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Home , Restoration ecology

OUP UNCORRECTED PROOF – FIRST PROOF, 01/23/12, SPi

CHAPTER 5.6 Applying Ecological Knowledge to Restore Services

S ara G . B aer , L iam H eneghan, and Valerie T . E viner

5.6.1 Introduction can take tens to thousands of years for some soil properties to develop through the interaction of Ecosystem degradation resulting from parent material, climate, topography, and organ- extraction, land-use change, and invasion by exotic isms (Jenny 1941 ). Soil degradation can result in the species alters numerous functions and services loss of or alteration to many soil properties and provided by intact and unexploited . functions. We defi ne soil legacy, as the physical, Ecological restoration is the human-facilitated chemical, and biological attributes and interactions improvement of a degraded ecosystem and repre- that remain following a signifi cant change to an sents an important means to repair or reinstate ecosystem. This defi nition of legacy is aligned with many natural services in degraded ecosystems ecological legacy (White & Jentsch 2004 ), but differs (Table 5.6.1 ). Restoration can be initiated from any from that which has been used to indicate the resid- point along a continuum of degradation and resto- ual effects of an abiotic or biotic on eco- ration goals can vary from focused improvements system properties (e.g. Reinhart & Callaway 2006 ). (e.g. amelioration of unsuitable pH, soil stabiliza- Thus, soil legacy is the degree to which soil proper- tion, and re-establishing rare species) to holistic ties (e.g. horizonation, porosity, texture, nutrient recovery of biological diversity, effi cient nutrient storage, organic matter content, aggregation, etc.) cycling, and complex energy fl ow pathways (Hobbs and functions (e.g. nutrient supply, infi ltration, etc.) & Harris 2001 ). Restorations that aim to re-establish at the onset of restoration refl ect characteristics ecosystem structure and function “prior to degra- before the degrading infl uence. dation” may fi nd that historical and extant targets The range of variation in ecosystem degradation can be diffi cult to defi ne considering the variability produces varying soil legacy at the onset of restora- of natural systems in space and time (White & tion. Heneghan et al. (2008 ) proposed that more soil Walker 1997 ) or unrealistic to attain if multiple abi- ecological knowledge (defi ned as the integrated otic and/or biotic factors have been highly modi- understanding of soil physical, chemical, and bio- fi ed through human activity ( Hilderbrand et al . logical factors and processes in the context of - 2005 ). Furthermore, restorations are now conducted soil feedback) may be required to restore complex under novel conditions including interactions following disturbance ( Fig. 5.6.1 ). This pressure, greater inputs of nutrients through atmos- chapter presents the utility of soil ecological knowl-

pheric deposition, and higher atmospheric CO2 lev- edge to the practice of ecological restoration along a els that may restrict the ability to restore systems to continuum of ecosystem degradation that results in some state in the past ( Hobbs & Harris 2001 ; Hobbs varying legacy of the plant and/or soil system. et al. 2009 ). Mineral resource extraction can result in severe eco- Degradation of terrestrial ecosystems is often system degradation that leaves little to no soil legacy, strongly refl ected as damage to the soil system. It initially. Restoration of these highly degraded lands

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378 SOIL AND ECOSYSTEM SERVICES

Table 5.6.1 Ecosystem goods and services provided through ecological restoration.

Ecosystem servicea How ecological restoration can provide ecosystem goods (functions) and services

Gas and climate regulation of degraded lands through , grassland establishment, and

restoration can aid in mitigating atmospheric CO2 ; conversion of agricultural land to perennial

vegetation can reduce N2 O emissions if nitrifi cation rates are reduced. Disturbance regulation Restoration of degraded lands to perennial vegetation increases transpiration of water to the atmosphere and improved promotes infi ltration to reduce runoff and provide fl ood control . Water supply Improved soil structure promotes infi ltration to groundwater; hydrologic manipulations in wetland and fl oodplain restorations can promote groundwater recharge . control and sediment retention Conversion of highly erodible arable lands and zones to perennial vegetation reduces erosion and traps sediment. Soil formation Restoration of degraded lands to perennial vegetation promotes organic matter accrual; organic acids from root exudates and can facilitate weathering and soil formation. Nutrient cycling Establishing associated with N-fi xing on degraded lands can promote N and organic matter accrual; developing root systems, microbial , and organic matter during restoration of degraded soil can promote nutrient conservation; conversion of agricultural lands to perennial vegetation reduces nutrient inputs and pollution. Biological control Perennial vegetation restored within agricultural landscapes can provide refugia for predators of crop pests. Pollination Floristically diverse restorations can provision pollinators for plant populations. Refugia Restoration can reduce landscape fragmentation and provide resources and/or reproduction requirements for local and transient populations. Food production Restoration of degraded land and with perennial vegetation can increase game populations. Raw materials can be managed for production of lumber; perennial grasslands can be used for biofuel and forage production. Genetic resources Restoration conducted with high fi delity to local gene pools represents a means to preserve genetic variation that could harbor genes for medicinal purposes or crop improvement (e.g. resistance to plant pathogens). Recreation Forest, wetland, and grassland restoration increase habitat for wildlife and opportunity for hunting; restoration can improve (increase clarity through less sedimentation, reduced eutrophication through nutrient abatement) and conditions for fi sh populations; restorations provide areas for hiking and nature appreciation. Cultural Restorations and re-creations, especially when conducted to achieve a historic , can preserve cultural heritage.

a List modifi ed from Costanzaet al . ( 1997 ).

has revealed the importance of physical, chemical, with the type of production system. Most develop- and biological properties of soil to revegetation, as ment and application of soil ecological knowledge well as the role of soil heterogeneity in providing to the restoration of agricultural systems has been refugia for the recolonization of soil biota. Addition gleaned from those that have been cultivated. of topsoil to mined land can rapidly increase the Although recovery of many aspects of soil structure amount of soil legacy at the onset of restoration and function can proceed passively following with consequence for improved ecosystem func- revegetation of cultivated , there are increasing tions. Restoration of agricultural systems often rep- efforts to restore biological and physical complexity resents moderate soil legacy at the onset of restoration. to better represent historic or extant systems. Restor- Soil degradation resulting from varies ing and structural complexity may

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APPLYING SOIL ECOLOGICAL KNOWLEDGE TO RESTORE ECOSYSTEM SERVICES 379

High I. II. III. Plant Factors and processes community in the soil system: P dynamics P = physical C = chemical r e B = biological v CB e g e time P r t e a v t P e i C B g o e n C t a t i Management Ecosystem functioning B o eutrophication n acidification invasion novel ecosystem

Low Low Soil system legacy High

Figure 5.6.1 Relationship between the degree of soil legacy in degraded lands (including interaction among physical (P), chemical (C), and biological (B) properties and processes) and ecosystem functioning in the context of restoration. Restoration is human-facilitated improvement of a degraded ecosystem’s functioning (indicated by stair-step line). Abiotic and biotic constraints may need to be alleviated to achieve proportionally greater improvement in ecosystem functioning (indicated by the dashed line within gray windows). Three general circumstances of varying soil legacy are depicted, although a continuum exists within the realm of each example (regions I-III). Region I represents restoration of highly disturbed lands (e.g. some mine land reclamations), where P, C, and/or B properties and processes may be disconnected, and reconnection of these components is required to restore vegetation. Region II depicts restoration from disturbance such as agriculture, where P, C, and B properties and processes may be present and interactive, but altered in status and/or composition. Revegetation drives the recovery of P, C, and B properties and processes towards a target state in region II, but recovery may be constrained by limited species, development of complex plant-soil feedback, or altered representation of higher trophic levels (e.g. aboveground ). Region III represents restoration scenarios where soil legacy may be signifi cant, but plant communities are undergoing dynamic change (invasion) or have been replaced by self-perpetuating assemblages of species that have never before coexisted (novel ecosystems). In Region III, aspects of soil biota, nutrient status, and/or biogeochemical processes may be highly altered, but P, C, and B properties and interactions are considered more complex and tightly coupled to aboveground community dynamics. (Modifi ed from Heneghanet al. 2008 , with permission from John Wiley & Sons.)

require more soil ecological knowledge and an resulting in new and self sustaining assemblages of explicit focus on plant–soil feedbacks (where a plant plant species that have never before coexisted and species or community alters soils in a way that perhaps a novel soil legacy. In these novel systems, impacts its own success as well as that of other spe- it is critical to recognize that the new communities cies). These plant–soil feedbacks can be a key mech- can provide ecosystem services (i.e. benefi cial func- anism by which species become invasive. Thus, tions, as defi ned by the Millennium Ecosystem ecosystems under dynamic change through inva- Assessment 2005 ), and attempts to restore these sion may contain high soil legacy that has become systems to a former state could result in failure and altered in subtle ways. Restoration of these systems compromise ecosystem services provided. may require considerable soil ecological knowledge Disturbance to the soil system has potential to to establish rare, historically dominant, or a diver- alter ecosystem services ( Daily et al. 1997 ) such as sity of species that depend on specifi c or historic effi cient and conservative nutrient cycling, soil for- soil properties, processes, and/or biotic composi- mation, and/or water holding capacity (to name tion (Heneghan et al. 2008 ; Eviner et al. 2010 ), par- only a few) that are provided by structurally hetero- ticularly if that system has attained a self-organizing geneous soils with a thriving biota. In the absence of (Suding et al. 2004 ). In no- restoration, soil structure and functioning, as well as analog environments, large shifts in the abiotic and ecosystem services, would remain in a degraded biotic components of an ecosystem have occurred, state, continue to decline, or recover slowly (Insam

00001497666.INDD001497666.INDD 337979 11/23/2012/23/2012 11:09:5411:09:54 PMPM OUP UNCORRECTED PROOF – FIRST PROOF, 01/23/12, SPi . . 2002 ) . . 2003 ); plant-dependent et al et al . . 2004 ). Carbon amendments have et al re required ( Franco & DeFaria 1997 ) ) 1997 & DeFaria Franco re required ( piled ( Mummey piled ( o time and/or re-development of macroag- land ( Paschke Paschke land ( c C and N in soil ( Kulmatiski & Beard 2006 ) ) 2006 & Beard Kulmatiski c C and N in soil ( particularly arbuscular mycorrhizae fungi in esources. Cutting followed by haying, cropping, cropping, Cutting followed by haying, esources. Marks 2007 ) and differentially affect recovery 2007 Marks communities of high conservation value ( Owen ( value communities of high conservation ver and promoted native perennial species al, and topsoil are used to increase organic matter al, P ( Walker Walker P ( ocesses, and/or biodiversity during restoration with and/or biodiversity during restoration ocesses, nd correlation with microbial biomass ( Andersen & nd correlation with microbial biomass ( ntent exhibit faster forest regrowth than those with round 6.5 contains the highest nutrient availability and round 6.5 contains the highest nutrient availability trient transformations during grassland restoration, with restoration, during grassland trient transformations tances to promote establishment of vegetation ( Ashby 1997 ) ) 1997 Ashby tances to promote establishment of vegetation ( m 1996 ) and biomass of coarsely defi ned microbial groups (i.e., based on ned microbial groups (i.e., and biomass of coarsely defi ers 2004 ); soil heterogeneity can decrease during restoration, soil heterogeneity can decrease during restoration, ); 2004 ers uminous tree species have been used to revegetate areas degraded uminous tree species have been used to revegetate areas degraded anges and successful response to manipulations; it should be recognized anges and successful response to manipulations; . . 2008 ) . . 2004 ) et al et al . . 2003 )

et al . . 2010 ) et al . . 2002 ) . . 2010 ) et al et al . . 1998 ; Bach et al . . 2010 ) et al . . 2009 ) et al . . 1999 ) ( Rowe Sparling 1995 ) and provide slow release nutrients in mined land ( Bradshaw & Chadwick 1980 ) ) 1980 & Chadwick Bradshaw and provide slow release nutrients in mined land ( micronutrients and K a gypsum, but supplements of rock phosphate, from mining in the tropics without addition of organic soils, of inert material has been used to reduce or dilute soil deep tillage and incorporation (ferric) sulfate, Al or Fe addition of Blumenthal N and non-target species ( been used to reduce available lowest metal toxicity ( Wong 2003 ) et al ) 2005 Lane & BassiriRad ants ( i.e. biota, particularly if it is imparted by transient as related t increases in response to establishing perennial vegetation on former cropland, phospholipid fatty acid biomarkers) Bach gregates ( of some plant species if topsoil is stock mycorrhizae) can be compromised with consequence for restoration soil organisms (e.g. rates of belowground ecosystem properties and processes during restoration ( Meyer ( of belowground ecosystem properties and processes during restoration rates higher sand content and nutrient leaching ( Lu higher sand content and nutrient leaching ( faster recovery occurring in soil with high clay content ( Baer faster recovery occurring in soil with high clay content ( restored grassland ( Jastrow Jastrow ( restored grassland Inoculating soil dominated by invasive species with soil derived from native plant communities can reduce invasive species co species with soil derived from native plant communities can reduce invasive Inoculating soil dominated by invasive Soil fauna have been used as indicators of recovery of other soil properties, e.g. recolonization of restored sites by ants a e.g. Soil fauna have been used as indicators of recovery other soil properties, Low levels of soil organic matter can result in inadequate nutrient supply to plants. Addition of organic waste, humus materi Addition of organic waste, Low levels of soil organic matter can result in inadequate nutrient supply to plants. Allelopathic compounds are often organic, and application activated C has been shown to effectively reduce extractable organi C has been shown to effectively reduce extractable and application activated Allelopathic compounds are often organic, often corresponds with low plant and by a few that monopolize r High nutrient availability Elemental S and pyritic peat have been used to lower pH of arable land to promote establishment of acid soil requiring plant Elemental S and pyritic peat have been used to lower pH of arable Flinn & uence plant community composition ( small-scale soil heterogeneity that may infl can dictate variation Microtopography of shrubs on mined Re-application of topsoil following mining aims to provide symbiotic microorganisms needed for restoration Manipulating soil heterogeneity can promote plant community heterogeneity ( Baer Manipulating soil heterogeneity can promote plant community ( Tropical forest regrowth is infl uenced by soil texture according to coarse soil classifi cations and soils with higher clay co uenced by soil texture according to coarse classifi forest regrowth is infl Tropical In soils degraded through cultivation, development of soil macroaggregates corresponds to soil microbial community recovery, development of soil macroaggregates corresponds to microbial community recovery, through cultivation, In soils degraded Knowledge or manipulation during ecological restoration. Knowledge or manipulation during ecological restoration. Knowledge or manipulation of soil properties associated with or used to promote revegetation, recovery of soil properties or pr Knowledge or manipulation of soil properties associated with used to promote revegetation,

Organic matter Huang & Cunningha of metals in contaminated soils ( Addition of organic chelators can enhance phytoextraction Nutrient status Leg soils may require nutrient inputs (fertilizer application) to support vegetation growth. Highly degraded pH soil with pH a In general, Soil pH affects plant growth through its regulation of nutrient solubility and metal toxicity. Heterogeneity Eijsack and fauna in contaminated sites ( ora Heterogeneous soils provide refugia for recolonization of soil fl Soil biota wSoil biota richness Total ). 1988 Insam & Domsch Microbial biomass increases in response to soil organic matter accrual ( Texture Texture and nu aggregate structure, microbial biomass, Soil texture modulates recovery of soil properties (C and N accrual), Structure Soil ripping has been applied in extreme circums ltration. infi Severe soil compaction limits root growth and water consequences for improving ecosystem functions and services listed in Table 5.6.1 . This list largely refl ects documented ch list largely refl This . 5.6.1 Table consequences for improving ecosystem functions and services listed in and inconsistent changes unsuccessful responses exist that failures commonly go unreported, Table 5.6.2 Table Soil property

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APPLYING SOIL ECOLOGICAL KNOWLEDGE TO RESTORE ECOSYSTEM SERVICES 381

& Haselwandter 1989 ). Restoration can facilitate residues, sewage sludge, etc.) are commonly added recovery or reinstate numerous ecosystem services to alleviate nutrients that are too limiting or to sus- in degraded lands and landscapes (Table 5.6.1 ) tain plant growth ( Bradshaw 1997 ; Wong 2003 ). largely through revegetation efforts, and establish- Knowledge of physico-chemical properties of soil ing key species that promote a specifi c ecosystem is required for restoration of extremely disturbed service ( Eviner & Chapin 2001 ). This chapter synthe- mined environments. Mining for coal and metal sizes how knowledge of soil processes, belowground ores results in the oxidation of sulphide minerals, heterogeneity, the roles of soil biota, the controls by and consequently, the severe acidifi cation of soil state factors, and potential feedbacks with the above- (pH 2–2.5; Bradshaw & Chadwick 1980 ; Bradshaw ground community can be applied to promote resto- 1997 ). Soil pH may be one of the most important ration success ( Table 5.6.2 ). properties to consider in mined because it infl uences the solubility of nutrients and 5.6.2 Low to high legacy: lessons from metal ions, activity of soil microorganisms and restoration of mined land fauna, and pH-sensitive processes such as nitrifi ca- tion and nitrogen fi xation ( van Breeman 2004 ). Soil ecological knowledge is an asset to restoring Remedying pH levels requires knowledge of soil severely disturbed ecosystems, particularly with acidity, the potential acidity that can arise from fur- respect to understanding the role of soil properties ther oxidation of metal sulfi des, and the acid neu- and plant–soil relationships that promote revegeta- tralizing capacity of the soil to determine effective tion. Mineral resources exist underground and their amounts of calcium carbonate (lime) to increase and extraction requires the removal of vegetation and maintain pH for successful restoration (Costigan soils, which can result in complete environmental et al. 1981 ). Waste materials produced from the min- destruction; however, a continuum of ecosystem ing for heavy metals can contain signifi cant amounts degradation from mining activity does exist. of residual metals, even after processing, with little Mine workings (materials produced by mining) to no potential for natural reduction (Li 2006 ). The generally contain factors that limit ecosystem devel- concentration and chemical forms of metals (as opment over the short term in the absence of recla- affected by pH) determine soil toxicity. Many heavy mation practices (Bradshaw 1997 ). In the extreme metal ions exhibit reduced solubility at low pH; case, deposition of waste rocks (i.e. host rock with however, the solubility (i.e. availability) of essential insuffi cient mineral amounts for extraction) and metals (e.g. molybdenum) decreases at high pH and mine tailings (i.e. remnant slurry post mineral results in metal defi ciencies ( van Breeman 2004 ). extraction) at the surface become sources of pollu- Organic acids derived from soil organic matter and tion, eliminate biodiversity, and drastically reduce root exudates represent important sources of chela- the production capacity of the environment. Resto- tors present at low levels in mined soils. Phytoex- ration of mine workings to an improved ecosystem traction of metals using hyperaccumulating plant state has elucidated that physically and chemically species, coupled with the addition of metal-chelat- altered soils are critical constraints to plant growth. ing compounds can enhance metal uptake and Transport of materials by heavy machinery inevita- removal to 1) render the soil more suitable for bly results in severe soil compaction, and mechani- growth of microorganisms that are inhibited by cal soil treatment (e.g. soil ripping) is commonly heavy metals (e.g. rhizobia), 2) promote develop- applied prior to revegetation to loosen compacted ment of symbioses important for plant growth (e.g. soil and promote root penetration ( Ashby 1997 ). legume-rhizobia), and 3) facilitate desired plant Mine waste materials can include weathered sub- communities ( Wong 2003 ). soils and overburdens with defi ciencies in mineral Heavy metals and persistent organic pollutants amounts or unweathered to the extent that nutri- can render soil toxic to belowground organisms ents cannot be released quickly enough to sustain and plants. Physicochemical properties of soil are plant growth. Thus, nutrient defi ciencies must be important for decontamination of organic pollutants, determined. Fertilizers or amendments (topsoil, which bind to soil constituents (e.g. clay, organic

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382 SOIL ECOLOGY AND ECOSYSTEM SERVICES

matter, organomineral complexes) over time. Once stock-piling) and absence of an associated plant contamination levels begin to decline, microbial community to sustain symbiotic microorganisms populations can recover. For example, Yin et al. during stock-piling imposed long-lasting effects on (2000 ) found that the arrival of bacterial species vegetation and the spatial organization of surface along a restoration gradient that included a mine mined land. Failure of native shrubs to establish in spoil, restored mined land, and undisturbed forest stock-piled and re-applied topsoil has consequences was dependent on time since disturbance. Refugia for root distribution, water relations, fi re cycles, and for soil organisms, provided by uncontaminated spatial resource heterogeneity in the restored mined soil microsites in physically heterogeneous soil, can lands. This study underscores the importance of serve as important sources of soil fl ora and fauna soil physicochemical properties and the soil micro- for recolonization ( Eijsackers 2004 ) that might be bial community to ecosystem restoration. In some necessary for some plants to establish. Delayed cases (e.g. shallow soil on steep terrain), topsoil arrival of microbial species could represent a con- conservation and replacement is diffi cult and mine straint to restoring plant diversity ( Harris 2003 ). spoils are substituted for topsoil, despite that their Recognition that soil health (vital whole soil) is properties and growth of therein can imperative for successful revegetation has resulted be different than native soils ( Showalter et al . 2010 ). in formulation of policy in many countries to con- serve and replace surface soils in mined sites. Mined land amended with topsoil can, in some cases, rep- 5.6.3 Moderate legacy: restoration of resent a rapid transition from a low to moderate or agricultural systems even high soil legacy starting point for restoration. Agriculture represents the most globally wide- Topsoil amendments provide biological or biologi- spread anthropogenic infl uence on ecosystems cally-produced soil components such as organic (Ellis & Ramankutty 2008 ). The degree of soil per- matter, stored nutrients, available nutrients, soil turbation through agriculture depends upon the biota, and plant propagules (Brenner et al. 1984 ). type of production system (e.g. annual crops, pas- Topsoil replacement, however, has resulted in var- ture, vs. timber) and historic management (e.g. till- ied restoration success and care must be taken in age and rotation regimes, fertilizer application, how it is managed. For example, if topsoil is added grazing intensity, and selective harvest, vs. forest to a dissimilar or compacted underlying material, clear-cutting). Restoration goals for agricultural then hydraulic discontinuity and instability can systems vary considerably, partly due to difference result (Bradshaw 1997 ). Depth of topsoil addition in historical communities (i.e. grassland, forest, or can also infl uence the trajectory of plant community wetland) that were converted to production sys- recovery. Paschke et al. ( 2003 ) reported that deep tems. Soil ecological knowledge is increasingly con- additions of topsoil encouraged grasses and forbs sidered to restore multiple ecosystem services in to outcompete mountain shrubs that are adapted to these systems, particularly those provided by the historically shallow soils. “Live handled” top- biodiversity. soil is preferred over stockpiled topsoil to provision restorations with symbiotic soil organisms. Preser- vation of topsoil through long-term stockpiling has 5.6.3.1 Restoration to grassland been shown to reduce soil organic matter (accord- ing to many of the same mechanisms as cultivation) Long-term conventionally cultivated systems gen- and be detrimental to symbiotic arbuscular mycor- erally contain no legacy of the historic plant com- rhizae fungi. These physical and biological changes munity and limited potential for colonization of to soil during long-term stockpiling, coupled with historic species from the regional due uniform re-application of the stored soil, has to few native propagules in landscapes dominated revealed important soil-related constraints to resto- by agriculture. It is well recognized that long-term ration. Mummey et al. (2002 ) documented that dis- cultivation degrades soil structure, promotes more turbance to soil structure (through removal and extreme wet–dry cycles, alters soil microbial com-

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APPLYING SOIL ECOLOGICAL KNOWLEDGE TO RESTORE ECOSYSTEM SERVICES 383

position, increases decomposition, and lowers soil nutrient enrichment. Manipulating has C and other nutrients to new equilibrium levels been explored as a restoration tool to increase plant (Dick 1992 ). Furthermore, long-term fertilized soils , reduce invasive or non-target spe- may exhibit high residual nutrient levels, high nitri- cies, and increase soil heterogeneity prior to, or dur- fi cation rates, and lower pH. In general, simple ing restoration. Walker et al. ( 2004 ) reviewed efforts goals such as revegetation to reduce erosion can be to reduce soil fertility in arable lands to re-create easily achieved and concomitant improvements in species-rich grasslands in Europe. Measures used to soil structure and functions can proceed passively. reduce phosphorus (P) availability include haying The development of perennial grass root systems in or cropping to promote off-take of P, addition of long-term conventionally cultivated soil can pro- aluminum (Al) or iron (Fe) (ferric) sulfate to increase mote soil macroaggregate formation, total microbial P adsorption capacity, and deep cultivation or addi- and fungal biomass, and soil C accrual in a decadal tion of inert or organic materials to dilute nutrient time scale ( Matamala et al . 2008 ), but recovery can pools. In the UK, nutrient reduction through deep vary drastically between highly contrasting soil tex- cultivation combined with seed addition has been tures (Bach et al . 2010 ; Baer et al . 2010 ). Paustian et al . shown to increase plant community similarity to ( 1998 ) reviewed the potential for agricultural sys- grassland targets with conservation value on a dec-

tems to mitigate increasing atmospheric CO2 and adal time scale (Walker et al. 2004 ). Carbon (C) addi- suggest that planting perennial vegetation such as tion can be an effective tool to reduce inorganic grass represents one of the most favorable scenarios nitrogen (N) availability by promoting the growth for increasing soil C stocks following land degrada- of the soil microbial biomass and immobilization of tion. Erosion reduces land and ability N, with signifi cant consequence for reducing non- to maintain or improve C stocks in biomass and target plant cover during grassland restoration soil; therefore, measures to reduce soil erosion are ( Baer et al. 2003 ; Blumenthal et al. 2003 ). Reducing N very important. Despite improvement in numerous availability at the onset of restoration may be a val- ecosystem services provided by simple erosion con- uable tool to prevent invasive species establish- trol measures (functional restorations), these prac- ment. However, the effectiveness of C additions is tices generally do not restore biodiversity and variable, and effi cacy of such manipulations should associated ecosystem services (e.g. genetic resources, be considered in the context of long-term temporal pollination, cultural heritage). dynamics of N availability modulated by develop- Restoration of some plant communities may ment of soil C stocks and increasing plant–soil feed- require specifi c soil conditions. In Europe, liming back over time ( Baer & Blair 2008 ). has been used to raise pH of naturally acidic soils to promote crop production; however, high soil pH limits restoration of grassland and heathland com- 5.6.3.2 Restoration to forest munities that prefer acidic soil conditions in these arable systems. Applications of elemental S and on agricultural land has generally pyritic peat have been used to effectively lower pH, employed fewer soil manipulations relative to promote establishment of species that require acidic restoring grasslands, but there are studies that dem- soil conditions, and reduce ( Owen onstrate soil properties affect on forest regrowth

et al. 1999 ). High residual soil fertility from nutrient and CO2 mitigation. Soil fertility and land-use his- management in agricultural soils can constrain the tory (as it has affected soil) are critical factors affect- restoration of diverse plant communities, particu- ing forest regrowth (Tucker et al . 1998 ). The tropics, larly if soil fertility results in asymmetric competi- in particular, are subject to a continuous cycle of for- tion for nutrients and dominance by a few species est clearing for agricultural purposes and socioeco- ( Baer et al. 2003 ). High species coexistence is gener- nomic constraints to maintaining the productive ally associated with low levels of soil fertility ( Jans- capacity of soil, which results in rapid soil degrada- sens et al. 1998 ), further supported by the consistent tion and further clearing for agriculture ( Lavelle phenomenon of declines in plant diversity under 1987 ). Soil C loss due to cultivation of tropical soils

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384 SOIL ECOLOGY AND ECOSYSTEM SERVICES

occurs at a much faster rate than in subhumid species distributions and the variable history of regions due to faster decomposition, and in moun- agricultural intensity and management among tainous regions exacerbated by erosion. Due to the sites. amount of C lost from forest land conversion to agriculture, Paustian et al . ( 1998 ) contended that the

most signifi cant opportunity for mitigating CO2 emissions is to reduce the rate of tropical land con- 5.6.3.3 Restoration to wetland version to agriculture. Change in soil C stocks in response to reforestation is not well documented Globally, over half of wetland area has been lost, for the tropics. Paul et al . ( 2010 ) found no change in most of which has been converted to agriculture. soil C pools in response to reforestation of rainforest Costanza et al. (1997 ) estimated that shallow waters species, but improvement in many other soil prop- which cover <2% of the Earth’s surface provide up erties (i.e., extractable inorganic N, plant available to 40% of global renewable ecosystem services; inorganic N, nitrifi cation rate, pH, and bulk den- thus, restoration of these systems has important sity) occurred on a decadal time scale. consequences for ecosystem services. Restoration of Our understanding of soil-related factors that wetland systems generally involves restoring or infl uence forest regrowth and/or composition on manipulating hydrology to promote recovery of former agricultural lands comes mostly from stud- ecosystem services such as disturbance regulation, ies of reforestation following land abandonment. water supply, sediment retention, and nutrient For example, differential rates of forest regrowth cycling (Table 5.6.1 ). However, the vast array of res- following agricultural land-use in the tropics have toration approaches coupled with variation in land- been attributed to a suite of soil properties associ- scape factors, disturbance regime, invasive species ated with different soil orders. Clay-rich Alfi sols pressure, historic seed banks, and nutrient supply hold more nutrients and support faster forest constrain predictability of wetland restorations to regrowth relative to Ultisols and Oxisols with achieve specifi c targets ( Zedler 2000 ). higher sand content and lower fertility ( Lu et al. Soil moisture, as affected by texture properties 2002 ). Over half of the forest cover throughout and hydrology, is probably the most frequently con- Europe and eastern North America occurs on former sidered soil-related factor in wetland restoration agricultural land (Vellend 2003 ). Residual impacts because it regulates biogeochemical processes and of cultivation on soil have been documented in sec- aboveground community structure. Wetting and ondary forest following 90–120 years of abandon- drying of soils affects nutrient dynamics. Nitrate ment from agriculture. Compton and Boone (2000 ) supplied by mineralization is stimulated by soil demonstrated that formerly cultivated secondary drying and removal of nitrate through denitrifi ca- forest sites contained less forest fl oor C, more min- tion requires temporary inundation to induce dis- eral soil N and P, lower C:N and C:P ratios, and similatory reduction of N conducted by facultative higher nitrifi cation rates relative to sites that were anaerobic microorganisms ( Venterink et al. 2002 ). selectively logged, with no cultivation disturbance Drained and cultivated agricultural soils are gener- to mineral soil. Furthermore, forest composition of ally considered to contain limited denitrifi cation abandoned agricultural lands can remain distinct potential due to aerobic conditions and depleted from forests that have never been cleared for agri- soil organic matter. Restoring wetlands has been culture for centuries, particularly with respect to promoted as a mechanism to reduce surface water herbaceous richness (Flinn & Marks 2007 ). Although nutrient loads through vegetation uptake and deni- the cause of compositional variation among second- trifi cation (Mitsch et al. 2001 ). However, Orr et al. ary forests on formerly cultivated soil and unculti- ( 2007 ) documented no change in actual and poten- vated soil has not been deduced, Flinn and Marks tial denitrifi cation rates in former agricultural soils (2007 ) speculated that soil properties and processes following cessation of agriculture and hydrologic play an important role, particularly in the reduction reconnection in a leveed fl oodplain. Thus, processes of microtopography, which infl uences small scale assumed to self-repair under certain conditions

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may not always develop from recreating the physi- ing species, 2) habitat, including soil-related factors cal template ( Hilderbrand et al. 2005 ). linked to facilitating or resisting invasion, 3) post- Water depth is a key controller of wetland struc- colonization effects of invasive species on a variety ture and function, and long-term drainage and till- of soil-mediated ecosystem processes, and 4) com- age of former wetland soils has led to organic matter plex interactions of all of these factors. There has loss and subsidence of soil (Verhoeven & Setter been a large effort to quantify how invasive plants 2010 ). In addition to hydrology, soil wetness and modify soil properties over the past two decades. variation in soil properties (e.g. texture, organic Recent studies have provided detailed information matter storage, and nutrient availability) are infl u- on the effects of invasive species (not only plants) enced by microtopography. Meyer et al. ( 2008 ) dem- on ecosystem productivity, decomposition, soil onstrated differential recovery rates of belowground nutrient dynamics, and soil food webs. Conse- structure and function in re-created sloughs con- quently, the prospect of developing a new array of taining slight topographic variation between the restoration tools to manage native communities or temporally inundated central channels of sloughs control invasive species is growing. and slightly elevated slough margins. Microtopo- Efforts to identify physiological or life history graphic variation also imparts variation in vegeta- traits that differentiate invasive from non-invasive tion composition (VivianSmith 1997 ). Thus, soil species, as well as highly invasible from less invasi- texture and moisture characteristics will need to be ble have been largely inconclusive. For considered (potentially re-created) to successfully instance, an analysis of 79 independent native ver- restore all species and assemblages associated with sus invasive plant comparisons revealed that invad- topographic transition. ers did not consistently have higher growth rates, Attaining biodiversity targets and associated competitive abilities, or fecundity ( Daehler 2003 ). In services through wetland restoration is generally fact, the success of non-native plants generally impeded under nutrient enrichment, as species depended on the growing conditions, illustrating richness is commonly low where nutrient supply is the importance of habitat and often soil-based fac- high (Green & Galatowitsch 2002 ). Managing nutri- tors infl uencing the invasion process. One particu- ents will require active consideration of inputs from larly infl uential hypothesis on habitat factors the landscape, as well as soil properties and proc- facilitating invasion has been the “fl uctuating esses that modulate nutrient availability through resource hypothesis” ( Davis et al. 2000 ). This adsorption and microbial transformations, respec- hypothesis suggests that plant invasion depends on tively. There can be a great disparity between recov- soil resource supply rates augmented by the avail- ery rates of vegetation and soil properties during ability of propagules of the invasive organism. wetland restoration ( Craft et al. 1999 ), which may Although there is some support for this hypothesis impose temporal constraints on realizing ecosystem ( Foster & Dickson 2004 ) other studies have been services that in-tact wetlands perform. either less emphatically supportive (Kercher et al. 2007 ) or conclude the contrary (Walker et al. 2005 ). 5.6.4 High legacy under dynamic change: The search for patterns of habitat susceptibility to preventing invasion and restoring invasion has also examined whether species-rich or invaded systems low resource environments are more resistant to invasion, but this potential mechanism has not been Invasion of natural and managed lands by new, supported consistently ( Lonsdale 1999 ; Funk & undesirable species can decrease diversity and Vitousek 2007 ). Thus, a variety of life history strate- impair ecosystem services provided by diverse gies can be associated with invasive species and communities (e.g. effi cient nutrient cycling, pollina- many habitat types are susceptible to invasion, tion, refugia, genetic resources, recreation, and cul- which limits the development of generalities about tural heritage). Efforts to understand biological the mechanisms driving biological invasion. invasion into habitats of biodiversity conservation Intimate knowledge of local systems and their concern have largely focused on 1) traits of invad- potential invaders remain essential to preventing

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invasion or restoring invaded environments. Such establishment of historical disturbance regimes (e.g. knowledge forms the basis for developing and fi re) may conserve and enhance native biodiversity implementing management. For instance, the inva- ( Hobbs & Huenneke 1992 ), there has been growing sion and establishment of Alliaria petiolata (garlic concern that methods exclusively targeted at physi- mustard), an herbaceous biennial prevalent in East- cal removal of an invader can leave a habitat vul- ern and Midwestern woodlands, is sensitive to both nerable to rapid reinvasion (Iannone & Galatowitsch woodland density and site fertility, but light avail- 2008 ). Furthermore, invasive species that share sim- ability appears to be the most important factor ilar physiological traits and response to manage- affecting the proliferation of this species (Meekins ment as native species can pose a real dilemma for & McCarthy 2000 ). Other biennial weeds in these managers ( Reed et al . 2005 ). systems, including Dipsacus sylvestris (common teasel) and Barbarea vulgaris (garden yellowrocket), fl ourish in response to soil disturbance ( Roberts 5.6.4.1 Using soil ecological knowledge to 1986 ). Managers can use these specifi c understand- control invasion ings to protect systems from a suite of potential invaders (e.g. managing for high woodland density On a large scale, there has been less active manage- with low light levels, coupled with minimal soil ment of soil properties and/or processes to prevent disturbance). Management based on well-under- or reduce biological invasion relative to active man- stood ecological traits of invasive species, potential agement of vegetation ( Callaham et al . 2008 ). Sev- novel invaders in the regional species pool, and eral studies have quantifi ed the effects of invaders environmental conditions are needed to predict and on soil properties and processes, and a few studies prevent invasion. have manipulated soil to reduce invasive species. Conservation programs should prioritize the pro- Because soil and plants interact, a modifi cation to tection of sites with high legacy of native biota, par- the plant community, such as invasion and domi- ticularly if restoring highly invaded areas is not cost nance by a new species, is expected to change soil effective and probability of success is low (Hobbs & conditions (Wardle et al . 2004 ; van der Putten et al . Harris 2001 ; Hobbs et al. 2009 ). For example, Chicago 2009 ). Common impacts of invaders on soil include Wilderness is a consortium of over 250 conserva- alteration to microbial communities and decompo- tion-oriented organizations that prioritizes habitat sition rates with consequence for nutrient supply conservation and management in exactly this man- ( Corbin & D’Antonio 2004b ; Belnap et al . 2005 ), and ner. Traditional management of areas retaining these changes may persist even after removal of some residual biodiversity has been to cut and invasive species (Ehrenfeld & Scott 2001 ). There is remove invasive plants, but other trophic levels that limited knowledge about the passive recovery of may interact with invasion processes are typically soil physical, chemical, and biological components not managed, such as non-native earthworms that following removal of invasive species. If feedback impact soil processes in ways that can have conse- processes develop, there is potential for the system quence for plant composition ( Heneghan et al. 2006 ). to persist in a self-organizing alternative stable state The success of invasive species removals is largely ( Suding et al . 2004 ). determined by the degree of invasion, as the “early Development of restoration strategies that amel- detection, rapid response” method to controlling iorate soil conditions modifi ed by invasive species invasive species is generally most effective. Ecosys- may be needed to successfully increase desired spe- tem changes resulting from invasion increase with cies. Invasive plants may benefi t themselves by time since invasion; therefore, restoration in the altering soil biota, soil structure, amount and qual- early stages of invasion is likely to be more success- ity of organic matter, form of available nutrients ful because there is less modifi cation to the historic (e.g. nitrate vs. ammonium), and/or by adding alle- system (or higher legacy) relative to long after an lochemicals to the soil. Soil restoration techniques invader has established ( Strayer et al. 2006 ). that mediate these changes to soil include: using Although removal of invaders followed by the re- plant species that reverse invader-cultured soil

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conditions, adding microbial-containing inoculums invasion, 4) quantify the impacts of invasion on bio- to soil, and adding charcoal to bind allelochemicals logical communities and soil, and 5) develop resto- ( Eviner et al. 2010 ). Kulmatiski and Beard ( 2006 ) ration tools that target reducing invasive species added 1% activated C to soil dominated by two and promoting desired communities, coupled with exotic species and found reduced extractable evaluating the effi cacy of those tools. Finally, report- organic C and N coupled with consistent shifts in ing the failures of soil-related manipulations to the community composition (reduction in cover of two management of invasive species can be just as valu- target exotic species and an increase in overall cover able as the knowledge that can be gained from res- of native species). Despite these promising results, toration success. not all native species responded positively and other (non-target) exotic species increased in cover. 5.6.5 Novel legacy: no-analog This example does illustrate the potential utility of ecosystems and environmental soil-based tools to reduce invasive species. conditions A common observation is that invasive plant spe- cies enhance the availability of limiting nutrients, N There is increasing recognition that restoring eco- and P in particular ( Ehrenfeld 2003 ). Numerous systems of the past may not be feasible under rapid methods, similar to ones used in restoring arable and widespread anthropogenic-driven changes to land, are used to reduce nutrient availability. Car- climate, land-use, disturbance regimes, and nutri- bon amendments (e.g. mulch, sawdust, and sugar ent deposition, as well as human-accelerated inva- additions) can be effective at reducing invasive spe- sion of non-native organisms and corresponding cies and promoting plant diversity. However, some loss of diversity ( Vitousek et al . 1997 ). These envi- studies found that benefi ts are short lived and that ronmental changes have led to the development of strategies targeted at manipulating propagule avail- novel ecosystems, defi ned as self-perpetuating ability were more effective (Morghan & Seastedt communities that contain no historic analog in 1999 ; Corbin & D’Antonio 2004a ). Rowe et al. ( 2009 ) terms of species composition and potential func- compared the effi cacy of nutrient reduction through tions (Fig. 5.6.1 , Scenario III) (Hobbs et al . 2009 ). sucrose application to inoculation of invaded com- Novel ecosystems represent a self-sustaining stable munities with soil from uninvaded communities state under new biotic and abiotic conditions. They and found that both tools reduced the focal invasive are often characterized by exotic (either actively or species’ cover and increased perennial native spe- formerly invasive) assemblages of species that have cies cover; however, nutrient reduction also never before coexisted. Because invasive plants increased non-native annual/biennial cover. commonly alter soils in ways that benefi t them- Although the tool box of soil-related approaches selves (Kulmatiski & Kardol 2008 ), they can facili- to reducing invasion is growing, there is not an tate the creation of a new stable state and partly immediate prospect of applicability, particularly on explain the of some novel ecosystems. a large spatial scale. Because invasive species can The development of novel ecosystems can be due alter many aspects of soil, the most promising man- to 1) shifts in abiotic conditions, which then drive agement option will vary with invader. Confl icting biotic changes, 2) shifts in the biotic community, outcomes of soil-based restoration tools to control which then alter abiotic conditions, or 3) interac- invasive species limit the ability to make general tions of biotic and abiotic changes (Hobbs et al. recommendations. To successfully manage ecosys- 2009 ). Increased N deposition in coastal sage scrub tems undergoing dynamic change through the inva- habitats of southern California has driven major sion of undesirable species it will be prudent to: shifts in the biological community, with conse- 1) study invasion on a local scale, 2) acquire com- quences for ecosystem structure and functioning. prehensive information on life history strategies Increased N inputs through deposition has increased and resource requirements (including interactions productivity, fuel loads, and fi re frequency, which with belowground fl ora and fauna) of local invad- has resulted in the conversion of native shrubland ers, 3) know the environmental conditions prior to to grasslands dominated by exotic species (reviewed

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in Fenn et al. 2003 ; Fenn et al. 2010 ). Even with efforts toric community composition, it does enhance eco- to control exotic grasses and planting native spe- system function and resilience of the system to cies, these grasslands persist over the long-term, drought. Conversely, Bai et al. ( 2010 ) reported that and represent a new stable state (Cox & Allen 2008 ). N addition to non-degraded perennial grasslands Wolkovich et al. ( 2010 ) reported that this grass-dom- in this region can lead to dominance by annual, inated system was nine times more productive than early successional species that may not establish in the native system, has lower erosion rates, and a a drought, and leave the system susceptible to mas- 1.4-fold greater C storage in soil and litter, which sive erosional losses and potentially desertifi cation has shifted the system from a variable C source to after a large windstorm. a C sink. In these novel systems, manipulations to soil Shifts in biotic composition can strongly affect structure, chemistry, and/or biota could be key to abiotic components of the ecosystem. In fact, the retaining some presence of historic species, but res- direct impacts of environmental changes on ecosys- toration to historic communities in most cases tem processes are often small compared to the indi- would require extreme modifi cation, potentially to rect ecosystem effects mediated by changes in the the extent of ecosystem degradation. Effective res- plant community (Chapin 2003 ). For example, in toration of novel ecosystems will require knowl- Western , clearing of native vegetation for edge of factors that caused state changes (Beisner agriculture leads to a shift from vegetation with et al. 2003 ). Threshold changes in system states are deep roots and high evapotranspiration rate, to more likely to occur in ecosystems with strong agricultural species that have shallower roots and interactions (Suding & Hobbs 2009 ). Thus, under- use less water. Low evapotranspiration rate in the standing abiotic and biotic interactions is critical to system dominated by agricultural species leads to a predict and manage state changes. Restoration of a rise in the water table and salinization of the soil novel system to some past condition may not be surface (with negative consequences for crop pro- feasible (or reasonable) if the environmental change duction). These abiotic shifts, caused by conversion needed to reverse a state change exceeds the envi- to agriculture, also limit which plant species can ronmental change that initiated the state change, a thrive in the surrounding (George phenomenon known as “hysteresis.” For example, et al. 1997 ). Furthermore, the saline soils can contain successful restoration of coastal wetlands may 2.5-fold lower soil organic C stocks, lower soil N, require large decreases in salinity, because salinity decreased soil aggregation, and increased bulk den- tolerance of establishing seedlings is much less than sity (Wong et al. 2008 ). Ecosystem transformations that of mature wetland plants ( Zedler 2005 ). such as these can also be mediated by species extinc- Key factors that contribute to the resilience/ tions, invasion of exotic species, and even shifts in recoverability of an ecosystem include regulation of dominant native species ( Hobbs et al. 2009 ). abiotic conditions (including nutrient supply), State changes can also be driven by an interaction regional/propagule species pool diversity, and between shifts in biotic and abiotic changes. For landscape connectivity. All of these factors must be example, 80–87% of Inner Mongolia’s grasslands considered to conserve and restore communities are undergoing desertifi cation due to , and ecosystem services. The role of regulating abi- which causes shifts in vegetation composition and otic conditions has proven critical for restoration of exacerbates drought ( Li et al. 2000 ). Plant composi- southern California salt marshes. In these systems, tion shifts resulting from overgrazing in Inner Mon- small patches cleared for restoration have been suc- golia’s grasslands leads to decreased plant diversity cessful, but larger-scale restoration efforts have and cover, increased erosion, decreased soil C and been unsuccessful because large expanses of bare nutrients, and increased aridity at a larger scale ( Li ground heat up and lead to high evaporation, high et al. 2000 ). Nitrogen additions to these degraded levels of salinity, and minimal plant establishment grasslands can enhance the prevalence of perennial ( Zedler 2005 ). Securing ecosystem functions will drought-tolerant species. Although N addition, in require a diversity of genotypes and species toler- this circumstance, does not necessarily restore his- ant of harsh abiotic conditions in the regional or

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restoration pool of propagules (Eviner & Chapin microbial populations, and nutrient cycling 2001 ). Finally, proximity and/or connectivity to in the trajectory of a target state within dec- remnant patches across the landscape can be critical ades, but full recovery is anticipated to take to maintaining abiotic conditions, providing much longer. resources and species through migration and prop- 2. Inconsistent patterns in plant community agule dispersal, and mediating disturbance regimes recovery are commonly attributed to soil critical to the maintenance of a system ( Bengtsson properties and/or processes altered through et al. 2003 ). disturbance, which can persist in some cases Rapidly changing environmental conditions may for over a century. necessitate shifting some restoration goals to main- 3. There is more knowledge about singular taining or reinstating ecosystem services. Restora- soil-related constraints to restoration than tion and management to some historic plant complex soil or plant-soil interactions. For communities could, in fact, promote species and example, soils with high nutrient availabil- communities that can no longer persist without ity typically support species-poor plant intense management (and elimination of the plants assemblages. Direct physical and chemical that can), potentially leading to a collapse in the pro- (reduction in P availability through dilution vision of multiple ecosystem services ( Hobbs et al. or adsorption) and indirect biological (car- 2009 ). Although novel ecosystems have received bon addition to increase microbial demand negative attention because they are often dominated for N) methods used to reduce nutrient by non-native species, in some scenarios, these com- availability can be effective. Similarly, pH munities have and will represent the most effective can be manipulated to alter metal toxicity to way to provide ecosystem services in greatly altered promote revegetation or promote restora- environmental conditions (Ewel & Putz 2004 ). tion of communities adapted to specifi c pH conditions. 4. The proverbial “black box” of species and 5.6.6 Conclusions their interactions with physical and chemi- cal properties of soil truly demonstrates that The importance of soil ecology in the science and a whole soil connected to the aboveground practice of ecological restoration has been repeat- community is greater than the sum of all the edly recognized ( Bradshaw 1999 ; Young et al . 2005 ; parts. This is evidenced by attempts to pre- Heneghan et al. 2008 ). To synthesize the role of soil serve vital topsoil for restoration, but dis- ecological knowledge in guiding restoration science covery that displacement, disconnection and practice is challenging because failures are (from plants), and replacement alters prop- commonly unreported and most published studies erties of soil enough to constrain restoration on ecological restoration refl ect only a brief excerpt of key species. (temporally and spatially discrete) from a continu- 5. Where substantial elements of the native ous process that inherently involves change over biota persist in ecosystems undergoing time. Furthermore, it is diffi cult to identify generali- dynamic change through biological inva- ties across varying types and degrees of disturbance sion, it may be prudent to integrate knowl- among an array of ecosystems that are restored and edge of the soil-factors that may promote managed in unique ways for different purposes. invasion (i.e. disturbance or nutrient sup- This summary of restoration from the soil legacy ply) and invasive species impacts on soil continuum perspective elucidates the following: into management. 1. Slower recovery of soil properties and proc- 6. In highly invaded sites, where the native esses relative to plant components (cover biota is vestigial, managers may want to and productivity) is consistent across many consider amelioration of soil properties and types of restored ecosystems. Many studies processes affected by the invasive species. report improved soil structure, C stocks, Novel tools have been tested in attempt to

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ameliorate allelopathic effects of invasive tures: implications for carbon mitigation and grassland species on soil and supply microbial prop- conservation. Ecosphere 1 : art. 5. agules through inoculum additions, but they Bai, Y., Wu, J., Clark, C.M., et al. (2010) Tradeoffs and are limited thus far in knowledge of their thresholds in the effects of nitrogen addition on biodi- effectiveness across ecosystems and at a versity and ecosystem functioning: evidence from inner Mongolia Grasslands. Global Change Biology 16 : large spatial scale. 358–72. 7. Under changing environmental conditions, Beisner, B.E., Haydon, D.T., & Cuddington, K. (2003) traditional restoration targets may need to Alternative stable states in ecology. Frontiers in Ecology be reconsidered, particularly if restoration and the Environment 1 : 376–82. requires continued intensive management to Belnap, J., Phillips, S.L., Sherrod, S.K., & Moldenke, A. sustain species that are no longer suitable for (2005) Soil biota can change after exotic plant invasion: the environment or restoration will require does this affect ecosystem processes? Ecology 86 : modifi cation to the extent that will cause eco- 3007–17. system degradation and compromise services. Bengtsson, J., Angelstam, P., Elmqvist, T., et al . (2003) Novel ecosystems may maintain ecosystem Reserves, resilience and dynamic landscapes. Ambio 32 : services such as productivity, nutrient provi- 389–96. Blumenthal, D.M., Jordan, N.R., & Russelle, M.P. (2003) sion and retention, erosion control, soil C stor- Soil carbon addition controls weeds and facilitates prai- age and protection, as well as the infi ltration, rie restoration. Ecological Applications 13 : 605–15. storage, and provision of water. Bradshaw, A.D. (1997) Reclamation of mined land—using Despite many uncertainties, the increasing number of natural processes. 8 : 255–69. studies that have monitored or manipulated soil prop- Bradshaw, A.D. (1999) The importance of soil ecology in restoration science. In: K.M. Urbanska, N.R. Webb, P.J. erties and processes (successfully or not) tangibly Edwards (eds.) and Sustainable Devel- achieve the ultimate goal of restoration ecology, which opment , pp. 33–64. Cambridge University Press, is to use ecological knowledge to steer restoration Cambridge. practice and test our basic understanding of ecology. Bradshaw, A.D., & Chadwick, M.J. (1980) The Restoration of Land: The Ecology and Reclamation of Derelict and Degraded References Land . Blackwell Scientifi c Publications, Berkeley, CA. Brenner, F.J., Werner, M., & Pike, J. (1984) Ecosystem devel- Andersen, A.N., & Sparling, G.P. (1995) Ants as indicators opment and natural succession in surface coal mine rec- of restoration success: relationship with soil microbial lamation. Minerals and the Environment 6:10–22. biomass in the Australian seasonal tropics. Restoration Callaham, M.A., Rhoades, C.C., & Heneghan, L. (2008) A Ecology 5 : 109–14. striking profi le: soil ecological knowledge in restoration Ashby, W.C. (1997) Soil ripping and herbicides enhance management and science. Restoration Ecology 16 : 604–7. shrub restoration on strip mines. Restoration Ecology 5 : Chapin, F.S., III. (2003) Effects of plant traits on ecosystem 169–77. and regional processes: a conceptual framework for pre- Bach, E.M, Baer, S.G., Meyer, C.K., & Six, J. (2010) Soil texture dicting the consequences of global change. Annals of affects microbial and structural recovery during grassland Botany 91 : 455–63. restoration. Soil Biology & Biochemistry 42 : 2182–91. Compton, J.A., & Boone, R.D. (2000) Long-term impacts of Baer, S.G., & Blair, J.M. (2008) Grassland establishment agriculture on soil carbon and nitrogen in New England under varying resource availability: a test of positive forests. Ecology 81 : 2314–30. and negative feedback. Ecology 89 : 1859–71. Corbin, J.D., & D’Antonio, C.M. (2004a) Can carbon addi- Baer, S.G., Blair, J.M., Collins, S.L., & Knapp, A.K. (2003) tion increase competitiveness of native grasses? A case Soil resources regulate productivity and diversity in study from California. Restoration Ecology 12 : 36–43. newly established tallgrass . Ecology 84 : 724–35. Corbin, J.D., & D’Antonio, C.M. (2004b) Effects of exotic Baer, S.G., Blair, J.M., Collins, S.L., & Knapp, A.K. (2004) species on soil nitrogen cycling: implications for restora- Plant community responses to resource availability and tion. Weed Technology 18 : 1464–7. heterogeneity during restoration. Oecologia 139 : 617–29. Costanza, R., dArge, R., Farber, S., et al . (1997) The value of Baer, S.G., Meyer, C.K., Bach, E.M., Klopf, R.P., & Six, J. the world’s ecosystem services and . (2010) Contrasting ecosystem recovery on two soil tex- Nature 387 : 253–60.

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