The effect and extent of heavy grazing by reindeer in oligotrophic pine heaths in northeastern Fennoscandia

Thirty and fifty years old exclosures established in northeastern Fennoscandia in lichen-rich oligotrophic pine Pinus sylvestris forests on podzolised soil were used to study the effect of reindeer grazing on pine fine roots, microbial activity, and on bryophyte, dwarf shrub and lichen biomasses. There were significantly less lichens, especially Cladina stellaris , at grazed than at ungrazed sites. Coverage of other lichens like C. arbuscula and C. rangiferina and bryophytes, especially Dicranum spp., benefitted from grazing. The biomass of vascular plants, mainly Calluna vulgaris, Empetrum nigrum and Vaccinium vitis-idaea , was reduced at grazed sites, although their coverage was not influenced. Microbial activity was significantly lower at grazed sites. The influence of grazing is most obviously mediated by reduced soil moisture during dry periods at grazed sites. Fine root parameters (per soil and stem volumes) were lower at grazed sites (pPCA = 0.072), the first principal component consisting of a number of fine root tips, length and weight. Grazing decreased all exchangeable nutrients by 30–60% in organic layer. Based on PCA the decrease was significant for exchangeable nutrients, although of individual elements only P and S showed statistically significant difference. The extent of heavy grazing in northeastern Fennoscandia coniferous forest was revealed by remote sensing. It revealed extensive area in which reindeer lichens are reduced in northeastern Finland. The Finnish-Russian border can be clearly distinguished in the satellite image composite.     

Growth of reindeer lichens and effects of reindeer grazing on ground cover vegetation in a Scots pine forest and a subarctic heathland in Finnish Lapland

Reindeer lichens are an important component of northern ecosystems. The aim of this study was to measure the growth rate of terricolous lichens as it is a key parameter involved in productivity of these ecosystems and an important part of lichen tolerance to reindeer grazing. Furthermore, the natural succession and the long-term effects of reindeer grazing on lichen community characteristics in two contrasting habitats were investigated as well as the interactions between lichen cover and mosses and vascular plants. Biomass and coverage measurements were conducted in a lichen woodland and in a subarctic heath with grazed and ungrazed areas in northern Finland. Measurements spanning over 13 yr of undisturbed development show that the growth rate of Cladina stellaris can be as high as >0.17 g g⁻¹ produced annually, although in average growth rates were much lower. During the succession of ground vegetation, C. stellaris , C. rangiferina , C. mitis and Cetraria nivalis increased in biomass in fenced areas and were reduced most in biomass by reindeer in unprotected areas. Reindeer grazing and trampling seem to change the vegetation towards a type that is dominated by small dwarf shrubs, bare soil and minute-cup lichens ( Cladonia spp.). Removing the lichen layer by reindeer may reduce natural regeneration of pine trees as implied by increasing numbers of pine seedlings with increasing lichen cover.     

Community food web, decomposition and nitrogen mineralisation in a stratified Scots pine forest soil

A soil community food web model was used to improve the understanding of what factors govern the mineralisation of nutrients and carbon and the decay of dead organic matter. The model derives the rates of C and N mineralisation by organisms by splitting their uptake rate of food resources into a rate at which faeces or prey remains are added to detritus, a rate at which elements are incorporated into biomass, and a rate at which elements are released by organisms as inorganic compounds. The functioning of soil organisms in the mineralisation of C and N was modelled in the soil horizon of a Scots pine forest. The organic horizon was divided into three distinct layers, representing successive stages of decay, i.e. litter, fragmented litter, and humus. Each of the layers had a different, quantitative, biota composition. For each layer the annual C and N mineralisation rates were simulated and compared to observed C and N mineralisation rates from organic matter in stratified litterbags. Simulated C and N mineralisation was relatively close to measured losses of C and N, but the fit was not perfect. Discrepancies between the observed and predicted mineralisation rates are discussed in terms of variation in model parameter values of those organisms that showed the highest contribution to mineralisation rates. The measured, and by the model predicted, significant decrease in mineralisation rates down the profile was not explained by the biomass of the primary decomposers and only partly by the total food web biomass. Modelling results indicated that indirect effects of soil fauna, due to trophic interactions with their resources, are an important explanatory factor. In addition, the analyses suggest that community food web structure is an important factor in the regulation of nutrient mineralisation. The model provided the means to evaluate the contribution of functionally defined groups of organisms, structured in a detrital food web, to losses of C and N from successive decay stages.     

The effect of fire on nutrients in a pine forest soil

The effect of a hot summer fire on soil nutrient contents in the upper 2 cm of Aleppo pine forest with a dense woody understory was studied from September 1985 to May 1986. In comparison with the adjacent unburned forest, total nitrogen decreased by 25% but available forms of nitrogen were much higher. In burned and unburned soils there was a similar trend to increase and decrease in NH ₄ ⁺ −N, However, while (NO ₂ ⁻ +NO ₃ ⁻ −N decreased in the unburned soil it rose rapidly in the burned ash soil. Total phosphorus increased by 300% after the fire but decreased again 2 months later. Also water-soluble P increased up to November and then decreased to the levels of the unburned soils. The same was true for electrical conductivity and pH, increasing immediately after the fire and then leveling off again. This increase in nutrient levels in the “ash soil” was reflected in the striking increase in shoot and root biomass and in the content of N, P, Mg, K, Ca, Zn and Fe in wheat and clover plants grown in pots in these soils. These nutrient levels were much higher in the wheat plants, which also produced 12 times more seeds in the “ash soil.” It seems that fire in these pine forests causes a short-term flush of the mineral elements in the upper “ash soil” layer which is reverted gradually via the herbaceous post-fire to the ecosystem.     

Nitrogen mineralization and H+ transfers in a Scots pine (Pinus sylvestris L.) forest soil as affected by liming

H⁺ production due to N uptake in a mature Scots pine stand subjected to high NH₄ ⁺ deposition was previously estimated to amount to approx. 2.2 kmol ha^-1 y^-1. The question whether H⁺ transfers related to N mineralization (ammonification and nitrification) offset or corroborate this proton production is investigated in the present research. To determine N mineralization, soil cores were used of which both ends were closed with layers of ion exchange resin (IER) to prevent influx and efflux of ions. The effect of liming on N mineralization and the resulting H⁺ production was investigated in 7 incubation periods of each ca. 8 wk. Because of its high mobility NO₃ accumulated in both IER layers at the expense of that in the incubated forest floor and mineral soil. Net N mineralization in the soil cores as a whole amounted to 40 and 77 kg N ha^-1 in 384 d in the control and limed plots, respectively. In both treatments ca. 65% of mineralized N was nitrified. H⁺ production due to N mineralization amounted to approx. 1.2 kmol ha^-1 y^-1 in the control and limed plots. Liming reduced the amount of C in the forest floor, but not forest floor mass, because of an increased mixing with mineral particles.     

Nutritional disturbances of young Scots pines caused by pine bark bugs in a dry heath forest

This study deals with the effect of pine bark bugs (Aradus cinnamomeus Panzer) on the nutrition of young Scots pines (Pinus sylvestris L.). Soil and needle samples for analytical purposes were collected from a young pine stand growing on a dry barren mineral soil afflicted by pine bark bugs.The damage to vascular tissues caused by pine bark bugs disturbed the nutrition of the trees, especially in the top part of the crown. The foliar calcium, magnesium, manganese and sulphur concentrations were highly reduced. Scots pines suffered from a lack of calcium, magnesium, nitrogen and phosphorus. These deficiencies were secondary by nature, because no differences were observed between the nutrient concentrations of the underlying soil of the healthy and affected trees. The symptoms of trees damaged by pine bark bugs resembled most of all calcium deficiency symptoms.     

The effects of mycorrhizal roots on litter decomposition, soil biota, and nutrients in a spodosolic soil

We studied the effects of mycorrhizal pitch pine (Pinus rigida) roots on litter decomposition, microbial biomass, nematode abundance and inorganic nutrients in the E horizon material of a spodosolic soil, using field microcosms created in a regenerating pitch pine stand in the New Jersey Pinelands. Pine roots stimulated litter decomposition by 18.7% by the end of the 29 month study. Both mass loss and N and P release from the litter were always higher in the presence of roots than in their absence. Nutrient concentrations in decomposing litter were similar, however, in the presence and absence of roots, which suggests that the roots present in the with-root treatment did not withdraw nutrients directly from the litter. The soil was slightly drier in the presence of roots, but there was no discernible effect on soil microbial biomass. The effects of roots on soil extractable inorganic nutrients were inconsistent. Roots, however, were consistently associated with higher numbers of soil nematodes. These results suggest that, in soils with low total C and N contents, roots stimulate greater activity of the soil biota, which contribute, in turn, to faster litter decomposition and nutrient release.Contribution No. 95-22 from the Institute of Marine and Coastal Sciences.Contribution No. 95-22 from the Institute of Marine and Coastal Sciences.     

Effects of reindeer grazing on understorey vegetation in dry Pinus sylvestris forests

Abstract. Data on floristic composition and environmental variables were collected in floristically homogeneous oligotrophic pine (Pinus sylvestris) forests with heath-like under- storey vegetation in eastern Fennoscandia, and ordinated by non-linear multidimensional scaling (NMDS) in order to study the effect of lichen grazing by reindeer on the understorey vegetation. The study sites included areas with varying grazing pressure, as well as 50-yr old grazing exclosures. Sites rich in respectively bryophytes and lichens were placed at opposite ends of the ordination axes, and heavily grazed sites were placed in between them. Reindeer grazing increased the abundance of bryophytes, especially Dicranum spp. and Pleurozium schreberi. Grazing changed the vegetation to the extent that it resembled more mesotrophic sites, but this did not show any relationship with tree volume or other site productivity indicators. This was observed both in the ordination and, in a more compelling way, when exclosures with adjacent grazed areas were compared. No such signs were evident at ungrazed sites, where especially Cladina spp. spatially replace Cladonia spp. and tiny bryophytes like Barbilophozia spp., Polytrichum spp. and Pohlia nutans during succession. Cladina stellaris had almost disappeared from the most intensively grazed sites. The soil at ungrazed sites was characterized by high Al and Fe concentrations and bryophyte-rich sites by high Mn concentrations. Shannon's diversity index, depth of humus layer and proportion of bare ground also increased in sites getting richer in bryophytes.     

Effects of grazing intensity on soil carbon stocks following deforestation of a Hawaiian dry tropical forest

The effects of forest-to-pasture conversion on soil carbon (C) stocks depend on a combination of climatic and management factors, but factors that relate to grazing intensity are perhaps the least understood. To understand the long-term impact of grazing in converted pastures, methods are needed that accurately measure the impact of grazing on recent plant inputs to soil C in a variety of pasture management and climate settings. Here, we present an analysis from Hawai'i of changes in vegetation structure and soil organic carbon (SOC) along gradients of grazing intensity and elevation in pastures converted from dry tropical forest 100 years ago. We used hyperspectral remote sensing of photosynthetic vegetation, nonphotosynthetic vegetation (NPV) and exposed substrate to understand the effects of grazing on plant litter cover, thus, estimating recent plant inputs to soils (the NPV component). Forest-to-pasture conversion caused a shift from C3 to C4 plant physiology, thus the δ ¹³C method was used in soil cores to measure the fraction of SOC accumulated from pasture vegetation sources following land conversion. SOC decreased in pasture by 5–9 kg C m⁻², depending upon grazing intensity. SOC derived from C3 (forest) sources was constant across the grazing gradient, indicating that the observed variation in SOC was attributable to changes in C inputs following deforestation. Soil C stocks were also reduced in pastures relative to forest soils. We found that long-term grazing lowers SOC following Hawaiian forest-to-pasture conversion, and that these changes are larger in magnitude that those occurring with elevation (climate). Further we demonstrate a relationship between remotely sensed measurements of surface litter and field SOC measurements, allowing for regional analysis of pasture condition and C storage where limited field data are available.     

Quantification of extraradical soil mycelium and ectomycorrhizas of Boletus edulis in a Scots pine forest with variable sporocarp productivity

The availability of most edible ectomycorrhizal mushrooms depends on their natural fructification. Sporocarp formation of these fungi is linked to habitat characteristics and climate conditions, but these data alone do not explain all the trends of fungal fruiting and dynamics. It could be hypothesized that the amount of soil mycelia could also be related to the production of carpophores. Soil samples (five cylinders of 250 cm³ per plot) were taken monthly, from September to November, in five fenced permanent plots (5 × 5 m) in Pinar Grande (Soria, Spain), a Pinus sylvestris stand situated in the north of the Sistema Ibérico mountain range. Plots were chosen to establish a gradient of Boletus edulis productivity from 0 to 38.5 kg/ha year, according to the mean fresh weight of sporocarps collected during the last 10 years. B. edulis ectomycorrhizal root tips were identified in each soil sample according to its morphology and counted. DNA extractions were performed with the PowerSoil^TM DNA Isolation Kit and quantification of extraradical soil mycelium by real-time polymerase chain reaction using specific primers and a TaqMan? probe. The concentration of soil mycelium of B. edulis (mg mycelium/g soil) did not differ significantly between plots (p = 0.1397), and sampling time (p = 0.7643) within the fructification period. The number of mycorrhizal short roots per soil volume showed significant differences between the plots (p = 0.0050) and the three sampling times (p < 0.0001). No significant correlation between the number of mycorrhizas and the productivity of the plot (kg of B. edulis/ha year) was detected (p = 0.615). A statistically significant positive correlation (p = 0.0481) was detected between the concentration of mycelia of B. edulis in the soil samples and the presence of short roots mycorrhizal with B. edulis in these samples. The productivity of the plots, in terms of sporocarps produced during the last 10 years, was not correlated either with the concentration of soil mycelium or with the presence or abundance of ectomycorrhizas.     

Diurnal patterns in Scots pine stem oleoresin pressure in a boreal forest

Coniferous tree stems contain large amounts of oleoresin under positive pressure in the resin ducts. Studies in North‐American pines indicated that the stem oleoresin exudation pressure (OEP) correlates negatively with transpiration rate and soil water content. However, it is not known how the OEP changes affect the emissions of volatile vapours from the trees. We measured the OEP, xylem diameter changes indicating changes in xylem water potential and monoterpene emissions under field conditions in mature Scots pine (Pinus sylvestris L.) trees in southern Finland. Contrary to earlier reports, the diurnal OEP changes were positively correlated with temperature and transpiration rate. OEP was lowest at the top part of the stem, where water potentials were also more negative, and often closely linked to ambient temperature and stem monoterpene emissions. However, occasionally OEP was affected by sudden changes in vapour pressure deficit (VPD), indicating the importance of xylem water potential on OEP as well. We conclude that the oleoresin storage pools in tree stems are in a dynamic relationship with ambient temperature and xylem water potential, and that the canopy monoterpene emission rates may therefore be also regulated by whole tree processes and not only by the conditions prevailing in the upper canopy.     

The effect of liming on quantity and chemical composition of soil organic matter in a pine forest in Berlin,Germany

The study was carried out in a 40-yr old pine plantation on a Cambic Arenosol within the urban area of Berlin. Lime application (6.1 t ha^-1) has led to a pH increase in the forest floor from 3.3 to 5.5 within one year and to a strong stimulation of macrofaunal and microbiological activity. Three years after liming, the C:N ratio of the forest floor decreased from 28 to 25 and P, Pb, Zn, Cu and Cd concentrations in organic matter increased significantly. The organic C pool of the forest floor was almost 7 t ha^-1 lower in the limed plot which is attributed to increased microbial respiration. In the mineral soil too, C-pools are lower in the limed plot, amounting to 13.2 t ha^-1 or 14% less than in the control. C:N ratios have narrowed significantly from 27–29 to 23 in 10–30 cm depth. The humic acid fraction is lower throughout the limed profile while the percentage of fulvic acids has increased significantly below 10 cm. The results point to severe losses of organic matter and to profound changes in its composition. This may be of consequences for site quality and leaching processes.     

Carbon sequestration and ecosystem respiration for 4 years in a Scots pine forest

The net exchange of CO₂ (NEE) between a Scots pine (Pinus sylvestris L.) forest ecosystem in eastern Finland and the atmosphere was measured continuously by the eddy covariance (EC) technique over 4 years (1999–2002). The annual temperature coefficient (Q₁₀) of ecosystem respiration (R) for these years, respectively, was 2.32, 2.66, 2.73 and 2.69. The light‐saturated rate of photosynthesis (A_max) was highest in July or August, with an annual average A_max of 10.9, 14.6, 15.3 and 17.1 μmol m^?2 s^?1 in the 4 years, respectively. There was obvious seasonality in NEE, R and gross primary production (GPP), exhibiting a similar pattern to photosynthetically active radiation (PAR) and air temperature. The integrated daily NEE ranged from 2.59 to ?4.97 g C m^?2 day^?1 in 1999, from 2.70 to ?4.72 in 2000, from 2.61 to ?4.71 in 2001 and from 5.27 to ?4.88 in 2002. The maximum net C uptake occurred in July, with the exception of 2000, when it was in June. The interannual variation in ecosystem C flux was pronounced. The length of the growing season, based on net C uptake, was 179, 170, 175 and 176 days in 1999–2002, respectively, and annual net C sequestration was 152, 101, 172 and 205 g C m^?2 yr^?1. It is estimated that ecosystem respiration contributed 615, 591, 752 and 879 g C m^?2 yr^?1 to the NEE in these years, leading to an annual GPP of ?768, ?692, ?924 and ?1084 g C m^?2 yr^?1. It is concluded that temperature and PAR were the main determinants of the ecosystem CO₂ flux. Interannual variations in net C sequestration are predominantly controlled by average air temperature and integrated radiation in spring and summer. Four years of EC data indicate that boreal Scots pine forest ecosystem in eastern Finland acts as a relatively powerful carbon sink. Carbon sequestration may benefit from warmer climatic conditions.     

模拟氮沉降对温带阔叶红松林地土壤氮素净矿化的影响

以长白山阔叶红松混交林为研究对象,于2006—2008年原位模拟不同形态氮((NH4)2SO4、NH4Cl和KNO3)沉降水平(22.5和45kgN·hm-2·a-1),利用树脂芯法技术(resin-core incubation technique)测定了表层(有机层0～7cm)和土层(0～15cm)土壤氮素净矿化、净氨化和净硝化通量的季节和年际变化规律。同时,结合前人报道的有关林地碳、氮过程及其环境变化影响的结果,力求有效预估森林生态系统中氮素年矿化通量对大气氮沉降量和水热条件等因子变化的响应。结果表明,长白山阔叶红松林地土壤氮素年净矿化通量为1.2～19.8kgN·hm-2·a-1,2008年不同深度的土壤氮素年净矿化通量均显著高于2006和2007年(P<0.05)。随着模拟氮沉降量增加,土壤氮素净矿化通量也随之增加,尤其外源NH4+-N输入对净矿化通量的促进作用更为明显(P<0.05),但随着施肥年限的延长,这种促进作用逐渐减弱。与林地0～15cm土壤相比,氮沉降增加对0～7cm有机层氮素净氨化和净矿化通量的促进作用更为明显,尤其NH4Cl处理的促进作用更大。结合前人报道的野外原位观测结果,土壤氮素年净矿化通量随氮素沉降量的增加而增大,氮沉降量对不同区域森林土壤氮素净矿化通量的贡献率约为52%;氮沉降量(x1)和pH值(x2)可以解释区域森林土壤氮素年净矿化通量(y)变化的70%(y=0.54x1-18.38x2-109.55,R2=0.70,P<0.0001)。前人研究结果仅提供区域年均温度,未考虑积温的影响,这可能是造成年净矿化通量与温度无关的原因。今后的研究工作应该加强区域森林土壤积温观测,进而更加准确地预估森林土壤氮素的年净矿化通量。     

Effects of livestock grazing intensity on soil biota in a semiarid steppe of Inner Mongolia

Intensive livestock is known to significantly affect soil physical and chemical parameters in steppe ecosystems. However, the effects on soil biological parameters still remain unknown. We hypothesized that intensive grazing would significantly decrease the size and diversity of soil biota due to deterioration of the soil environment and reduction in vegetation cover, while the adapted grazing intensity would improve the biological parameters. Soil samples were collected from five sites with different grazing intensities and history in a semiarid steppe of Inner Mongolia in August 2005. Two sites were long-term ungrazed since 1979 (UG79) and 1999 (UG99), one had been moderately grazed in winter (WG), one continuously grazed moderately (CG) and one long-term site was heavily grazed (HG). Soil microbial biomass carbon (C), basal respiration (BR), catabolic diversity of soil microbial communities, protozoa and nematodes abundance were measured. Soil physicochemical variables were also measured to establish the relationships between soil biological parameters and key soil physical and chemical properties. Soil microbial biomass C, BR, biomass specific respiration (qCO₂) and soil protozoa abundance were significantly lower at the HG site compared to the UG79 site, but no clear differences were found in the other sites. However, soil nematodes abundance increased with increasing grazing intensity, and the abundance of soil amoeba were greater in CG than in the other sites. Principal component analysis (PCA) of Biolog data revealed large differences in catabolic capacity of soil microbial communities between UG79, HG and UG99, WG, CG. However, Shannon??s diversity index did not indicate marked effects of grazing intensity on substrate catabolic community structure. In conclusion, heavy grazing negatively affected soil microbial biomass, activity and protozoan abundance, but positively influenced soil nematodes abundance and did not affect soil microbial catabolic diversity. Based on these results, CG may provide an appropriate grazing intensity to be used in the long term in the semiarid steppe of Inner Mongolia.     

Mites (Acari,Mesostigmata) in boreal Scots pine forest floors: effect of distance to stumps

Coarse woody debris (CWD) is a basic component of forest ecosystems and it plays a crucial role in species-poor boreal forests. Generally, previous studies have focused on differences between the forest floor and decaying logs of various tree species. The impact of distance to CWD has been investigated mainly for forest-floor snails and some groups of macrofauna, but not yet for mesostigmatid mites communities. We hypothesized that the effect of CWD decreases with increasing distance from CWD. To test this hypothesis we conducted a study in relatively species-poor Finnish boreal forest (at ca. 100 km northwest of Helsinki). In total, 81 samples were collected in 2007 from nine Scots pine (Pinus sylvestris) stumps, three microhabitats (CWD, soil/litter at 0.5 m from a stump and soil/litter at 1.5 m from a stump) and in three main directions (9 stumps × 3 microhabitats × 3 directions). Overall, 1965 mesostigmatid mites were collected representing 24 species. The mean number of mite species collected was significantly different between decaying stumps and forest litter; however, there was no significant difference between the litter samples at 0.5 and 1.5 m distance. The evenness index was significantly lower for samples collected from stumps than for litter in close (0.5 m) or far (1.5 m) distance. The most frequently encountered mite species were Veigaia nemorensis, Parazercon radiatus and Zercon zelawaiensis.     

Abundance, distribution and feeding relations of root/fungal feeding nematodes in a Scots pine forest

In a 15–20 yr old forest of Pinus sylvestris, Tylenchus spp., Paratylemchus spp? and Malenchus tantulus were the most abundant stylet-bearing nematodes. Soil layers and ground cover affected the abundance. Mycophagy was observed for Tylenchus sp., M. tantulus, Aphelenchoides sp. and Tylencholaimus stecki. Feeding on roots of the forest grass, Deschampsia flexuosa was recorded for W. tantulus, Paratylenchus spp? and Rotylenchus sp. The latter fed also on pine roots. Connections occurred between feeding habits and field distribution. The facultative feeding habit of M. tantulus probably allowed this species to reach higher numbers under D. flexuosa. The importance, in the field, of the grass Calamagrostis arundinacea for Paratylenchus spp? was confirmed by laboratory experiments. As pine was a non-host, these nematodes were probably grass feeders. Paratylenchus spp? dominated the obligate root feeders with forest grass being a key-factor for the group abundance. Suspected high densities of Paratylenchus spp? on clearings should not affect the growth of pine seedlings and although Rotylenchus sp. to some extent reduced seedling growth its pathogenic status on P. sylvestris was doubtful.     

Effects of a great cormorant colony on wood-inhabiting fungal communities in a coastal Scots pine forest

We studied the changes in the wood-inhabiting fungal communities on dead wood of Scots pine (Pinus sylvestris) under varying duration and intensity of cormorant nesting. The activities of the birds created large amounts of dead wood, changed wood chemistry and altered stand characteristics. Analyses based on records of fruiting structures of fungi showed that fungal communities in the cormorant colony differed from communities in unaffected forest. Ornithogenic factors impacted least on generalist species and promoted the occurrence of fungi uncharacteristic of coniferous wood or wood in general. The rate of ornithogenic change was rapid: a shift in fungal assemblages was evident by the third year of expansion of the bird colony into unaffected forest. It featured the disappearance of tremelloid fungi, a decline of corticioid fungi and an increase of hyphomycetoid fungi. The proportion of coniferous wood specialists decreased and that of generalists increased. It is expected that the changes in fungal communities will persist until the forest is fully regenerated.