Feedbacks Between Climate and Freshwater Ecosystem Engineers

Microplastic impacts on soil and sediment bioturbation: insights from microcosm experiments across diverse ecosystems.

Invasive species are a key stressor in freshwater ecosystems. When these species are also ecosystem engineers, their impacts are exacerbated because they modulate resource availability for a wide range of other species. The aim of this review is to synthesise existing knowledge of the impacts of invasive ecosystem engineers in freshwaters and identify knowledge gaps requiring further research. The four questions explored in this review are: (1) What are the trends in research into invasive ecosystem engineers? (2) What are common negative effects of invasive ecosystem engineers in freshwater? (3) Do all impacts of invasive ecosystem engineers have negative consequences for biodiversity? (4) What happens when multiple ecosystem engineers interact? Four literature searches in Web of Science have been used to identify articles for the review and to estimate relative research effort between terrestrial, marine and freshwater ecosystems. The number of research articles focusing on ecosystem engineers across all ecosystem types is increasing. Despite well‐known examples of ecosystem engineer species in freshwaters (e.g. beaver), more research has focussed on terrestrial environments and invasive species. The effects of invasive ecosystem engineers in freshwater systems are varied and often context dependent. Their effects on biodiversity or native ecosystem engineers are often shown to be negative; however, not all effects associated with these species are deleterious to native species. For instance, some invasive ecosystem engineers support native species through the provision of food or refuges. Although freshwater ecosystems are often influenced by multiple species of ecosystem engineers (including native, invasive or both), little is known about interactions between these species or the combined effects of multiple ecosystem engineers. More research is also needed that relates the results of laboratory experiments to the field and develops methods for measuring factors that govern the impact of engineers on ecosystems. Understanding the spatial variability of the impacts of invasive ecosystem engineers as well as their interaction with anthropogenic stressors (e.g. hydrologic modification) is also necessary. The lag in research surrounding invasive ecosystem engineers in freshwater compared to other biomes is concerning, as freshwater ecosystems support biodiversity disproportionate to the area they occupy. Creating predictive models of the impacts of freshwater ecosystem engineers would help anticipate the effects of invasive ecosystem engineers in freshwater and add to the broader understanding of their effects in other biomes.

In both terrestrial and aquatic ecosystems, invasive species are a major driver of global change that is increasing in scope and impact due to the increase in global trade, habitat modification and climate change. Among invasive species, those that are known as ‘ecosystem engineers’ are considered a specific, highly influential, type of invaders, where an invader significantly alters the new environment, either physically or chemically. In this context, this perspective review discusses the different types of possible impacts of invasive ecosystems engineers (IEE) in terrestrial, freshwater, and marine ecosystems. Scanning the relevant literature on the topic, we find a 12‐year lag in the use of terms relevant to the concept of ecosystem engineers in invasion‐related publications since the first publication of the concept in 1994, with a steep increase in use between 2006 and 2014. A bibliometric mapping showed a high level of connectedness between related terms and clusters, suggesting an ample flow of concepts, ideas and knowledge between realms, ecosystems and regions, and researchers that study them. Throughout this essay, we illustrate with recent examples the context‐dependency of their (positive and negative) impacts in the three realms. We review the distinction between autogenic (altering the environment for other species with their body) and allogenic (altering the environment for other species with their actions) ecosystem engineering in the context of alien species. We also put a spotlight on the well‐studied engineering effects of IEE plants and macroalgae, terrestrial and aquatic bioturbators and burrowers as well as highly effective consumers. We finalize with discussing how IEE can strongly affect ecosystem services for human wellbeing and explore the possible contribution of IEE in restoring functions and services in the face of climate change in highly invaded and fast‐warming systems like the southeastern Mediterranean Sea or areas that face frequent fires for example. We claim that the last topic has received little attention from the scientific community and should be given priority in future studies. Read the free Plain Language Summary for this article on the Journal blog.

Large-bodied predators are well represented among the world's threatened and endangered species. A significant body of literature shows that in terrestrial and marine ecosystems large predators can play important roles in ecosystem structure and functioning. By contrast, the ecological roles and importance of large predators within freshwater ecosystems are poorly understood, constraining the design and implementation of optimal conservation strategies for freshwater ecosystems. Conservationists and environmentalists frequently promulgate ecological roles that crocodylians are assumed to fulfil, but often with limited evidence supporting those claims. Here, we review the available information on the ecological importance of crocodylians, a widely distributed group of predominantly freshwater-dwelling, large-bodied predators. We synthesise information regarding the role of crocodylians under five criteria within the context of modern ecological concepts: as indicators of ecological health, as ecosystem engineers, apex predators, keystone species, and as contributors to nutrient and energy translocation across ecosystems. Some crocodylians play a role as indicators of ecosystem health, but this is largely untested across the order Crocodylia. By contrast, the role of crocodylian activities in ecosystem engineering is largely anecdotal, and information supporting their assumed role as apex predators is currently limited to only a few species. Whether crocodylians contribute significantly to nutrient and energy translocation through cross-ecosystem movements is unknown. We conclude that most claims regarding the importance of crocodylians as apex predators, keystone species, ecosystem engineers, and as contributors to nutrient and energy translocation across ecosystems are mostly unsubstantiated speculation, drawn from anecdotal observations made during research carried out primarily for other purposes. There is a paucity of biological research targeted directly at: understanding population dynamics; trophic interactions within their ecological communities; and quantifying the short- and long-term ecological impacts of crocodylian population declines, extirpations, and recoveries. Conservation practices ideally need evidence-based planning, decision making and justification. Addressing the knowledge gaps identified here will be important for achieving effective conservation of crocodylians.

For more than five decades, research has been conducted at Ny-Ålesund, in Svalbard, Norway, to understand the structure and functioning of High-Arctic ecosystems and the profound impacts on them of environmental change. Terrestrial, freshwater, glacial and marine ecosystems are accessible year-round from Ny-Ålesund, providing unique opportunities for interdisciplinary observational and experimental studies along physical, chemical, hydrological and climatic gradients. Here, we synthesize terrestrial and freshwater research at Ny-Ålesund and review current knowledge of biodiversity patterns, species population dynamics and interactions, ecosystem processes, biogeochemical cycles and anthropogenic impacts. There is now strong evidence of past and ongoing biotic changes caused by climate change, including negative effects on populations of many taxa and impacts of rain-on-snow events across multiple trophic levels. While species-level characteristics and responses are well understood for macro-organisms, major knowledge gaps exist for microbes, invertebrates and ecosystem-level processes. In order to fill current knowledge gaps, we recommend (1) maintaining monitoring efforts, while establishing a long-term ecosystem-based monitoring programme; (2) gaining a mechanistic understanding of environmental change impacts on processes and linkages in food webs; (3) identifying trophic interactions and cascades across ecosystems; and (4) integrating long-term data on microbial, invertebrate and freshwater communities, along with measurements of carbon and nutrient fluxes among soils, atmosphere, freshwaters and the marine environment. The synthesis here shows that the Ny-Ålesund study system has the characteristics needed to fill these gaps in knowledge, thereby enhancing our understanding of High-Arctic ecosystems and their responses to environmental variability and change.

After centuries of extinction due to human persecution, Eurasian beavers Castor fiber L. have been released to Southern Europe in the last decades. Being ecosystem engineers, beavers have attracted great attention regarding restoration of aquatic and terrestrial ecosystems. Nonetheless, the effects of the species on aquatic invertebrates known to date are not univocal and mostly refer to central European riverine systems. Here, we evaluated the effects of beaver presence on aquatic macroinvertebrates for the first time in a Mediterranean riverine ecosystem, by applying a sound control‐impact sampling design and controlling for seasonal variation in macroinvertebrate assemblage composition. A significant variation in response to season was evident for macroinvertebrate communities, revealing distinct assemblages during spring and summer. Furthermore, the presence of beavers was also identified as a significant driver of species composition, as samples near the beaver dam showed significant variation from control sites. Macroinvertebrate community traits changed according to the relative position to the beaver dam, as control sites featured on average higher abundances of taxa with higher values of bioindication score, larger size, lower adaptation to drag, and were less frequently of introduced origins. Yet, these differences were strongly taxon‐ and season‐specific in their intensity and direction. Differences across sites were mainly driven by the relative abundances of few taxa—including both alien species and high environmental quality indicators—such as those from genera Potamopyrgus, Baetis, Habrophlebia, Ephemerella, Leuctra, and Radix, which explained about 70% of the observed divergence among conditions. Our results indicate that beavers and their engineering activity may induce highly variable species‐specific responses in macroinvertebrates, thus possibly representing a driver of environmental heterogeneity along Mediterranean rivers, and that both bioindicators and alien species may exploit such heterogeneity.

Ecosystem Engineers in Freshwater Ecosystems

Rethinking foundation species in a changing world: The case for Rhododendron maximum as an emerging foundation species in shifting ecosystems of the southern Appalachians

Beavers, Castor canadensis in North America and Castor fiber in Eurasia, are widely referred to as nature's engineers due to their ability to rapidly transform diverse landscapes into dynamic wetland ecosystems. Few other organisms exhibit the same level of control over local geomorphic, hydrologic, and ecological conditions. Though freshwater ecosystems are particularly vulnerable to changing climate, beavers and their wetland homes have persisted throughout the Northern Hemisphere during numerous prior periods of climatic change. Some research suggests that the need to create stable, climate-buffered habitats at high latitudes during the Miocene directly led to the evolution of dam construction. As we follow an unprecedented trajectory of anthropogenic warming, we have the unique opportunity to describe how beaver ecosystem engineering ameliorates climate change today. Here, we review how beavers create and maintain local hydroclimatic stability and influence larger-scale biophysical ecosystem processes in the context of past, present, and future climate change.

North American beavers (Castor canadensis) are ecosystem engineers that create novel habitats in stream ecosystems. Although driven to the edge of extinction by historical over‐exploitation, beavers are recolonizing much of their former range. In the Southern Appalachian Mountains beavers alter habitats occupied by diverse fish communities across a mosaic of changing forest cover and management priorities. Fishes were sampled from nine streams with active beaver ponds and from seven streams with inactive ponds in the Southern Appalachians during summer 2019. The status of beaver ponds was among the most important factors affecting fish communities; however, both active and inactive beaver ponds affected fish community structure, α‐diversity, and local contributions to β‐diversity (LCBD). Fish α‐diversity was lower in reaches from streams with active ponds and the magnitude of this reduction was dependent on the position relative to a pond. However, both active and abandoned beaver ponds and pond tail races had higher LCBD values compared with free‐flowing reaches. Arcsin‐transformed forest cover did not improve the model fit for any dependent variables. Indicator species analysis found that two species had an association with active beaver ponds, whereas eight species were associated with inactive beaver ponds or free‐flowing reaches. The effects of beaver impoundments on fish communities in Appalachian streams are dependent on the pond activity level and spatial context. Although diversity and species richness may be reduced in individual habitats, beaver ponds contribute to higher biodiversity at a regional spatial scale. An increased understanding of beaver effects on Southern Appalachian fish communities will enable stakeholders to make informed decisions regarding the management of sensitive or economically important fishes. Mitigating conflicts between sensitive resources and these ecosystem engineers will become an increasingly important management concern as beaver populations recover in the Southern Appalachians.

The impact of beaver dams on distribution of waterborne Escherichia coli and turbidity in an agricultural landscape.

Megaherbivores perform vital ecosystem engineering roles, and have their last remaining stronghold in Africa. Of Africa's remaining megaherbivores, the common hippopotamus (Hippopotamus amphibius) has received the least scientific and conservation attention, despite how influential their ecosystem engineering activities appear to be. Given the potentially crucial ecosystem engineering influence of hippos, as well as mounting conservation concerns threatening their long-term persistence, a review of the evidence for hippos being ecosystem engineers, and the effects of their engineering, is both timely and necessary. In this review, we assess, (i) aspects of hippo biology that underlie their unique ecosystem engineering potential; (ii) evaluate hippo ecological impacts in terrestrial and aquatic environments; (iii) compare the ecosystem engineering influence of hippos to other extant African megaherbivores; (iv) evaluate factors most critical to hippo conservation and ecosystem engineering; and (v) highlight future research directions and challenges that may yield new insights into the ecological role of hippos, and of megaherbivores more broadly. We find that a variety of key life-history traits determine the hippo's unique influence, including their semi-aquatic lifestyle, large body size, specialised gut anatomy, muzzle structure, small and partially webbed feet, and highly gregarious nature. On land, hippos create grazing lawns that contain distinct plant communities and alter fire spatial extent, which shapes woody plant demographics and might assist in maintaining fire-sensitive riverine vegetation. In water, hippos deposit nutrient-rich dung, stimulating aquatic food chains and altering water chemistry and quality, impacting a host of different organisms. Hippo trampling and wallowing alters geomorphological processes, widening riverbanks, creating new river channels, and forming gullies along well-utilised hippo paths. Taken together, we propose that these myriad impacts combine to make hippos Africa's most influential megaherbivore, specifically because of the high diversity and intensity of their ecological impacts compared with other megaherbivores, and because of their unique capacity to transfer nutrients across ecosystem boundaries, enriching both terrestrial and aquatic ecosystems. Nonetheless, water pollution and extraction for agriculture and industry, erratic rainfall patterns and human-hippo conflict, threaten hippo ecosystem engineering and persistence. Therefore, we encourage greater consideration of the unique role of hippos as ecosystem engineers when considering the functional importance of megafauna in African ecosystems, and increased attention to declining hippo habitat and populations, which if unchecked could change the way in which many African ecosystems function.

A Note on Trophic Complexity and Community Stability

Facilitation processes constitute basic elements of vegetation dynamics in harsh systems. Recent studies in tropical alpine environments demonstrated how pioneer plant species defined as “ecosystem engineers” are capable of enhancing landscape-level richness by adding new species to the community through the modification of microhabitats, and also provided hints about the alternation of different ecosystem engineers over time. Nevertheless, most of the existing works analysed different ecosystem engineers separately, without considering the interaction of different ecosystem engineers. Focusing on the altitudinal limit of Peruvian Dry Puna vegetation, we hypothesized that positive interactions structure plant communities by facilitation cascades involving different ecosystem engineers, determining the evolution of the microhabitat patches in terms of abiotic resources and beneficiary species hosted. To analyze successional mechanisms, we used a “space-for-time” substitution to account for changes over time, and analyzed data on soil texture, composition, and temperature, facilitated species and their interaction with nurse species, and surface area of engineered patches by means of chemical analyses, indicator species analysis, and rarefaction curves. A successional process, resulting from the dynamic interaction of different ecosystem engineers, which determined a progressive amelioration of soil conditions (e.g. nitrogen and organic matter content, and temperature), was the main driver of species assemblage at the community scale, enhancing species richness. Cushion plants act as pioneers, by starting the successional processes that continue with shrubs and tussocks. Tussock grasses have sometimes been found to be capable of creating microhabitat patches independently. The dynamics of species assemblage seem to follow the nested assemblage mechanism, in which the first foundation species to colonize a habitat provides a novel substrate for colonization by other foundation species through a facilitation cascade process.

The Climate Science for Ecological Forecasting symposium