Ecosystem feedbacks and cascade processes: understanding their role in the responses of Arctic and alpine ecosystems to environmental change

Global environmental change, related to climate change and the deposition of airborne N‐containing contaminants, has already resulted in shifts in plant community composition among plant functional types in Arctic and temperate alpine regions. In this paper, we review how key ecosystem processes will be altered by these transformations, the complex biological cascades and feedbacks that might result, and some of the potential broader consequences for the earth system. Firstly, we consider how patterns of growth and allocation, and nutrient uptake, will be altered by the shifts in plant dominance. The ways in which these changes may disproportionately affect the consumer communities, and rates of decomposition, are then discussed. We show that the occurrence of a broad spectrum of plant growth forms in these regions (from cryptogams to deciduous and evergreen dwarf shrubs, graminoids and forbs), together with hypothesized low functional redundancy, will mean that shifts in plant dominance result in a complex series of biotic cascades, couplings and feedbacks which are supplemental to the direct responses of ecosystem components to the primary global change drivers. The nature of these complex interactions is highlighted using the example of the climate‐driven increase in shrub cover in low‐Arctic tundra, and the contrasting transformations in plant functional composition in mid‐latitude alpine systems. Finally, the potential effects of the transformations on ecosystem properties and processes that link with the earth system are reviewed. We conclude that the effects of global change on these ecosystems, and potential climate‐change feedbacks, cannot be predicted from simple empirical relationships between processes and driving variables. Rather, the effects of changes in species distributions and dominances on key ecosystem processes and properties must also be considered, based upon best estimates of the trajectories of key transformations, their magnitude and rates of change.     

On carbon sequestration in desert ecosystems

Recent reports of net ecosysytem production >100 gCm⁻² yr⁻¹ in deserts are incompatible with existing measurements of net primary production and carbon pools in deserts. The comparisions suggest that gas exchange measurements should be used with caution and better validation if they are expected to indicate the magnitude of carbon sink in these ecosysytems.     

Change in CO2 balance under a series of forestry activities in a cool-temperate mixed forest with dense undergrowth

To evaluate the effects on CO₂ exchange of clearcutting a mixed forest and replacing it with a plantation, 4.5 years of continuous eddy covariance measurements of CO₂ fluxes and soil respiration measurements were conducted in a conifer-broadleaf mixed forest in Hokkaido, Japan. The mixed forest was a weak carbon sink (net ecosystem exchange, −44 g C m⁻² yr⁻¹), and it became a large carbon source (569 g C m⁻² yr⁻¹) after clearcutting. However, the large emission in the harvest year rapidly decreased in the following 2 years (495 and 153 g C m⁻² yr⁻¹, respectively) as the gross primary production (GPP) increased, while the total ecosystem respiration (RE) remained relatively stable. The rapid increase in GPP was attributed to an increase in biomass and photosynthetic activity of Sasa dwarf bamboo, an understory species. Soil respiration increased in the 3 years following clearcutting, in the first year mainly owing to the change in the gap ratio of the forest, and in the following years because of increased root respiration by the bamboo. The ratio of soil respiration to RE increased from 44% in the forest to nearly 100% after clearcutting, and aboveground parts of the vegetation contributed little to the RE although the respiration chamber measurements showed heterogeneous soil condition after clearcutting.     

Linking above- and belowground responses to global change at community and ecosystem scales

Cryptic belowground organisms are difficult to observe and their responses to global changes are not well understood. Nevertheless, there is reason to believe that interactions among above- and belowground communities may mediate ecosystem responses to global change. We used grassland mesocosms to manipulate the abundance of one important group of soil organisms, arbuscular mycorrhizal (AM) fungi, and to study community and ecosystem responses to CO₂ and N enrichment. Responses of plants, AM fungi, phospholipid fatty acids and community-level physiological profiles were measured after two growing seasons. Ecosystem responses were examined by measuring net primary production (NPP), evapotranspiration, total soil organic matter (SOM), and extractable mineral N. Structural equation modeling was used to examine the causal relationships among treatments and response variables. We found that while CO₂ and N tended to directly impact ecosystem functions (evapotranspiration and NPP, respectively), AM fungi indirectly impacted ecosystem functions by influencing the community composition of plants and other root fungi, soil fungi and soil bacteria. We found that the mycotrophic status of the dominant plant species in the mesocosms determined whether the presence of AM fungi increased or decreased NPP. Mycotrophic grasses dominated the mesocosm communities during the first growing season, and the mycorrhizal treatments had the highest NPP. In contrast, nonmycotrophic forbs were dominant during the second growing season and the mycorrhizal treatments had the lowest NPP. The composition of the plant community strongly influenced soil N, and the community composition of soil organisms strongly influenced SOM accumulation in the mesocosms. These results show how linkages between above- and belowground communities can determine ecosystem responses to global change.     

Strong links between teleconnections and ecosystem exchange found at a Pacific Northwest old-growth forest from flux tower and MODIS EVI data

Variability in three Pacific teleconnection patterns are examined to see if net carbon exchange at a low‐elevation, old‐growth forest is affected by climatic changes associated with these periodicities. Examined are the Pacific Decadal Oscillation (PDO), Pacific/North American Oscillation (PNA) and El Niño‐Southern Oscillation (ENSO). We use 9 years of eddy covariance CO₂, H₂O and energy fluxes measured at the Wind River AmeriFlux site, Washington, USA and 8 years of tower‐pixel remote sensing data from the Moderate Resolution Imaging Spectroradiometer (MODIS) to address this question. We compute a new Composite Climate Index (CCI) based on the three Pacific Oscillations to divide the measurement period into positive‐ (2003 and 2005), negative‐ (1999 and 2000) and neutral‐phase climate years (2001, 2002, 2004, 2006 and 2007). The forest transitioned from an annual net carbon sink (NEP=+217 g C m^?2 yr^?1, 1999) to a source (NEP=?100 g C m^?2 yr^?1, 2003) during two dominant teleconnection patterns. Net ecosystem productivity (NEP), water use efficiency (WUE) and light use efficiency (LUE) were significantly different (P<0.01) during positive (NEP=?0.27 g C m^?2 day^?1, WUE=4.1 mg C g^?1 H₂O, LUE=0.94 g C MJ^?1) and negative (NEP=+0.37 g C m^?2 day^?1, WUE=3.4 mg C g^?1 H₂O, LUE=0.83 g C MJ^?1) climate phases. The CCI was linked to variability in the MODIS Enhanced Vegetation Index (EVI) but not to MODIS Fraction of absorbed Photosynthetically Active Radiation (FPAR). EVI was highest during negative climate phases (1999 and 2000) and was positively correlated with NEP and showed potential for using MODIS to estimate teleconnection‐driven anomalies in ecosystem CO₂ exchange in old‐growth forests. This work suggests that any increase in the strength or frequency of ENSO coinciding with in‐phase, low frequency Pacific oscillations (PDO and PNA) will likely increase CO₂ uptake variability in Pacific Northwest conifer forests.     

Evaluating different harvest intensities over understory plant diversity and pine seedlings,in a <Emphasis Type="Italic">Pinus pinaster</Emphasis> Ait. natural stand of Spain

Although modern forestry takes into consideration the analysis of the effects of forest management on plant structure, diversity and seedlings, little is known about how those parameters respond to harvest techniques in the Mediterranean region. We investigated the effect of three different harvest intensities, respect to uncut controls, on understory plant species functional groups, richness, diversity and pine seedlings in a natural Maritime pine stand in Spain, three years after harvesting. The harvest treatments produced a reduction of the number of Pinus pinaster seedlings and woody species cover, and an increase of species richness (total and of annual species) and plant cover of annual species respect to control plots (CO). The Shannon diversity values showed no differences between treatments. These results emphasize that the tree harvest treatments analyzed are not suitable for the management of this P. pinaster stand. Otherwise, the reduction of pine seedling density by harvest treatments and the changes in richness and cover of functional groups would not induce the natural regeneration of this stand maintaining the understory plant layer.     

Reducing horizontal and vertical diversity in a foodweb triggers extinctions and impacts functions

Species loss can result in secondary extinctions and changes in ecosystem functions at distant trophic levels. Such effects of species loss are predicted to be affected by both the number of species lost within a trophic level (horizontal diversity) and the number of trophic levels lost (vertical diversity). We experimentally manipulated horizontal and vertical diversity within an aquatic insect community, and examined responses throughout the food web. Horizontal and vertical diversity both impacted ciliates: reduction of detritivorous insect diversity resulted in secondary extinctions and decreased density of ciliates, but only when an insect predator was simultaneously absent. Horizontal and vertical diversity differed in their effect on other foodweb processes, including detrital processing, predator growth, and densities of rotifers, flagellates and flatworms. These results caution that foodweb effects of multitrophic species loss may not be reliably predicted from manipulations of just one dimension of diversity.     

Understanding relationships among multiple ecosystem services

Ecosystem management that attempts to maximize the production of one ecosystem service often results in substantial declines in the provision of other ecosystem services. For this reason, recent studies have called for increased attention to development of a theoretical understanding behind the relationships among ecosystem services. Here, we review the literature on ecosystem services and propose a typology of relationships between ecosystem services based on the role of drivers and the interactions between services. We use this typology to develop three propositions to help drive ecological science towards a better understanding of the relationships among multiple ecosystem services. Research which aims to understand the relationships among multiple ecosystem services and the mechanisms behind these relationships will improve our ability to sustainably manage landscapes to provide multiple ecosystem services.