Non-indigenous invasive bivalves as ecosystem engineers

Several non-indigenous bivalve species have been colonising aquatic ecosystems worldwide, in some cases with great ecological and economic impacts. In this paper, we focus on the ecosystem engineering attributes of non-indigenous invasive bivalves (i.e., the capacities of these organisms to directly or indirectly affect the availability of resources to other species by physically modifying the environment). By reviewing the ecology of several invasive bivalves we identify a variety of mechanisms via which they modify, maintain and/or create habitats. Given the usually high densities and broad spatial distributions of such bivalves, their engineering activities can significantly alter ecosystem structure and functioning (e.g., changes in sediment chemistry, grain size, and organic matter content via bioturbation, increased light penetration into the water column due to filter feeding, changes in near bed flows and shear stress due to the presence of shells, provision of colonisable substrate and refuges by shells). In addition, changes in ecosystem structure and functioning due to engineering by invasive bivalves often have very large economic impacts. Given the worldwide spread of non-indigenous bivalves and the varied ways in which they physically modify habitats, their engineering effects should receive more serious consideration in restoration and management initiatives.     

Characterizing ecosystem-level consequences of biological invasions: the role of ecosystem engineers

Invasions by exotic species represent both threats to ecosystems as well as opportunities to learn more about them. Among the invaders that will have the largest impacts are those that directly modify ecosystems and thus have cascading effects for resident biota. Exotics can affect ecosystems by altering system-level flows, availability, or quality of nutrients, food, and physical resources (e.g. living space, water, heat or light). The invader-mediated control of physical resources, typically achieved through the modification of habitats, has received limited attention in invasion biology. This reflects a general trend in ecology, and only recently has the concept of ecosystem engineering been developed to account for the role of species that shape habitats. Plants and animals in terrestrial and aquatic systems can both create and destroy structure. When introduced into ecosystems, these exotic engineers cause physical state changes with effects that ramify throughout the system. Although the consequences of these modifications are varied and complex, insight gained from general ecological principles offers an opportunity to predict what invaders will do upon their integration into systems. Examples from the literature suggest that introduced ecosystem engineers that increase habitat complexity or heterogeneity tend to cause abundances and/or species richness to rise, while those that decrease complexity tend to have the reverse effect. In assessing such patterns, however, it is critical to also consider spatial scales and the life habits of resident organisms. In addition to providing predictive power, recognition of engineering as a major means by which invasive species affect ecosystems provides a unifying theme for invasion biology and offers a chance to consider more fully the general role of species in ecosystems.     

A mathematical model of plants as ecosystem engineers

Understanding the structure and dynamics of plant communities in water-limited systems often calls for the identification of ecosystem engineers--key species that modify the landscape, redistribute resources and facilitate the growth of other species. Shrubs are excellent examples; they self-organize to form patterns of mesic patches which provide habitats for herbaceous species. In this paper we present a mathematical model for studying ecosystem engineering by woody plant species in drylands. The model captures various feedbacks between biomass and water including water uptake by plants' roots and increased water infiltration at vegetation patches. Both the uptake and the infiltration feedbacks act as mechanisms for vegetation pattern formation, but have opposite effects on the water resource; the former depletes the soil-water content under a vegetation patch, whereas the latter acts to increase it. Varying the relative strength of the two feedbacks we find a trade-off between the engineering capacity of a plant species and its resilience to disturbances. We further identify two basic soil-water distributions associated with engineering at the single patch level, hump-shaped and ring-shaped, and discuss the niches they form for herbaceous species. Finally, we study how pattern transitions at the landscape level feedback to the single patch level by affecting engineering strength.     

Branching archaeocyaths as ecosystem engineers during the Cambrian radiation

The rapid origination and diversification of major animal body plans during the early Cambrian coincide with the rise of Earth's first animal-built framework reefs. Given the importance of scleractinian coral reefs as ecological facilitators in modern oceans, we investigate the impact of archaeocyathan (Class Archaeocyatha) reefs as engineered ecosystems during the Cambrian radiation. In this study, we present the first high-resolution, three-dimensional (3D) reconstructions of branching archaeocyathide (Order Archaeocyathida) individuals from three localities on the Laurentian paleocontinent. Because branched forms in sponges and corals display phenotypic plasticity that preserve the characteristics of the surrounding growth environment, we compare morphological measurements from our fossil specimens to those of modern corals to infer the surface conditions of Earth's first reefs. These data demonstrate that archaeocyaths could withstand and influence the flow of water, accommodate photosymbionts, and build topographically complex and stable structures much like corals today. We also recognize a stepwise increase in the roughness of reef environments in the lower Cambrian, which would have laid a foundation for more abundant and diverse coevolving fauna.     

Range shifts and species diversity in marine ecosystem engineers: patterns and predictions for European sedimentary habitats

Aim Ecosystem engineering polychaetes in the genus Diopatra are undergoing range shifts in western Europe. Here we: (1) assess the species diversity underlying these shifts; (2) link biogeographic patterns to sea surface temperature patterns; and (3) predict possible ecosystem‐level outcomes of Diopatra's northward expansion. Location Western Europe. Methods We use molecular phylogenetic and morphological evidence to assess species diversity and biogeographic ranges. Using regression tree analyses, we assess thermal limits for two Diopatra species. We compare biogeographic patterns with historical sea surface temperature patterns to draw links between range shifts and climate change. Finally, we review published data to predict potential ecological changes as Diopatra invades new habitats. Results The native Diopatra neapolitana range has contracted 130 km to the south. A cryptogenic species, Diopatra sp. A, has extended the northern limit of the genus 350 km to the southern Brittany Peninsula. Both shifts can be explained by historical sea surface temperature anomalies. The Diopatra sp. A expansion is predicted to continue into the English Channel and the North Sea, introducing large tube structures to sheltered sedimentary habitats that currently lack such structures. Main conclusions As climate change intensifies, the sediment‐stabilizing Diopatra sp. A will invade habitats dominated by the bioturbating lugworm Arenicola marina. The resulting interaction between functionally different ecosystem engineers will probably cause ecological changes in northern European coastal waters. Existing data for Diopatra species and arenicolids suggest that the diversity and biomass of macroalgae, vascular plants, infauna and epibenthic fauna may increase, while microbial activity may decrease. Net changes in productivity will depend on the relative rates of these changes.     

Secondhand homes: The multilayered influence of woodpeckers as ecosystem engineers

Ecosystem engineers alter, and can be influenced in turn by, the ecosystems they live in. Woodpeckers choose foraging and nesting sites based, in part, on food availability. Once abandoned, these cavities, particularly within areas of high forage, may be crucial to secondary cavity‐nesting birds otherwise limited by cavities formed through decay. Our study examined factors that influence the nesting success of primary cavity nesters and the subsequent impact on secondary cavity‐nesting birds. Using 5 years of point count data, we monitored the outcomes of cavity‐nesting birds in South Texas. We used logistic‐exposure models to predict daily survival rates based on cavity metrics and used woodpecker foraging trends and insect surveys to determine if nesting where woodpeckers actively forage benefits secondary cavity‐nesting birds. Both woodpeckers and secondary cavity nesters shared predictors of daily survival; nests were more successful in cavities with small openings in minimally decayed trees. All secondary cavity nesters had higher probabilities of success when nesting in an abandoned woodpecker cavity, opposed to ones formed by decay. Woodpeckers tended to forage in areas with higher‐than‐average levels of the insect orders Coleoptera, Hymenoptera, and Orthoptera, and secondary cavity nesters had higher rates of success when nesting in these areas. Our results suggest abandoned woodpecker cavities may be constructed in a way that directly benefit secondary cavity nesters. Additionally, we suggest an interplay between these ecosystem engineers, food availability, and secondary cavity nesters: Woodpeckers engineer superior nesting cavities in areas where food is more abundant, and the resultant cavities in areas of high forage may benefit local secondary cavity nesters. Our findings indicate that there is still much to be explored in the role of ecosystem engineers, and how they influence local communities on multiple trophic levels.     

Arbuscular mycorrhizal fungi as (agro)ecosystem engineers

Symbiotic interactions have been shown to facilitate shifts in the structure and function of host plant communities. For example, parasitic plants can induce changes in plant diversity through the suppression of competitive community dominants. Arbuscular mycorrhizal (AM) fungi have also be shown to induce shifts in host communities by increasing host plant nutrient uptake and growth while suppressing non-mycorrhizal species. AM fungi can therefore function as ecosystem engineers facilitating shifts in host plant communities though the presumed physiological suppression of non-contributing or non-mycorrhizal plant species. This dichotomy in plant response to AM fungi has been suggested as a tool to suppress weed species (many of which are non-mycorrhizal) in agro-ecosystems where mycorrhizal crop species are cultivated. Rinaudo et al. (2010), this issue, have demonstrated that AM fungi can suppress pernicious non-mycorrhizal weed species including Chenopodium album (fat hen) while benefiting the crop plant Helianthus annuus (sunflower). These findings now suggest a future for harnessing AM fungi as agro-ecosystem engineers representing potential alternatives to costly and environmentally damaging herbicides.     

European rabbits as ecosystem engineers: warrens increase lizard density and diversity

Mammals that build extensive open burrow systems are often classified as ecosystem engineers, since they have the potential to modulate the availability of resources for themselves and other organisms. Lizards may benefit from the heterogeneity created by these structures, especially if coupled with an increased offer of sites for refuge and thermoregulation. However, information about these engineering effects by burrowing animals is scarce. We investigated the influence of European rabbit burrows on several parameters of a Mediterranean lizard community (abundance, density, diversity and body condition) in three different habitats (open pastures, holm oak and scrub patches). We found that lizards were positively associated with burrows, and that burrows determined lizard presence at otherwise unfavourable habitats. Moreover, community parameters such as density and species richness were higher in sites with burrows. Burrows influenced lizard species in different ways, and were also relevant for other Mediterranean vertebrates, as revealed by questionnaires to experts. We also explored the possible resources provided by burrows for lizards. Warrens offer relatively abundant prey and appropriate retreat sites for refuge and thermoregulation. Warrens may have further implications within the ecosystem, acting as stepping stones, allowing lizards to reach otherwise inaccessible habitat patches. This study shows that European rabbit warrens have a positive influence on lizard density and diversity, and confirms the role of rabbits as ecosystem engineers. This reinforces the need for appropriate conservation measures for rabbits, especially given their threatened status in the Iberian Peninsula. Furthermore, our study highlights that taking into account the influence of engineering activities increases our awareness of species interactions, and may translate into more adequate conservation measures for the preservation of biodiversity.     

Fungal biodiversity in aquatic habitats

Fungal biodiversity in freshwater, brackish and marine habitats was estimated based on reports in the literature. The taxonomic groups treated were those with species commonly found on submerged substrates in aquatic habitats: Ascomycetes (exclusive of yeasts), Basidiomycetes, Chytridiomycetes, and the non-fungal Saprolegniales in the Class Oomycetes. Based on presence/absence data for a large number and variety of aquatic habitats, about 3,000 fungal species and 138 saprolegnialean species have been reported from aquatic habitats. The greatest number of taxa comprise the Ascomycetes, including mitosporic taxa, and Chytridiomycetes. Taxa of Basidiomycetes are, for the most part, excluded from aquatic habitats. The greatest biodiversity for all groups occurs in temperate areas, followed by Asian tropical areas. This pattern may be an artifact of the location of most of the sampling effort. The least sampled geographic areas include Africa, Australia, China, South America and boreal and tropical regions worldwide. Some species overlap occurs among terrestrial and freshwater taxa but little species overlap occurs among freshwater and marine taxa. We predict that many species remain to be discovered in aquatic habitats given the few taxonomic specialists studying these fungi, the few substrate types studied intensively, and the vast geographical area not yet sampled.     

Burrowing owls and burrowing mammals: are ecosystem engineers interchangeable as facilitators?

Terrestrial vertebrates exhibit dynamic, positive interactions that form and dissolve under different circumstances, usually with multiple species as participants. Ecosystem engineers are important facilitators of other species because they cause physical changes in the environment that alter resource availability. Although a species can be associated with more than one partner, facilitators may not be interchangeable if they differ in abundance, behavioral characteristics, or interactions with other factors in ways that condition the outcome of the association. We examined interactions between burrowing owls (Athene cunicularia) and two burrowing mammals, hairy armadillos (Chaetophractus villosus) and plains vizcachas (Lagostomus maximus), and determined whether these ecosystem engineers are interchangeable for owls. We examined reproductive success for owls nesting in these mammal burrows, constructed a logistic regression model to identify habitat characteristics associated with owl nests, and examined the engineering activities of the mammals. Data on reproduction and habitat indicate that armadillos and vizcachas are not interchangeable for owls. Thirty-five percent of the nests in vizcacha burrows produced fledglings; no fledglings were produced from nests outside vizcachas colonies, even though owls nest successfully in armadillo burrows in other parts of Argentina. Vizcachas facilitate burrowing owls by construction of burrows and by producing open understory vegetation through herbivory. In contrast, armadillos do not alter vegetation, and their burrows are suitable for nest sites only when they occur in recently burned areas or areas maintained by anthropogenic disturbance. Our habitat model also suggests that fire plays a key role in maintaining owl populations because fire is the only natural process that reduces shrubs to the level required by owls. Current management practices of eradication of vizcachas and fire suppression in shrublands could have strong negative consequences for burrowing owls.     

Exergy as an indicator: Observations of an aquatic ecosystem model

Exergy is considered as a goal function or ecological orientor. Normally at the edge of oscillation exergy reaches to its maximum value when the ecosystem had no adaptation on it. To study the variation of exergy in different states of ecosystem, a simple three species (phytoplankton, zooplankton and fish) food chain model has been considered. From the model it is shown that the system moves from steady state to chaotic state by decreasing zooplankton body volume in turn increasing its grazing rate. By the property of self-adaptability the system tries to overcome this situation. Two such possible processes are described here: (i) by the toxic effect of phytoplankton and (ii) by reducing half saturation constant of fishes. In both this cases exergy value reduces and the system reaches to stable state. Through the analysis of exergy variation in all these situations this paper shows that the system chose the process in which the reduction of exergy will be the minimum.     

Animal ecosystem engineers modulate the diversity-invasibility relationship

Background

Invasions of natural communities by non-indigenous species are currently rated as one of the most important global-scale threats to biodiversity. Biodiversity itself is known to reduce invasions and increase stability. Disturbances by ecosystem engineers affect the distribution, establishment, and abundance of species but this has been ignored in studies on diversity-invasibility relationships.

Methodology/Principal Findings

We determined natural plant invasion into 46 plots varying in the number of plant species (1, 4, and 16) and plant functional groups (1, 2, 3, and 4) for three years beginning two years after the establishment of the Jena Experiment. We sampled subplots where earthworms were artificially added and others where earthworm abundance was reduced. We also performed a seed-dummy experiment to investigate the role of earthworms as secondary seed dispersers along a plant diversity gradient. Horizontal dispersal and burial of seed dummies were significantly reduced in subplots where earthworms were reduced in abundance. Seed dispersal by earthworms decreased with increasing plant species richness and presence of grasses but increased in presence of small herbs. These results suggest that dense vegetation inhibits the surface activity of earthworms. Further, there was a positive relationship between the number of earthworms and the number and diversity of invasive plants. Hence, earthworms decreased the stability of grassland communities against plant invasion.

Conclusions/Significance

Invasibility decreased and stability increased with increasing plant diversity and, most remarkably, earthworms modulated the diversity-invasibility relationship. While the impacts of earthworms were unimportant in low diverse (low earthworm densities) and high diverse (high floral structural complexity) plant communities, earthworms decreased the stability of intermediate diverse plant communities against plant invasion. Overall, the results document that fundamental processes in plant communities like plant seed burial and invader establishment are modulated by soil fauna calling for closer cooperation between soil animal and plant ecologists.     

Mussels as ecosystem engineers: Their contribution to species richness in a rocky littoral community

Mussels are important ecosystem engineers in marine benthic systems because they aggregate into beds, thus modifying the nature and complexity of the substrate. In this study, we evaluated the contribution of mussels (Brachidontes rodriguezii, Mytilus edulis platensis, and Perna perna) to the benthic species richness of intertidal and shallow subtidal communities at Cerro Verde (Uruguay). We compared the richness of macro-benthic species between mussel-engineered patches and patches without mussels but dominated by algae or barnacles at a landscape scale (all samples), between tidal levels, and between sites distributed along a wave exposition gradient. Overall, we found a net increase in species richness in samples with mussels (35 species), in contrast to samples where mussels were naturally absent or scarce (27 species). The positive trend of the effect did not depend upon tidal level or wave exposition, but its magnitude varied between sites. Within sites, a significant positive effect was detected only at the protected site. Within the mussel-engineered patches, the richness of all macro-faunal groups (total, sessile and mobile) was positively correlated with mussel abundance. This evidence indicates that the mussel beds studied here were important in maintaining species richness at the landscape-level, and highlights that beds of shelled bivalves should not be neglected as conservation targets in marine benthic environments.     

Consumer movement dynamics as hidden drivers of stream habitat structure: suckers as ecosystem engineers on the night shift

Ecosystem engineering can control the spatial and temporal distribution of resources and movement by engineering organisms within an ecosystem can mobilize resources across boundaries and distribute engineering effects. Movement patterns of fishes can cause physical changes to aquatic habitats though nesting or feeding, both of which often vary in space and time. Here we present evidence of ecosystem engineering by the Sonora sucker Catostomus insignis, a dominant fish in streams of the southwestern United States, and show how cryptic nocturnal movement patterns and bioturbation activities control heterogeneity in benthic substrates, and in sediment and carbon export. Sonora suckers exhibit distinct diel movement patterns, spending daylight hours in refuge habitats (typically deep pools) while moving into shallow habitats at night to feed. Feeding by suckers creates substantial disturbance in soft sediments that are patchy in space and time. These disturbances moved up to 2.4 × 10⁴ cm³ of sediment per square meter per week in locations that are up to hundreds of meters away from sucker daytime refuges. The diel cycles in feeding activity (i.e. nocturnal digging in benthic substrates) caused nighttime pulses in suspended sediment that comprised up to 32% of the daily suspended load and organic matter transport of a stream reach. During the daytime, this particulate transport settles in habitats beyond the location of the initial disturbance, thus redistributing both sediment and organic matter. Our data indicate that cryptic movement by ecosystem engineers can distribute their effects in space and time generating heterogeneity in resources and suggest that habitat modifications restricting consumer movement may alter the impact of engineering activities.     

Overexploiting marine ecosystem engineers: potential consequences for biodiversity

Overfishing is a major environmental problem in the oceans. In addition to the direct loss of the exploited species, the very act of fishing, particularly with mobile bottom gear, destroys habitat and ultimately results in the loss of biodiversity. Furthermore, overfishing can create trophic cascades in marine communities that cause similar declines in species richness. These effects are compounded by indirect effects on habitat that occur through removal of ecological or ecosystem engineers. Mass removal of species that restructure the architecture of habitat and thus increase its complexity or influence the biogeochemistry of sediments could have devastating effects on local biodiversity and important water–sediment processes. The possible overexploitation of engineering species requires more attention because the consequences extend beyond their own decline to affect the rest of the ecosystem. This is particularly problematic in the deep ocean, where oil and gas exploration and fishing pressure are likely to increase.