Invasive macroalgae in native seagrass beds: vectors of spread and impacts.

BACKGROUND AND AIMS: Worldwide, invasive species are spreading through marine systems at an unprecedented rate with both positive and negative consequences for ecosystems and the biological functioning of organisms. Human activities from shipping to habitat damage and modification are known vectors of spread, although biological interactions including epibiosis are increasingly recognized as potentially important to introduction into susceptible habitats. METHODS: We assessed a novel mechanism of spread - limpets as transporters of an invasive alga, Sargassum muticum, into beds of the seagrass Zostera marina - and the physiological impact of its invasion. The association of S. muticum with three limpet species and other habitats was assessed using intertidal surveys on rocky shores and snorkelling at two seagrass sites in the UK. A 4-year field study tested the effect of S. muticum on Z. marina shoot density, dry weight and phenolic compounds (caffeic and tannic acid) content, and a laboratory experiment tested the impact of S. muticum on nutrient partitioning (C/H/N/P/Si), photosynthetic efficiency (Fv/Fm) and growth of Z. marina. RESULTS: On rocky shores 15 % of S. muticum occurrences were attached to the shells of live limpets. In seagrass beds 5 % of S. muticum occurrences were attached to the shells of dead limpets. The remainder were attached to rock, to cobblestones, to the seagrass matrix or embedded within the sand. Z. marina density and phenolics content was lower when S. muticum co-occurred with it. Over 3 years, photosynthetic responses of Z. marina to S. muticum were idiosyncratic, and S. muticum had no effect on nutrient partitioning in Z. marina. CONCLUSIONS: Our results show limpets support S. muticum as an epibiont and may act as a previously unreported transport mechanism introducing invaders into sensitive habitats. S. muticum reduced production of phenolics in Z. marina, which may weaken its defensive capabilities and facilitate proliferation of S. muticum. The effect of S. muticum on Z. marina photosynthesis requires further work but having no effect on the capacity of Z. marina to sequester nutrients suggests a degree of resilience to this invader.

Positive role of plateau pika (Ochotona coronae) on environmental quality at low and moderate density on the Tibetan plateau: Evidence from a meta-analysis.

The roles of plateau pika (Ochotona coronae) in the Tibetan Plateau are often controversial, because it is often regarded as a destructive pest or an ecosystem engineer. Here a meta-analysis using 72 paired observations was conducted to examine whether the impacts of plateau pika on environmental quality (i.e., plant and soil properties) depend on population density in the Tibetan Plateau. Pika population density was used as a proxy for disturbance intensity. The pika disturbance intensity was divided into five groups based on the number of burrows, including low disturbance intensity (LD) (9-30 burrows per ha), medium disturbance intensity (MD) (31-100 burrows per ha), high disturbance intensity (HD) (101-170 burrows per ha), extreme disturbance intensity (ED) (171-240 burrows per ha) and uncontrolled (or excessive) disturbance intensity (UD) (>241 burrows per ha). Given that sample sizes in some of the groups are small (especially for the HD), we further pooled the disturbance groups including the LD-MD and HD-UD. Overall, relative to control (i.e., no disturbing), there was a great increase (80.3%) in aboveground biomass under the LD-MD, whereas a decrease of 41.1% occurred under the HD-UD. At the same time, plant coverage, species richness, height, and belowground biomass greatly decreased only in the HD-UD. Furthermore, the effect size of plant coverage, species richness, and aboveground biomass also declined with pika burrow density significantly. With regard to soil properties, there was a significant increase in soil organic carbon, ammonium nitrogen, and soil organic carbon stock under the LD-MD, whereas a decrease under the HD-UD. In addition, soil total nitrogen, total potassium, and nitrate nitrogen increased at the LD-MD and HD-UD. Nevertheless, the effect size of these soil properties (with >20 observations) was not related to pika burrow density. In summary, there is an implication that the low and moderate disturbance of pikas is beneficial to maintain and promote ecosystem functioning in the Tibetan grasslands. In the future pikas' eradication policy should be reconsidered in alpine grassland management.

An apex predator engineers wetland food-web heterogeneity through nutrient enrichment and habitat modification.

The potential for animals to modify spatial patterns of nutrient limitation for autotrophs and habitat availability for other members of their communities is increasingly recognized. However, net trophic effects of consumers acting as ecosystem engineers remain poorly known. The American Alligator Alligator mississippiensis is an abundant predator capable of dramatic modifications of physical habitat through the creation and maintenance of pond-like basins, but its role in influencing community structure and nutrient dynamics is less appreciated. We investigated if alligators engineer differences in nutrient availability and changes to community structure by their creation of 'alligator ponds' compared to the surrounding phosphorus (P)-limited oligotrophic marsh. We used a halo sampling design of three distinct habitats extending outward from 10 active alligator ponds across a hydrological gradient in the Everglades, USA. We performed nutrient analysis on basal food-web resources and quantitative community analyses, and stoichiometric analyses on plants and animals. Our findings demonstrate that alligators act as ecosystem engineers and enhance food-web heterogeneity by increasing nutrient availability, manipulating physical structure and altering algal, plant and animal communities. Flocculent detritus, an unconsolidated layer of particulate organic matter and soil, showed strong patterns of P enrichment in ponds. Higher P availability in alligator ponds also resulted in bottom-up trophic transfer of nutrients as evidenced by higher growth rates (lower N:P) for plants and aquatic consumers. Edge habitats surrounding alligator ponds contained the most diverse communities of invertebrates and plants, but low total abundance of fishes, likely driven by high densities of emergent macrophytes. Pond communities exhibited higher abundance of fish compared to edge habitat and were dominated by compositions of small invertebrates that track high nutrient availability in the water column. Marshes contained high numbers of animals that are closely tied to periphyton mats, which were absent from other habitats. Alligator-engineered habitats are ecologically important by providing nutrient-enriched 'hotspots' in an oligotrophic system, habitat heterogeneity to marshes, and refuges for other fauna during seasonal disturbances. This work adds to growing evidence that efforts to model community dynamics should routinely consider animal-mediated bottom-up processes like ecosystem engineering.

Sea-level rise and the emergence of a keystone grazer alter the geomorphic evolution and ecology of southeast US salt marshes.

Keystone species have large ecological effects relative to their abundance and have been identified in many ecosystems. However, global change is pervasively altering environmental conditions, potentially elevating new species to keystone roles. Here, we reveal that a historically innocuous grazer-the marsh crab Sesarma reticulatum-is rapidly reshaping the geomorphic evolution and ecological organization of southeastern US salt marshes now burdened by rising sea levels. Our analyses indicate that sea-level rise in recent decades has widely outpaced marsh vertical accretion, increasing tidal submergence of marsh surfaces, particularly where creeks exhibit morphologies that are unable to efficiently drain adjacent marsh platforms. In these increasingly submerged areas, cordgrass decreases belowground root:rhizome ratios, causing substrate hardness to decrease to within the optimal range for Sesarma burrowing. Together, these bio-physical changes provoke Sesarma to aggregate in high-density grazing and burrowing fronts at the heads of tidal creeks (hereafter, creekheads). Aerial-image analyses reveal that resulting "Sesarma-grazed" creekheads increased in prevalence from 10 ± 2% to 29 ± 5% over the past <25 y and, by tripling creek-incision rates relative to nongrazed creekheads, have increased marsh-landscape drainage density by 8 to 35% across the region. Field experiments further demonstrate that Sesarma-grazed creekheads, through their removal of vegetation that otherwise obstructs predator access, enhance the vulnerability of macrobenthic invertebrates to predation and strongly reduce secondary production across adjacent marsh platforms. Thus, sea-level rise is creating conditions within which Sesarma functions as a keystone species that is driving dynamic, landscape-scale changes in salt-marsh geomorphic evolution, spatial organization, and species interactions.

Litter, Plant Competition, and Ecosystem Dynamics: A Theoretical Perspective.

AbstractCommunity structure depends jointly on species' responses to, and effects on, environmental factors. Many such factors, including detritus, are studied in ecosystem ecology. Detritus in terrestrial ecosystems is dominated by plant litter (nonliving organic material), which, in addition to its role in material cycling, can act as a niche factor modulating interactions among plants. Litter thus links traditional community and ecosystem processes, which are often studied separately. We explore this connection using population dynamics models of two plant species and a litter pool. We first find conditions determining the outcome of interactions between these species, highlighting the role that litter plays and the role of broader ecosystem parameters, such as decomposition rate. Species trade-offs in tolerance to direct competition and litter-based interference competition allow for coexistence, provided the litter-tolerant species produces more litter at the population level; otherwise, priority effects may result. When species coexist, litter-mediated interactions between plants disrupt the traditional relationship between biomass accumulation and decomposition. Increasing decomposition rate may have no effect on standing litter density and, in some cases, may even increase litter load. These results illustrate how ecosystem variables can influence community outcomes that then feed back to influence the ecosystem.

Unrecognized threat to global soil carbon by a widespread invasive species.

Most of Earth's terrestrial carbon is stored in the soil and can be released as carbon dioxide (CO2 ) when disturbed. Although humans are known to exacerbate soil CO2 emissions through land-use change, we know little about the global carbon footprint of invasive species. We predict the soil area disturbed and resulting CO2 emissions from wild pigs (Sus scrofa), a pervasive human-spread vertebrate that uproots soil. We do this using models of wild pig population density, soil damage, and their effect on soil carbon emissions. Our models suggest that wild pigs are uprooting a median area of 36,214 km2 (mean of 123,517 km2 ) in their non-native range, with a 95% prediction interval (PI) of 14,208 km2 -634,238 km2 . This soil disturbance results in median emissions of 4.9 million metric tonnes (MMT) CO2 per year (equivalent to 1.1 million passenger vehicles or 0.4% of annual emissions from land use, land-use change, and forestry; mean of 16.7 MMT) but that it is highly uncertain (95% PI, 0.3-94 MMT CO2 ) due to variability in wild pig density and soil dynamics. This uncertainty points to an urgent need for more research on the contribution of wild pigs to soil damage, not only for the reduction of anthropogenically related carbon emissions, but also for co-benefits to biodiversity and food security that are crucial for sustainable development.

Density-dependence and seasonal variation in reproductive output and sporophyte production in the kelp, Ecklonia radiata.

The kelp, Ecklonia radiata, is an abundant subtidal ecosystem engineer in southern Australia. Density-dependent changes in the abiotic environment engineered by Ecklonia may feedback to affect reproduction and subsequent recruitment. Here, we examined: 1) how the reproductive capacity of Ecklonia individuals in the field (zoospores released · mm-2 reproductive tissue) varied with adult density and time, and 2) how the recruitment of microscopic gametophytes and sporophytes was influenced by zoospore density at two times. Zoospore production did not vary with adult density, with only one month out of ten sampled over a 2-y period showing a significant effect of density. However, zoospore production varied hugely over time, being generally highest in mid-autumn and lowest in mid-late summer. There were strong effects of initial zoospore density on gametophyte and sporophyte recruitment with both a minimum and an optimum zoospore density for sporophyte recruitment, but these varied in time. Almost no sporophytes developed when initial zoospore density was <6.5 · mm-2 in spring or <0.5 · mm-2 in winter with optimum densities of 90-355 · mm-2 in spring and 21-261 · mm-2 in winter, which resulted in relatively high recruitment of 4-7 sporophytes · mm-2 . Sporophyte recruitment declined at initial zoospore densities >335 · mm-2 in spring and >261 · mm-2 in winter and was zero at very high zoospore densities. These findings suggest that although adult Ecklonia density does not affect per-capita zoospore production, because there is a minimum zoospore density for sporophyte production, a decline in population-level output could feedback to impact recruitment.

Reference genome for the California ribbed mussel, Mytilus californianus, an ecosystem engineer.

The California ribbed mussel, Mytilus californianus, is an ecosystem engineer crucial for the survival of many marine species inhabiting the intertidal zone of California. Here, we describe the first reference genome for M. californianus and compare it to previously published genomes from three other Mytilus species: M. edulis, M. coruscus, and M. galloprovincialis. The M. californianus reference genome is 1.65 Gb in length, with N50 sequence length of 118 Mb, and an estimated 86.0% complete single copy genes. Compared with the other three Mytilus species, the M. californianus genome assembly is the longest, has the highest N50 value, and the highest percentage complete single copy genes. This high-quality genome assembly provides a foundation for population genetic analyses that will give insight into future conservation work along the coast of California.

Whole-organism responses to constant temperatures do not predict responses to variable temperatures in the ecosystem engineer Mytilus trossulus.

Understanding and predicting responses of ectothermic animals to temperature are essential for decision-making and management. The thermal performance curve (TPC), which quantifies the thermal sensitivity of traits such as metabolism, growth and feeding rates in laboratory conditions, is often used to predict responses of wild populations. However, central assumptions of this approach are that TPCs are relatively static between populations and that curves measured under stable temperature conditions can predict performance under variable conditions. We test these assumptions using two latitudinally matched populations of the ecosystem engineer Mytilus trossulus that differ in their experienced temperature variability regime. We acclimated each population in a range of constant or fluctuating temperatures for six weeks and measured a series of both short term (feeding rate, byssal thread production) and long-term (growth, survival) metrics to test the hypothesis that performance in fluctuating temperatures can be predicted from constant temperatures. We find that this was not true for any metric, and that there were important interactions with the population of origin. Our results emphasize that responses to fluctuating conditions are still poorly understood and suggest caution must be taken in the use of TPCs generated under constant temperature conditions for the prediction of wild population responses.

Natural selection on a carbon cycling trait drives ecosystem engineering by Sphagnum (peat moss).

Sphagnum peat mosses have an extraordinary impact on the global carbon cycle as they control long-term carbon sequestration in boreal peatland ecosystems. Sphagnum species engineer peatlands, which harbour roughly a quarter of all terrestrial carbon, through peat accumulation by constructing their own niche that allows them to outcompete other plants. Interspecific variation in peat production, largely resulting from differences in tissue decomposability, is hypothesized to drive niche differentiation along microhabitat gradients thereby alleviating competitive pressure. However, little empirical evidence exists for the role of selection in the creation and maintenance of such gradients. In order to document how niche construction and differentiation evolved in Sphagnum, we quantified decomposability for 54 species under natural conditions and used phylogenetic comparative methods to model the evolution of this carbon cycling trait. We show that decomposability tracks the phylogenetic diversification of peat mosses, that natural selection favours different levels of decomposability corresponding to optimum niche and that divergence in this trait occurred early in the evolution of the genus prior to the divergence of most extant species. Our results demonstrate the evolution of ecosystem engineering via natural selection on an extended phenotype, of a fundamental ecosystem process, and one of the Earth's largest soil carbon pools.

Foraging by an avian ecosystem engineer extensively modifies the litter and soil layer in forest ecosystems.

Ecosystem engineers physically modify their environment, thereby altering habitats for other organisms. Increasingly, "engineers" are recognized as an important focus for conservation and ecological restoration because their actions affect a range of ecosystem processes and thereby influence how ecosystems function. The Superb Lyrebird Menura novaehollandiae is proposed as an ecosystem engineer in forests of southeastern Australia due to the volume of soil and litter it turns over when foraging. We measured the seasonal and spatial patterns of foraging by Lyrebirds and the amount of soil displaced in forests in the Central Highlands, Victoria. We tested the effects of foraging on litter, soil nutrients and soil physical properties by using an experimental approach with three treatments: Lyrebird exclusion, Lyrebird exclusion with simulated foraging, and non-exclusion reference plots. Treatments were replicated in three forest types in each of three forest blocks. Lyrebirds foraged extensively in all forest types in all seasons. On average, Lyrebirds displaced 155.7 Mg/ha of litter and soil in a 12-month period. Greater displacement occurred where vegetation complexity (<50 cm height) was low. After two years of Lyrebird exclusion, soil compaction (top 7.5 cm) increased by 37% in exclusion plots compared with baseline measures, while in unfenced plots it decreased by 22%. Litter depth was almost three times greater in fenced than unfenced plots. Soil moisture, pH, and soil nutrients showed no difference between treatments. The enormous extent of litter and soil turned over by the Superb Lyrebird is unparalleled by any other vertebrate soil engineer in terrestrial ecosystems globally. The profound influence of such foraging activity on forest ecosystems is magnified by its year-round pattern and widespread distribution. The disturbance regime that Lyrebirds impose has implications for diverse ecosystem processes including decomposition and nutrient cycling, the composition of litter- and soil-dwelling invertebrate communities, the shaping of ground-layer vegetation patterns, and fire behavior and post-fire ecosystem recovery. Maintaining Lyrebird populations as a key facilitator of ecosystem function is now timely and critical as unprecedented wildfires in eastern Australia in summer 2019-2020 have severely burned ~12 million ha of forest, including ~30% of the geographic range of the Superb Lyrebird.

Linking Bacterial Communities Associated with the Environment and the Ecosystem Engineer Orchestia gammarellus at Contrasting Salt Marsh Elevations.

The digestive tract of animals harbors microbiota important for the host's fitness and performance. The interaction between digestive tract bacteria and soil animal hosts is still poorly explored despite the importance of soil fauna for ecosystem processes. In this study, we investigated the interactions between the bacterial communities from the digestive tract of the litter-feeding, semi-terrestrial crustacean Orchestia gammarellus and those obtained from the environment; these organisms thrive in, i.e., soil and plant litter from salt marshes. We hypothesized that elevation is an important driver of soil and litter bacterial communities, which indirectly (via ingested soil and litter bacteria) influences the bacterial communities in the digestive tract of O. gammarellus. Indeed, our results revealed that elevation modulated soil and litter bacterial community composition along with soil organic matter content and the C:N ratio. Soil and plant litter differed in alpha diversity indexes (richness and diversity), and in the case of plant litter, both indexes increased with elevation. In contrast, elevation did not affect the composition of bacterial communities associated with O. gammarellus' digestive tract, suggesting selection by the host, despite the fact that a large component of the bacterial community was also detected in external sources. Importantly, Ca. Bacilloplasma and Vibrio were highly prevalent and abundant in the host. The taxonomic comparison of Ca. Bacilloplasma amplicon sequence variants across the host at different elevations suggested a phylogenetic divergence due to host habitat (i.e., marine or semi-terrestrial), thus supporting their potential functional role in the animal physiology. Our study sheds light on the influence of the environment on soil animal-bacteria interactions and provides insights into the resilience of the O. gammarellus-associated bacteria to increased flooding frequency.

Biogeomorphological eco-evolutionary feedback between life and geomorphology: a theoretical framework using fossorial mammals.

Engineer organisms not only adapt to pre-existing environmental conditions but also co-construct their physical environment. By doing so, they can subsequently change selection pressures for themselves and other species, as well as change community and ecosystem structures and functions. Focusing on one representative example, i.e., fossorial mammals, we show that geomorphological Earth system components are crucial for understanding and quantifying links between evolutionary and ecosystem dynamics and that feedbacks between geomorphology and engineer organisms constitute a major driver of geomorphological organization on the Earth's surface. We propose a biogeomorphological eco-evolutionary feedback synthesis from the gene to the landscape where eco-evolutionary feedbacks are mediated by the geomorphological dimensions of a niche that are affected by engineer organisms, such as fossorial mammals. Our concept encompasses (i) the initial responses of fossorial mammals to environmental constraints that enhance the evolution of their morphological and biomechanical traits for digging in the soil; (ii) specific adaptations of engineer fossorial mammals (morphological, biomechanical, physiological and behavioural feedback traits for living in burrows) to their constructed geomorphological environment; and (iii) ecological and evolutionary feedbacks diffusing at the community and ecological levels. Such a new perspective in geomorphology may lead to a better conceptualization and analysis of Earth surface processes and landforms as parts of complex adaptive systems in which Darwinian selection processes at lower landscape levels lead to self-organization of higher-level landforms and landscapes.

Intertidal Canopy-forming Seaweeds Modulate Understory Seaweed Photoprotective Compounds.

Foundation species provide physical structure that enhances the diversity and abundance of associated organisms. Canopy-forming seaweeds are known to act as foundation species on rocky shores by lowering temperature and desiccation stress. Direct solar radiation, including ultraviolet (UV) light, can also reduce photosynthetic rates in algae, cause oxidative stress and DNA damage. The reduction in UV exposure provided by an algal canopy could therefore be important for understory organisms, including the red alga Chondrus crispus on New England's (USA) rocky shores, and this relationship may be more important at higher tidal elevations with increased low-tide exposure time. In field experiments, we investigated the relationship between tidal elevation and an index of C. crispus UV exposure, the concentration of UV-absorbing pigments. Low on the shore, C. crispus grew without a canopy. Higher on the shore, in the mid-intertidal zone, C. crispus was found under the canopy-forming rockweed, Fucus distichus subsp. evanescens. At this elevation, C. crispus was shaded (>50%; >1 m above MLLW). We performed a canopy removal experiment that spanned the mid-zone where C. crispus and F. distichus subsp. evanescens co-occur and the low-zone (no canopy). Following canopy removal in the mid-zone, UV pigment concentrations increased with tidal elevation. After accounting for the effect of elevation, removal of the algal canopy resulted in UV-protective pigment concentrations 2-fold higher than in un-manipulated control plots. These results suggest that amelioration of solar UV exposure might be another mechanism by which canopy seaweeds, acting as foundation species, facilitate understory seaweeds on rocky shores.

Demography of the Intertidal Fucoid Hormosira banksii: Importance of Recruitment to Local Abundance.

Canopy-forming macroalgae form the basis of diverse coastal ecosystems globally. The fucoid Hormosira banksii is often the dominant canopy-forming macroalga in the temperate intertidal of southern Australia and New Zealand, where it is commonly associated with an understory of coralline turf. Hormosira banksii is susceptible to both natural and anthropogenic disturbance and despite its abundance, few studies have examined the demography of this important species. This study determined the demographic response of H. banksii to different gradients of disturbance to both its canopy and to the understory coralline turf. We established plots in which the density of H. banksii and/or understory coralline turf was manipulated in a pulse perturbation to simulate a disturbance event. The manipulated plots contained eight treatments ranging from 100% removal of H. banksii to 100% removal of the understory coralline turf. We then measured recruitment and followed individual recruits for up to 18 months to determine growth and survivorship. We found that H. banksii recruitment was seasonally variable throughout the experiment and highest over summer, survivorship of recruits was generally high, and the species was slow-growing and long-lived. Moreover, the level of disturbance did not seem to affect recruitment, growth, or survivorship and post-recruitment mortality was independent of H. banksii density. In this system, it appears that H. banksii is a relatively long-lived perennial species whose demography is density-independent which appears to allow recovery from disturbance.

Linking Physiology To Ecological Function: Environmental Conditions Affect Performance And Size Of The Intertidal Kelp Hedophyllum Sessile (Laminariales, Phaeophyceae).

For autogenic ecosystem engineers, body size is an aspect of individual performance that has direct connections to community structure; yet the complex morphology of these species can make it difficult to draw clear connections between the environment and performance. We combined laboratory experiments and field surveys to test the hypothesis that individual body size was determined by disparate localized physiological responses to environmental conditions across the complex thallus of the intertidal kelp Hedophyllum sessile, a canopy-forming physical ecosystem engineer. We documented substantial (> 40%) declines in whole-thallus photosynthetic potential (as Maximum Quantum Yield, MQY) as a consequence of emersion, which were related to greater than 10-fold increases in intra-thallus MQY variability (as Coefficient of Variation). In laboratory experiments, desiccation and high light levels during emersion led to lasting impairment of photosynthetic potential and an immediate > 25% reduction in area due to tissue contraction, which was followed by complete loss of structural integrity after three days of submersion. Tissue exposed to desiccation and high light during emersion had higher nitrogen concentrations and lower phlorotannin concentrations than tissue in control treatments (on average 1.36 and 0.1x controls, respectively), suggesting that conditions during emersion have the potential to affect food quality for consumers. Our data indicate that the complex thallus morphology of H. sessile may be critical to this kelp's ability to persist in the intertidal zone despite the physiological challenges of emersion and encourage a more nuanced view of the concept of "sub-lethal stress" on the scale of the whole individual.

One ring to rule them all: an ecosystem engineer fungus fosters plant and microbial diversity in a Mediterranean grassland.

Species coexistence in grasslands is regulated by several environmental factors and interactions with the soil microbial community. Here, the development of the Basidiomycetes fungus Agaricus arvensis, forming fairy rings, in a species-rich Mediterranean grassland, is described. Effects of the mycelial front on plants, fungi and bacteria were assessed by vegetation survey and next generation sequencing approaches. Our results showed a fungal-dependent shift in the community structure operated by a wave-like spread of fairy rings that decreased plant, fungal and bacterial diversity, indicating a detrimental effect of fairy rings on most species. The fairy rings induced successional processes in plants that enhanced the replacement of a community dominated by perennial plants with short-living and fast-growing plant species. In parallel, fungal and bacterial communities showed evident differences in species composition with several taxa associated within distinct sampling zone across the fairy rings. Notably, bacteria belonging to the Burkholderia genus and fungi of the genus Trichoderma increased in response to the advancing mycelium of A. arvensis. The profound changes in community composition and the overall increase in taxa diversity at ecosystemic scale suggest that fairy ring-forming fungi may act as ecosystem engineer species in Mediterranean grasslands.

Cancer Cells and M2 Macrophages: Cooperative Invasive Ecosystem Engineers.

Many aspects of cancer can be explained utilizing well-defined ecological principles. Applying these principles to cancer, cancer cells are an invasive species to a healthy organ ecosystem. In their capacity as ecosystem engineers, cancer cells release cytokines that recruit monocytes to the tumor and polarize them to M2-like protumor macrophages. Macrophages, recruited by the cancer cells, act as a secondary invasive species. The ecosystem engineering functions of M2-macrophages in turn support and stimulate cancer cell survival and proliferation. The cooperative ecosystem engineering of both the primary invasive species of the cancer cell and the secondary invasive species of the M2-macrophage thus creates a vicious cycle of tumor promotion. Targeting a specific aspect of this tumor-promoting ecosystem engineering, such as blocking efferocytosis by M2-like macrophages, may improve the response to standard-of-care anticancer therapies. This strategy has the potential to redirect cooperative protumor ecosystem engineering toward an antitumor ecosystem engineering strategy.

Bioturbation by a reintroduced digging mammal reduces fuel loads in an urban reserve.

Digging animals may alter many characteristics of their environment as they disrupt and modify the ground's surface by creating foraging pits or burrows. Extensive disturbance to the soil and litter layer changes litter distribution and availability, potentially altering fuel loads. In many landscapes, including peri-urban areas, fire management to reduce fuel loads is complex and challenging. The reintroduction of previously common digging animals, many of which are now threatened, may have the added benefit of reducing fuel loads. We experimentally examined how the reintroduction of a marsupial bandicoot, quenda (Isoodon fusciventer), altered surface fuel loads in an urban bush reserve in Perth, Western Australia. Foraging activities of quenda (where they dig for subterranean food) were substantial throughout the reserve, creating a visibly patchy distribution in surface litter. Further, in open plots where quenda had access, compared to fenced plots where quenda were excluded, quenda foraging significantly reduced litter cover and litter depth. Similarly, estimated surface fuel loads were nearly halved in open plots where quenda foraged compared to fenced plots where quenda were absent (3.6 cf. 6.4 Mg/ha). Fire behavior modeling, using the estimated surface fuel loads, indicated the predicted rate of spread of fire were significantly lower for open plots where quenda foraged compared to fenced plots under both low (29.2 cf. 51.4 m/h; total fuels) and high (74.3 cf. 130.4 m/h; total fuels) fire conditions. Although many environments require fire, including the bushland where this study occurred, fire management can be a considerable challenge in many landscapes, including urban bushland reserves, which are usually small and close to human infrastructure. The reintroduction of previously common digging species may have potential value as a complimentary tool for reducing fuel loads, and potentially, fire risk.

Parasitism in ecosystem engineer species: A key factor controlling marine ecosystem functioning.

Although parasites represent a substantial part of marine communities' biomass and diversity, their influence on ecosystem functioning, especially via the modification of host behaviour, remains largely unknown. Here, we explored the effects of the bopyrid ectoparasite Gyge branchialis on the engineering activities of the thalassinid crustacean Upogebia pusilla and the cascading effects on intertidal ecosystem processes (e.g. sediment bioturbation) and functions (e.g. nutrient regeneration). Laboratory experiments revealed that the overall activity level of parasitized mud shrimp is reduced by a factor 3.3 due to a decrease in time allocated to burrowing and ventilating activities (by factors 1.9 and 2.9, respectively). Decrease in activity level led to strong reductions of bioturbation rates and biogeochemical fluxes at the sediment-water interface. Given the world-wide distribution of mud shrimp and their key role in biogeochemical processes, parasite-mediated alteration of their engineering behaviour has undoubtedly broad ecological impacts on marine coastal systems functioning. Our results illustrate further the need to consider host-parasite interactions (including trait-mediated indirect effects) when assessing the contribution of species to ecosystem properties, functions and services.