Central Finland

The post-drainage changes in vegetation composition and carbon balance were studied on four site types (from minero- to ombrotrophic conditions) in Lakkasuo mire, central Finland, by directly comparing undrained and drained parts (30 years ago) of the mire. Drainage had drastically changed the species composition of the sites, especially at the minerotrophic sites, where almost all Sphagna had been replaced by forest mosses. On the ombrotrophic sites much of the mire vegetation still remained 30 years after drainage. Drainage had decreased the C stores in ground vegetation on the minerotrophic sites but increased them on the ombrotrophic sites. The changes were, however, very small compared to the changes in the tree stand, where the C stores had increased at all sites (increasing with nutrient level). The change in peat C balance over the 30-year post-drainage period was negative on the most nutrient-rich site, and positive on the others, increasing with lower nutrient levels. The decrease in the peat C balance on the most nutrient-rich site was compensated by the greater increase in the tree stand C stores and the changes in the total C balance (peat+tree stand+ground vegetation) remained positive on all sites. This revised version was published online in June 2006 with corrections to the Cover Date.     

Fungal associations of roots of dominant and sub-dominant plants in high-alpine vegetation systems with special reference to mycorrhiza

Summary Types of root infection were analysed in healthy dominant and sub-dominant plants of zonal and azonal vegetation above the timberline in the Central and Northern Calcareous Alps of Austria. In the open nival zone vegetation, infection by fungi of the Rhizoctonia type was predominant, vesicular-arbuscular mycorrhizal infection, which was mostly of the fine endophyte (Glomus tenuis) type, being light and mainly restricted to grasses in closed vegetation patches. More extensive Glomus tenuis infection was found in the alpine grass heath, but in Carex, Rhizoctonia was again the most important fungus. The ericaceous plants of the dwarf shrub heath have typical ericoid infection, but quantitative analysis reveals a decrease of infection intensity with increase of altitude. The possible function of the various types of root infection are discussed, and the status of Rhizoctonia as a possible mycorrhizal fungus is considered.     

Hydro-ecological analysis of the fen system Lieper Posse,eastern Germany

The Lieper Posse is a calcareous, ground water-fed fen system, situated in the end-moraine area of northeastern Germany. Nowadays the fen is partly drained and the vegetation adversely affected; hence regeneration measures are considered. Here we report a pilot survey of the actual vegetation cover in relation to the hydrological conditions of the system. The vegetation in the central part of the fen system can be assigned to the Caricetum lasiocarpae, with small areas of Eleocharitetum quinqueflorae. The southern part of the system includes a Ledo-Pinetum sylvestris bog. Along the edges eutrophic forest types are present. Peat analysis revealed that the system started as an open lake and subsequently changed into a rheophilous mire, ‘percolating mire’. The southern bog has only recently come into being. Hydrological investigations showed that the original mineral-rich ground water had to a large extent been replaced by acid rainwater; this is likely due to the construction of a ditch before 1850. The effects of this drainage on hydrological conditions and some aspects of soil fertility are discussed. Possible consequences for the restoration of calciphilous vegetation types are being considered. It is concluded that the vegetation development is predictable only to a limited extent, which is mainly due to a lack of knowledge on hysteresis effects in both vegetation and soil.     

The age of Calluna stands moderates post‐fire regeneration rate and trends in northern Calluna heathlands

Questions: Does stand age influence the direction and rate of post‐fire successional dynamics in coastal Calluna heaths and can old degraded heath vegetation be restored through reintroduction of fire? Location: Coastal heaths in the Tarva archipelago, central Norway. Methods: We investigated revegetation dynamics after experimental fires set in young (8 years since last fire) and old (>50 years since last fire) grazed heath stands. A repeated measures design was used, with floristic data recorded in permanent plots in the post‐fire successions (n=12) over a 7‐year period. The data were analysed using multivariate ordination techniques (PCA, RDA and PRC) and mixed effects models. Results: The age of Calluna stands strongly influenced post‐fire succession, different trends due to age explained 10.4% of variation in floristic data. Young heath showed faster succession towards pre‐fire community composition than old heath, and this could partially be explained by succession‐related factors: young heath had lower cover of mosses and lichens in the pre‐burned vegetation, and lower cover of litter early in succession. Young heath had a less pronounced overall community response to fire than old heath. Vegetative regeneration of C. vulgaris was absent in both old and young heath, but Calluna still re‐established as the dominant species within 5–7 years in both young and old stands. Regeneration dynamics were also affected by habitat conditions, different trends due to habitat explained 6% of variation. Conclusions: Our study demonstrates that old stands do develop characteristic heathland vegetation and structure after fire, and while potential invasives into the system such as trees and rhizomatous species are present, they do not impair Calluna regeneration or vegetation development towards the target heathland community composition and structure. Further, as our young stands are only in their second fire rotation after restoration, we suggest that characteristic dynamics of managed heathlands can re‐establish relatively rapidly, even in severely degenerated sites (>50 years since last fire). Site‐specific factors also need to be considered. We conclude that there is restoration potential in old heaths, despite slow dynamics in the first rotation.     

The application of multivariate land classification to vegetation survey in the Wicklow Mountains,Ireland

Multivariate land classification and land cover mapping by aerial photographic interpretation were used to model spatial variation of land cover in the Wicklow Mountains, Ireland and to structure a stratified random sampling programme of upland blanket bog vegetation. The total area of blanket bog with gully-erosion features was estimated as 33% of the area studied. Vegetation with hand peat-cutting patterns was estimated at 5%, and there was 35% undissected (intact) vegetation. There were differences between land classes in the estimated area of land cover with gully-erosion features or hand peat-cutting patterns.Sample vegetation quadrats, stratified by land class and aerial photographic land cover type, were grouped by their plant species composition. The groups represented ombrotrophic mire, soligenous mire and shrub heath vegetation. There was significant association between vegetation group and land class, related to variation in regional landscape type, but no significant association between vegetation group and the aerial photographic land cover types, undissected (intact) and dissected (gullied and cut-over) peats. It is proposed that the similarity of vegetation between undissected and dissected blanket bog is related to vegetation regeneration. The need to consider differences in vegetation distribution, composition and dynamics in ecological management strategies is emphasised. The study demonstrated the value of stratified random field sampling for cost-efficient regional ecological assessment in upland blanket bog landscapes typified by the Wicklow mountains, Ireland.     

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.     

Photosynthetic traits of Sphagnum and feather moss species in undrained,drained and rewetted boreal spruce swamp forests

In restored peatlands, recovery of carbon assimilation by peat‐forming plants is a prerequisite for the recovery of ecosystem functioning. Restoration by rewetting may affect moss photosynthesis and respiration directly and/or through species successional turnover. To quantify the importance of the direct effects and the effects mediated by species change in boreal spruce swamp forests, we used a dual approach: (i) we measured successional changes in moss communities at 36 sites (nine undrained, nine drained, 18 rewetted) and (ii) photosynthetic properties of the dominant Sphagnum and feather mosses at nine of these sites (three undrained, three drained, three rewetted). Drainage and rewetting affected moss carbon assimilation mainly through species successional turnover. The species differed along a light‐adaptation gradient, which separated shade‐adapted feather mosses from Sphagnum mosses and Sphagnum girgensohnii from other Sphagna, and a productivity and moisture gradient, which separated Sphagnum riparium and Sphagnum girgensohnii from the less productive S. angustifolium, S. magellanicum and S. russowii. Undrained and drained sites harbored conservative, low‐production species: hummock‐Sphagna and feather mosses, respectively. Ditch creation and rewetting produced niches for species with opportunistic strategies and high carbon assimilation. The direct effects also caused higher photosynthetic productivity in ditches and in rewetted sites than in undrained and drained main sites.     

Variations in mountain vegetation use by reindeer (Rangifer tarandus) affects dry heath but not grass heath

Question: Are differences in landscape use of semi‐domesticated reindeer reflected in the vegetation of summer grazing grounds? Location: Alpine heaths, central east Sweden. Methods: Dry heath and grass heath vegetation plots with inferred grazing intensities (high, intermediate and low) were selected a priori from an interpolated pellet count map compiled in 2002. In each plot, faecal pellets were counted, environmental variables measured and vegetation sampled by listing presence and absence. Species composition was compared with a detrended correspondence analysis, and a canonical correspondence analysis was used to infer relations between species composition and environmental variables. Plots were also clustered to provide groupings for an indicator species analysis. Results: Significant differences in faecal pellet count were present between the highest and lowest grazing intensities for both vegetation types, showing that the pattern in the interpolated pellet maps was robust. Differences in species composition between grazing intensities were found for the dry heath only. Here, there was an apparent grazing gradient, with lichens and mosses in the low‐use plots and grasses and herbs in the high‐use plots. No such gradient was found for the grass heath. Conclusions: Within the dry heath vegetation type, grazing levels had a subtle effect on the vegetation, while no effects were seen in the grass heath, probably as a result of the dominance of more grazing‐tolerant graminoids. Even in the dry heath, species richness did not differ between grazing levels, but the relative abundances of species differed.     

The vegetation of the Øvre Heimdalen valley

The catchment area of Øvre Heimdalsvatn reaches from the upper half of the subalpine birch forest zone to the lowermost part of the high-alpine subzone within the alpine zone. The units dominating the area include mountain birch forest, tall herb meadow, mire vegetation, bilberry heath, snowbed communities, chionophobous heath vegetation, and areas without a continuous vegetation layer. It is shown that the potential natural vegetation corresponds closely to the actual vegetation distribution.     

Soil seed banks of woodland, heathland, grassland, mire and montane communities, Cairngorm Mountains, Scotland

The size and species composition of soil seed banks were assessed at 111 altitudinally diverse sites in the Cairngorm Mountains. Mean densities of germinable seeds varied from 83 000 m^–2 in Scots pine (Pinus sylvestris L.) woodland at 230–490 m to 200 m^–2 in moss (Racomitrium lanuginosum (Hedw.) Brid.) heath at 1000–1120 m. Seed banks were dominated by Calluna vulgaris (L.) Hull, not only wherever it was prominent in the vegetation, but also at some sites with less than 5% cover of parent plants in the ground vegetation. Many species conspicuous in the vegetation were under-represented in or absent from the seed bank and surface vegetation generally was more species rich than was the underlying seed bank, especially in high montane communities. Multiple regression was used to examine the relationship between the density of buried Calluna seeds and the abundance of parent plants in the vegetation, site altitude and the organic matter content of the soil. The model fitted to woodland communities accounted for 95% of the variation in seed density. The heathland model was less predictive but still explained 52% of the variation in seed bank size. In mire communities there was no relationship, collective or individual, between buried seed density and the measured environmental variables, possibly due to variations in the duration and frequency of waterlogging at these sites. The potential role of seed banks for initiating the recolonisation of disturbed ground is discussed. Densities of buried seeds at most Calluna-dominant sites were probably sufficient to generate successful recolonisation but the prospects for recovery were poor at other sites, particularly in graminaceous communities at 800 m or higher.     

Forest floor photosynthesis and respiration in a drained peatland forest in southern Finland

Background: Although forest floor forms a large biomass pool in forested peatlands, little is known about its role in ecosystem carbon (C) dynamics.

Aim: We aimed to quantify forest floor photosynthesis (P _FF) and respiration (R _FF) as a part of overall C dynamics in a drained peatland forest in southern Finland.

Methods: We measured net forest floor CO₂ exchange with closed chambers and reconstructed seasonal CO₂ exchange in the prevailing plant communities.

Results: The vegetation was a mosaic of plant communities that differed in CO₂ exchange dynamics. The reconstructed growing season P _FF was highest in the Sphagnum community and lowest in the feather moss communities. On the contrary, R _FF was highest in the feather moss communities and lowest in the Sphagnum community. CO₂ assimilated by the forest floor was 20–30% of the total CO₂ assimilated by the forest. The forest floor was a net CO₂ source to the atmosphere, because respiration from ground vegetation, tree roots and decomposition of soil organic matter exceeded the photosynthesis of ground vegetation.

Conclusions: Tree stand dominates C fluxes in drained peatland forests. However, forest floor vegetation can have a noticeable role in the C cycle of peatlands drained for forestry. Similarly to natural mires, Sphagnum moss-dominated communities were the most efficient assimilators of C.