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1.
Abstract. Harsh conditions in arid and semi‐arid environments make seedling establishment rare. Plant recruitment in arid environments often occurs only in years with above average rainfall or in safe sites under the canopy of nurse plants that provide shelter from high temperatures and low moisture. Associations of establishing seedlings with adult plants are referred to as nurse‐protégé interactions and are thought to be commensalisms in which seedlings benefit from the micro‐environment created by adult plants with no effect for the latter. This phenomenon is thought to be more frequent in harsh than in mild environments and appears to occur frequently in deserts and arid and semi‐arid biomes. Here, we investigate whether nurse‐protégé interactions are more common in arid environments by searching the published literature from the previous 92 years using the terms nurse plants, protégé plants, facilitation, nucleation and facultative mutualism. We then quantitatively compared these reports from arid zones to other environments. A total of 296 papers were found which referred to nurse‐protégé interactions. More than half (158) focused on arid and semi‐arid zones. This information was also used to explore hypotheses of potential causative forces that might have selected for such interactions in the arid zones such as seed trapping, nutrient, moisture, protection from browsing or trampling and support availability. Because of the large number of different nurse species (147, from 98 genera and 40 families) and protégé species (429, from 273 genera and 84 families), described across a diversity of environments, we suggest that there may be more than one causative factor selecting for nurse‐protégé interactions in arid and semi‐arid environments.  相似文献   

2.
Gymnosperms and angiosperms can co-occur within the same habitats but key plant traits are thought to give angiosperms an evolutionary competitive advantage in many ecological settings. We studied ontogenetic changes in competitive and facilitative interactions between a rare gymnosperm (Dioon sonorense, our target species) and different plant and abiotic neighbours (conspecific-cycads, heterospecific-angiosperms, or abiotic-rocks) from 2007 to 2010 in an arid environment of northwestern Mexico. We monitored survival and growth of seedlings, juveniles, and adults of the cycad Dioon sonorense to evaluate how cycad survival and relative height growth rate (RHGR) responded to intra- and interspecific competition, canopy openness, and nearest neighbour. We tested spatial associations among D. sonorense life stages and angiosperm species and measured ontogenetic shifts in cycad shade tolerance. Canopy openness decreased cycad survival while intraspecific competition decreased survival and RHGR during early ontogeny. Seedling survival was higher in association with rocks and heterospecific neighbours where intraspecific competition was lower. Shade tolerance decreased with cycad ontogeny reflecting the spatial association of advanced stages with more open canopies. Interspecific facilitation during early ontogeny of our target species may promote its persistence in spite of increasing interspecific competition in later stages. We provide empirical support to the long-standing assumption that marginal rocky habitats serve as refugia from angiosperm competition for slow-growing gymnosperms such as cycads. The lack of knowledge of plant–plant interactions in rare or endangered species may hinder developing efficient conservation strategies (e.g. managing for sustained canopy cover), especially under the ongoing land use and climatic changes.  相似文献   

3.
The importance of ontogeny and the degree of abiotic stress in determining the interplay between facilitation and competition is well known. However, their joint effect on the outcome of plant interactions remains poorly understood, especially when a continuous gradient of abiotic stress is considered. Our objective was to evaluate the frequency of association of individuals of Clusia criuva with typical coastal dune species across a gradient of water stress and how this association affects the growth of juveniles and sub-adults. The study was performed in a coastal dune region in South Brazil, where the sandy soil promotes severe water stress. One-year growth of 293 individuals and their distance to the closest humid slacks were measured. This distance is a good surrogate for water stress, since slacks represent proximity to groundwater. The proportion of associated individuals increased with abiotic stress in both ontogenetic stages, but was always greater for juveniles. This suggests that association is progressively more important to guarantee survival as abiotic stress increases. Nonetheless, the benefit of neighbors to growth decreased with abiotic stress, and associated plants grew less than isolated ones in harsher environments. This was mainly true for juveniles, since the height growth of sub-adults was not affected by association or abiotic stress. In our study, facilitation became more intense with environmental severity, increasing survival, although competition also became more influent, reducing growth particularly for younger plants. This demonstrates that ontogenetic stage and abiotic stress must be considered simultaneously in order to better understand interactions among plants.  相似文献   

4.
A nurse–protégé relationship is a frequent facilitation interaction in deserts that allows the recruitment of new individuals of many species. Our aim was to evaluate the relationship of Echinocereus enneacanthus during its life cycle and its nurse plants in the brousse tigrée (tiger bush) vegetation of the Chihuahuan Desert. The population structure of the cactus is skewed toward adult stages, which are commonly found in the bare areas of the brousse tigrée, whereas juveniles are located in the vegetated bands. The vegetation structure and the nurse–protégé relationship were determined using censuses of 19 permanent plots monitored between 2008 and 2019. The differential association of E. enneacanthus with potential nurse plants was evaluated using an interaction network, under the hypothesis of strict association of the seedlings and juveniles with nurse species. In addition, χ2 tests and standardized residuals were fitted among likely nurse species, weighted by the cover of the nurse and of the areas without vegetation. The study species, as well as the nurse plants, had an aggregated distribution pattern. The interaction network showed that the frequency of the association with nurse plants decreased as the size of the protégé individuals increased. At the same time, there were differences in the establishment of E. enneacanthus under particular nurse plants; Prosopis glandulosa and Hilaria mutica were the most important. The nurse–protégé relationship for seeds and seedlings is integral to the vegetation bands of the brousse tigrée.  相似文献   

5.
Elgersma  Kenneth J.  Yu  Shen  Vor  Torsten  Ehrenfeld  Joan G. 《Plant and Soil》2012,352(1-2):341-351

Background and Aims

In line with the Stress Gradient Hypothesis, studies of facilitation have tended to focus on plant–plant interactions (biotic nurses), while the relative role of abiotic nurses has been little studied. We assessed the role of biotic and abiotic nurses, and their interaction, on soil enhancement and the consequential performance of a native annual grass, Dactyloctenium radulans.

Methods

We used a growth chamber study with two levels of water application to compare the performance of D. radulans growing in soil from foraging pits of the Short-beaked echidna (Tachyglossus aculeatus; abiotic nurse) and non-pit soil from either under tree canopies (biotic nurse) or surrounding open areas.

Results

All measures of plant performance were more pronounced under the high than the low water treatment. The greatest differences between pit and surface Microsites occurred under the low water application, reinforcing our view that facilitatory effects are greater in resource-limited environments. Despite tree canopy soil having greater N, there was no significant effect on plant performance, nor any significant interaction with Microsite.

Conclusions

Our study provides strong evidence that foraging pits enhance soil properties and this soil, in turn, facilitates plant growth; and supports previous work documenting the positive effect of nurse-protégé interactions under greater levels of abiotic stress.  相似文献   

6.
Positive and negative plant–plant interactions are major processes shaping plant communities. They are affected by environmental conditions and evolutionary relationships among the interacting plants. However, the generality of these factors as drivers of pairwise plant interactions and their combined effects remain virtually unknown. We conducted an observational study to assess how environmental conditions (altitude, temperature, irradiance and rainfall), the dispersal mechanism of beneficiary species and evolutionary relationships affected the co‐occurrence of pairwise interactions in 11 Stipa tenacissima steppes located along an environmental gradient in Spain. We studied 197 pairwise plant–plant interactions involving the two major nurse plants (the resprouting shrub Quercus coccifera and the tussock grass S. tenacissima) found in these communities. The relative importance of the studied factors varied with the nurse species considered. None of the factors studied were good predictors of the co‐ocurrence between S. tenacissima and its neighbours. However, both the dispersal mechanism of the beneficiary species and the phylogenetic distance between interacting species were crucial factors affecting the co‐occurrence between Q. coccifera and its neighbours, while climatic conditions (irradiance) played a secondary role. Values of phylogenetic distance between 207–272.8 Myr led to competition, while values outside this range or fleshy‐fruitness in the beneficiary species led to positive interactions. The low importance of environmental conditions as a general driver of pairwise interactions was caused by the species‐specific response to changes in either rainfall or radiation. This result suggests that factors other than climatic conditions must be included in theoretical models aimed to generally predict the outcome of plant–plant interactions. Our study helps to improve current theory on plant–plant interactions and to understand how these interactions can respond to expected modifications in species composition and climate associated to ongoing global environmental change.  相似文献   

7.
A popular hypothesis for tree and grass coexistence in savannas is that tree seedlings are limited by competition from grasses. However, competition may be important in favourable climatic conditions when abiotic stress is low, whereas facilitation may be more important under stressful conditions. Seasonal and inter-annual fluctuations in abiotic conditions may alter the outcome of tree–grass interactions in savanna systems and contribute to coexistence. We investigated interactions between coolibah (Eucalyptus coolabah) tree seedlings and perennial C4 grasses in semi-arid savannas in eastern Australia in contrasting seasonal conditions. In glasshouse and field experiments, we measured survival and growth of tree seedlings with different densities of C4 grasses across seasons. In warm glasshouse conditions, where water was not limiting, competition from grasses reduced tree seedling growth but did not affect tree survival. In the field, all tree seedlings died in hot dry summer conditions irrespective of grass or shade cover, whereas in winter, facilitation from grasses significantly increased tree seedling survival by ameliorating heat stress and protecting seedlings from herbivory. We demonstrated that interactions between tree seedlings and perennial grasses vary seasonally, and timing of tree germination may determine the importance of facilitation or competition in structuring savanna vegetation because of fluctuations in abiotic stress. Our finding that trees can grow and survive in a dense C4 grass sward contrasts with the common perception that grass competition limits woody plant recruitment in savannas.  相似文献   

8.
Alpine snowbeds are habitats where the major limiting factors for plant growth are herbivory and a small time window for growth due to late snowmelt. Despite these limitations, snowbed vegetation usually forms a dense carpet of palatable plants due to favourable abiotic conditions for plant growth within the short growing season. These environmental characteristics make snowbeds particularly interesting to study the interplay of facilitation and competition. We hypothesised an interplay between resource competition and facilitation against herbivory. Further, we investigated whether these predicted neighbour effects were species‐specific and/or dependent on ontogeny, and whether the balance of positive and negative plant–plant interactions shifted along a snowmelt gradient. We determined the neighbour effects by means of neighbour removal experiments along the snowmelt gradient, and linear mixed model analyses. The results showed that the effects of neighbour removal were weak but generally consistent among species and snowmelt dates, and depended on whether biomass production or survival was considered. Higher total biomass and increased fruiting in removal plots indicated that plants competed for nutrients, water, and light, thereby supporting the hypothesis of prevailing competition for resources in snowbeds. However, the presence of neighbours reduced herbivory and thereby also facilitated survival. For plant growth the facilitative effects against herbivores in snowbeds counterbalanced competition for resources, leading to a weak negative net effect. Overall the neighbour effects were not species‐specific and did not change with snowmelt date. Our finding of counterbalancing effects of competition and facilitation within a plant community is of special theoretical value for species distribution models and can explain the success of models that give primary importance to abiotic factors and tend to overlook interrelations between biotic and abiotic effects on plants.  相似文献   

9.
Question: A number of studies have demonstrated that the interaction between nurse plants and their protégés changes from one of facilitation to interference as life history stage progresses. Here, I aimed to assess relative fitness of the protégé plant Ziziphus mucronata (Rhamnaceae) under the subcanopy of Acacia erioloba (Leguminaceae), versus in the open, at various stages of the lifecycle. Location: Southern Kalahari, southern Africa. Methods: Germination was compared to assess the effects of shade, maternal origin (under A. erioloba or in the matrix) and soil. Seedlings were transplanted into the field under trees and in the open to ascertain establishment rates. Flower production, seed set and mistletoe infestation were compared between the two microhabitats. Results: Seeds from maternal Z. mucronata plants growing beneath A. erioloba had higher germination rates than those from maternal plants growing in the open. Germination was higher in full sun and in matrix soil, although this result was found in experimental conditions mimicking good rainfall and may not hold under dry or medium rainfall conditions. Z. mucronata demography suggested that the A. erioloba subcanopy is important for establishment, but seedlings transplanted into the matrix had higher survival rates than those under trees. Causes of seedling mortality were different: desiccation in the matrix, but invertebrate herbivory and trampling beneath trees. Mistletoe infestation was higher for adult plants growing in the open. Seed set was not significantly different between subcanopy and matrix environments, but once mistletoe infestation rates and parent plant size were accounted for, seed set was lower for adult plants growing in the open than those growing beneath trees. Conclusions: It appears that the nurse tree environment remains important for Z. mucronata throughout its lifecycle.  相似文献   

10.
There is increasing recognition that both competition and facilitation are important drivers of plant community dynamics in arid and semi-arid environments. Decades of research have provided a litany of examples of the potential for shrubs as nurse plants for establishment of desirable species, especially in water-limited environments. However, interactions with the existing understory community may alter the outcome of interactions between shrubs and understory plants. A manipulative experiment was conducted to disentangle interactions between a native forb species (Penstemon palmeri A. Gray), a native shrub (Artemisia tridentata Nutt.), and a diverse understory of exotic and native forbs and grasses in a semi-arid shrubland of Northern Utah, USA. Seedlings of P. palmeri were transplanted in a factorial design: (1) beneath shrub canopies or into their interspaces and (2) with understory interactions retained or removed. Transplant survival was tracked for roughly 1 year. Shrubs appeared to facilitate P. palmeri survival while interactions with the existing understory community were equivalently negative, leading to overall neutral interactions. Further, positive shrub interactions and negative understory interactions appeared to operate independently and simultaneously. While the debate over the importance of facilitation and competition in driving plant community dynamics continues, our observations strongly suggest that both have considerable effects on plant establishment in A. tridentata communities. Furthermore, our results inform the conservation and restoration of P. palmeri populations, and suggest the utility of nurse shrubs and/or understory thinning as strategies for increasing the diversity of desirable species in the arid and semi-arid western United States shrublands.  相似文献   

11.
Changes in plant interactions along a gradient of environmental stress   总被引:29,自引:0,他引:29  
A combination of competition and facilitation effects operating simultaneously among plant species appears to be the rule in nature, where these effects change along productivity gradients often in a non-proportional manner. We investigated changes in competition and facilitation between a leguminous shrub, Retama sphaerocarpa , and its associate understorey species along an environmental gradient in semi-arid southeast Spain. Our results show a change in the net balance of the interaction between the shrub and several of its associated species, from clearly positive in the water-stressed, infertile environment to neutral or even negative in the more fertile habitat. There was a weakening of facilitation along the fertility gradient as a consequence of improved abiotic conditions. Competition was the most intense for below-ground resources in the less fertile environment while total competition tended to increase towards the more productive end of the gradient. Changes in the balance of the interaction between and among different plant species along the gradient of stress were caused by a decline in facilitation rather than by a change in competition. As both competition intensity and facilitation change along gradients of resource availability, plant interactions are best viewed as dynamic relationships, the outcome of which depends on abiotic conditions.  相似文献   

12.
Biotic interactions play an important role in ecosystem function and structure in the face of global climate change. We tested how plant–plant interactions, namely competition and facilitation among grassland species, respond to extreme drought and heavy rainfall events. We also examined how the functional composition (grasses, forbs, legumes) of grassland communities influenced the competition intensity for grass species when facing extreme events. We exposed experimental grassland communities of different functional compositions to either an extreme single drought event or to a prolonged heavy rainfall event. Relative neighbour effect, relative crowding and interaction strength were calculated for five widespread European grassland species to quantify competition. Single climatic extremes caused species specific shifts in plant–plant interactions from facilitation to competition or vice versa but the nature of the shifts varied depending on the community composition. Facilitation by neighbouring plants was observed for Arrhenatherum elatius when subjected to drought. Contrarily, the facilitative effect of neighbours on Lotus corniculatus was transformed into competition. Heavy rainfall increased the competitive effect of neighbours on Holcus lanatus and Lotus corniculatus in communities composed of three functional groups. Competitive pressure on Geranium pratense and Plantago lanceolata was not affected by extreme weather events. Neither heavy rainfall nor extreme drought altered the overall productivity of the grassland communities. The complementary responses in competition intensity experienced by grassland species under drought suggest biotic interactions as one stabilizing mechanism for overall community performance. Understanding competitive dynamics under fluctuating resources is important for assessing plant community shifts and degree of stability of ecosystem functions.  相似文献   

13.
Niche changes during a species’ lifespan are known as ontogenetic niche shifts. These shifts reflect changes in resource availability, requirements, organisms’ foraging ability and/or size‐dependent biotic interactions. In the plant kingdom, however, this issue remains poorly covered. We investigated nutritional niche shifts over the ontogeny of 23 temperate tree species (among nine phylogenetic families) by a synchronic approach. We used 1963 temporary phytoecological surveys conducted throughout metropolitan French forests. The realised niches of three life‐history stages (seedling: <0.5 m; sapling: >0.5 m and < 8 m; tree: >8 m) in each tree species were modelled on the basis of presence/absence data and the main factors of species distribution (energy, water and nutritional resources). We computed the nutritional optima and amplitudes by bootstrapping partial response curves for the C:N ratio and base saturation rate. We assessed changes in these niche parameters over ontogeny and also evaluated the relative importance of ontogenetic shifts in the differentiation of nutritional niche among the selected tree species. The tree stage was found to occur mainly at higher nutrient availability than the seedling (+16.3% on the nutritional gradient) or sapling (+11.1%) stages. In addition, nutritional niches of tree species exhibited, successively, a niche enlargement in eutrophic conditions and a niche restriction in oligotrophic conditions during growth. These global nutritional niche shifts observed over the species’ lifespan contributed moderately but significantly to the niche separation in temperate tree communities (up to 4.5%). We interpreted niche shifts as a response to an increase in nutritional requirements over ontogeny, leading to an intra‐specific selection where individuals established in eutrophic soils have the maximal fitness. Biotic interactions and temporal changes in the environment may secondarily enhance or counteract the process. The importance of ontogenetic niche shifts requires consideration in the study of species autecology and plant community organisation.  相似文献   

14.
There is currently considerable evidence support that plant community structures are driven by plant-plant interactions (e.g., competition and facilitation). In contrast, there is also evidence demonstrating that plant community structure is affected by the impact of consumer pressure (e.g., grazing). In this study, 15 and 10 Acacia gerrardii nurse plants were selected inside and outside Sudyrah natural reserve (protected) area in western Saudi Arabia, respectively. The understory vegetation abundance (e.g. cover and density) was measured among quadrats around the nurse trees in both protected and unprotected areas to examine the impact of grazing and the positive interaction on the understory species. I found that understory vegetation associated with nurse trees (A. gerrardii) has been driven by both the positive impact of nurse plant and the grazing. Although the understory vegetation was positively affected by the impact of facilitation, the composition of such vegetation has been changed due to the impact of herbivory.  相似文献   

15.
Many biotic interactions influence community structure, yet most distribution models for plants have focused on plant competition or used only abiotic variables to predict plant abundance. Furthermore, biotic interactions are commonly context‐dependent across abiotic gradients. For example, plant–plant interactions can grade from competition to facilitation over temperature gradients. We used a hierarchical Bayesian framework to predict the abundances of 12 plant species across a mountain landscape and test hypotheses on the context‐dependency of biotic interactions over abiotic gradients. We combined field‐based estimates of six biotic interactions (foliar herbivory and pathogen damage, fungal root colonization, fossorial mammal disturbance, plant cover and plant diversity) with abiotic data on climate and soil depth, nutrients and moisture. All biotic interactions were significantly context‐dependent along temperature gradients. Results supported the stress gradient hypothesis: as abiotic stress increased, the strength or direction of the relationship between biotic variables and plant abundance generally switched from negative (suggesting suppressed plant abundance) to positive (suggesting facilitation/mutualism). For half of the species, plant cover was the best predictor of abundance, suggesting that the prior focus on plant–plant interactions is well‐justified. Explicitly incorporating the context‐dependency of biotic interactions generated novel hypotheses about drivers of plant abundance across abiotic gradients and may improve the accuracy of niche models.  相似文献   

16.

Background and Aims

Competition drives self-thinning (density-dependent mortality) in crowded plant populations. Facilitative interactions have been shown to affect many processes in plant populations and communities, but their effects on self-thinning trajectories have not been investigated.

Methods

Using an individual-based ‘zone-of-influence’ model, we studied the potential effects of the size symmetry of competition, abiotic stress and facilitation on self-thinning trajectories in plant monocultures. In the model, abiotic stress reduced the growth of all individuals and facilitation ameliorated the effects of stress on interacting individuals.

Key Results

Abiotic stress made the log biomass – log density relationship during self-thinning steeper, but this effect was reduced by positive interactions among individuals. Size-asymmetric competition also influenced the self-thinning slope.

Conclusions

Although competition drives self-thinning, its course can be affected by abiotic stress, facilitation and competitive symmetry.  相似文献   

17.
Background and Aims The development of plant secondary metabolites during early life stages can have significant ecological and evolutionary implications for plant–herbivore interactions. Foliar terpenes influence a broad range of ecological interactions, including plant defence, and their expression may be influenced by ontogenetic and genetic factors. This study investigates the role of these factors in the expression of foliar terpene compounds in Eucalyptus globulus seedlings.Methods Seedlings were sourced from ten families each from three genetically distinct populations, representing relatively high and low chemical resistance to mammalian herbivory. Cotyledon-stage seedlings and consecutive leaf pairs of true leaves were harvested separately across an 8-month period, and analysed for eight monoterpene compounds and six sesquiterpene compounds.Key Results Foliar terpenes showed a series of dynamic changes with ontogenetic trajectories differing between populations and families, as well as between and within the two major terpene classes. Sesquiterpenes changed rapidly through ontogeny and expressed opposing trajectories between compounds, but showed consistency in pattern between populations. Conversely, changed expression in monoterpene trajectories was population- and compound-specific.Conclusions The results suggest that adaptive opportunities exist for changing levels of terpene content through ontogeny, and evolution may exploit the ontogenetic patterns of change in these compounds to create a diverse ontogenetic chemical mosaic with which to defend the plant. It is hypothesized that the observed genetically based patterns in terpene ontogenetic trajectories reflect multiple changes in the regulation of genes throughout different terpene biosynthetic pathways.  相似文献   

18.
The intensity of competitive interactions between fishes is partly determined by prey use and ontogenetic niche shifts. In a wetland where distinct habitat shifts are missing we compared prey use of three generalist benthivorous sunfishes to look for evidence of ontogenetic, interspecific, and “seasonal” variation in prey composition. Diet analysis revealed evidence of diet ontogeny in warmouth (Lepomis gulosus, 30–152 mm standard length, SL), but not in bluespotted sunfish (Enneacanthus gloriosus, 30–47 mm SL) or dollar sunfish (Lepomis marginatus, 30–60 mm SL). Bluespotted and dollar sunfishes consumed small dipteran and amphipod prey and had similar diets in both seasons suggesting a potential for strong interspecific competition. In the dry season, warmouth shifted from using smaller insect prey to larger decapod and fish prey with increasing size. This shift to prey types that were little used by the other species reduced dietary niche overlap with the other sunfishes. After drought and re-flooding (in the wet season), decapods and small fish were less abundant in the wetland and the warmouth ontogenetic shift was less distinct. When matched for gape width, prey composition differed between warmouth and both dollar and bluespotted sunfishes in the wet season, suggesting differences in sunfish foraging modes, but prey use differences were less clear in the dry season when prey were abundant. Both warmouth ontogenetic diet shifts and seasonal variation in prey use (probably mediated by prey abundance) had strong influences on diet overlap and therefore the potential for intra- and interspecific competition between sunfishes in this wetland ecosystem.  相似文献   

19.
Desertification can be an irreversible process due to positive feedback among degraded plant and soil dynamics. The recovery of semiarid degraded ecosystems may need human intervention. In restoration practices, the abiotic conditions often need to be improved to overcome the positive plant–soil feedback loops. Using nurse‐plants to improve abiotic conditions for introduced individuals (facilitation) has been suggested as an alternative to direct abiotic amelioration. Here, we compared direct abiotic amelioration and facilitation as tools for restoration of semiarid grasslands in Spain. Seedlings and seeds of Lygeum spartum and Salsola vermiculata were planted and sown in a stably degraded semiarid area in Northeast Spain. Two levels of direct abiotic amelioration (ploughing and damming) and indirect abiotic amelioration through facilitation by Suaeda vera nurse shrubs were compared with a control with no amelioration treatment. The control treatment showed low plant establishment, confirming the practical irreversibility of the degraded state. Plant establishment was significantly higher in the three treatments with interventions than in the control treatment. The best treatment depended on the plant trait considered, but damming was in most cases better than plant facilitation. However, facilitation maintained the nutrient‐rich topsoil layer. Given the relative success of facilitation, revegetation using the facilitative effect of nurse‐plants would, in principle, be recommended for restoring semiarid grasslands. Direct abiotic amelioration would be needed under extreme degradation or harsh climatic conditions.  相似文献   

20.
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