Positive Species Interactions Research Articles

Positive species interactions structure rhodolith bed communities at a global scale

Positive species interactions structure rhodolith bed communities at a global scale

The Biodiversity and Climate Variability Experiment (BioCliVE): Quantifying the role of biodiversity in buffering ecosystems against climatic variability

Extreme climate events such as floods and droughts are becoming increasingly frequent and intense across the world. Future climate scenarios predict both an increase in individual extreme events, as well as chronic changes in climatic seasonality. Yet, the combined and relative effects of these pressures on ecosystems remain unknown. Concurrently, human-induced ecological disruption is accelerating species extinction rates, which are estimated to be 100 to 1000 times greater than pre-human levels. This is alarming as greater biological diversity is thought to buffer ecosystem functioning against extreme climate events, thereby safeguarding the provisioning of essential ecological services that contribute to human well-being. However, how and to what extent biodiversity buffers ecosystems against climate variability remains unclear. We recently constructed experimental grassland communities in a mesocosm-based field design representing a realistic gradient of plant diversity. Both extreme events (drought and flood) and a change in seasonality of precipitation are manipulated in a full factorial design to quantify the effects of future seasonal shifts and extremes in precipitation. We will: 1) determine to what extent higher biological diversity ensures that grasslands can continue to provide multiple ecosystem services even in the context of climate change and 2) unravel the fundamental mechanisms by which this is achieved including species asynchrony and positive species interactions. Results of our experimental approach will advance our understanding of the buffering potential of plant diversity and contribute to the development of strategies for sustainable service provisioning of our ecosystems in the face of climate change.

Oaks enhance early life stage longleaf pine growth and density in a subtropical xeric savanna.

The interplay of positive and negative species interactions controls species assembly in communities. Dryland plant communities, such as savannas, are important to global biodiversity and ecosystem functioning. Sandhill oaks in xeric savannas of the southeastern United States can facilitate longleaf pine by enhancing seedling survival, but the effects of oaks on recruitment and growth of longleaf pine have not been examined. We censused, mapped, and monitored nine contiguous hectares of longleaf pine in a xeric savanna to quantify oak-pine facilitation, and to examine other factors impacting recruitment, such as vegetation cover and longleaf pine tree density.We found that newly recruited seedlings and grass stage longleaf pines were more abundant in oak-dominated areas where densities were 230% (newly recruited seedlings) and 360% (grass stage) greater from lowest to highest oak neighborhood densities. Longleaf pine also grew faster under higher oak density. Longleaf pine recruitment was lowest under longleaf pine canopies. Mortality of grass stage and bolt stage longleaf pine was low (~1.0%yr-1) in the census interval without fire. Overall, our findings highlight the complex interactions between pines and oaks-two economically and ecologically important genera globally. Xeric oaks should be incorporated as a management option for conservation and restoration of longleaf pine ecosystems.

Combining ecology and technology to kick‐start oyster reef restoration

Techniques that enhance the recruitment of foundation species to restoration sites can inform the ecological development of the restored habitat. However, techniques are often considered in isolation, potentially overlooking synergies from combining them. Native oyster reefs have been lost worldwide, resulting in restoration efforts in systems that are often recruitment limited, or where recruiting oysters must spatially compete with opportunistic species. Here, we present a field‐based study that combines ecological knowledge on positive species interactions with novel acoustic technology, both of which are demonstrated to boost oyster recruitment in isolation, to test whether their interaction synergistically enhances the early larval recruitment that drives oyster reef development. At three sites across a 20 ha oyster reef restoration in southern Australia, we used self‐made speakers to broadcast healthy reef soundscapes that attract oysters and combine this with artificial kelp that facilitates oyster recruitment by suppressing competitive species (turfing algae). The combination of acoustic enrichment and artificial kelp increased oyster recruitment to the topside of substrate (326.98% increase), whereas only acoustic enrichment increased recruitment to the underside of substrate (126.95% increase). Our findings suggest that the combination of multiple techniques and their interactive effects might boost the early stages of reef development, providing proof‐of‐concept that these approaches can help oysters to build and bind reefs (i.e. recruit to the topside and underside, respectively). By combining ecology with technology during the first stages of a developing reef restoration, we show the potential value of these novel approaches to kick‐start the recovery of lost oyster reefs.

Different components of biodiversity mediate two ecosystem functions derived from naturally assembled filter‐feeder communities

Positive biodiversity–ecosystem functioning (BEF) relationships observed in experiments can be challenging to identify in natural communities. Freshwater animal communities are disproportionately harmed by global change that results in accelerated species loss. Understanding how animal‐mediated ecosystems functions may change as a result of global change can help determine whether biodiversity or species‐specific conservation will be effective at maintaining function. Unionid mussels represent half of imperiled species in freshwater ecosystems globally and perform important ecological functions such as water filtration and nutrient recycling. We explored BEF relationships for 22 naturally assembled mussel aggregations spanning three river basins. We used the Price equation to partition the contributions of species richness, composition, and context dependent interactions to two functions of interests: spatially‐explicit standing‐stock biomass (indirect proxy for function) and species‐specific nitrogen (N) excretion rates (direct measure of N recycling). Random and non‐random species loss each reduced biomass and N recycling. Many rare species with low contributions to biomass contributed to standing‐stock biomass in all basins. Widespread species had variable function across sites, such that context dependent effects (CDEs) outweighed richness effects on standing‐stock biomass in two basins, and were similar to richness effects in the third. Richness effects outweighed CDEs for N recycling. Thus, many species contributed a low proportion to overall N‐recycling; a product we attribute to the high evenness and functional effect trait diversity associated with these communities. The loss of low‐functioning species reduced the function of persisting species. This novel result using observational data adds evidence that positive species interactions, such as interspecific facilitation, may be a mechanism by which biodiversity enhances ecosystem functions. Our work stresses the importance of evaluating species‐specific contributions to functions in diverse systems, such as nutrient cycling when maintaining specific animal‐mediated functions is a management goal because indirect proxies may not completely characterize BEF relationships.

Interspecific sociality alters the colonization and extinction rates of birds on subtropical reservoir islands.

Island biogeography theory has proved a robust approach to predicting island biodiversity on the assumption of species equivalency. However, species differ in their grouping behaviour and are entangled by complex interactions in island communities, such as competition and mutualism. We here investigated whether intra- and/or interspecific sociality may influence biogeographic patterns, by affecting movement between islands or persistence on them. We classified bird species in a subtropical reservoir island system into subcommunities based on their propensity to join monospecific and mixed-species flocks. We found that subcommunities which had high propensity to flock interspecifically had higher colonization rates and lower extinction rates over a 10-year period. Intraspecific sociality increased colonization in the same analysis. A phylogenetically corrected analysis confirmed the importance of interspecific sociality, but not intraspecific sociality. Group-living could enable higher risk crossings, with greater vigilance also linked to higher foraging efficiency, enabling colonization or long-term persistence on islands. Further, if group members are other species, competition can be minimized. Future studies should investigate different kinds of island systems, considering positive species interactions driven by social behaviour as potential drivers of community assembly on islands. This article is part of the theme issue 'Mixed-species groups and aggregations: shaping ecological and behavioural patterns and processes'.

Testing the stress gradient hypothesis in soil bacterial communities associated with vegetation belts in the Andean Atacama Desert

BackgroundSoil microorganisms are in constant interaction with plants, and these interactions shape the composition of soil bacterial communities by modifying their environment. However, little is known about the relationship between microorganisms and native plants present in extreme environments that are not affected by human intervention. Using high-throughput sequencing in combination with random forest and co-occurrence network analyses, we compared soil bacterial communities inhabiting the rhizosphere surrounding soil (RSS) and the corresponding bulk soil (BS) of 21 native plant species organized into three vegetation belts along the altitudinal gradient (2400–4500 m a.s.l.) of the Talabre–Lejía transect (TLT) in the slopes of the Andes in the Atacama Desert. We assessed how each plant community influenced the taxa, potential functions, and ecological interactions of the soil bacterial communities in this extreme natural ecosystem. We tested the ability of the stress gradient hypothesis, which predicts that positive species interactions become increasingly important as stressful conditions increase, to explain the interactions among members of TLT soil microbial communities.ResultsOur comparison of RSS and BS compartments along the TLT provided evidence of plant-specific microbial community composition in the RSS and showed that bacterial communities modify their ecological interactions, in particular, their positive:negative connection ratios in the presence of plant roots at each vegetation belt. We also identified the taxa driving the transition of the BS to the RSS, which appear to be indicators of key host-microbial relationships in the rhizosphere of plants in response to different abiotic conditions. Finally, the potential functions of the bacterial communities also diverge between the BS and the RSS compartments, particularly in the extreme and harshest belts of the TLT.ConclusionsIn this study, we identified taxa of bacterial communities that establish species-specific relationships with native plants and showed that over a gradient of changing abiotic conditions, these relationships may also be plant community specific. These findings also reveal that the interactions among members of the soil microbial communities do not support the stress gradient hypothesis. However, through the RSS compartment, each plant community appears to moderate the abiotic stress gradient and increase the efficiency of the soil microbial community, suggesting that positive interactions may be context dependent.

Three-way species interactions reverse the positive pairwise effects of two natives on an exotic invader

The disruptive effects of tertiary species on otherwise positive pairwise species interactions (e.g. context-dependent parasitism in pollinator syndromes) is well-known. However, few—if any—studies have investigated how invasive plants affect interactions between facilitative plants and their native plant communities. Further, if tertiary invasive species can change interactions among native species from positive to negative, then a tertiary native should be capable of the same phenom for pairwise interactions between natives and invasives. Our previous research indicates invasive black mustard (Brassica nigra) changes interaction signs for otherwise positive species interactions between the dominant, native facilitator California buckwheat (Eriogonum fasciculatum) and its co-dominant beneficiary California sagebrush (Artemisia californica) in semi-arid California coastal sage scrub habitat. Here, E. fasciculatum and A. californica seedlings increased B. nigra shoot growth in pairwise species interactions in the greenhouse. However, in three-way species interactions, E. fasciculatum and A. californica together reduced B. nigra SLA, height, and reproductive potential while not increasing shoot DW. In three-way species interactions, B. nigra did not significantly reduce E. fasciculatum facilitation of A. californica. Also surprisingly, light competition with B. nigra resulted in an increase in A. californica height, which reduced the negative effects of A. californica light competition on shade-intolerant E. fasciculatum. In an additive field experiment, A. californica protected E. fasciculatum from facilitating germination and growth of B. nigra when water competition was minimized. Taken together, this study demonstrates the importance of studying species interactions between competitive, native perennials in the current ecological context of invaded ecosystems.

Refining the stress gradient hypothesis for mixed species groups of African mammals

Species interactions such as facilitation and predation influence food webs, yet it is unclear how they are mediated by environmental gradients. Here we test the stress gradient hypothesis which predicts that positive species interactions increase with stress. Drawing upon spatially-explicit data of large mammals in an African savanna, we tested how predation risk and primary productivity mediate the occurrence of mixed species groups. Controlling for habitat structure, predation risk by lions and primary productivity affected the frequency of mixed species groups in species-specific ways, likely reflecting distinct stress perceptions. To test whether mixed species groups indicate positive interactions, we conducted network analyses for specific scenarios. Under predation risk, dyadic associations with giraffes were more pronounced and metrics of animal networks changed markedly. However, dyadic association and network metrics were weakly mediated by primary productivity. The composition of mixed species groups was associated with similarities in prey susceptibility but not with similarities in feeding habits of herbivores. Especially predation risk favoured the frequency of mixed species groups and pronounced dyadic associations which dilute predation risk and increase predator detection. While our results provide support for the stress gradient hypothesis, they also highlight that the relative importance of stressors is context-dependent.

A large invasive consumer reduces coastal ecosystem resilience by disabling positive species interactions

Invasive consumers can cause extensive ecological damage to native communities but effects on ecosystem resilience are less understood. Here, we use drone surveys, manipulative experiments, and mathematical models to show how feral hogs reduce resilience in southeastern US salt marshes by dismantling an essential marsh cordgrass-ribbed mussel mutualism. Mussels usually double plant growth and enhance marsh resilience to extreme drought but, when hogs invade, switch from being essential for plant survival to a liability; hogs selectively forage in mussel-rich areas leading to a 50% reduction in plant biomass and slower post-drought recovery rate. Hogs increase habitat fragmentation across landscapes by maintaining large, disturbed areas through trampling of cordgrass during targeted mussel consumption. Experiments and climate-disturbance recovery models show trampling alone slows marsh recovery by 3x while focused mussel predation creates marshes that may never recover from large-scale disturbances without hog eradication. Our work highlights that an invasive consumer can reshape ecosystems not just via competition and predation, but by disrupting key, positive species interactions that underlie resilience to climatic disturbances.

Mapping species niche and fitness differences for communities with multiple interaction types

Modern coexistence theory (MCT) holds the potential to study the ability of species to avoid extinction (i.e. to persist) across community types but is rarely applied beyond pairs of competing species. Here, we show that this limitation can be overcome by mapping species according to their niche () and fitness differences (). This application provides three main benefits to study processes of multispecies persistence across trophic levels. First,mapping introduces a novel categorization of species and communities according to the high‐level processes at play: frequency dependence (negative or positive), the occurrence of positive species interactions (facilitation and mutualism) and whether persistence is possible without the presence of other species because of trophic interactions, such as herbivory or predation. Therefore, this mapping can be seen as a toolbox to describe how species persistence depends on species interactions. Second,mapping facilitates studying how species persistence responds to environmental changes that shift intrinsic growth rates and the strength and sign of species interactions. Third,mapping has the potential to foster synthesis across community types because it can accommodate co‐occurrence of positive, negative and neutral interactions between species. We, therefore, argue thatmapping can promote collaboration across sub‐fields, as it provides a common concept to link disparate ecological communities.

Positive species interactions strengthen in a high-CO2 ocean.

Negative interactions among species are a major force shaping natural communities and are predicted to strengthen as climate change intensifies. Similarly, positive interactions are anticipated to intensify and could buffer the consequences of climate-driven disturbances. We used in situ experiments at volcanic CO2 vents within a temperate rocky reef to show that ocean acidification can drive community reorganization through indirect and direct positive pathways. A keystone species, the algal-farming damselfish Parma alboscapularis, enhanced primary productivity through its weeding of algae whose productivity was also boosted by elevated CO2. The accelerated primary productivity was associated with increased densities of primary consumers (herbivorous invertebrates), which indirectly supported increased secondary consumers densities (predatory fish) (i.e. strengthening of bottom-up fuelling). However, this keystone species also reduced predatory fish densities through behavioural interference, releasing invertebrate prey from predation pressure and enabling a further boost in prey densities (i.e. weakening of top-down control). We uncover a novel mechanism where a keystone herbivore mediates bottom-up and top-down processes simultaneously to boost populations of a coexisting herbivore, resulting in altered food web interactions and predator populations under future ocean acidification.

Positive species interactions shape species' range limits

The relationship between niche and distribution, and especially the role of biotic interactions in shaping species' geographic distributions, has gained increasing interest in the last two decades. Most ecological research has focused on negative species interactions, especially competition, predation and parasitism. Yet the relevance of positive interactions – mutualisms and commensalisms – have been brought to the fore in recent years by an increasing number of empirical studies exploring their impact on range limits. Based on a review of 73 studies from a Web of Science search, we found strong evidence that positive interactions can influence the extent of species' geographic or ecological ranges through a diversity of mechanisms. More specifically, we found that while obligate interactions, and especially obligate mutualisms, tend to constrain the ranges of one or both partners, facultative positive interactions tend to widen ranges. Nonetheless, there was more variation in effects of facultative interactions on range limits, pointing to important context‐dependencies. Therefore, we propose that conceptual development in this field will come from studying ecological interactions in the context of networks of many species across environmental gradients, since pairwise interactions alone might overlook the indirect and environmentally‐contingent effects that species have on each other in communities of many interacting species. Finally, our study also revealed key data gaps that limit our current understanding of the pervasiveness of effects that positive interactions have on species' ranges, highlighting potential avenues for future theoretical and experimental work.

Aquatic worms (Tubificidae) facilitate productivity of macrophyte Azolla filiculoides in a wastewater biocascade system

Due to high waste sludge disposal at wastewater treatment plants and increasing phosphorus scarcity, there is a need to combine waste removal and nutrient recovery. One way to achieve this is by incorporating ecological processes into wastewater treatment. Positive species interactions, such as facilitation, are critical to increase productivity of biomass and nutrient recovery. In this study we showed the potential of using ecological principles including interspecific facilitation processes of aquatic plants (Azolla filiculoides) and worms (Oligochaeta, Tubificidae) in waste recovery and biomass production. This was investigated by developing a biocascade with monocultures of plants and aquatic worms that was fed on activated sludge. Tubificidae had an average relative growth rate of 0.02 g g−1 DW d−1 whereby sludge predation resulted in 45% sludge reduction. When Tubificidae were present in the biocascade, A. filiculoides biomass production significantly increased to a relative growth rate of 0.15 g g−1 DW d−1. The activity of Tubificidae mostly affected total suspended solids, chemical oxygen demand and ammonium concentration in the first compartment of the biocascade. Additionally, nitrification rates increased and the water acidified, leading to increased carbon dioxide concentrations and dissolved phosphorus-binding metals (zinc, iron, aluminium and manganese) that stimulated A. filiculoides growth. The high sludge reduction (45%) and phosphorus sequestration (133 mmol m−3 d−1) show a strong potential of the biocascade for combined sludge waste reduction and phosphorus recovery from wastewater.

Positive and Negative Species Interactions Shape Recruitment Patterns of a Range Expanding Native Alga

A species’ ability to spread is in part governed by the extent to which recipient habitats either resist, tolerate or promote the species’ recruitment. In disturbed marine environments, there is a general trend for the loss of algal canopies, thought to resist invasion, toward algal turf or rock barrens habitat. This study tested whether the spread of the range-expanding native macroalga Caulerpa filiformis was resisted by algal canopies but facilitated by algal turf or barrens habitats. Large-scale field surveys generally supported the predicted recruitment patterns, with C. filiformis recruits being most abundant on turf (but not barrens) and absent under kelp canopies. However, a discrepancy existed between different structural forms of turf, with the positive association holding true only for geniculate corallines, not filamentous turf. Secondly, a laboratory experiment tested whether the physical structure and/or the sediment-trapping properties of coralline turf influenced the recruitment success of C. filiformis. Whilst the structural complexity of turf aided overall recruitment performance (i.e., increased rhizoid production, attachment speed, success, and strength), a positive influence of turf-derived sediment on recruits’ growth was less obvious, at least over 10 days. The high morphological plasticity of C. filiformis propagules resulted in possible benefits of faster or stronger attachment of more developed propagules being only temporary, and that recruitment may be regulated in accordance with habitat preference. Finally, a field experiment confirmed the observed positive role of turf and the negative influence of algal canopies in the short-term, however, adverse environmental conditions in the longer-term resulted in the loss of most fragments. In conclusion, this study demonstrated the importance of both positive and negative species interactions for the recruitment success of a native alga, suggesting that a shift from kelp to turf algae can initiate further community change.

Physiological responses to light explain competition and facilitation in a tree diversity experiment

Abstract Ecologists often invoke interspecific facilitation to help explain positive biodiversity–ecosystem function relationships in plant communities, but seldom test how it occurs. One mechanism through which one species may facilitate another is by ameliorating abiotic stress. Physiological experiments show that a chronic excess of light can cause stress that depresses carbon assimilation. If shading by a plant's neighbours reduces light stress enough, it may facilitate that plant's growth. If light is instead most often a limiting factor for photosynthesis, shading may have an adverse, competitive effect. In a temperate tree diversity experiment, we measured stem growth rates and photosynthetic physiology in broadleaf trees across a gradient of light availability imposed by their neighbours. At the extremes, trees experienced nearly full sun (monoculture), or were shaded by nearby fast‐growing conifers (shaded biculture). Most species had slower growth rates with larger neighbours, implying a net competitive effect. On the other hand, the two most shade‐tolerant species (Tilia americana and Acer negundo) and the most shade‐intolerant one (Betula papyrifera) had faster stem growth rates with larger neighbours. The two shade‐tolerant species had the greatest increases in photoinhibition (reduced dark‐acclimated Fv/Fm) across the gradient of increasing light availability, which suggests they are more vulnerable to chronic light stress. While most species had lower carbon assimilation rates in the shaded biculture treatment, T. americana had rates up to 25% higher. T. americana also dropped its leaves 3–4 weeks earlier in monocultures, curtailing its growing season. We conclude that although large neighbours can cause light limitation in shade‐intolerant species, they can also increase growth through abiotic stress amelioration in shade‐tolerant species. Finally, in shade‐intolerant B. papyrifera, we find a pattern of stem elongation in trees with larger neighbours, which suggests that a shade avoidance response may account for the apparent positive trend in stem volume. Synthesis. Both positive and negative species interactions in our experiment can be explained in large part by the photosynthetic responses of trees to the light environment created by their neighbours. We show that photosynthetic physiology can help explain the species interactions that underlie biodiversity–ecosystem function relationships. The insights that ecologists gain by searching for such physiological mechanisms may help us forecast species interactions under environmental change.

Optimal Planting Distance in a Simple Model of Habitat Restoration With an Allee Effect

Ecological restoration is emerging as an important strategy to improve the recovery of degraded lands and to combat habitat and biodiversity loss worldwide. One central unresolved question revolves around the optimal spatial design for outplanted propagules that maximizes restoration success. Essentially, two contrasting paradigms exist: the first aims to plant propagules in dispersed arrangements to minimize competitive interactions. In contrast, ecological theory and recent field experiments emphasize the importance of positive species interactions, suggesting instead clumped planting configurations. However, planting too many propagules too closely is likely to waste restoration resources as larger clumps have less edges and have relatively lower spread rates. Thus, given the constraint of limited restoration efforts, there should be an optimal planting distance that both is able to harness positive species interactions but at the same time maximizes spread in the treated area. To explore these ideas, here we propose a simple mathematical model that tests the influence of positive species interactions on the optimal design of restoration efforts. We model the growth and spatial spread of a population starting from different initial conditions that represent either clumped or dispersed configurations of planted habitat patches in bare substrate. We measure the spatio-temporal development of the population, its relative and absolute growth rates as well as the time-discounted population size and its dependence on the presence of an Allee effect. Finally, we assess whether clumped or dispersed configurations perform better in our models and qualitatively compare the simulation outcomes with a recent wetland restoration experiment in a coastal wetland. Our study shows that intermediate clumping is likely to maximize plant spread under medium and high stress conditions (high occurrence of positive interactions) while dispersed designs maximize growth under low stress conditions where competitive interactions dominate. These results highlight the value of mathematical modeling for optimizing the efficiency of restoration efforts and call for integration of this theory into practice.

Do positive interactions between marine invaders increase likelihood of invasion into natural and artificial habitats?

Positive species interactions such as facilitation are important for enabling species to persist, especially in stressful conditions, and the nature and strength of facilitation varies along physical and biological gradients. Expansion of coastal infrastructure is creating hotspots of invasive species which can spillover into natural habitats, but the role of positive species interactions associated with biological invasions remains understudied. Theory suggests that stronger biotic pressure in natural habitats inhibits invasion success. In space‐limited marine systems, sessile organisms can overcome this limiting resource by settling as an epibiont on a substrate organism – basibiont. Using a series of spatially extensive surveys, we explored the role of invasive and native basibionts in providing habitat for other invasive and native epibionts, and tested whether environmental context (i.e. if the receiving habitat was natural or artificial), altered ecological outcomes. Overall, provision of space by basibionts was more important for invasive epibionts than for native epibionts but was dependent on the environmental context. Invasive basibionts facilitated invasive epibionts in natural habitats, and appeared to be more important for native epibionts in artificial habitats respectively. Native basibionts facilitated invasive, but not native epibionts in both natural and artificial habitats. These results advance our understanding of facilitation and highlight the idiosyncratic nature of biofouling and epibiosis, and the potentially important influence of environmental context. The degree to which native habitat‐forming species versus invasive habitat‐forming species either do or do not facilitate other native or non‐native species is a rich area for investigation. Experimental work is required to disentangle the processes underpinning these patterns.

Positive cascading effects of epiphytes enhance the persistence of a habitat‐forming macroalga and the biodiversity of the associated invertebrate community under increasing stress

AbstractEnhanced persistence of stress sensitive species in harsh environments due to amelioration of physical stress by habitat‐forming species has been widely documented. By contrast, less attention has been given to positive species interactions that may enhance stress tolerance in habitat‐forming species, with positive cascading effects on biodiversity and ecosystem functioning.We used a combination of field surveys and experiments to assess the effects of the epiphyteJania rubens, on its subtidal macroalgal hostHalopithys incurva, and their cascading effects on the associated invertebrate community, along a depth gradient associated with increasing stress (i.e. light stress at shallower depths).Correlative studies showed that the biomass and photosynthetic efficiency ofH. incurvaincreased with depth, along with a decrease of epiphyte biomass. The experimental manipulation of epiphytes (removal in shallow water and addition in deep water) revealed that their effects onH. incurvaphysiology varied from positive in shallow habitats to negative in deep habitats. These results suggest that reduction of light/UV stress byJ. rubensallowsH. incurvato persist in shallow‐water habitats that would be otherwise unsuitable. In addition, colonization by large clumps ofJ. rubensincreased the abundance and diversity of associated invertebrates in shallow habitat, suggesting that positive feedbacks betweenH. incurva, acting as the primary habitat‐former, andJ. rubens, acting as a secondary habitat‐former, generate a facilitation cascade.Synthesis. Our study suggests that the persistence of some habitat‐forming species and their ability to support biodiversity may rely upon physical stress reduction by other species (e.g. epiphytes in our case) that are generally neglected when planning conservation strategies. A deeper understanding of context dependency in positive species interactions is essential for predicting patterns of species distribution under increasingly stressful climate scenarios.

Rapid evolution destabilizes species interactions in a fluctuating environment.

Positive species interactions underlie the functioning of ecosystems. Given their importance, it is crucial to understand the stability of positive interactions over evolutionary timescales, in both constant and fluctuating environments; e.g., environments interrupted with periods of competition. We addressed this question using a two-species microbial system in which we modulated interactions according to the nutrient provided. We evolved in parallel four experimental replicates of species growing in isolation or together in consortia for 200 generations in both a constant and fluctuating environment with daily changes between commensalism and competition. We sequenced full genomes of single clones isolated at different time points during the experiment. We found that the two species coexisted over 200 generations in the constant commensal environment. In contrast, in the fluctuating environment, coexistence broke down when one of the species went extinct in two out of four cases. We showed that extinction was highly deterministic: when we replayed the evolution experiment from an intermediate time point we repeatably reproduced species extinction. We further show that these dynamics were driven by adaptive mutations in a small set of genes. In conclusion, in a fluctuating environment, rapid evolution destabilizes the long-term stability of positive pairwise interactions.