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Ecosystem services are typically valued for their immediate material or cultural benefits to human wellbeing, supported by regulating and supporting services. Under climate change, with more frequent stresses and novel shocks, 'climate adaptation services', are defined as the benefits to people from increased social ability to respond to change, provided by the capability of ecosystems to moderate and adapt to climate change and variability. They broaden the ecosystem services framework to assist decision makers in planning for an uncertain future with new choices and options. We present a generic framework for operationalising the adaptation services concept. Four steps guide the identification of intrinsic ecological mechanisms that facilitate the maintenance and emergence of ecosystem services during periods of change, and so materialise as adaptation services. We applied this framework for four contrasted Australian ecosystems. Comparative analyses enabled by the operational framework suggest that adaptation services that emerge during trajectories of ecological change are supported by common mechanisms: vegetation structural diversity, the role of keystone species or functional groups, response diversity and landscape connectivity, which underpin the persistence of function and the reassembly of ecological communities under severe climate change and variability. Such understanding should guide ecosystem management towards adaptation planning.
The original publication is available at: Global Change Biology 21, 1231 (2015)
Ecological mechanisms underpinning climate adaptation services
Sandra Lavorel1*, Matthew J. Colloff2, Sue McIntyre2, Michael D. Doherty2, Helen T. Murphy3, Daniel J. Metcalfe4,
Michael Dunlop2, Richard J. Williams5, Russell M. Wise2, Kristen J. Williams2
1Laboratoire d’Ecologie Alpine, UMR 5553 CNRS Université Joseph Fourier, BP 53, 38041 Grenoble, Cedex 9, France
2CSIRO Ecosystem Sciences, Canberra, ACT 2601, Australia
3CSIRO Ecosystem Sciences, Atherton, Qld 4883, Australia
4CSIRO Ecosystem Sciences, Dutton Park, Qld 4102, Australia
5CSIRO Ecosystem Sciences, Winnellie, NT 0822, Australia
*Corresponding author. E-mail:
Ecosystem services are typically valued for their immediate material or cultural benefits to human wellbeing,
supported by regulating and supporting services. Under climate change, with more frequent stresses and novel
shocks, ’climate adaptation services’, are defined as the benefits to people from increased social ability to respond
to change, provided by the capability of ecosystems to moderate and adapt to climate change and variability. They
broaden the ecosystem services framework to assist decision makers in planning for an uncertain future with new
choices and options. We present a generic framework for operationalising the adaptation services concept. Four
steps guide the identification of intrinsic ecological mechanisms that facilitate the maintenance and emergence of
ecosystem services during periods of change, and so materialise as adaptation services. We applied this framework
for four contrasted Australian ecosystems. Comparative analyses enabled by the operational framework suggest
that adaptation services that emerge during trajectories of ecological change are supported by common
mechanisms: vegetation structural diversity, the role of keystone species or functional groups, response diversity
and landscape connectivity, which underpin the persistence of function and the reassembly of ecological
communities under severe climate change and variability. Such understanding should guide ecosystem
management towards adaptation planning.
Keywords: climate change adaptation, ecosystem service, fire, functional diversity, functional traits, landscape
configuration,littoral rainforest, livestock grazing, Murray-Darling Basin, resilience
Ecosystem services (ES) are defined as the benefits that humans derive from ecosystems. They have become
prominent in informing natural resource management and policy (Perrings et al., 2011; Crossman et al., 2013).
Practice of ecosystem service valuation commonly focuses on supply of immediate, direct benefits for human
wellbeing - provisioning and cultural services (Millennium Ecosystem Assessment, 2005) - often without
considering underpinning regulating and supporting services (Abson & Termansen, 2011). However, shifts towards
maintaining regulating and supporting services are increasingly advocated given the threats posed by climate
change to the continued supply of provisioning and cultural services. Such shifts are exemplified by the
development of adaptation responses to climate change (Prober et al., 2012), including increased investment in
ecosystem-based adaptation (EBA) (World Bank, 2010).
Ecosystem-based adaptation approaches ‘harness the capacity of nature to buffer human communities against
the adverse impacts of climate change through the sustainable delivery of ecosystem services’ and include ‘the
potential for natural infrastructure to provide. . . disaster risk reduction, food security, sustainable water
management and livelihood diversification’ (Jones et al., 2012). Examples include local economic benefits from
supply of nontimber products from tropical forests, regulation of local and regional climate and hydrology,
mitigation of riparian erosion and coastal protection from storms (Jones et al., 2012; Pramova et al., 2012; Arkema
et al., 2013). In most cases, the benefits are incremental, focused on proximate causes of vulnerability, and
assessments are based on assumptions that ecosystem characteristics and services, and societal preferences for
these, will remain largely unchanged. However, empirical data and modelled projections indicate ecosystems and
landscapes will undergo fundamental, unpredictable changes in struc- ture, composition and functions in response
to novel, global-scale warming of >+3 °C (Schellnhuber et al., 2012). Therefore, applications of the ES concept that
assume stationarity of current bundles of ES that support livelihoods and economies, and societal preferences for
these, will likely lead to maladaptive future actions (Stafford Smith et al., 2011).
The concept of climate adaptation services (hereafter ‘adaptation services’) can be proposed to complement
the ecosystem services approach and help people develop choices for adaptation to climate change. This concept
highlights the prospect of substantial ecosystem change and stresses the importance of option and insurance
values of services not currently considered important for human wellbeing, but which may prove critical in the
future. Societal responses to climate change will vary from short-term incremental management aimed at
maintaining existing ecosystem characteristics, through to transformational change in values, rules and knowledge
that promote resilience and resistance of novel ecosystems. In this context, adaptation services are defined as the
benefits to people from increased social ability to respond to change, provided by the capacity of ecosystems to
moderate and adapt to climate change and variability. Adaptation services include the buffering capacity of
ecosystems against change and incorporate some currently valued ecosystem services such as coastal protection
or crop diversification for food security. Also, the capacity of ecosystems to transform in composition, structure
and function in response to climate change or new management regimes will result in new provisioning services
such as firewood supply after grassland has transitioned to woodland. Such new services form the basis for choices
and options to support social climate adaptation.
To operationalize the adaptation services concept we need to explore its applicability and usefulness across a
range of social-ecological systems. Here, considering the ecological dimensions of this endeavour, we propose a
methodological framework to reveal testable hypotheses on biological traits and ecological mechanisms that
underpin adaptation services, and to facilitate comparison and synthesis. To illustrate this capability, we present
one case study in detail and three in summary for Australian ecosystems that are vulnerable to climate change. We
synthesize common features of these case studies that relate to the main types of adaptation services and their
trait-based mechanisms. We identify knowledge and research gaps in the operational framework and conclude by
identifying the implications for management.
The operational framework
The operational framework involves the identification of adaptation services under different scenarios of climate
and management change, based on our understanding of the ecological mechanisms that underpin the supply of
ecosystem services (Bennett et al., 2009; Luck et al., 2009) (Fig. 1). Climate, other abiotic factors and management
are proximate drivers that determine the ecosystem state at a particular time and place. Each state has specific
characteristics of biodiversity and functioning that contribute to the supply of ecosystem services.
Scenarios may involve change of ecosystem state due to direct climate impacts, including increased
temperatures, droughts and storms, and indirect impacts via management responses such as changed grazing
regimes or water flows in regulated rivers. Adaptation services can then be identified that support human well-
being under these new conditions. Finally, ecosystem properties that underpin adaptation services need to be
managed for, including, where possible, the conservation of existing biodiversity and ecosystem services. The four
steps in the operational framework are outlined below.
Step 1: Initial system characterization
1a: Characterization of ecosystem dynamics under historical and current climate allows projections of future states
and properties. Appropriate models may be conceptual or quantitative (Scheffer, 2009), state-and-transition
(McIntyre & Lavorel, 2007; Zweig & Kitchens, 2009), or succession-based (Noble & Slatyer, 1980; Dickie et al.,
2011). Alternative states may be represented by vegetation composition and associated biodiversity, biomass
pools, ecosystem functions and disturbance regimes.
1b: For each ecosystem state, identification of bundles of ecosystem services supplied under current climate
helps define benefits desirable for the long-term. We emphasize bundles of services (Bennett et al., 2009;
Crossman et al., 2013), rather than single services, because bundles covary though space and time, dependent on
shared ecosystem properties, processes and environmental drivers. Thus, management approaches need to
consider functional linkages between all adaptation services.
Step 2: Climate change effects
To identify adaptation services (Step 3), it is necessary to describe and, where possible, quantify biodiversity and
ecosystem responses to climate change scenarios, including recent empirical or modelled trends, like changed
frequency and magnitude of extreme events. Scenarios integrate direct climate impacts and indirect effects of land
use and management (Fig. 2, inset diagrams).
Ecosystem responses involve changes affecting supply of current ecosystem services and autonomous
ecological adaptation: community structure, composition, functional groups and traits, plus landscape changes
that affect ecosystem responses and persistence. Methods for characterization include the assessment of altered
supply of ecosystem services under climate- and land-use change by building on state-and-transition models as
done for rangelands (Havstad et al., 2007), or alpine grasslands (Lamarque et al., 2014). In the latter, vegetation
states-and-transitions(Quétier et al., 2007) were linked to plant and microbial functional traits-based models
(Lavorel et al., 2011; Grigulis et al., 2013) to project effects of climate and management change on ecosystem
service bundles.
Step 3. Adaptation services
Identification of adaptation services. Ecosystem properties that facilitate societal climate adaptation by
supporting current ecosystem service bundles, supplying novel services and moderating or enabling ecological
transformation are identified as adaptation services. These different functions contribute to the different facets of
social adaptation: providing time for societies to change, slowing down ecological responses to climate change, or
providing novel livelihoods. In contrast with authors who determined adaptation services a priori, subsequently
relating them to ecosystem characteristics (Jones et al., 2012; Pramova et al., 2012), we take a bottom-up
approach by identifying intrinsic ecological mechanisms required to support future bundles of ecosystem services.
Identification of mechanisms underlying adaptation services. Ecological mechanisms and processes that
support adaptation services and their management include traits of organisms, biodiversity effects on
biogeochemical cycling and biotic moderation of resource availability, and landscape properties. Assessment of
functional diversity and redundancy of relevant traits is required to anticipate ecosystemservice responses to
climate change, and how species range shifts may affect service supply.
Step 4. Management of adaptation services
Changes in ecosystem properties and supply of adaptation services are likely to be emergent, requiring an adaptive
management approach. Management may target abiotic drivers such as water flows and disturbance regimes, or
biotic components including keystone taxa and functional groups, through conservation, restoration and
translocation. Intervention objectives should be to retain the ecosystem properties that supply those ecosystem
services that support societal adaptation to climate change.
Four case studies
In this section, we apply the operational framework to four case studies from climate-sensitive Australian social-
ecological systems representing different climatemanagement interactions related to ecosystem service supply
(Fig. 2). We provide a detailed account for grassy woodlands and summaries for littoral rainforests, temperate
montane forests and floodplain ecosystems.
Grassy eucalypt woodlands in south-eastern Australia
Box 1 Five alternative states identified by the stateand-transition model for grassy woodlands of southeastern
Australia (based on McIntyre & Lavorel, 2007; McIntyre, 2008)
Reference: Our understanding of pre-European vegetation relies on historical accounts depicting well-spaced
trees, tall thick grass with abundant forbs and variable, but often low, shrub densities (Gammage 2011). Available
water, hunting and dingoes would have regulated marsupial grazing.
Native pasture: Permanent settlement and introduced domestic livestock (sheep and cattle) increased grazing
pressures and transformed the composition of the grassland to shorter, more-grazing tolerant native grasses and
forbs. While still perennial-dominated, the annual component of the vegetation (native and exotic) is higher owing
to the presence of livestock. Native plant diversity (of grazing-tolerant species) is high, and trees and shrubs
generally persist, with some impacts of browsing.
Fertilized pasture: The addition of nutrients through fertilizers and legume N-fixation, elevates nitrogen and
phosphorus and allows higher grazing pressures than in native pastures. Exotic and native perennial grasses can be
maintained with grazing management, but annual grasses and forbs are abundant and often dominate. Native
plant diversity is low, and conditions for tree regeneration are poor.
Sown pasture and crops: Cultivation and fertilization, with the addition of legumes and sown grasses or crop
species result in almost complete loss of native diversity and perennial species.
Enriched grassland: Any fertilized or otherwise nutrient enriched grassland that is not grazed. Common
situations are roadsides or pastures fenced for tree establishment. Large perennial grasses dominate,
accumulating thick litter with a very low diversity of forbs.
Step 1a: Ecosystem description. Temperate grassy woodlands occupying subhumid regions of eastern Australia, are
associated with soils of inherently low fertility (McIntyre, 2011), and since European settlement have been
modified by livestock grazing, tree clearing and cropping. Transformation to exotic plant dominance was hastened
in the mid-20th century though the use of phosphorus fertilizers and introduced annual legumes, encouraging the
replacement of perennial native species with annual exotics (Dorrough et al., 2006, 2011). Crops and fertilized
pastures have reduced the regenerative capacity of remaining trees. Soils are prone to erosion, acidification and
salinization (Hobbs & Yates, 2000).
Grassy woodlands have characteristics that make them resistant to warming and drying, owing to the highly
perennial, stress-tolerant nature of their native plants. Some climatic adaptability may be found in the flora as the
geographical range of many species spans large rainfall and temperature gradients (McIntyre, 2011). The flora is
also stress-tolerant and slow-growing as an adaptation to naturally low soil fertility, while low pH and poor soil
structure make large areas vulnerable to erosion. These features of the ecosystem make it vulnerable to
agricultural impacts that lead to losses of perennial native vegetation and displacement by ruderal species under
conditions of disturbance and enrichment.
Five major ground-layer vegetation states are recognized (McIntyre & Lavorel, 2007; McIntyre, 2008; Fig. 3; Box
1). Reference and native pasture states have high resistance to grazing and fire, which maintain diversity, but low
resistance to fertilization. Transformation from native to exotic dominance occurs when fertilization raises soil
phosphorus above ~20 mg kg-1 (Dorrough et al., 2006; McIntyre, 2008). Established trees persist, but are
vulnerable to insect attack, root damage and disrupted regeneration. Decline of trees and shrubs is associated with
decline in fauna (Lindenmayer, 2011).
Dominant perennial tussocks are the keystone plant functional group in the ground layer, with the greatest
influence on ecosystem properties and processes (Mokany et al., 2008). Fertilization disrupts the tussock matrix
and diverse forbs are replaced with exotics. Available phosphorus declines over decades following cessation of
fertilization (Sharpley, 1995), but nitrate pools remain seasonally elevated when annuals dominate, thus
reinforcing the prevalence of exotic species (Prober & Lunt, 2009).
Step 1b: Current ecosystem services. Grassy woodlands are valuable production systems, with a trade-off between
intensifying or extending production and overall sustainability in terms of preserving vegetation, soils and
biodiversity (Smith et al., 2013). We undertook a literature review to identify ecosystem attributes that underpin
ecosystem services supplied by grassy woodlands (Tables S1 and S2, Supporting Information). We quantified each
ecosystem service using a 5-level rating (Table S3, Supporting Information). Star diagrams summarize the varying
provision of ecosystem services between states (Fig. 3). A principal component analysis of Table S3 (not shown)
revealed three distinct bundles of services, i.e. services co-occuring in given grassland states.
Reference and native pasture states share high multifunctionality with high values for regulating and cultural
services and biodiversity indicators. The incorporation of production, albeit low, in native pastures comes at a
small cost to these other services (Fig. 3). In contrast, the fertilized pasture, sown pasture and cropping states
show radical shifts towards provisioning services at the expense of all others (except aesthetics) due to loss of
perenniality and native plant diversity, tree decline and ultimately loss. Ceasing or reducing production in fertilized
pastures restores some regulating services by allowing recovery of a largely exotic perennial ground layer, but not
of biodiversity or cultural services specific to reference and native pastures.
Step 2: Climate change effects. Climate change projections indicate a trend of high temperature, reduced or warm
rather than cool-season precipitation precipitation, and more extreme rainfall and drought events (CSIRO &
Bureau of Meteorology 2012). These changes are depicted under four scenarios of increasingly severe climate
change and predicted land use responses (Table 1).
Reduced capacity for cropping is predicted, and cropping is likely to become more marginal (scenario 3) or be
replaced by extensive grazing (scenario 4; Nidumolu et al., 2012). A history of extensive cropping in a landscape
may limit the development of native perennial vegetation suitable for rangeland grazing, and decrease its ability to
retain soil and sustain biomass. Abandoned cropland is likely to become annual grassland, with extensive areas of
exposed soil in summer, vulnerable to water and wind erosion.
In reference and native pastures there is potential to support assembly of novel perennial diverse semi-arid
rangelands combining some original grassy woodland, exotic and semi-arid species. We consider that the plant
diversity currently available in these states provides a sufficient array of ecological strategies in order to support
responses to scenario 3, and to a lesser extent scenario 4. The latter may require some assisted dispersal of semi-
arid native species.
Step 3: Adaptation services. We assume that, similar to current use in semi-arid and arid Australia, livestock grazing
will remain a primary future land use. Other objectives may be biodiversity conservation and maintenance of
vegetation cover as appropriate for natural resource conservation under more severe scenarios.
We identified a bundle of adaptation services that are essential to support viable medium-term livestock
production (Fig. 4), including soil regulation services underpinning stable plant production, microclimate regulation
for welfare of livestock and the provision of water for livestock and people. During the transformation of the flora
towards novel semi-arid communities (Step 2), several ecosystem attributes will increasingly support these
adaptation services (Fig. 4). Perennial grasses, the keystone functional group supporting current landscape
multifunctionality, will have increasing importance in soil protection and primary productivity. Additional
ecosystem attributes supporting adaptation services for continued livestock production (Table 1) include structural
and life form diversity, and large plant functional diversity of natives and exotics in the ground layer. These have a
large response diversity to climate variability, to management interventions, and disturbance from drought and
fire. The proportion of stress-tolerant species, already a major component of these ecosystems, is also expected to
increase. The persistence of all these attributes under changing climate requires landscape scale biodiversity and
connectivity to support interactions and trophic complexity.
As climate changes, many species will not persist. Their contribution to response diversity will decline and be
replaced by species from less intensively managed states, with wide geographic distributions (Figure S1,
Supporting Information). We assume gradual community re-assembly with immigration of semi-arid species under
more severe scenarios 3 and 4, but translocation may be required to maintain the matrix structure of perennial
grasses and enable the adaptation service of response diversity. For this, landscape connectivity will be an
essential attribute. Climate regulation is expected to emerge as a novel adaptation service as climate change
becomes more severe.
Importantly, the ecosystem properties that are likely to contribute to future adaptation services currently
support ecosystem services from conservatively grazed native pastures (Fig. 3). They comprise many of the
attributes that maintain viable livestock production as part of the bundle of services provided by reference
grasslands and native pastures.
Step 4: Management implications. Currently, the highest priority is maintaining perennial vegetation to reduce the
risk of future desertification. While short-term returns from the expansion of annual cropping may be attractive,
the long-term functionality and productivity of the landscape will be at risk from erosion and declining soil health.
Expanding cropping will further reduce plant and animal diversity, thus risking loss of species tolerant of
environmental stress, with potentially important, yet presently unrealized, adaptation service roles. As aridity
increases, management will need to be fundamentally different in type and operation. Errors in over-harvesting
and intensification are likely as new climate extremes are experienced. It will become increasingly important that
agricultural extension incorporates perspectives from semi-arid areas and implements landscape-integrated
Littoral rainforest
Step 1a: Ecosystem description. Consistent with global patterns, and exacerbated by the scarcity of inland
resources, Australian coasts have been, and are increasingly, the focus of human settlement and urbanization. The
Littoral Rainforest and Coastal Vine Thickets of eastern Australia (henceforth littoral rainforest) represent a
complex of rainforests and coastal vine thickets structured by processes including effects of salt spray and on-
shore winds, tidal inundation and storms, saltwater intrusion of groundwater and unstable, dynamic substrates. In
the Wet Tropics, due to the protection provided by the fringing reefs, these communities only exist within 2 km of
the coast and <10 m above sea level (Metcalfe et al., 2013). Littoral rainforest is naturally distributed as a series of
disjunct, localized stands which provide important stepping stones for migratory shorebirds.
Littoral rainforests are highly dynamic ecosystems, often in a state of regeneration following repeated storm
disturbance. Structure is typically a closed canopy of trees which may include emergents. Patches regularly
exposed to disturbance may have many canopy gaps. Whilst canopy species are well adapted to coastal exposure,
the canopy protects less tolerant species and propagules in the understorey.
Because of their coastal location, littoral rainforests are highly vulnerable to interacting effects of climate
change and sea-level rise, along with existing threats such as invasion by transformer weeds, which alter structure
and function, and fragmentation due to coastal development.
Step 1b: Current ecosystem services. Littoral rainforest occurs at the interface between terrestrial and aquatic
systems, where it protects land from erosion, filters sediments, nutrients and pollutants, mitigates the effects of
flooding and wind during storms and provides habitat for biodiversity (Fig. 5). Foreshore vegetation and natural
dunes also provide protection to coastal communities, beaches and infrastructure including roads, marinas,
agricultural (e.g. sugarcane, fruit and nut production) and aquacultural industries. Tree height, vegetation
structure, stem density and ground cover all influence the ability of littoral rainforest to stabilize soils, attenuate
waves and act as wind breaks. Other ecosystem services include the provision of shade, nesting sites and food
resources for fauna, migration capacity for endemic and iconic species, and cultural and aesthetic services.
Step 2: Climate change effects. Four scenarios of increasing severity, and outcomes for species, ecosystems,
landscapes and adaptation services, are summarized in Table S4 (Supporting Information). Climate change
predictions for the tropical rainforest habitats of Queensland (Suppiah et al., 2010) indicate air temperature
increases and declining rainfall particularly during the ‘dry season’, resulting in seasonal drought (Williams et al.,
2012), increase in extreme rainfall events across the coastal high rainfall zone, and more intense cyclones (Suppiah
et al., 2010). Projections of sea level rise are of particular significance for littoral rainforest.
Under less severe scenarios, minor changes in community composition and diversity can be expected as some
native species reach their tolerances to heat and low water availability. Rising sea levels and greater storm
intensity will increase exposure to inundation and disturbance, which will gradually increase fragmentation and
create opportunities for invasion by exotic species.
As climate change increases in rate and severity across scenarios, a simplification of community structure is
expected. Loss of emergent trees and physical canopy damage from more intense storms effects will reduce
vegetation height and create canopy gaps (Kellner & Asner, 2009). Increasing internal and external fragmentation
of littoral rainforest patches will continue to facilitate exotic species invasions and place further stress on native
species, resulting in losses of biodiversity.
Transformative changes in ecosystem structure, function and composition are likely with increased sea level
rise, storm surges, floods and cyclone intensity, or with co-incident extreme events like fire during the dry season.
Littoral rainforests may be lost or transform to mangroves, depending on location and level of inundation. In many
coastal margins, natural features and urban development prevent shoreward migration. Sea-level rise will result in
narrower distributions (intensifying the effect of coastal squeeze), smaller patch size and increasing fragmentation.
These effects may result in littoral rainforest patches being lost or dominated by transformer weeds, highly
invasive taxa with the potential themselves to significantly alter the structure and function of the community.
Step 3: Adaptation services. Littoral rainforests confer adaptation services by (i) mitigating storm surge, inundation
and wind impacts on infrastructure and people by providing a barrier to wind, salt spray and debris mobilized by
cyclones (Wamsley et al., 2010; Shepard et al., 2011; Arkema et al., 2013); and (ii) buffering impacts, enabling
continued ecosystem function after extreme events (Fig. 5). High diversity of response traits in the community
allows important functions to be maintained, including provision of food, shade, and habitat for endemic and
iconic fauna targeted by conservation and tourism. Littoral rainforest provides a barrier to wind and salt spray.
The adaptation service of erosion regulation will be important in mitigating the impact of sea level rise on
nearby settlements and ecosystems. Eroded coastal dune systems and saltwater intrusion into freshwater
wetlands will negatively affect public uses such as tourism and recreation (Environment Planning, 2011), including
the loss of public assets like beaches. Critical ecological mechanisms supporting these adaptation services are an
intact vegetation structure and a diversity of plant life forms including emergent canopy trees, sub-canopy and
understorey layers, a diversity of disturbance and regeneration response traits, and the extent and connectivity of
littoral rainforest patches.
Step 4: Management implications. Maintenance of intact, diverse, connected forest stands of good quality is the
key management requirement to support adaptation services. Assisted regeneration to maintain size and quality
may be required in vulnerable, small, isolated patches following major disturbance. Under more severe climate
change scenarios, assisted regeneration and restoration may be necessary if autonomous regeneration is
compromised. Landscape and development planning that facilitates the inland movement of dunes and littoral
rainforest will become increasingly important with rising sea levels combined with greater intensity of storm
surges and inundation events.
Engineering solutions may substitute for some services, but are usually not as cost-effective as natural
foreshore vegetation at protecting coastal infrastructure (Jones et al., 2012). Infrastructure engineering
approaches may also impair recreation and aesthetic values, enhance seaward erosion of littoral habitats, degrade
water quality and impair aquatic system function (Arkema et al., 2013).
Temperate montane forest
Step 1a: Ecosystem description. The Australian Alps and Southeastern Highlands Bioregions in south-eastern
Australia include large areas of montane forest (Thackway & Cresswell, 1995) (Fig. 2). The dynamics of
sclerophyllous montane forests are driven by infrequent high-intensity fire. Continuing supply of ecosystem
services, such as erosion protection and water quality hinges upon the ability of these forests to recover from
periodic severe disturbances. Montane forest communities broadly comprise two disturbance-response types: (i)
forests dominated by fire-killed eucalypt ‘ash’ species (Eucalyptus delegatensis and E. regnans) which as obligate
seeders are sensitive to inter-fire intervals shorter than the ~20-years required to reach maturity; and (ii) forest
dominated by resprouting eucalypts (E. dalrympleana, E. robertsonii, E. fastigata and E. obliqua) which recover
within a few years from even high intensity fire.
Step 1b: Current ecosystem services. The major land uses associated with montane forests are catchment
protection (to provide water to towns), forestry and nature conservation with diverse forested vegetation types
supporting bundles of ecosystem services including soil conservation, water supply, carbon sequestration, nutrient
cycling, timber provision, genetic diversity, biomass production, fauna habitat, landscape aesthetics and recreation
(Fig. 6).
Because most of the forested areas are public lands, management of timber harvests, fuel-reduction burns to
protect adjacent or nearby property, stock grazing and the mix of production and conservation uses are hotly
contested (Lindenmayer, 1995). For example, after extensive fire events in E. regnans forests, streamflow and
water yields reach a maximum reduction 2025 years after fire, only reaching their prefire levels after 100150
years in the absence of further disturbance (Kuczera, 1987). After large fire events, salvage logging has occurred
within prescriptions designed to protect water catchment values (O’Shaughnessy & Jayasuriya, 1991). These
prescriptions do not account for multiple values of standing timber, including provision of breeding and roosting
hollows, and carbon storage (Mackey et al., 2008).
Step 2: Climate change effects. Under climate change, the number of extreme fire weather days is expected to
increase in south-eastern Australia (Hennessy et al., 2005; Lucas et al., 2007) with potential for greater number,
extent and return time of large fires. There is a direct feedback between fire as a driver, the ensuing ecosystem
state and maintenance of ecosystem services. Increased frequency and extent of high intensity fires may lead to a
change in forest structure (scenario 2), the loss of mature forests (scenario 3), or the loss of forest and substitution
with shrubby vegetation (scenario 4; Fig. 6), with alternative state-and-transition pathways dependent on the
dominant eucalypt species (Table S5).
Resprouting forests are likely to maintain composition but may change structure, forming coppiced low
woodlands (Stephens et al., 2013). Ultimately, resprouters may be replaced by seedlings, but only over hundreds
of years, with a mixture of age cohorts present at any time point. In contrast, Ash-type eucalypt forests may be
buffered from short-term climate and fire regime change because of their mesic understorey and low background
fire frequency. However, under more severe change scenarios, with increased recurrence of catastrophic fire, a
transformation is expected due to the local extirpation of these obligate seeders. The beginnings of transformation
have occurred in the southern Alps (Bowman et al., 2014). Ash-type eucalypt forests lack endogenous capacity to
adapt locally under these fire regime scenarios.
Step 3: Adaptation services. Resprouting is a key functional trait supporting autonomous adaptation, enabling
persistence with an increase in fire frequency, recovery after high intensity fire (Burrows, 2013; Clarke et al., 2013)
and underpinning the supply of ecosystem services. Species lacking this trait are far more sensitive to a shortened
inter-fire interval and, where they dominate, landscapes are vulnerable to losses of ecosystem services under a
future drier, hotter climate. Conversely, the ecosystem services from ash-type eucalypt forests are vulnerable to
changing climate and fire regimes. These forests could become the focus of expensive, maladaptive management if
there is a mismatch between the type of forest that is currently desired and the type of forest that can persist
under a range of future climate and fire regime scenarios.
Step 4: Management implications. The risk to ecosystem services is severe because montane forests occur in upper
catchments that supply water to cities and regional centres. High intensity fire in E. regnans forests reduces water
yield, timber and biodiversity habitat. In E. delegatensis forests, fires in 2003, 2007, 2009, and 2013 have created
patches with no regeneration (Bowman et al., 2014). These areas have been aerially reseeded at a cost of ~A$
3000 ha-1, but if anticipated fire regime change converts larger tracts to this denuded state, the cost of re-seeding
with ash species may become prohibitive. The inherent adaptation service provided by resprouting eucalypt
forests requires no active management. This means in the short-term a greater management focus on fire-
sensitive Ash-type eucalypt forests, including fire suppression, fuel reduction and reseeding. However, novel
approaches to management may need to be considered in the future, such as translocating seed from resprouting
montane species rather than fire-sensitive ash species. The management of these systems need to be formulated
within the context of multiple scenarios and competing or conflicting ecosystem service values, particularly
between water yield, timber harvests and wildlife habitat (Lindenmayer & Likens, 2009).
Inland floodplains and wetlands
Step 1a: Ecosystem description. The Murray-Darling Basin covers 14% of Australia and generates 45% of the
irrigated agricultural production, worth ~$5.5 bn p.a. (MDBA, 2010). Westerly flowing rivers receive runoff from
high rainfall regions to the east, undergoing irrigation diversions and high evaporative losses over long traverses
across semi-arid floodplains before terminating in extensive wetlands that drain into the Darling and Murray rivers.
Floodplain woodlands and forests, consisting of few flood- and drought-tolerant Eucalyptus and Acacia species
(river red gum Eucalyptus camaldulensis, black box E. largiflorens, coolibah E. coolabah, river cooba Acacia
stenophylla), give way to riparian woodland corridors hemmed by chenopod shrubland and grassland in more arid
regions. Historically, river flows were highly variable, driven by inter-decadal ENSO cycles, with prolonged drought
preceding extensive floods (Chiew et al., 2008). The biota has adapted accordingly, and dry periods are needed for
key ecosystem processes (Baldwin et al., 2013). Due to 50100 years of river regulation and water resource
development, floods are now of lower volume, duration, extent and frequency (Sims et al., 2012) and many
wetlands are in poor and declining condition (Davies et al., 2012).
Step 1b: Current ecosystem services. Floodplains supply grazing and cropping. Wetlands supply critical habitat for
vegetation, waterbirds and fish, and are important for tourism, recreation, cultural and spiritual values. Rivers
supply water for irrigated agriculture and domestic use and wetlands buffer the impact of floods on surrounding
land and regulate nutrients, sediments and microclimate. Supply of ecosystem services from wetlands dependent
on regulated river flows is via natural floods and managed releases of ‘environmental flows’ from dams. The
impact of historical water diversions has been increased provisioning services from irrigated agriculture at the
expense of biodiversity/habitat and regulating services (Fig. 7).
Step 2: Climate change effects. Climate change adds to stressors from water diversions. Four scenarios of climate
change are considered, that represent increasing levels of rainfall reduction and aridity, frequency of extreme
floods, and their consequences for cropping and grazing, all the way to a shift to rangeland grazing under the most
severe scenario. Detailed information on their climate impacts and adaptation services is presented in Table S6
(Supporting Information). Floodplain vegetation is structured by flood frequency and duration, with the most
flood-requiring species located on the lower floodplain. Responses to climate change include contraction and
compositional shifts in plant communities, fewer flood-induced breeding and regeneration events and loss of
habitat for biodiversity. There will be fewer, smaller, saltier wetlands, filled less often (Nielsen & Brock, 2009).
Water-demanding plants will be replaced with water-conserving ones. Areas of E. camaldulensis forest will thin to
open woodland and be partly replaced by slower-growing, drought-tolerant E. largiflorens and E. coolabah.
Floodplain extent is likely to contract, with upper zones transitioning to terrestrial grassy woodland and chenopod
shrubland. Impacts on ecosystem services include declining irrigated agriculture and some increases in regulating,
habitat/biodiversity and cultural services (Fig. 7).
Step 3: Adaptation services. Floodplain ecosystems are likely to persist under climate change, though with reduced
extent and altered vegetation structure, due to the following attributes: (i) high response diversity of trees and
understorey to flood and drought; (ii) drought-resistant life-cycle stages, with long-lived propagule banks; (iii) rapid
growth and regeneration after rainfall and flooding (Colloff & Baldwin, 2010); (iv) resources produced during
floods are sequestered to provision dry-phase biotic activity (Baldwin et al., 2013); (v) high connectivity via riparia
for recolonization and propagule transport. These adaptation services underpin bundles of regulating services,
including flood mitigation and erosion prevention from rare, extreme rainfall events, salinity control and provision
of shade and shelter. The fewer, smaller wetlands that can be maintained with environmental flows are likely to be
of high biodiversity conservation value and increasingly important for bundled cultural services of tourism,
recreation, heritage and spiritual values. Some wetlands will persist in a low-diversity state, with rare floods driving
regeneration from propagule banks, but capable of rapid, high-productivity responses to rainfall and floods.
Fodder for livestock grazing is an adaptation service likely to become increasingly important from upper
floodplains that transition to drought- and salt-tolerant grassy woodland and chenopod shrubland.
Step 4: Management implications. To counter ecological decline, limits on water diversions and an objective of
2750 gigalitres of water restored to the environment is planned (MDBA, 2010). For scenarios 3 and 4 of an
increasingly arid climate associated with much reduced river flows, along with rare, intense floods (Table S6,
Supporting Information), prioritising wetlands by their likelihood of autonomous adaptation or capacity to shift to
alternate stable states will be needed. Future environmental flow management will include cessation of delivery to
wetlands unlikely to persist long-term and inclusive, adaptive community and governance arrangements. As less
irrigation water is available, dryland agriculture, grazing and mixed income approaches will emerge and their
impacts be managed. Lake Eyre, west of the Murrray-Darling Basin, is a dry salt lake most of the time. It filled in
20102011, stimulating high-value tourism and recreation (Lockyer, 2012), providing an example of how wetlands
enhance cultural values, even in arid landscapes.
The four case studies illustrate the operational application of the adaptation services concept (Fig. 1) to
contrasting ecosystems and climatic regions. They highlight the versatility of the framework across bioclimatic
conditions, land use types, ecosystem service bundles, and likely ecological response pathways to anticipated
climate change. Beyond their diversity, they illustrate common functional mechanisms that we hypothesize are
generic to adaptation services. Below we synthesize these mechanisms before considering their implications for
the selection of methods to quantify adaptation services. Lastly, we discuss how the new concept of adaptation
services and knowledge about intrinsic biophysical mechanisms can inform management and decision-making in
adapting to a variable and uncertain future.
Trajectories of ecosystem change under worsening climate change and types of adaptation services
The four case studies illustrate trajectories for ecosystems with different initial resistance and disturbance regimes
(Fig. 2). Grassy woodlands, inland floodplains and fire-driven montane forests have already beenmodified during
historical and recent times (e.g. the 19972010 Millennium Drought). Grassy woodlands are expected to gradually
transform under climate changes and ultimately reach critically dry conditions, with vegetation only suitable for
rangeland grazing. Floodplains are expected to reach a tipping point where water availability no longer meets
environmental requirements, so that wetlands contract in area and upper floodplains transform to terrestrial
ecosystems. Resprouting montane forests may gradually change structurally while maintaining their characteristic
species, but the probability and timing of transformation to a complex mosaic comprising native trees and shrubs,
and even exotic grass-dominated vegetation need to be considered as core uncertainties. In contrast, pathways for
stands dominated by fire-killed Ash are highly sensitive to management. If regenerating stands in surrounding
forests are protected then gradual change would be possible, but if fire recurs within the 20-year maturation
period then an abrupt transformation to shrubland would be inevitable. Contrasting with these cases of initial
gradual dynamics, littoral rainforest can readily shift to an alternative community state with a highly simplified
structure and low diversity, particularly in the presence of exotic transformer weeds, as soon as an unprecedented
combination of disturbance events occurs, such as a series of cyclones in rapid succession or cyclone followed by
fire. These alternative states may provide dune stabilization and erosion prevention, but at the expense of
biodiversity/habitat services and cultural and aesthetic services.
The four case studies highlight three types of adaptation services, i.e. benefits to people from increased social
ability to respond to change, provided by the capacity of ecosystems to moderate and adapt to climate change and
variability: attributes that (i) allow ecosystems to resist or cope in the face of direct and indirect impacts of climate
change; or (ii) transform autonomously to a state that supports social adaptation, especially by being responsive to
management that fosters new bundles of ecosystem services and (iii) support new valuable ecosystem services,
including ones currently supplied but less-valued or in demand. The first type prevails across all case studies and is
expected to be an important feature of climate change adaptation across many ecosystems. As detailed in the next
section, it is underpinned by mechanisms relating to ecological resilience (Holling, 1973). Examples for the second
type are provided by resprouting trees replacing Ash-type eucalypt stands in montane forest, and the role of other
keystone functional groups such as perennial tussocks in grassy woodlands. Landscape connectivity also recurs as a
critical process for this second type. Here, adaptation services rely on ecological transformability rather than
resilience mechanisms (Walker et al., 2004). Instances of the third type include climate regulation in grassy
woodlands, coastal buffering by littoral rainforest and the increasing importance of regulating services and novel
cultural values and grazing resources on floodplains. More generally, and as noted in previous ecosystem-based
adaptation literature (Jones et al., 2012; Pramova et al., 2012) the value of regulating services will increase and
new food, fibre and fuel sources will become available to societies in various ecosystems undergoing climate-
driven transformation. In this last case, not only are ecological resilience and transformability necessary, but also
social adaptation (Adger et al., 2005) enabling new values for existing or emerging ecosystem attributes.
Functional mechanisms underpinning adaptation services
Table 2 summarizes five recurring functional mechanisms across the four case studies. Vegetation structure, its
complexity and structural diversity appeared critical in all case studies, except floodplains, for the future
maintenance of current ecosystem services by buffering climate change effects.
The dynamics of all systems hinge on the persistence of keystone species or functional groups carrying a
syndrome of traits which shape the ecosystem across trophic levels and have strong effects on nearly all current
ecosystem services. Examples include perennial tussocks in grassy woodlands, eucalypt resprouters in montane
forests as well as canopy-forming species in littoral rainforest and floodplains. We assume that they would be
critical to adaptation by supporting ecological resilience, while their local extinction would readily lead to
transformation. Thus, a high risk is associated with their climate responses and possible range shifts, especially in
the case of one or a few keystone species such as the drought-, flood- and salinity-tolerant trees on floodplains.
Conversely, their response diversity to climate through range shifts among species or replacement by more arid-
adapted species could be a strong component of coping and transition. Overall, we hypothesize that in all systems,
response diversity to climate, combined with functional redundancy for ecosystem services results in greater
ecological resilience (Walker et al., 1999), and thereby supports the realization of adaptation services. This
hypothesis could be tested across a greater number of case studies, and by analyzing responses to recent
increased climate variability or to historical extremes.
We noted that disturbance response diversity, i.e. the presence of species with diverse survival and
regeneration responses, was involved in acclimation or coping. Relevant traits include high diversity of germination
cues (all systems), varying fire tolerance and resprouting abilities (all systems), response diversity to cyclones in
littoral rainforest, or to management and drought in grassy woodlands and floodplains. Response diversity can
mediate ecological transformation in several ways: (i) through shifting contributions of different species groups
(Table 1, Figure S1 for grassy woodlands; see also Tables S4 for littoral rainforests and S6 for floodplains); (ii)
through response diversity within keystone functional groups of grassy woodlands, montane forests and
floodplains and (iii) through generic response diversity in the absence of keystone functional groups (littoral
Landscape connectivity is already adaptive currently and sustains multiple ecosystem services (Mitchell et al.,
2013). It is expected to play a key role in ecological transformation of fragmented systems like grassy woodlands
and littoral rainforests through its effects on propagule flows that are necessary for disturbance responses and for
migration of climatically suitable species. Connectivity is critical for floodplains whereby carbon, nutrients and
propagules are distributed via water flows (Baldwin et al., 2013). Alternatively, increased connectivity in relation to
fire in montane forests would result in greater rather than lesser impacts where obligate-seeders dominate.
Functional assembly mechanisms supporting adaptation services
Assembly of future communities remains problematic for prediction of future biodiversity and ecosystem
functioning (Boulangeat et al., 2012). Elucidating assembly mechanisms will be critical for understanding and
quantifying adaptation services. New communities will assemble following species range shifts, first within
response groups like resprouters in fire-driven systems or gradual integration of semi-arid species into grassy
woodland communities. In these cases, functional redundancy within these response groups will support
adaptation services. Next, in contrast with gradual increase or decrease of specific life forms or functional groups
(e.g. transition to fire-tolerant species in rainforest or montane forests under changed fire regimes),
transformation will require immigration by, or dominance of, species with novel traits where previously dominant
traits have been lost. Novel ecosystems containing exotic species, following extinction of dominant native species,
are likely to become commonplace (Hobbs et al., 2006). Also, ecosystems may be engineered for adaptation using
dominant species with traits that support desired ecosystem services, for example using a species framework
approach (Tucker & Murphy, 1997) as done for restoration of littoral rainforests (Goosem & Tucker, 2013).
Quantifying adaptation services
Quantifying adaptation services will be required for scenario predictions and adaptive management. Knowledge of
ecological mechanisms and functional traits is likely to facilitate this challenging task. Below we identify three
avenues for the development of quantitative approaches.
Step 1 in the operational framework highlights the importance of historical context. Its analysis identifies
whether adaptation services are present or have been compromised by past management or recent climate
change (e.g. a mean 0.9 °C increase for Australia since 1910; CSIRO & Bureau of Meteorology 2012) or, in the case
of floodplains, major water diversions for irrigation. Consideration of history informs which adaptation services
should be maintained or restored under climate change. For instance, for grassy woodlands, adaptation services
are supported by relatively intact perennial-dominated communities, but these have largely been lost through
historical management (Fig. 3). Once adaptation services have been identified (Step 3), they can be quantified by
transfer of values depending on ecosystem state, or by modelling supporting services and processes like net
primary productivity and nutrient cycling (Crossman et al., 2013). Considering the covarying nature of ecosystem
services, and their linkages to supporting services (Bennett et al., 2009), we recommend systemic approaches to
quantify bundles of ecosystem services that include adaptation services. An intermediate level of modelling
sophistication could be achieved by predictive mapping of landscape ‘hot spots’ that are critical for adaptation
services (see e.g. Qiu & Turner, 2013 for relevant methodology), for example when linked to particular functional
groups such as Ash eucalypt species in montane forests.
Lastly, landscape configuration effects will need to be considered. These can be implicit through the
proportions of different vegetation states, which can be linked to connectivity and the functional quality of the
landscape [Smith et al. (2013) for grassy woodlands; Fahrig et al. (2011) for agricultural landscapes], or the
presence of refugia. Spatially explicit modelling will be required to account for adaptation services linked to
processes like fire propagation, river flows, storm energy interception or species migration.
Management of adaptation services
Implicit to increasing severity of climate change is the shifting nature of risk profiles which will be increasingly
characterized by extreme events, unpredictability and coincident novel events. The unpredictable, changing nature
of risks will necessitate that approaches to planning, development and management change focus from optimising
returns to managing those risks, including management of ecosystems for their adaptation services. Since many
changes in ecosystem properties and adaptation services will be emergent, an adaptive learning and management
approach is necessary, as suggested for managing future ecosystem services (Cowling et al., 2008). With our four
case studies we identified three types of management for adaptation services.
As the first priority, management should identify and protect existing adaptation services such as those that
occur in ecosystems in good condition. Secondly, degraded but existing adaptation services can be restored by
supporting establishment of key functional groups of the vegetation matrix and restoring drivers and regimes. The
former will serve biodiversity conservation objectives (McIntyre, 2008) and adaptation needs simultaneously. The
latter applies to the management of environmental flows, as outlined by the Murray-Darling Basin Plan (MDBA,
2010) or altered fire regimes (Bradstock et al., 2012). Lastly, novel management will be needed for emergent
adaptation services, e.g. adaptive management of environmental water to high-priority wetlands likely to persist
under climate change. Beyond this sequence, the novel context of climate change will require prioritization in
adaptation service management, for instance managing fire in dry forest to protect humid, sensitive Ash-type
eucalypt forest patches. For adaptation service management to be successful, social adaptation is required
including specific actions for managers, new knowledge and technology and transformation of whole industries.
Adaptation service management may be incorporated into biodiversity conservation strategies when it has
benefits for nonutilitarian biodiversity values (e.g. response diversity within non-iconic species groups), or when
managing for iconic biodiversity (e.g. vertebrates in montane forests, birds and bats in littoral rainforest) and
promotes ecosystem states commensurate with adaptation services.
Finally, the focus on adaptation services should help avoid maladaptive management trajectories, for example
the use of environmental water allocations for floodplains that cannot be sustained in the long-term because of
likely transition to terrestrial ecosystems, or the reseeding of fire-killed eucalypt Ash stands when risk from
recurring fire increases. In montane forests, a hierarchy of strategies may be required such as prioritizing
controlled fire management in areas from timber production to water production, managing dryer forest types
more intensively than wet types, or potentially reseeding resprouting species with desirable adaptive properties
into some fire killed Ash stands.
The broadened concept of adaptation services we present moves beyond that currently proposed in the literature
under Ecosystem Based Adaptation (EBA) by focusing on the understanding of key ecological mechanisms and
traits supporting the intrinsic capacity of ecosystems to adapt to change. While EBA highlighted how ecosystems
can support new valuable ecosystem services under climate change, we have demonstrated the additional value of
ecosystems in good condition for: (i) allowing ecosystems to resist or cope in the face of climate change direct and
indirect impacts or (ii) transforming autonomously to a state that supports social adaptation. This novel
perspective provides direction and focus for climate change research and ecosystem modelling relevant for the
exploration of social adaptation pathways. While this article provides a deliberate focus on biophysical knowledge,
next steps will also need to consider economic, institutional and social mechanisms for adaptation services.
This research was supported by a CSIRO OCE Distinguished Visiting Scientist fellowship to SL and by the CSIRO
Climate Adaptation National Research Flagship Theme 1157. It contributes to OPERAs FP7-ENV-2012-two-stage-
308393 and BiodivERsA project REGARDs with funding from the French Agence Nationale pour la Recherche (ANR).
We thank Drs Sarina Macfadyen and Saul Cunningham (CSIRO Ecosystem Sciences) for constructive reviews of the
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Supporting Information (available at:
Appendix S1. Ecosystem attributes, functions and services associated with different vegetation states in the temperate eucalypt
grassy woodlands of southeastern Australia.
Table S1. Summary of ecosystem functions and attributes. References relevant to more than two states are listed under the
function name.
Table S2. Summary of ecosystem services that may be associated with five major land uses in the temperate grassy eucalypt
ecosystems. Values are those associated with the functional state of the ground layer vegetation. References relevant to
more than two states are listed under the value name.
Table S3. Semi-quantitative translation of data from Table S2 using a categorical scale of increasing provision (15).
Figure S1. Hypothetical response of plant groups to increasing aridity in the eucalypt grassy woodland biome. Native species are
predicted to increase initially with increasing drought stress, while more mesic adapted exotics may decline. At a critical level
of heat and dryness, locally adapted species could potentially be replaced by semi-arid species in adjoining regions assuming
dispersal is not limiting.
Appendix S2. Data for the adaptation service assessment in littoral rainforests, montane forests and the Murray-Darling Basin
Table S4. Description for the littoral rainforest system of climate and management scenarios, impacts on ecosystem state and
processes, adaptation services and their supporting attributes and mechanisms, and management needs for each of the four
types of increasing severity. Arrows indicate increasing importance of specific attributes across scenarios.
Table S5. Description for the montane forests system of climate and management scenarios, impacts on ecosystem state and
processes, adaptation services and their supporting attributes and mechanisms, and management needs for each of the four
change types of increasing severity. Arrows indicate increasing importance of specific attributes across scenarios.
Table S6. Description for the inland floodplains and wetlands of climate and management scenarios, impacts on ecosystem
state and processes, adaptation services and their supporting attributes and mechanisms, and management needs for each
of the four change types of increasing severity. Arrows indicate increasing importance of specific attributes across scenarios.
Table 1. Description of the grassy woodlands system under climate and management scenarios, impacts on ecosystem state and processes, adaptation services and their
supporting attributes and mechanisms, and management needs for each of the four change types of increasing severity.
Scenario 1 (moderate effects over
regional scale and longer timeframe)
Scenario 2 (moderate effects over
regional scale and shorter timeframe)
Scenario 3 (severe, widespread effects
over longer timeframe)
Scenario 4 (severe, widespread effects
over shorter timeframe)
Periods of severe dryness
Mixed cropping, fertilized pastures,
native pastures and conserved
woodlands. Fertilization in wet periods.
Rotational grazing (recent)
Record drought
Increasing periods of extremely high
grazing pressure.
Reduction in fertilizer use due to
diminishing returns.
Declining rainfall averages.
Transition to opportunistic cropping.
Greater grazing pressure on diminishing
average forage production. Continued
reduction in fertilizer use; lag effects of
past use
Declining rainfall averages.
Threshold of dryness and temperature,
transforming to semi-arid climate.
Transformation to rangeland use -
dryness prohibits cropping
Species, ecosystem and
landscape effects
Landscape pattern
More diversified landscape mosaic
A mosaic of exotic-dominated
vegetation and re-assembling native
Trees and shrub
Negative effects of intensification and
Eucalypt mortality increasing, recruitment of a few semi-arid tree and shrub species
Ground layer - native
Negative effects on grazing-sensitive
and nutrient-sensitive spp.,
Local extinctions of some species,
increase in native grazing-tolerant
Climate and increasing grazing pressure
causes turnover in dominants.
Reduced plant diversity.
Perennial grass dominance varies with grazing and fertilization
Annual dominance varies with grazing
and fertilization
Opportunities for annual flushes,
including infrequent species.
Increasing native ephemerals
Groundlayer - exotic
Exotics increasing
Some exotic species vulnerable
Stress tolerant ruderal exotics increasing
Annual exotics increasing
Negative effects of intensification and
Bare ground and erosion increasing
Landscape-scale processes
Local climate effects leading to positive feedback reinforcing more arid local climate
Ecosystem properties
supporting adaptation
Original flora of grassy
woodlands (exotics and
Presence of trees and shrubs and perennial grasses, current diversity of plant and animals, (native and some exotic). Many hemicryptophytes and stress-tolerant
spp.Diversity of strategies and species associated with states with relatively less intensification; and/or with diversified exotic flora in un-fertilized statesSpecies with wide
geographical ranges
Responses of original flora
to climate change and
management adaptation
Increasing importance of stress-tolerant plant species
Decreasing contribution of original species pool to response diversity
Greater contribution of flora from reference and native pasture states to re-
Desirable for adaptation
Immigrating or translocated species
from semi-arid regions
Adaptation services
Increasing importance of stability of production under increasing inter-annual climate variability, maintenance of soil structure, erosion protection, rural microclimatic
regulation, amount of abstracted surface water
Increasing importance of: global climate regulation, local and regional climate
regulation, response diversity to climate change and variability, landscape
Relevant traits and
Functional composition and
Increasing importance of perenniality, presence of trees and shrubs, diversity of life forms, diversity of species within life forms providing response diversity
Increasing importance of amount and connectivity of the above
Management response to
support adaptation
Landscape planning to create a matrix of native dominated pastures and native vegetation
Increasing value of assisted dispersal of semi-arid native plant species
Transfer of arid grazing knowledge to
the east.
Extension on improved grazing and soil
management practices.
Transfer of arid grazing knowledge to
the east.
Up-skilling with respect to agronomic
practices, extension and ‘ecologically
acceptable’ technology development.
Increase in farm sizes.
Support the transition economic base
from grain production to livestock.
Increase in farm sizes.
Table 2. Synthesis of functional mechanisms underpinning adaptation services
Case study
Grassy woodlands
Mountain forests
Littoral rainforests
Structural / life
form diversity
Keystone species /
functional groups
Perennial tussock
Keystone acacia,
eucalypt species
Resprouter /
Climate response
diversity within:
All life forms and
especially perennial
Keystone acacia,
eucalypt and
understorey spp.
Resprouting species
All life forms
response diversity
Diversity of
Diversity of survival
and persistence
Diversity of
Seeders: No
Diversity of
responses to wind
and inundation and
Propagule flow
Propagule flow
Figure 1. The operational framework for the identification and quantification of adaptation services. Four steps
are identified: (1) Characterisation of the system and its bundles of ecosystem services, (2) Describing climate
change direct and indirect impacts, (3) Identifying adaptation services, and (4) Proposing management for
adaptation services.
Figure 2. The four case studies: distribution ranges and typical landscapes for littoral rainforests, the Murray-Darling Basin (MDB) floodplains, grassy woodlands and
montane forests. Inserts for each case study picture the most important drivers and feedbacks for each case study: climate (C), management (E), exotic species (Ex),
ecosystem properties (EP) and ecosystem services (ES)
Figure 3. Summary of Step 1 for grassy woodlands: initial system characterization of system dynamics by a
state-and-transition model and ecosystem service characterization The staes are described in Box 1
Provisioning services
Regulation services
Forage & fodder
Fire hazard
Quality of fodder
C sequestration
Surface water quantity
Alleviation of GHG emissions
Production stability
Erosion protection + maintenance of soil fertility
Rural microclimatic regulation
Salinity control
Biodiversity component
Potential for P level decrease
Eucalypt health
Control of soil acidification
Native ground layer regen
Surface water quality
Genetic resources
Restoration potential
Cultural services
Resistance to new weed invasions
Woodland aesthetics
Figure 4. Bundles of adaptation services associated with increasing climate change and variability from
historical and current trends (top) to more severe transition to a semi-arid climate (bottom)
Figure 5. Characterisation of the ecosystem services, and semi-quantitative assessment of their relative
importance, for littoral rainforests, comparing intact forest with the effects of alternative drivers in scenarios
Figure 6. Characterisation of the ecosystem services, and semi-quantitative assessment of their relative
importance, for montane ash forests of the Australian Alps and Southeastern Highlands comparing current
values of ash forests with values under transition (scenario 3) and transformation (scenario 4) scenarios with
greatly increased fire frequency
Figure 7. Characterisation of the ecosystem services, and semi-quantitative assessment of their relative
importance, for floodplains and wetlands of the Murray-Darling Basin under three scenarios of river flow
regimes: historical, without river regulation and water resource development; under current regulation and
development and future, under climate change including Basin Plan water. Dots = adaptation services related
to future options
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Social-ecological systems underpinning nature-based solutions (NbS) must be resilient to changing conditions if they are to contribute to long-term climate change adaptation. We develop a two-part conceptual framework linking social-ecological resilience to adaptation outcomes in NbS. Part one determines the potential of NbS to support resilience based on assessing whether NbS affect key mechanisms known to enable resilience. Examples include social-ecological diversity, connectivity, and inclusive decision-making. Part two includes adaptation outcomes that building social-ecological resilience can sustain, known as nature's contributions to adaptation (NCAs). We apply the framework to a global dataset of NbS in forests. We find evidence that NbS may be supporting resilience by influencing many enabling mechanisms. NbS also deliver many NCAs such as flood and drought mitigation. However, there is less evidence for some mechanisms and NCAs critical for resilience to long-term uncertainty. We present future research questions to ensure NbS can continue to support people and nature in a changing world. Expected final online publication date for the Annual Review of Environment and Resources, Volume 47 is October 2022. Please see for revised estimates.
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The concept of resilience has evolved considerably since Holling's (1973) seminal paper. Different interpretations of what is meant by resilience, however, cause confusion. Resilience of a system needs to be considered in terms of the attributes that govern the system's dynamics. Three related attributes of social-ecological systems (SESs) determine their future trajectories: resilience, adaptability, and transformability. Resilience (the capacity of a system to absorb disturbance and reorganize while undergoing change so as to still retain essentially the same function, structure, identity, and feedbacks) has four components-latitude, resistance, precariousness, and panarchy-most readily portrayed using the metaphor of a stability landscape. Adaptability is the capacity of actors in the system to influence resilience (in a SES, essentially to manage it). There are four general ways in which this can be done, corresponding to the four aspects of resilience. Transformability is the capacity to create a fundamentally new system when ecological, economic, or social structures make the existing system untenable. The implications of this interpretation of SES dynamics for sustainability science include changing the focus from seeking optimal states and the determinants of maximum sustainable yield (the MSY paradigm), to resilience analysis, adaptive resource management, and adaptive governance.
Eucalypt ash type forests comprise 10% of the total forest estate in Victoria. On a unit area basis average annual streamflows are about four times higher than those from mixed eucalypt species forest which comprises about 80% of the forest estate. Research conducted by the Melbourne and Metropolitan Board of Works (MMBW) has shown that the streamflow yield from ash type forest is correlated with stand age. Theoretical case studies for three catchments carrying ash type forests show, on the basis of revenue received in royalty for timber and payments for bulk water, that the benefits of long rotation length can be greater than the costs incurred by loss of wood production. Victorian government policies for the timber industry emphasise that water production values must be considered during the planning and operational stages of timber production. Past policies of excluding commercial timber harvesting from the catchments directly managed by the Melbourne and Metropolitan Board of Works are being maintained with the aim of stream flow yield and quality protection. -from Authors
Eucalypts encounter a wide range of severe disturbances such as extensive defoliation by insects, major structural damage from cyclonic winds, as well as foliage and bark loss during drought and fire. Most healthy, mature eucalypts are not killed by these events, but regenerate vegetatively. With increasing intensity of disturbance, resprouting first occurs from the accessory buds in the small-diameter branchlets of the crown, followed by the epicormic buds in the medium- and large-diameter branches and stems, and then from the buds of the lignotuber. All these modes of regeneration are ultimately dependent on preventitious buds and, thus, the present review concentrates on axillary buds, their subsequent development into epicormic or lignotuber buds and their degree of protection from fire. The eucalypts have remarkably abundant, well protected and anatomically distinctive bud-forming structures in their leaf axils, branches, stems and lignotubers. These structures are quite consistent across this large genus, but are generally different from resprouting structures in many other plants. From an anatomical perspective, these structures seem best adapted to regeneration after fire, rather than damage from insects, storms or drought and this also correlates with ecological observations. On a worldwide basis, the eucalypts are some of the most successful post-fire resprouters, especially epicormic resprouting after medium- and high-intensity fires. Given the apparent ecological advantages of epicormic resprouting (the rapid reestablishment of extensive leaf area while simultaneously shading basal resprouters and seedlings) this could be an important factor in the success of eucalypts in Australia. Recent phylogenetic analysis has indicated a long relationship between eucalypts, fire and bud structures that facilitate resprouting.