Species‐specific positive effects in an annual plant community

Plant facilitation studies commonly test the nurse‐plant hypothesis wherein an adult shrub species enhances the establishment of associated herbaceous species under its canopy. Using field and glasshouse experiments, this hypothesis is extended by testing the following four predictions: (1) nurse‐plant effects can occur between species with similar life‐forms and phenologies (2) positive effects are species specific, (3) the outcome of interactions is life‐stage dependent, and (4) facilitative interactions among annuals are primarily commensal. In the Negev Desert in Israel, the response of an annual plant community to removal of relatively larger annuals, Erodium laciniatum, Erucaria pinnata and Trifolium tomentosum, was tested in the field and in the glasshouse. Removal of these dominants was applied early in the growing season, immediately after germination but before establishment of seedlings, and again mid‐season following establishment and growth to adults. In both the field and glasshouse, the presence of E. laciniatum increased establishment and survival to reproduction of neighbouring plants. These positive effects were life‐stage dependent with more positive effects occurring early in the season, and there was no cost of facilitation to E. laciniatum. This positive effect was species specific in that neither E. pinnata nor T. tomentosum had an effect on the plant community. There was also a cost of association with the nurse E. laciniatum in that biomass of neighbours was reduced. These experiments demonstrate that the positive effects typically detected at larger scales between species of different life‐forms are also occurring at finer spatial scales amongst annuals. This study clearly supports the predictions made in the facilitation literature that effects are species‐specific and highly life‐stage dependent.     

Occurrence patterns of facilitation by shade along a water gradient are mediated by species traits

In disturbed habitats, shade often has facilitative effects on plants by ameliorating water and thermal stresses. Facilitation by shade tends to increase as water availability decreases. At the same time, several studies have suggested that facilitation by shade is not affected by water status or collapses under extremely dry conditions. We hypothesized that traits of beneficiary plants, specifically, the flexibility in the allocation of biomass between shoots and roots, would mediate variation in the relationship between facilitation by shade and water status. To test this hypothesis, we examined the responses of two bog species to shade under various water conditions in a post-mined peatland. The seeds of Rhynchospora alba and Moliniopsis japonica were sown under three water levels (dry: 53% peat water content, wet: 77%, and control: 71%) × two shading levels (50% shaded and unshaded). The survival, biomass, and biomass allocation between the shoots and roots of the two species were monitored for two years. Shade increased the survival and biomass of both species. However, the facilitation of R. alba by shade was independent of water level, whereas the strength of the facilitative effects on M. japonica increased as water content decreased. R. alba preferentially allocated biomass to roots under dry conditions and was highly drought tolerant. M. japonica did not alter the allocation of its biomass in response to either shade or water level and was drought intolerant. Our results suggest that flexibility in biomass allocation of beneficiary plants mediates occurrence patterns of facilitation by shade along a water gradient. The facilitation of species with inflexible biomass allocation by shade through the amelioration of water stress increases as water availability decreases, whereas the facilitation of species with flexible biomass allocation is independent of water status. Such species-specific facilitation would promote the coexistence of diverse species in a community.     

Advanced snowmelt causes shift towards positive neighbour interactions in a subarctic tundra community

Positive and negative species interactions are important factors in structuring vegetation communities. Studies in many ecosystems have focussed on competition; however, facilitation has often been found to outweigh competition under harsh environmental conditions. The balance between positive and negative species interactions is known to shift along spatial, temporal and environmental gradients and thus is likely to be affected by climate change. Winter temperature and precipitation patterns in Interior Alaska are rapidly changing and could lead to warmer winters with a shallow, early melting snow cover in the near future. We conducted snow manipulation and neighbour removal experiments to test whether the relative importance of positive and negative species interactions differs between three winter climate scenarios in a subarctic tundra community. In plots with ambient, manually advanced or delayed snowmelt, we assessed the relative importance of neighbours for survival, phenology, growth and reproduction of two dwarf shrub species. Under ambient conditions and after delayed snowmelt, positive and negative neighbour effects were generally balanced, but when snowmelt was advanced we found overall facilitative neighbour effects on survival, phenology, growth and reproduction of Empetrum nigrum, the earlier developing of the two target species. As earlier snowmelt was correlated with colder spring temperatures and a higher number of frosts, we conclude that plants experienced harsher environmental conditions after early snowmelt and that neighbours could have played an important role in ameliorating the physical environment at the beginning of the growing season.     

Do biotic interactions shape both sides of the humped‐back model of species richness in plant communities?

A humped-back relationship between species richness and community biomass has frequently been observed in plant communities, at both local and regional scales, although often improperly called a productivity-diversity relationship. Explanations for this relationship have emphasized the role of competitive exclusion, probably because at the time when the relationship was first examined, competition was considered to be the significant biotic filter structuring plant communities. However, over the last 15 years there has been a renewed interest in facilitation and this research has shown a clear link between the role of facilitation in structuring communities and both community biomass and the severity of the environment. Although facilitation may enlarge the realized niche of species and increase community richness in stressful environments, there has only been one previous attempt to revisit the humped-back model of species richness and to include facilitative processes. However, to date, no model has explored whether biotic interactions can potentially shape both sides of the humped-back model for species richness commonly detected in plant communities. Here, we propose a revision of Grime's original model that incorporates a new understanding of the role of facilitative interactions in plant communities. In this revised model, facilitation promotes diversity at medium to high environmental severity levels, by expanding the realized niche of stress-intolerant competitive species into harsh physical conditions. However, when environmental conditions become extremely severe the positive effects of the benefactors wane (as supported by recent research on facilitative interactions in extremely severe environments) and diversity is reduced. Conversely, with decreasing stress along the biomass gradient, facilitation decreases because stress-intolerant species become able to exist away from the canopy of the stress-tolerant species (as proposed by facilitation theory). At the same time competition increases for stress-tolerant species, reducing diversity in the most benign conditions (as proposed by models of competition theory). In this way our inclusion of facilitation into the classic model of plant species diversity and community biomass generates a more powerful and richer predictive framework for understanding the role of plant interactions in changing diversity. We then use our revised model to explain both the observed discrepancies between natural patterns of species richness and community biomass and the results of experimental studies of the impact of biodiversity on the productivity of herbaceous communities. It is clear that explicit consideration of concurrent changes in stress-tolerant and competitive species enhances our capacity to explain and interpret patterns in plant community diversity with respect to environmental severity.     

The interplay between species' positive and negative interactions shapes the community biomass–species richness relationship

We used an individual-based spatially-explicit model to assess the role of facilitation and plant strategies in shaping the 'community biomass–species richness' relationship. Facilitation had few impacts on community's richness under both the most benign (high community biomass) and the most severe (low community biomass) environments where its intensity was weak. From medium to high environmental severity, facilitation increased community richness, because all plant strategies were facilitated. In contrast, from low to medium environmental severity facilitation decreased community richness, because only the most competitive species were facilitated, which induced a decrease in the richness of the stress-tolerant species overwhelming the increase in richness of the competitive species. Above all, our simulations show how 'strategy-dependent' interactions among species combine to shape the humped-back biomass–species richness relationship. It also demonstrates that facilitative effects might have long-term negative effects on species richness, which result is not included in current facilitation models.     

Burial disturbance leads to facilitation among coastal dune plants

There is growing evidence that interactions among plants can be facilitative as well as competitive, but knowledge of how disturbances influence these interactions and how they vary with species diversity is lacking. We manipulated plant density, species diversity (richness), and a burial disturbance in a controlled, complete factorial experiment to test theories about the relationships among species interactions, disturbance, and richness. The hypotheses tested were 1) burial disturbance reduces plant performance at all levels of density and richness, 2) burial disturbance can cause net plant interactions to become more facilitative, and 3) facilitation increases with species richness. Burial decreased plant survival by 60% and biomass by 50%, supporting the hypothesis that burial reduces plant performance. In the control (unburied) treatment, there was no difference in proportion survival or per plant biomass between low and high density plots, meaning that neither competition nor facilitation was detected. In the buried treatment, however, high density plots had significantly greater survival and greater per plant biomass than the low density plots, indicating net facilitative interactions. Thus facilitation occurred in the buried treatment and not in the unburied control plots, supporting the hypothesis that facilitation increases with increasing disturbance severity. The hypothesis that facilitation increases with increasing species richness was not supported. Richness did not affect survival or biomass, and there was no richness by burial treatment interaction, indicating that richness did not influence the response of the community to burial. The influence of the disturbance on plant interactions was thus consistent across levels of richness, increasing the generality of the relationship between disturbance and facilitation.     

Understorey plant and soil responses to disturbance and increased nitrogen in boreal forests

Question: How do N fertilization and disturbance affect the understorey vegetation, microbial properties and soil nutrient concentration in boreal forests? Location: Kuusamo (66°22′N; 29°18′E) and Oulu (65°02′N; 25°47′E) in northern Finland. Methods: We conducted a fully factorial experiment with three factors: site (two levels), N fertilization (four levels) and disturbance (two levels). We measured treatment effects on understorey biomass, vegetation structure, and plant, soil and microbial N and C concentrations. Results: The understorey biomass was not affected by fertilization either in the control or in the disturbance treatment. Fertilization reduced the biomass of deciduous Vaccinium myrtillus. Disturbance had a negative effect on the biomass of V. myrtillus and evergreen Vaccinium vitis‐idaea and decreased the relative proportion of evergreen species. Fertilization and disturbance increased the biomass of grass Deschampsia flexuosa and the relative proportion of graminoids. The amount of NH₄⁺ increased in soil after fertilization, and microbial C decreased after disturbance. Conclusions: Our results suggest that the growth of slow‐growing Vaccinium species and soil microbes in boreal forests are not limited by N availability. However, significant changes in the proportion of dwarf shrubs to graminoids and a decrease in the biomass of V. myrtillus demonstrate the susceptibility of understorey vegetation to N enrichment. N enrichment and disturbance seem to have similar effects on understorey vegetation. Consequently, increasing N does not affect the rate or the direction of recovery after disturbance. Moreover, our study demonstrates the importance of understorey vegetation as a C source for soil microbes in boreal forests.     

Establishment, competition and the distribution of native grasses among Michigan old-fields

1 In this study the potential role of competition in influencing the distribution of three displaced native perennial grasses across complex gradients of plant productivity and species composition was investigated in Michigan old-fields. To do this plant removal and propagule addition experiments were conducted at nine old-field sites to examine the effects of living plant neighbours and litter on seedling establishment and growth of target species in relation to community biomass.
2 For two target species, Andropogon gerardi and Schizachyrium scoparium , living plant neighbours suppressed establishment from seed at most sites, and suppressed the growth of transplants at all sites.
3 Plant litter strongly inhibited the seedling establishment of both Andropogon and Schizachyrium at sites of high community biomass and litter accumulation, but had little impact on the growth rate of transplants at any of the sites.
4 The total suppressive effect of the plant community on seedling establishment and transplant growth of both Andropogon and Schizachyrium increased in magnitude in a non-linear fashion with community biomass. These effects increased in magnitude more rapidly across sites of low to medium biomass than sites of medium to high biomass.
5 The results suggest that these native grasses may be restricted to low productivity habitats within this landscape because of strong competitive interference with establishment by the existing vegetation in the most productive sites.     

Counterbalancing effects of competition for resources and facilitation against grazing in alpine snowbed communities

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.     

Species removal and experimental warming in a subarctic tundra plant community

Neighbor interactions are likely to play an important role in subarctic plant communities. We conducted experiments in Interior Alaska in which we crossed species removal with greenhouse warming manipulations. We examined changes in community biomass, and in plant survival and growth of individual species in response to experimental warming and to: (1) removal of whole species versus an equivalent amount of biomass across many species, and (2) removal of subdominant (locally common) versus minor (locally uncommon) plants. Community biomass indicated compensation in growth after removal of minor species and after biomass removal without elimination of entire species, but under-compensation after removal of subdominants. Growth and survival of individual species showed facilitation between some species. Warming increased growth of dominant vascular plants, but at the same time reduced survival, and these impacts were greater for larger, more mesic species than for the smaller species associated with drier habitats. Growth of mosses was reduced by the warming. Removal effects did not differ between warming and ambient conditions. The results indicate that common species are able to reduce resources for others (competitive effect) and increase their growth after neighbor removal, whereas locally uncommon species are not able to respond rapidly to increased resources made available by neighbor removal. Therefore, the impact of the presence of common species on locally uncommon species was facilitative overall, but not vice versa. The balance between disturbances such as changes in temperature and species losses from the community will likely be crucial in determining shifts in subsequent community composition.     

Facilitation by a dwarf shrub enhances plant diversity of human-valued species at high elevations in the Himalayas of Nepal

Facilitation is a global phenomenon that occurs when one species promotes the growth, survival, or reproduction of another species, mostly in stressful environments. However, the importance of facilitation by shrubs in maintaining plant community diversity is not well evaluated in the Himalayas, especially for the richness and conservation of medicinal and human-valued species. Therefore, we aimed to explore the facilitative role of a dwarf shrub species, Berberis angulosa, in maintaining plant composition and richness of human-valued species in the Langtang valley of Nepal's Himalayas. We censused plant species in open patches and beneath Berberis during monsoon and post-monsoon (dry) seasons at three elevations.Total species richness and richness of human-valued species were significantly higher inside the Berberis canopy than in gaps; the former being 39% and the latter 46% greater under shrubs than in open sites. Facilitation by Berberis shrubs promoted plant community diversity irrespective of season and elevation; however, higher differences in mean species richness for both total plant species and human-valued species during the dry season and at high elevation indicated increased facilitation intensity under more stressful conditions. The facilitative effect of Berberis shrubs increased, combining both seasons, overall plant diversity by 19% (total=105), and human-valued species by 16% (total=56). Our results show the importance of facilitation by nurse shrubs in structuring plant communities and protecting medicinal and socio-ecologically important plants, thus enriching ecosystem services in the Himalayas. These results suggest nurse plant species should be incorporated into conservation policies and management strategies for effective biodiversity conservation and sustainability, especially in the face of climate change.     

Microbial activity and soil respiration under nitrogen addition in Alaskan boreal forest

Climate warming could increase rates of soil organic matter turnover and nutrient mineralization, particularly in northern high‐latitude ecosystems. However, the effects of increasing nutrient availability on microbial processes in these ecosystems are poorly understood. To determine how soil microbes respond to nutrient enrichment, we measured microbial biomass, extracellular enzyme activities, soil respiration, and the community composition of active fungi in nitrogen (N) fertilized soils of a boreal forest in central Alaska. We predicted that N addition would suppress fungal activity relative to bacteria, but stimulate carbon (C)‐degrading enzyme activities and soil respiration. Instead, we found no evidence for a suppression of fungal activity, although fungal sporocarp production declined significantly, and the relative abundance of two fungal taxa changed dramatically with N fertilization. Microbial biomass as measured by chloroform fumigation did not respond to fertilization, nor did the ratio of fungi : bacteria as measured by quantitative polymerase chain reaction. However, microbial biomass C : N ratios narrowed significantly from 16.0 ± 1.4 to 5.2 ± 0.3 with fertilization. N fertilization significantly increased the activity of a cellulose‐degrading enzyme and suppressed the activities of protein‐ and chitin‐degrading enzymes but had no effect on soil respiration rates or ¹⁴C signatures. These results indicate that N fertilization alters microbial community composition and allocation to extracellular enzyme production without affecting soil respiration. Thus, our results do not provide evidence for strong microbial feedbacks to the boreal C cycle under climate warming or N addition. However, organic N cycling may decline due to a reduction in the activity of enzymes that target nitrogenous compounds.     

Community traits affect plant–plant interactions across climatic gradients

Plant abundances and demography often vary along gradients of environmental stress, and neighboring plants can amplify or diminish such variation. We asked to what degree the effects of neighboring plants on a focal species can be explained by the traits and abundances of species in the surrounding community. We studied a common understory herb, Trientalis latifolia, across climatic gradients created by topography in the Siskiyou Mountains, southwestern Oregon. We compared Trientalis fitness along these gradients with and without neighbor removal, and asked whether the effects of neighboring plants could be predicted by their community‐weighted trait values and abundances. Environmental conditions alone did not explain whether neighbors had competitive or facilitative effects on Trientalis. However, the environment interacted with neighbor traits and biomass to influence neighbor effects: at cool, higher elevations, high neighbor biomass was associated with stronger facilitative effects, while at warm, lower elevations, high neighbor dissimilarity from Trientalis was associated with stronger competitive effects. We suggest that covariation and interactions among environmental and community characteristics are key to understanding species performance along climatic gradients.     

施肥和刈割对亚高山草甸物种多样性与生产力及其关系的影响

施肥和刈割分别对植物群落物种多样性和生产力有重要的影响。采用不同施肥水平和刈割频度, 研究了施肥和刈割对亚高山草甸植物群落物种多样性与生产力及其关系的影响。结果显示, 5年的施肥和刈割处理对群落地上生物量均有显著影响, 但对物种多样性影响不显著; 物种多样性与生产力之间的关系因施肥和刈割次数的不同而不同, 有负相关、二次函数关系和不相关等几种类型, 多数为不相关。此结果表明施肥和刈割并不总是一致性地影响群落物种多样性与生产力。因此, 不同施肥和刈割处理下的亚高山草甸植物群落生产力与物种多样性之间并没有确定的关系。     

The importance of facilitation in the zonation of shrubs along a coastal salinity gradient

Question: What are the interactive roles of abiotic stress and plant interactions in mediating the zonation of the shrub Tamarix chinensis along a salinity gradient? Location: Yellow River estuary (37°46′N, 119°09′E), northeast China. Methods: We surveyed the zonation of T. chinensis along a salinity gradient and quantified its salt tolerance using a pot experiment. In two field experiments, we transplanted T. chinensis seedlings into salt marsh, transitional zone and upland habitats, manipulated neighbours and quantified survivorship and biomass to examine neighbour effects. We also quantified vegetation effects on abiotic conditions in each zone. Results: Tamarix chinensis dominated the transitional zone, but was absent in upland and salt marsh habitats. In the pot experiment, T. chinensis grew well in freshwater treatments, but was inhibited by increasing salinity. Field experiments revealed that competition from neighbours limited T. chinensis growth in the uplands, while T. chinensis transplants were limited, with or without neighbours, in the salt marsh by high soil salinity. In the transitional zone, however, T. chinensis transplants performed better with than without neighbours. Vegetation removal significantly elevated soil salinity in the transitional zone, but not in other zones. Conclusions: Competition, facilitation and abiotic stress are all important in mediating the zonation of T. chinensis. Within its physiological stress tolerance range, or fundamental niche, it is limited by plant competition in low salinity habitats, and facilitated by neighbours in high salt stress habitats, but cannot survive in salt marshes having salinities above its salt stress tolerance limit. Our results have implications for understanding the relationships between facilitation and stress gradients.     

The influence of vines on an oligohaline marsh community: results of a removal and fertilization study

The effects of competitive suppression by vines on the non-vine plant community have received little attention in temperate habitats. This study investigated the impact vines have on their herbaceous hosts in a wetland community at two soil fertility levels. Plots in an oligohaline marsh were treated in a 2 × 2 factorial design with vine removal and fertilization over two growing seasons. There was no significant interaction between removal and fertilization treatments on any of the measured variables. Vine removal initially caused an increase in light penetration through the canopy, but by the end of the study, plots with vines removed had less light due to a 25% increase in biomass by the plants released from competition with vines. For plots with vines removed, species richness was higher during a brief period in the spring of the second year, but by the end of the study, richness in removal plots decreased relative to controls. Fertilization caused a 40% increase in biomass overall, although only two species, Sagittaria lancifolia L. and Polygonum punctatum Ell., showed dramatic increases. Despite fertilization causing a 40% decrease in light penetration to the ground, no change in species richness was observed. Overall, these results show that vine cover in this wetland suppresses non-vine species and reduces community biomass. Removal of vines increased biomass of non-vine dominants but resulted in only an ephemeral change in species richness. Fertilization did not increase the effects of vines on the non-vine community. Received: 14 November 1996 / Accepted: 10 June 1997     

Effects of fertilization and clipping on species diversity,productivity and their relationship in subalpine meadow

施肥和刈割分别对植物群落物种多样性和生产力有重要的影响。采用不同施肥水平和刈割频度, 研究了施肥和刈割对亚高山草甸植物群落物种多样性与生产力及其关系的影响。结果显示, 5年的施肥和刈割处理对群落地上生物量均有显著影响, 但对物种多样性影响不显著; 物种多样性与生产力之间的关系因施肥和刈割次数的不同而不同, 有负相关、二次函数关系和不相关等几种类型, 多数为不相关。此结果表明施肥和刈割并不总是一致性地影响群落物种多样性与生产力。因此, 不同施肥和刈割处理下的亚高山草甸植物群落生产力与物种多样性之间并没有确定的关系。     

Effects of nutrient availability and neighbours on shoot growth,resprouting and flowering of Erica multiflora

Abstract. To test if low soil fertility and competition limit the performance of Mediterranean shrubs, and if the effects of competition on plant performance were modified by soil fertility, we subjected shrubs of Erica multiflora to a factorial field experiment of fertilization and removal of neighbours around target plants. After 18 months of treatment, fertilization had stimulated the growth of pre-existent sprouts and biomass allocation to stems into new sprouts, but decreased the frequency of sprout flowering. Removal of neighbours increased the number and biomass of new sprouts, the probability of sprout flowering and the biomass of flowers. Fertilization slightly enhanced sprout recruitment and the probability of sprout flowering when neighbours were removed, but did not modify the other parameters of plant performance. According to our results, both low soil fertility and competition limited plant performance. Competition was slightly more intense in fertilized plants, but only in determining sprout and flowering bud stimulation.     

Role of biotic interactions and physical factors in determining the distribution of marsh species along an estuarine salinity gradient

Understanding the mechanisms that shape plant distribution patterns is a major goal in ecology. We investigated the role of biotic interactions (competition and facilitation) and abiotic factors in creating horizontal plant zonation along salinity gradients in the Elbe estuary. We conducted reciprocal transplant experiments with four dominant species from salt and tidal freshwater marshes at two tidal elevations. Ten individuals of each species were transplanted as sods to the opposing marsh type and within their native marsh (two sites each). Transplants were placed at the centre of 9‐m² plots along a line parallel to the river bank. In order to disentangle abiotic and biotic influences, we set up plots with and without neighbouring vegetation, resulting in five replicates per site. Freshwater species (Bolboschoenus maritimus and Phragmites australis) transplanted to salt marshes performed poorly regardless of whether neighbouring vegetation was present or not, although 50–70% of the transplants did survive. Growth of Phragmites transplants was impaired also by competition in freshwater marshes. Salt marsh species (Spartina anglica and Puccinellia maritima) had extremely low biomass when transplanted to freshwater marshes and 80–100% died in the presence of neighbours. Without neighbours, biomass of salt marsh species in freshwater marshes was similar to or higher than that in salt marshes. Our results indicate that salt marsh species are precluded from freshwater marshes by competition, whereas freshwater species are excluded from salt marshes by physical stress. Thus, our study provides the first experimental evidence from a European estuary for the general theory that species boundaries along environmental gradients are determined by physical factors towards the harsh end and by competitive ability towards the benign end of the gradient. We generally found no significant impact of competition in salt marshes, indicating a shift in the importance of competition along the estuarine gradient.