Ecosystem engineering affects ecosystem functioning in high-Andean landscapes

Ecosystem engineers are organisms that change the distribution of materials and energy in the abiotic environment, usually creating and maintaining new habitat patches in the landscape. Such changes in habitat conditions have been widely documented to affect the distributions and performances of other species but up to now no studies have addressed how such effects can impact the biotically driven physicochemical processes associated with these landscapes, or ecosystem functions. Based on the widely accepted positive relationship between species diversity and ecosystem functions, we propose that the effects of ecosystem engineers on other species could have an impact on ecosystem functions via two mutually inclusive mechanisms: (1) by adding new species into landscapes, hence increasing species diversity; and (2) by improving the performances of species already present in the landscape. To test these hypotheses, we focused on the effects of a high-Andean ecosystem engineer, the cushion plant Azorella monantha, by comparing the accumulation of plant biomass and nitrogen fixed in plant tissues as species richness increases in landscapes with and without the engineer species. Our results show that both ecosystem functions increased with species richness in both landscape types, but landscapes including A. monantha cushions reached higher outcomes of plant biomass and nitrogen fixed in plant tissues than landscapes without cushions. Moreover, our results indicate that such positive effects on ecosystem functions could be mediated by the two mechanisms proposed above. Then, given the conspicuousness of ecosystem engineering in nature and its strong influence on species diversity, and given the well-known relationship between species diversity and ecosystem function, we suggest that the application of the conceptual framework proposed herein to other ecosystems would help to advance our understanding of the forces driving ecosystem functioning. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Ecosystem engineering across ecosystems: do engineer species sharing common features have generalized or idiosyncratic effects on species diversity?

Aim  To integrate the effects of ecosystem engineers (organisms that create, maintain or destroy habitat for other species) sharing the same archetype on species diversity, and assess whether different engineer species have generalized or idiosyncratic effects across environmentally similar ecosystems.
Location  High-Andean habitats of Chile and Argentina, from 23° S to 41° S.
Methods  We measured and compared the effects of eight alpine plants with cushion growth-form on species richness, species diversity (measured as the Shannon–Wiener index) and evenness of vascular plant assemblages across four high-Andean ecosystems of Chile and Argentina.
Results  The presence of cushion plants always increased the species richness, diversity (measured as the Shannon–Wiener index) and evenness of high-Andean plant assemblages. However, while the presence of different cushion species within the same ecosystem controlled species diversity in the same way, these effects varied between cushion species from different ecosystems.
Main conclusions  Results consistently supported the idea that increases in habitat complexity due to the presence of ecosystem engineers, in this case cushion plants, would lead to higher community diversity. Results also indicate that effects of the presence of different cushion species within the same ecosystem could be generalized, while the effects of cushion species from different ecosystems should be considered idiosyncratic.     

Engineering role models: do non-human species have the answers?

A shift from traditional engineering approaches to ecologically-based techniques will require changing societal values regarding ‘how and what’ is defined as engineering and design. Non-human species offer many ecological engineering examples that are often beneficial to ecosystem function and other biota. For example, organisms known as ‘ecosystem engineers’ build, modify, and destroy habitat in their quest for food and survival. Similarly, ‘keystone species’ have greater impacts on community or ecosystem function than would be predicted from their abundance. The capacity of these types of organisms to affect ecosystems is great. They exert controlling influences over ecosystems and communities by altering resource allocation, creating habitats and modifying relative competitive advantages.Species’ effects in ecosystems, although context-dependent, can be evaluated as ‘beneficial’ or ‘detrimental’. The evaluation depends on whether effects on other species or ecosystem function are more or less desirable from a given perspective. Organisms with beneficial impacts facilitate the presence of other species, employ efficient nutrient cycling, and are sometimes characterized by specific mutualisms. In contrast, many cases of detrimental engineering are found from introduced (i.e., exotic) species and are characterized by a loss of species richness, a lack of nutrient retention and the degradation of ecosystem integrity. Species’ impacts on ecosystems and community traits have been quantified in ecological studies and can be used similarly to understand, design and model human engineering structures and impacts on the landscape. Emulation of species with beneficial impacts on ecosystems can provide powerful guidance to the goals of ecological engineering. Using role model organisms that have desirable effects on species diversity and ecosystem function will be important in developing alternatives to traditional engineering practices.     

Impacts of ecosystem engineers on community attributes: effects of cushion plants at different elevations of the Chilean Andes

Ecosystem engineers are organisms able to modulate environmental forces and, hence, may change the habitat conditions for other species. In so doing, ecosystem engineers may affect both species richness and evenness of communities and, in consequence, change species diversity. If these changes in community attributes are related to the magnitude of the habitat changes induced by the engineers, it seems likely that engineer species will have greater effects on diversity in sites where they cause larger habitat changes. We addressed this issue by evaluating the effects of three alpine cushion plants on species richness, evenness, and diversity of high-Andean plant communities. Given that the difference in microclimatic conditions between cushions and the external environment increases with elevation, we proposed that these organisms should have greater effects on community attributes at higher than at lower elevation sites. Results showed that the three cushion species had positive effects on species richness, diversity, and evenness of plant communities. It was also observed that the magnitude of these effects changed with elevation: positive effects on species richness and diversity increased towards upper sites for the three cushions species, whereas positive effects on evenness increased with elevation for one cushion species but decreased with elevation for other two cushion species. These results suggest that the presence of cushions is important to maintain plant diversity in high-Andean communities, but this positive effect on diversity seems to increase as the difference in environmental conditions between cushions and the external environment increases with elevation.     

Macroalgal blooms alter community structure and primary productivity in marine ecosystems

Eutrophication, coupled with loss of herbivory due to habitat degradation and overharvesting, has increased the frequency and severity of macroalgal blooms worldwide. Macroalgal blooms interfere with human activities in coastal areas, and sometimes necessitate costly algal removal programmes. They also have many detrimental effects on marine and estuarine ecosystems, including induction of hypoxia, release of toxic hydrogen sulphide into the sediments and atmosphere, and the loss of ecologically and economically important species. However, macroalgal blooms can also increase habitat complexity, provide organisms with food and shelter, and reduce other problems associated with eutrophication. These contrasting effects make their overall ecological impacts unclear. We conducted a systematic review and meta‐analysis to estimate the overall effects of macroalgal blooms on several key measures of ecosystem structure and functioning in marine ecosystems. We also evaluated some of the ecological and methodological factors that might explain the highly variable effects observed in different studies. Averaged across all studies, macroalgal blooms had negative effects on the abundance and species richness of marine organisms, but blooms by different algal taxa had different consequences, ranging from strong negative to strong positive effects. Blooms' effects on species richness also depended on the habitat where they occurred, with the strongest negative effects seen in sandy or muddy subtidal habitats and in the rocky intertidal. Invertebrate communities also appeared to be particularly sensitive to blooms, suffering reductions in their abundance, species richness, and diversity. The total net primary productivity, gross primary productivity, and respiration of benthic ecosystems were higher during macroalgal blooms, but blooms had negative effects on the productivity and respiration of other organisms. These results suggest that, in addition to their direct social and economic costs, macroalgal blooms have ecological effects that may alter their capacity to deliver important ecosystem services.     

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.     

Effects of Tea Plantations on Stream Invertebrates in a Global Biodiversity Hotspot in Africa

Tropical stream ecosystems in montane forest watersheds are important centers of endemism and diversity and provide essential ecosystem services. These habitats are subject to a variety of stressors, including the conversion of adjacent terrestrial habitats from forest to agriculture, but the impacts of these anthropogenic effects are largely unknown because of the paucity of studies in these systems. In montane habitats in the wet tropics, large-scale cultivation of tea is common and can represent an important source of income at local and national scales. However, little is known about how tea cultivation impacts adjacent stream ecosystems. In this study, we examine stream macroinvertebrate assemblages in a biodiversity hotspot the East Usambara Mountains, Tanzania. Specifically, we compare diversity of macroinvertebrate assemblages found on cobbles in stream riffles in watersheds dominated by forest with those surrounded by tea cultivation. We found that streams surrounded by tea were characterized by significantly lower dissolved oxygen and had lower total estimated species richness and number of families. Furthermore, the richness of invertebrate taxa known to be sensitive to anthropogenic disturbance were substantially reduced in tea streams and general assemblage-level analysis shows significant differences in the composition of macroinvertebrate assemblages between tea and forested streams. Our results suggest that tea cultivation may reduce stream habitat quality and biodiversity in the East Usambaras. Further research is needed to evaluate the effects of tea cultivation on streams over longer times scales and to address methods for minimizing negative effects of agriculture on montane stream communities.     

An ecosystem engineer,the beaver,increases species richness at the landscape scale

Ecosystem engineering - the physical modification of habitats by organisms - has been proposed as an important mechanism for maintaining high species richness at the landscape scale by increasing habitat heterogeneity. Dams built by beaver (Castor canadensis) dramatically alter riparian landscapes throughout much of North America. In the central Adirondacks, New York, USA, ecosystem engineering by beaver leads to the formation of extensive wetland habitat capable of supporting herbaceous plant species not found elsewhere in the riparian zone. We show that by increasing habitat heterogeneity, beaver increase the number of species of herbaceous plants in the riparian zone by over 33% at a scale that encompasses both beaver-modified patches and patches with no history of beaver occupation. We suggest that ecosystem engineers will increase species richness at the landscape scale whenever there are species present in a landscape that are restricted to engineered habitats during at least some stages of their life cycle.     

Shrubs as ecosystem engineers across an environmental gradient: effects on species richness and exotic plant invasion

Ecosystem-engineering plants modify the physical environment and can increase species diversity and exotic species invasion. At the individual level, the effects of ecosystem engineers on other plants often become more positive in stressful environments. In this study, we investigated whether the community-level effects of ecosystem engineers also become stronger in more stressful environments. Using comparative and experimental approaches, we assessed the ability of a native shrub (Ericameria ericoides) to act as an ecosystem engineer across a stress gradient in a coastal dune in northern California, USA. We found increased coarse organic matter and lower wind speeds within shrub patches. Growth of a dominant invasive grass (Bromus diandrus) was facilitated both by aboveground shrub biomass and by growing in soil taken from shrub patches. Experimental removal of shrubs negatively affected species most associated with shrubs and positively affected species most often found outside of shrubs. Counter to the stress-gradient hypothesis, the effects of shrubs on the physical environment and individual plant growth did not increase across the established stress gradient at this site. At the community level, shrub patches increased beta diversity, and contained greater rarified richness and exotic plant cover than shrub-free patches. Shrub effects on rarified richness increased with environmental stress, but effects on exotic cover and beta diversity did not. Our study provides evidence for the community-level effects of shrubs as ecosystem engineers in this system, but shows that these effects do not necessarily become stronger in more stressful environments.     

Grazing weakens temporal stabilizing effects of diversity in the Eurasian steppe

Many biodiversity experiments have demonstrated that plant diversity can stabilize productivity in experimental grasslands. However, less is known about how diversity–stability relationships are mediated by grazing. Grazing is known for causing species losses, but its effects on plant functional groups (PFGs) composition and species asynchrony, which are closely correlated with ecosystem stability, remain unclear. We conducted a six‐year grazing experiment in a semi‐arid steppe, using seven levels of grazing intensity (0, 1.5, 3.0, 4.5, 6.0, 7.5, and 9.0 sheep per hectare) and two grazing systems (i.e., a traditional, continuous grazing system during the growing period (TGS), and a mixed one rotating grazing and mowing annually (MGS)), to examine the effects of grazing system and grazing intensity on the abundance and composition of PFGs and diversity–stability relationships. Ecosystem stability was similar between mixed and continuous grazing treatments. However, within the two grazing systems, stability was maintained through different pathways, that is, along with grazing intensity, persistence biomass variations in MGS, and compensatory interactions of PFGs in their biomass variations in TGS. Ecosystem temporal stability was not decreased by species loss but rather remain unchanged by the strong compensatory effects between PFGs, or a higher grazing‐induced decrease in species asynchrony at higher diversity, and a higher grazing‐induced increase in the temporal variation of productivity in diverse communities. Ecosystem stability of aboveground net primary production was not related to species richness in both grazing systems. High grazing intensity weakened the temporal stabilizing effects of diversity in this semi‐arid grassland. Our results demonstrate that the productivity of dominant PFGs is more important than species richness for maximizing stability in this system. This study distinguishes grazing intensity and grazing system from diversity effects on the temporal stability, highlighting the need to better understand how grazing regulates ecosystem stability, plant diversity, and their synergic relationships.     

Ecosystem engineers maintain a rare species of butterfly and increase plant diversity

We evaluated whether ecosystem engineers can accomplish two conservation goals simultaneously: (1) indirectly maintain populations of an endangered animal through habitat modification and (2) increase riparian plant diversity. We tested for effects of a prominent ecosystem engineer, the beaver Castor canadensis, on populations of St. Francis' satyr butterfly Neonympha mitchellii francisci and plant species richness and composition. We performed our test by surveying riparian vegetation communities in all stages of beaver‐influenced wetland succession. We found that beavers created wetland habitats that supported plant species not found elsewhere in riparian zones and increased plant species diversity across the landscape by creating a novel combination of patch types. Our results confirmed what others have found about engineering effects on plant diversity, but these results further demonstrated a case where ecosystem engineers indirectly maintain populations of rare animals by modifying the composition and diversity of plant communities within wetlands. Our research demonstrates how an ecosystem engineer can influence habitat availability and composition of plant communities important for an endangered insect, and maintain overall plant species diversity by increasing habitat heterogeneity.     

Phylogenetic diversity does not capture body size variation at risk in the world's mammals

Mammals contribute to important ecosystem processes and services, but many mammalian species are threatened with extinction. We compare how global patterns in three measures of mammalian diversity—species richness, phylogenetic diversity (PD) and body mass variance (BMV)—would change if all currently threatened species were lost. Given that many facets of species'' ecology and life history scale predictably with body mass, the BMV in a region roughly reflects the diversity of species'' roles within ecosystems and so is a simple proxy for functional diversity (FD). PD is also often considered to be a proxy for FD, but our results suggest that BMV losses within ecoregions would be much more severe than losses of PD or species richness, and that its congruence with the latter two measures is low. Because of the disproportionate loss of large mammals, 65 per cent of ecoregions would lose significantly more BMV than under random extinction, while only 11 per cent would lose significantly more PD. Ecosystem consequences of these selective losses may be profound, especially throughout the tropics, but are not captured by PD. This low surrogacy stresses a need for conservation prioritization based on threatened trait diversity, and for conservation efforts to take an ecosystem perspective.     

Insects as stem engineers: interactions mediated by the twig-girdler Oncideres albomarginata chamela enhance arthropod diversity

Background

Ecosystem engineering may influence community structure and biodiversity by controlling the availability of resources and/or habitats used by other organisms. Insect herbivores may act as ecosystem engineers but there is still poor understanding of the role of these insects structuring arthropod communities.

Methodology/Principal Findings

We evaluated the effect of ecosystem engineering by the stem-borer Oncideres albomarginata chamela on the arthropod community of a tropical dry forest for three consecutive years. The results showed that ecosystem engineering by O. albomarginata chamela had strong positive effects on the colonization, abundance, species richness and composition of the associated arthropod community, and it occurred mainly through the creation of a habitat with high availability of oviposition sites for secondary colonizers. These effects cascade upward to higher trophic levels. Overall, ecosystem engineering by O. albomarginata chamela was responsible for nearly 95% of the abundance of secondary colonizers and 82% of the species richness.

Conclusions/Significance

Our results suggest that ecosystem engineering by O. albomarginata chamela is a keystone process structuring an arthropod community composed by xylovores, predators and parasitoids. This study is the first to empirically demonstrate the effect of the ecosystem engineering by stem-boring insects on important attributes of arthropod communities. The results of this study have important implications for conservation.     

Interactive effects of keystone rodents on the structure of desert grassland arthropod communities

Certain species play particularly large roles in ecosystems, and are often referred to as keystones. However, little is known about the interactive effects of these species where they co-occur. Prairie dogs ( Cynomys spp.) and banner-tailed kangaroo rats Dipodomys spectabilis are commonly considered keystone species of grassland ecosystems, creating a mosaic of unique habitats on the landscape through ecosystem engineering and herbivory. We examined the separate and interactive effects of these species on the structure of grassland arthropod communities. We conducted a cross-site study at two locations in the northern Chihuahuan Desert, and evaluated the impacts of these rodents on ground-dwelling arthropod and grasshopper communities in areas where prairie dogs and kangaroo rats co-occurred compared to areas where each rodent species occurred alone. Our results demonstrate that prairie dogs ( C. gunnisoni and C. ludovicianus ) and banner-tailed kangaroo rats had keystone-level impacts on arthropod communities both separately and interactively. Their burrow systems provided important habitats for multiple trophic and taxonomic groups of arthropods, and increased overall arthropod abundance and species richness. Many arthropods also were attracted to the aboveground habitats around the mounds and across the landscapes where the rodents occurred. Detritivores, predators, ants, grasshoppers, and rare rodent burrow inhabitants were especially associated with prairie dog and kangaroo rat activity. The impacts of prairie dogs and kangaroo rats were unique, and the habitats they created supported different assemblages of arthropods. Where both rodent species co-occurred, there was greater heterogeneity and arthropod diversity on the landscape. Our results suggest that the interaction of multiple keystones, especially those with engineering roles, results in unique and more diverse communities in time and space.     

Ecosystem engineering by invasive exotic beavers reduces in-stream diversity and enhances ecosystem function in Cape Horn,Chile

Species invasions are of global significance, but predicting their impacts can be difficult. Introduced ecosystem engineers, however, provide an opportunity to test the underlying mechanisms that may be common to all invasive engineers and link relationships between changes in diversity and ecosystem function, thereby providing explanatory power for observed ecological patterns. Here we test specific predictions for an invasive ecosystem engineer by quantifying the impacts of habitat and resource modifications caused by North American beavers (Castor canadensis) on aquatic macroinvertebrate community structure and stream ecosystem function in the Cape Horn Biosphere Reserve, Chile. We compared responses to beavers in three habitat types: (1) forested (unimpacted) stream reaches, (2) beaver ponds, and (3) sites immediately downstream of beaver dams in four streams. We found that beaver engineering in ponds created taxonomically simplified, but more productive, benthic macroinvertebrate assemblages. Specifically, macroinvertebrate richness, diversity and number of functional feeding groups were reduced by half, while abundance, biomass and secondary production increased three- to fivefold in beaver ponds compared to forested sites. Reaches downstream of beaver ponds were very similar to natural forested sections. Beaver invasion effects on both community and ecosystem parameters occurred predominantly via increased retention of fine particulate organic matter, which was associated with reduced macroinvertebrate richness and diversity (via homogenization of benthic microhabitat) and increased macroinvertebrate biomass and production (via greater food availability). Beaver modifications to macroinvertebrate community structure were largely confined to ponds, but increased benthic production in beaver-modified habitats adds to energy retention and flow for the entire stream ecosystem. Furthermore, the effects of beavers on taxa richness (negative) and measures of macroinvertebrate biomass (positive) were inversely related. Thus, while a generally positive relationship between diversity and ecosystem function has been found in a variety of systems, this work shows how they can be decoupled by responding to alterative mechanisms. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Partitioning the effects of algal species identity and richness on benthic marine primary production

Influential research in terrestrial habitats indicates that several ecosystem processes are related to plant biodiversity, yet these links remain poorly studied in marine ecosystems. We conducted one field and one mesocosm experiment to quantify the relative effects of macroalgal species identity and richness on primary production in coral reef macroalgal communities off the north coast of Jamaica. We measured production as the net accumulation of algal biomass in the absence of consumers and as photosynthetic rate using oxygen probes in sealed aquaria. We used two recently developed techniques to attribute deviations in expected relative yield to components associated with species identity or diversity and then to further partition diversity effects into mechanistic components based on dominance, trait-dependent complementarity, and trait-independent complementarity. Our results indicate that algal identity had far greater effects on absolute net growth and photosynthesis than richness. The most diverse mixture of macroalgae did not outperform the most productive monoculture or the average monoculture in either measure of primary production (i.e. we did not find evidence of either transgressive or non-transgressive overyielding). Trait-independent complementarity effects were positive but dominance and trait-dependent complementarity were both negative and became stronger when richness was increased. Thus the potentially positive influence of species interactions and niche partitioning on production were negated by dominance and other negative selection effects. These results demonstrate that the counteracting influence of component effects can diminish the net richness effects on production. This could explain frequently observed weak net richness effects in other aquatic and terrestrial systems and suggests that life history tradeoffs greatly reduce the potential for ecologically relevant plant biodiversity effects on ecosystem properties.     

A global synthesis of the effects of diversified farming systems on arthropod diversity within fields and across agricultural landscapes

Agricultural intensification is a leading cause of global biodiversity loss, which can reduce the provisioning of ecosystem services in managed ecosystems. Organic farming and plant diversification are farm management schemes that may mitigate potential ecological harm by increasing species richness and boosting related ecosystem services to agroecosystems. What remains unclear is the extent to which farm management schemes affect biodiversity components other than species richness, and whether impacts differ across spatial scales and landscape contexts. Using a global metadataset, we quantified the effects of organic farming and plant diversification on abundance, local diversity (communities within fields), and regional diversity (communities across fields) of arthropod pollinators, predators, herbivores, and detritivores. Both organic farming and higher in‐field plant diversity enhanced arthropod abundance, particularly for rare taxa. This resulted in increased richness but decreased evenness. While these responses were stronger at local relative to regional scales, richness and abundance increased at both scales, and richness on farms embedded in complex relative to simple landscapes. Overall, both organic farming and in‐field plant diversification exerted the strongest effects on pollinators and predators, suggesting these management schemes can facilitate ecosystem service providers without augmenting herbivore (pest) populations. Our results suggest that organic farming and plant diversification promote diverse arthropod metacommunities that may provide temporal and spatial stability of ecosystem service provisioning. Conserving diverse plant and arthropod communities in farming systems therefore requires sustainable practices that operate both within fields and across landscapes.     

The impacts of invasive ecosystem engineers in freshwaters: A review

1. 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.
2. 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.
3. 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.
4. 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.
5. 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.
6. 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.

     

Meta‐analysis of the effects of small mammal disturbances on species diversity,richness and plant biomass

The disturbance activities of many small mammals, including building burrows, mounds, trails and tunnels, and herbivory, can have significant impacts on their ecosystems, both through trophic and non‐trophic interactions. Some species have large enough impacts through their disturbances to be classed as ecosystem engineers and/or keystone species. Others have negative or null effects. However, at present it is difficult to predict whether the disturbances created by a given species will have significant effects on common measures of ecosystem response such as species richness, diversity and biomass. We ask whether variables characterizing disturbance type, responding species, disturbance‐making species and the environment can predict changes in magnitude and direction of effects on biomass, richness and diversity. We test these predictions with a meta‐analysis of 106 data entries in a database derived from 63 papers, representing 40 small mammal species. We find that small mammal disturbances in general increase biomass, and both increase and decrease richness and diversity. We also identify individual environmental, disturbance‐related, and species‐related variables associated with these changes in magnitude and direction. We discuss the likely interactions between these variables, and how current proxy measures of disturbance impact could be replaced by more accurate direct measures. We recommend that future studies focus on conditions characterized by combinations of variables we identify as significant, in order to understand how these variable interactions (which cannot be analysed through meta‐analysis) affect disturbance outcomes. Based on the gaps in our database and results, we also recommend that future studies directly measure disturbance impact, measure disturbance effects on animal and well as plant taxa, and take measurements on multiple scales.     

A perfect storm: two ecosystem engineers interact to degrade deciduous forests of New Jersey

Ecosystem engineers play a large role in physically structuring the ecosystem in which they are embedded. The focus of much of the research surrounding these species is to document the impacts of a single engineer on community composition and ecosystem processes. However, most ecosystems harbor multiple engineering species that interact in complex ways and rarely have the dynamics of such species been fully investigated. We look at how two ecosystem engineers, the white-tailed deer (Odocoileus virginianus) and the invasive plant Japanese stilt grass (Microstegium vimineum), interact to completely alter the structure and composition of the subcanopy within northern deciduous forests. This interaction has wide-ranging repercussions on forest food webs which we explore through a case study of breeding woodland birds in the state of New Jersey, USA. We show that the guilds of birds that rely on the subcanopy have experienced greater declines from 1980 to 2005 than birds that specialize on the intact upper canopy of impacted forests. This dynamic is not restricted to immediate temporal effects and may act to derail the long-term successional pathway of northern deciduous forests. It is no longer prudent to set aside tracts of forest and expect them to retain their native biodiversity without active management.