Rosa multiflora’s performance under water stress: the role of positive and negative density-dependent intraspecific interactions

It is well known that biotic interactions may significantly alter plant population and community dynamics in natural ecosystems. Multiple studies have reported that density-dependent positive interactions can modify the effect of extreme stress on plant performance (in line with predictions by the stress gradient hypothesis, SGH). However, the performance of invasive species and the role of intraspecific density dependence, either negative or positive, in successful invasions are not well understood. In the present study, we tested if monocultures of Rosa multiflora, a highly invasive shrub, experience a group advantage under extremely low water availability. Using a manipulative greenhouse study, we tested the growth response of R. multiflora seedlings to a 4-level conspecific density treatment and a 4-level water availability gradient. Overall, our results provide preliminary evidence that biotic interactions under stressful conditions between R. multiflora individuals do not experience a group advantage, as predicted. Instead, our study indicates that seedling performance was strongly driven by negative density dependence, in which individuals grown at higher conspecific densities displayed lower plant performance, regardless of water availability. This demonstrates that R. multiflora experiences intense intraspecific competition even under a stressful, low resource environment. Thus, management efforts of R. multiflora need to ensure that re-sprouting of individuals after treatment is eliminated, as release from negative density dependence could lead the invader population to rebound.

     

Shift from facilitative to neutral interactions by the cushion plant Silene acaulis along a primary succession gradient

Background

The stress‐gradient hypothesis predicts a shift from facilitative to competitive plant interactions with decreasing abiotic stress. This has been supported by studies along elevation and temperature gradients, but also challenged by the hypothesis of a facilitation collapse at extremely harsh sites. Although facilitation is known to be important in primary succession, few studies have examined these hypotheses along primary succession gradients.

Aim

To examine whether there is a relationship between the presence of the circumpolar cushion plant Silene acaulis and other species, and if so, whether there is a shift between positive and negative interactions along a primary succession gradient in a glacier foreland.

Location

Finse, southern Norway.

Methods

We examined the performance of the common alpine forb Bistorta vivipara, species richness of vascular plants, bryophytes and lichens, and the number of seedlings and fertile vascular plants in S. acaulis cushions, and control plots without S. acaulis, along a succession gradient with increasing distance from a glacier front, and thus decreasing abiotic stress. To examine if S. acaulis cushions modify the abiotic environment, we recorded soil temperature, moisture, organic content and pH in cushions and control plots.

Results

Bistorta vivipara performed better, as shown by bigger leaves in S. acaulis cushions compared to control plots in the harshest part of the gradient close to the glacier. There were few differences in B. vivipara performance between cushion and control plots in the more benign environment further away from the glacier. This suggests a shift from facilitative to mainly neutral interactions by S. acaulis on the performance of B. vivipara with decreasing abiotic stress. A trend, although not significant, of higher vascular species richness and fertility inside S. acaulis cushions along the whole gradient, suggests that S. acaulis also facilitates community‐level species richness. The causal mechanism of this facilitation is likely that the cushions buffer extreme temperatures.

Conclusions

Our results support the stress‐gradient hypothesis for the relationship between the cushion plant S. acaulis and the performance of a single species along a primary succession gradient in a glacier foreland. S. acaulis also tended to increase vascular plant species richness and fertility regardless of stress level along the gradient, suggesting facilitation at the community level. We found no collapse of facilitation at the most stressful end of the gradient in this alpine glacier foreland.     

Competition and coexistence of rhizomatous perennial plants along a nutrient gradient

I studied competition and coexistence of three tall clonal perennial plant species, Calamagrostis epigejos (L.) Roth, Solidago canadensis L., and Tanacetum vulgare L. along a gradient of soil productivity over five years. A replacement series field experiment was conducted with high, moderate and low fertility levels in 1m×1m plots. There were significant effects of soil type on ramet density (P<0.001), mean height (P<0.01), and total biomass (P<0.01). Ramet density, mean height, and total biomass increased with increasing soil fertility. There were also significant effects of mixture on ramet density (P<0.01), but not on mean height and total biomass for all species. Significant neighbor effects on ramet density and total biomass (P<0.01) were found for Solidago, showing that it is important whether Tanacetum or Calamagrostis is its neighbor within mixtures. During the five years there was only one case of competitive exclusion: Calamagrostis excluded Solidago on the most fertile substrate in the fifth growing season. In most cases species coexisted over the five years. Each of the three species was able to dominate in at least one combination of substrate type and mixture. The experiment showed that asymmetric competition for light on substrates of high fertility, symmetric competition for nutrients on nutrient-poor soil and positive interactions especially on substrates of intermediate fertility played a role. A founder effect was evident in aggregated mixtures of Calamagrostis and Solidago on the nutrient-rich substrate. A conceptual model of the relative importance of root competition for soil nutrients, shoot competition for light, and positive interactions along the fertility gradient is presented. The model emphasizes that positive interactions play an important role over a broad range of the productivity scale with a peak at intermediate levels of fertility. On the substrate of high productivity shoot competition for light is more important than positive interactions and root competition for soil nutrients as well. The competitive superiority of Calamagrostis on the most productive substrate was evident only in the long run. Rare events like extreme summer drought or selective herbivore pressure caused a switch in dominance in mixtures with Solidago, respectively Tanacetum. The guerrilla growth strategy of Calamagrostis and interference competition through a dense cover of aboveground biomass and litter could further cause competitive exclusion.     

Competitive displacement and predation between introduced and native mud snails

Experimental field and laboratory studies indicate that Cerithidea californica, a native mud snail, is restricted to only a portion of its normal habitat range in San Francisco Bay as a result of direct interactions with an introduced ecological equivalent, Ilyanassa obsoleta. The native snail typically inhabits marsh pans, tidal creeks and mudflats in estuaries along the Pacific coast. However, in San Francisco Bay it is confined to pans for most of the year, while the non-native snail inhabits the creeks and mudflats. Experiments and field monitoring demonstrate that this abnormal distribution pattern is caused by 1) interference competition for space in the form of an adult-adult behavioral avoidance by C. californica in the presence of invading I. obsoleta, and 2) predation by I. obsoleta on the eggs and juveniles of C. californica. The competitive exclusion of C. californica by I. obsoleta has not led to the extinction of the native snail because of the existence of a refuge for C. californica in pan habitats, beyond the physiological tolerances of I. obsoleta. As a consequence of the seasonal migrations of both species and changes in abiotic factors along the habitat gradient, repeated competitive displacements, rather than a one-time competitive exclusion, are observed between these two species. This is the first documented case of the competitive displacement of an endemic marine intertidal species by an introduced ecological equivalent.     

Intra-generic competition among Nothofagus in New Zealand's primary indigenous forests

Competitive interactions between New Zealand's four Nothofagus or southern beech species were analysed using an extensive dataset describing the composition of natural forests, supplemented by environmental estimates describing both climate and landform. Using multiple regression models of progressively increasing complexity, the analysis first accounted for variation in tree abundance attributable to both environment and regional-scale distributional disjunctions of likely historic origin. Intra-generic competition, expressed as variation in tree abundance dependent on the presence or absence of each congener, was then assessed by adding (1) simple terms to assess the magnitude of gross changes in abundance, and (2) interaction terms to assess variation in abundance along the dominant temperature gradient given different competitive contexts. Results indicate the presence of substantial intra-generic interactions, with simple interaction terms giving marginal increases in explained deviance equal to that explained by initial regressions using environment alone. Addition of interaction terms brought about smaller improvements in model fit, but confirm that variation in abundance along the dominant annual temperature gradient is strongly influenced by the competitive context provided by the remaining congeners. Such results are consistent with current understanding of the niche concept, and underline the difficulty inherent in using current species limits to predict likely changes in species distributions consequent on global warming.     

Interspecific Interactions between Phragmites australis and Spartina alterniflora along a Tidal Gradient in the Dongtan Wetland,Eastern China

The invasive species Spartina alterniora Loisel was introduced to the eastern coast of China in the 1970s and 1980s for the purposes of land reclamation and the prevention of soil erosion. The resulting interspecific competition had an important influence on the distribution of native vegetation, which makes studying the patterns and mechanisms of the interactions between Spartina alterniora Loisel and the native species Phragmites australis (Cav.) Trin ex Steud in this region very important. There have been some researches on the interspecific interactions between P. australis and S. alterniora in the Dongtan wetland of Chongming, east China, most of which has focused on the comparison of their physiological characteristics. In this paper, we conducted a neighbor removal experiment along a tidal gradient to evaluate the relative competitive abilities of the two species by calculating their relative neighbor effect (RNE) index. We also looked at the influence of environmental stress and disturbance on the competitive abilities of the two species by comparing interaction strength (I) among different tidal zones both for P. australis and S. alterniora. Finally, we measured physiological characteristics of the two species to assess the physiological mechanisms behind their different competitive abilities. Both negative and positive interactions were found between P. australis and S. alterniora along the environmental gradient. When the direction of the competitive intensity index for P. australis and S. alterniora was consistent, the competitive or facilitative effect of S. alterniora on P. australis was stronger than that of P. australis on S. alterniora. The interspecific interactions of P. australis and S. alterniora varied with environmental conditions, as well as with the method used, to measure interspecific interactions.     

Community and species responses to water level fluctuations with reference to soil layers in different habitats of mid-boreal mire complexes

The present paper discusses water level fluctuations in different parts of boreal mire complexes (eleven localities), mainly aapa mire complexes, on the basis of measurements performed by means of shallow observation wells and a few deeper observation tubes (piezometers) in the coastal half of the southern aapa mire zone in Finland. The sites represented intact vegetation from 12 different habitat types (communities), which were divided a priori into habitats with a stable surface moisture status (stable habitats) and into habitats with an unstable surface moisture status (unstable habitats). In stable habitats water level fluctuations took place according to the acrotelm–catotelm model, but the unstable habitats clearly deviated from the general pattern: water level fluctuations in them were not at all concentrated to the surficial, porous peat layer. Direct gradient analysis was used for arranging the communities along the water level fluctuation gradient. Variability of the water table, using 80% amplitude of water table residence, was used for the arrangement. The gradient was split into three groups: (1) habitats with a slightly fluctuating water table, (2) habitats with a considerably fluctuating water table and (3) habitats with an extremely fluctuating water table. The last group nearly corresponded to aro wetlands, and represented a very special habitat type. Indirect gradient analysis (NMDS ordination) also revealed the water level fluctuation gradient along with the gradient of traditional water level categories. According to the results of direct and indirect gradient analysis, the water level fluctuation seems to be an independent and important vegetation gradient. In peatlands, it occurs alongside with the traditional gradient of water level categories reflecting the mean water table. The responses of species to the range of water level fluctuations seem to reflect their tolerance to disturbances and evidently to seasonal drought. Most Sphagnum species are absent or in poor condition in habitats with extremely fluctuating water table. Vascular plant species that experienced most extreme water level fluctuations (Carex nigra, Juncus filiformis) have earlier been regarded as disturbance indicators. In addition, the difference between the piezometric water level and simultaneously measured water table depth reached the highest values within the habitats of those species (i.e., within Polytrichum commune aro wetlands) showing the downward direction of water movement in sandy mineral soil.     

Plant Community Structure and Conservation of a Northern Peru Sclerophyllous Forest

The vegetation near El Bosque Petrificado Piedra Chamana, in the northern Peruvian Andes, is evergreen sclerophyllous forest with significant shrub, epiphyte, and mat components. Important/characteristic genera include Dodonaea, Polylepis, Oreopanax, Oreocallis, Myrcianthes, and the mat-forming orchid Pleurothallis. A vegetation survey including 12 transects and 240 plots in high- and low-grazed areas documented 96 plant species. Compared with low-grazed areas, high-grazed areas had significantly fewer tree species, more herbs, and higher density of individuals. Both grazing categories exhibited high connectedness (as seen in network metrics) and positive biotic associations (nestedness), suggesting facilitation of some species by others, but high-grazed areas showed greater indications of positive associations (as seen in the C-score and V-ratio). These positive biotic associations may relate to the harsh environment and the role of keystone taxa such as Dodonaea viscosa, canopy trees, and mat-forming elements in moderating conditions and promoting species establishment. Only in the low-grazed areas was there any indication of competitive interactions (negative C-score/ less than expected species-pair occurrence). The shift in sign of the C-score, from negative in low-grazed areas to positive in high-grazed areas, indicates a loss of competitive interactions as a factor influencing community structure where grazing pressure is higher. Conservation of the area's natural resources would be advanced by protection of areas where the vegetation structure is more intact, better controls on grazing animals, and identification of development alternatives that would reduce pressure on the area's unique vegetation.     

Differential herbivory tolerance of dominant and subordinate plant species along gradients of nutrient availability and competition

We tested whether differences in the herbivory tolerance of plant species is related to their abundance in grassland communities and how herbivory and nutrient availability affect competitive balances among plant species through changes in their tolerance. The experimental approach involved a simulated grazing treatment (clipping) of two competitive grass species (Arrhenatherum elatius and Holcus lanatus) and two subordinate forb species (Prunella vulgaris and Lotus corniculatus) along a gradient of nutrient availability and under conditions of competition. Total standing, aboveground, root, and regrowth biomass were evaluated at the end of the experiment as an estimate of the capacity to compensate for twice removing aboveground biomass at different nutrient levels (NPK). Although clipping had a more pronounced negative effect on dominant plant species (Arrhenatherum and Holcus) than on subordinate species, the negative effects on dominant species were offset by the application of fertilizer. The combined effect of fertilizer and competition had more negative effects on the performance of Lotus and Prunella than on the dominant species. In terms of competition, the regrowth ability of Arrhenatherum and Holcus increased with the application of fertilizer, while the opposite pattern was observed for Lotus and Prunella. The addition of fertilizer has a positive effect on both grass species in terms of growth in clipped pots and competition, while subordinate species did not respond to the addition of fertilizer to the clipped pots and were negatively affected by competition with both grass species. The results suggest (1) that species replacement towards subordinate species as a function of herbivory is partially dependent on the herbivory tolerance of that species, (2) competitive relations between competitive grass species and subordinate forb species change under different environmental conditions, and (3) although grazing disturbance significantly influences competitive relations in favor of less competitive species, increasing nutrient levels counteract the negative effect of grazing on dominant competitive plant species.     

Abiotic conditions shape the relationship between indigenous and exotic species richness in a montane biodiversity hotspot

Montane ecosystems are more prone to invasions by exotic plant species than previously thought. Besides abiotic factors, such as climate and soil properties, plant-plant interactions within communities are likely to affect the performance of potential invaders in their exotic range. The biotic resistance hypothesis predicts that high indigenous species richness hampers plant invasions. The biotic acceptance hypothesis, on the other hand, predicts a positive relationship between indigenous and exotic species richness. We tested these two hypotheses using observational data along an elevational gradient in a southern African biodiversity hotspot. Species composition data of indigenous and exotic plants were recorded in 20 road verge plots along a gradient of 1775–2775 m a.s.l. in the Drakensberg, South Africa. Plots were 2?×?50 m in size and positioned at 50 m elevational intervals. We found a negative correlation between indigenous and exotic richness for locations with poorly developed mineral soils, suggesting biotic resistance through competitive interactions. A strong positive correlation for plots with very shallow soils at high elevations indicated a lack of biotic resistance and the possibility of facilitating interactions in harsher environments. These results suggest that biotic resistance is restricted to the lower and mid elevations while biotic acceptance prevails in presence of severe abiotic stress, potentially increasing the risk of plant invasions into montane biodiversity hotspots.

     

Invasive species and climate change: <Emphasis Type="Italic">Conyza canadensis</Emphasis> (L.) Cronquist as a tool for assessing the invasibility of natural plant communities along an aridity gradient

The predicted reduction in precipitation in the eastern Mediterranean due to climate change may expose the natural plant communities to invasive species. We assessed whether natural plant communities along an aridity gradient in Israel were resistant to invasion by considering differences in abiotic conditions and community characteristics in these regions. We considered Conyza canadensis as a model plant as it is a common invader in the region. We examined the mechanisms and functional traits of both the plant communities and C. canadensis that promote or discourage invasion. Study sites represented a rainfall gradient with four ecosystem types: mesic Mediterranean, Mediterranean, semiarid and arid. Our results showed that the mechanisms of community invasion resistance varied along the aridity gradient. At the arid and semiarid sites, water deficiency impaired the establishment of C. canadensis. At the mesic Mediterranean site, plant competition had a negative effect on C. canadensis performance, thus greatly reducing the likelihood of its establishment. We conclude that a decrease in regional precipitation due to climate change may not affect intrinsic resistance characteristics of natural plant communities to invasion in the area.     

Determinants of coastal treeline and the role of abiotic and biotic interactions

Recent die-off of coastal forests has been attributed primarily to the effects of sea level rise by correlation with tide-gage records. Due to the temporal and spatial scales involved, direct monitoring of sea level rise impacts is challenging and its attribution can be confounded by both land-use history and species interactions. Here we present experimental evidence for a micro-tidal, oligohaline estuarine system that the location of coastal treeline is determined by both environmental controls and positive and negative species interactions. We conducted field surveys and a transplant experiment to determine the controls on pine seedling establishment and survival along a salinity and flooding gradient. Using a two-way changepoint model we determined that sawgrass cover (Cladium jamaicense) and salinity concentrations interact to control natural pine regeneration (Pinus taeda). The long-term removal of sawgrass resulted in increased soil salinity and high rates of (planted) pine seedling mortality. In contrast, pine seedlings planted directly under sawgrass were able to survive at the same level as upland forest plots because of reduced salinity levels. This research provides evidence that sawgrass can facilitate pine seedling survival, but also suggests that either competitive exclusion by sawgrass or dispersal limitations prevent initial pine seedling regeneration. We propose that forest dynamics are closely coupled to fire, which consumes sawgrass biomass and reduces competition between pine seedlings and grass. Following fire, pine seedling establishment and the regrowth of sawgrass facilitates long-term pine seedling survival. Under this scenario, recent marsh invasion into coastal forests may not necessarily represent a permanent state change in locations where abiotic stress is not the only determinant for community composition.     

Changes in algal assemblages along observed and experimental phosphorus gradients in a subtropical wetland, U.S.A.

• 1 We wanted to determine if changes in algae in the Everglades were due to increased phosphorus (P) loading. Epiphytic algae, water chemistry, and surface sediment chemistry were characterized from 32 sloughs along a P gradient in the Everglades and changes in the algal assemblages along the P gradient were compared with those along an experimental P gradient of in situ mesocosms. The sloughs are the wettest open water habitats characterized by floating and submerged aquatic plants in the Everglades.
• 2 Algal species composition was much more sensitive to P concentration than algal biomass. The diatom species variance among sloughs, captured by 1st ordination axis, was more highly correlated with total P (TP) in surface sediments (r = ‐ 0.79), than soluble reactive P (SRP) (r = ‐ 0.08) and TP (r = ‐ 0.48) in the water column. Algal biomass (µg chl a cm^‐2) was not significantly correlated with P (SRP: r = 0.22, TP: r = 0.19, sediment TP: r = 0.07) along the P gradient in the Everglades. Cluster analysis classified diatom species assemblages in 32 sloughs into three groups (TWIN I, II, III), which corresponded to three zones along the P gradient. Dominant diatom species shifted from Mastogloia smithii (40.3%), Cymbella scotica (22.3%), and Fragilaria synegrotesca (21.8%) in TWIN I to Nitzschia amphibia (22.4%) and C. microcephala (12.4%) in TWIN III. TP in surface sediments and TP in epiphyton assemblages increased 4‐ and 5‐fold from TWIN I to TWIN III, respectively.
• 3 Patterns in epiphytic assemblages along the experimental P gradient in the mesocosms were very similar to those along the Everglades P gradient. Shannon diversity indices and species richness significantly increased along both P gradients. TN : TP ratio in epiphyton assemblages significantly decreased as sediment TP increased along both P gradient. Ordination analysis showed that diatom assemblages in the impacted zone (TWIN III) were ordinated closely to the assemblages from the highest P treatments in the mesocosms. The assemblages from the less impacted zone (TWIN I) were ordinated closely to the assemblages from controls in the mesocosms.
• 4 Concurrence between results of our survey and experiments suggest that changes in epiphytic assemblages along the P gradient in the Everglades are caused by increases in P concentrations.

     

Grazing intensity influences the strength of an associational refuge on temperate reefs

Recent studies have emphasized the role of positive interactions in ecological communities, but few have addressed how positive interactions are mediated by abiotic stress and biotic interactions. Here, I investigate the effect of a facilitator species on the abundance of macroalgae over a gradient of herbivory. Grazing by sea urchins can be intense on temperate reefs along the California coast, with benthic macroalgae growing exclusively in physical refuges and interspersed within colonies of the strawberry anemone, Corynactis californica. Field experiments indicated that the net effect of C. californica on turf algae was strongly nonlinear over a gradient in density of sea urchins. At low intensities of urchin grazing, the anemone and macroalgae competed for space, with algae capable of overgrowing C. californica. At intermediate grazing intensities, C. californica provided a refuge for turf algae but not for juvenile kelp. Neither turf algae nor kelp benefited from the presence of C. californica at the highest levels of grazing intensity, as sea urchins consumed nearly all macroalgae. The hump-shaped effect observed for C. californica contrasts with the prevailing view in ecological theory that positive interactions are more common in harsh environmental conditions. The results reported here qualify this view and underscore the need to evaluate positive interactions over a range of abiotic stress and consumer pressure.     

Competitive interactions in macroinfaunal animals of exposed sandy beaches

The influence of biotic interactions in structuring macroinfaunal communities of exposed sandy beaches, an unstable habitat characterized by strong physical forces, is generally considered negligible. We investigated the hypothesis that competitive interactions during burrowing could potentially affect the intertidal distribution and abundance of macroinfaunal animals of sandy beaches using two species of invertebrates, a hippid crab, Emerita analoga, and a bivalve, Mesodesma donacium, common along the coast of Chile. Spatial overlap in the intertidal distributions of these species was dynamic, varying with abundance, location, time of year and tide. Highest density zones of each species were often distinctly separated at low tide and spatial overlap in their distributions decreased significantly with increasing density, suggesting density dependence of the interactions. Negative relationships between densities of the two species at the smallest spatial scale examined also suggested active interactions among individuals. Over a tidal cycle, peak densities of the two species overlapped suggesting that interactions could occur frequently. Burrowing performance of E. analoga varied between size classes in three experimental densities of clams (5, 10 and 15 clams 0.008 m^-2) and in controls with no clams. Burrowing times of large crabs were significantly longer (~twofold) in all densities of clams than in controls, while those of small crabs did not differ significantly among treatments and controls. Large crabs also displaced clams from the sand while burrowing suggesting that two mechanisms of direct interference can occur, both of which could increase exposure of individuals involved to active swash and transport across or along the beach with potentially negative consequences. Our results suggest that competitive interactions capable of affecting zonation and population and community biology on a number of scales can occur among burrowing macroinfauna on exposed sandy beaches. Those interactions could be more ecologically significant than previously appreciated and may contribute to patterns observed in community structure and zonation on sandy beaches. Our results illustrate the potential importance of negative biological interactions in a physically stressful environment.     

Testing the Stress-Gradient Hypothesis at the Roof of the World: Effects of the Cushion Plant Thylacospermum caespitosum on Species Assemblages

Many cushion plants ameliorate the harsh environment they inhabit in alpine ecosystems and act as nurse plants, with significantly more species growing within their canopy than outside. These facilitative interactions seem to increase with the abiotic stress, thus supporting the stress-gradient hypothesis. We tested this prediction by exploring the association pattern of vascular plants with the dominant cushion plant Thylacospermum caespitosum (Caryophyllaceae) in the arid Trans-Himalaya, where vascular plants occur at one of the highest worldwide elevational limits. We compared plant composition between 1112 pair-plots placed both inside cushions and in surrounding open areas, in communities from cold steppes to subnival zones along two elevational gradients (East Karakoram: 4850–5250 m and Little Tibet: 5350–5850 m). We used PERMANOVA to assess differences in species composition, Friedman-based permutation tests to determine individual species habitat preferences, species-area curves to assess whether interactions are size-dependent and competitive intensity and importance indices to evaluate plant-plant interactions. No indications for net facilitation were found along the elevation gradients. The open areas were not only richer in species, but not a single species preferred to grow exclusively inside cushions, while 39–60% of 56 species detected had a significant preference for the habitat outside cushions. Across the entire elevation range of T. caespitosum, the number and abundance of species were greater outside cushions, suggesting that competitive rather than facilitative interactions prevail. This was supported by lower soil nutrient contents inside cushions, indicating a resource preemption, and little thermal amelioration at the extreme end of the elevational gradient. We attribute the negative associations to competition for limited resources, a strong environmental filter in arid high-mountain environment selecting the stress-tolerant species that do not rely on help from other plants during their life cycle and to the fact the cushions do not provide a better microhabitat to grow in.     

Local interactions, dispersal, and native and exotic plant diversity along a California stream

Although the species pool, dispersal, and local interactions all influence species diversity, their relative importance is debated. I examined their importance in controlling the number of native and exotic plant species occupying tussocks formed by the sedge Carex nudata along a California stream. Of particular interest were the factors underlying a downstream increase in plant diversity and biological invasions. I conducted seed addition experiments and manipulated local diversity and cover to evaluate the degree to which tussocks saturate with species, and to examine the roles of local competitive processes, abiotic factors, and seed supply in controlling the system‐wide patterns.
Seeds of three native and three exotic plants sown onto experimentally assembled tussock communities less successfully established on tussocks with a greater richness of resident plants. Nonetheless, even the most diverse tussocks were somewhat colonized, suggesting that tussocks are not completely saturated with species. Similarly, in an experiment where I sowed seeds onto natural tussocks along the river, colonization increased two‐ to three‐fold when I removed the resident species. Even on intact tussocks, however, seed addition increased diversity, indicating that the tussock assemblages are seed limited. Colonization success on cleared and uncleared tussocks increased downstream from km 0 to km 3 of the study site, but showed no trends from km 3 to km 8. This suggests that while abiotic and biotic features of the tussocks may control the increase in diversity and invasions from km 0 to km 3, similar increases from km 3 to km 8 are more likely explained by potential downstream increases in seed supply. The effective water dispersal of seed mimics and prevailingly downstream winds indicated that dispersal most likely occurs in a downstream direction. These results suggest that resident species diversity, competitive interactions, and seed supply similarly influence the colonization of native and exotic species.     

Negative and neutral marsh plant interactions dominate in early life stages and across physical gradients in an Oregon estuary

Conceptual models and some empirical studies in plant systems show that species interactions can shift from competitive under low stress and high productivity conditions to positive under high stress and low productivity conditions. In this study, we explore the relative strength and direction of marsh plant interactions at early life stages across a stress and productivity gradient in South Slough, Coos Bay, Oregon, USA. Germination and survival of five plant species (Atriplex patula L., Distichlis spicata (L.), Plantago maritima (Lam.) A. Gray, Sarcocornia pacifica (Standl.) A. J. Scott, Triglochin maritimum L.) were examined in the presence and absence of neighboring vegetation at three intertidal levels in each of three marsh sites along an estuarine gradient. We found that many of the interactions measured across these gradients were negative or neutral. The direction of the interactions depended on life stage, with neighboring plants having mostly negative and neutral effects on seed germination, and mostly neutral and rarely positive effects on seedling survival especially at lower intertidal locations. The exception was in the high intertidal at all sites, where competition was common. We found that the intensity of the interactions varied depending on marsh site, intertidal elevation, and plant species identity and that life stage and regional climate may be responsible for the general lack of positive interactions. We suggest that measuring species interactions across complex gradients of physical stress at different life stages can help refine our conceptual models and lead to better predictions of the factors controlling community structure.     

On the relationship between abiotic stress and co-occurrence patterns: an assessment at the community level using soil lichen communities and multiple stress gradients

The stress‐gradient hypothesis (SGH) predicts a shift from predominant competition to facilitation as abiotic stress increases. Most empirical tests of the SGH have evaluated the interactions between a single or a few pairs of species, have not considered the effects of multiple stress factors, and have not explored these interactions at nested spatial scales. We sampled 63 0.25‐m² plots, each subdivided into 100 5×5 cm and 25 10×10 cm sampling squares, in a semi‐arid Mediterranean environment to evaluate how co‐occurrence patterns among biological soil crusts (BSC)‐forming lichens changed along natural stress gradients driven by water and nutrient (N, P, K) availability. According to the SGH, we tested the hypothesis that the fine‐scale spatial arrangement of BSC‐forming lichens should shift from prevailing interspecific segregation to aggregation as abiotic stress increases. Co‐occurrence patterns ranged from significant species segregation to aggregation at the two spatial scales studied. When using the 5×5 cm grid, more plots showed significant species segregation than aggregation. At this sampling scale, co‐occurrence increased as water and nutrient availability decreased and increased, respectively. Small increases in soil pH promoted species co‐occurrence. Interspecific segregation was promoted as the cover of highly competitive species, such as Diploschistes diacapsis, increased. No significant relationships between co‐occurrence and the surrogates of abiotic stress were observed when data was arranged in a 10×10 cm grid. Our co‐occurrence analyses partially supported predictions from the SGH, albeit the results obtained were dependent on the type of abiotic stress and the spatial scale considered. They show the difficulties of predicting how co‐occurrence patterns change along complex stress gradients, and highlight the need of incorporating the effects of abiotic stress promoted by different resources, such as water and nutrients, into the conceptual framework of the SGH.     

Multi‐species interactions in competitive hierarchies: New methods and empirical test

Question: Are competitive hierarchies, which are typically based on the results of pair‐wise competition experiments, sensitive to the level of species interaction in the underlying competition experiments? Location: Controlled greenhouse study using vegetation typical of old‐fields in East Tennessee, USA. Methods: We extend traditional competitive effect/response methods to incorporate data from competition experiments featuring any level of species interaction (i. e., 2, 3, …, n species interacting simultaneously) and develop an ordinal technique that makes hierarchies more robust to variation in the numerical values of relative yield. We apply these methodological techniques to empirical data from a greenhouse experiment wherein four old‐field plant species were grown in pair‐wise and tri‐wise combination. We also demonstrate how resampling can be used to determine the variability of data and its consequences for development of competitive hierarchies. Results: Different hierarchies were produced when we used different evaluation methods, different levels of species interaction, and different levels of replication. More acute resampling distributions and wider ranges of target/neighbor scores revealed that higher levels of species interaction lead to more distinct hierarchies. Conclusions: Hierarchies developed from interactions among subsets of species may inadequately characterize relationships among the full community because of indirect or higher‐order interactions within multi‐species assemblages. Different evaluation methods can yield different hierarchies, and resampling is an effective tool to determine the sensitivity of resultant hierarchies to the level of replication. In sum, our new methodology can be used to control uncertainty in poorly‐replicated experiments.