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Spatial Restoration Ecology: Placing

Restoration in a Landscape Context
BEN L. GILBY , ANDREW D. OLDS, ROD M. CONNOLLY, CHRISTOPHER J. HENDERSON, AND
THOMAS A. SCHLACHER

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Researchers on conservation planning and practice have increasingly recognized and adopted the pivotal role of landscape attributes in shaping
the effectiveness of protected areas. However, the degree to which these concepts have been integrated into habitat restoration projects has not
been quantified. We reviewed the global literature and found that landscape context was considered in fewer than one in eight restoration
projects in the selection of restoration sites (11% of 472 projects). This figure was remarkably similar across terrestrial (10% of 243 projects),
marine (13% of 89), and freshwater (13% of 164) ecosystems. Of the 54 restoration projects in which landscape context was considered in site
selection, in just over half (56%), animal populations were reported to be larger or more diverse than in control areas. Tighter integration of
concepts from spatial ecology and systematic conservation planning into restoration practice could improve the design, optimize placement, and
enhance the ecological effectiveness of restoration projects in all ecosystems.

Keywords: connectivity, design, effectiveness, habitat restoration, landscape ecology

E stablishing nature reserves and restoring  

ecosystems   are complementary approaches in con-
servation (Soule 1985, Holl et  al. 2003, Pressey et  al. 2007,
are, therefore, of higher ecological value for animals than
others because they differ in terms of their food resources,
refuge value from weather or predators, accessibility to dis-
Palmer et al. 2016) that require significant financial invest- persal pathways, and numerous other ecological properties
ment (Wilson et al. 2006, De Groot et al. 2013). Maximizing that help to shape the fitness of individuals, demographics of
the net returns of these investments, in terms of both bio- populations, and the composition of assemblages (Pittman
diversity and ecosystem protection, is therefore sensible et al. 2011).
(Halpern et  al. 2013, Possingham et  al. 2015). Common Globally, across multiple ecosystems and realms, it has
strategies for enhancing conservation returns are to place been established that placing conservation areas strategi-
reserves in areas that are threatened, support high biodi- cally within landscapes can have synergistic benefits for
versity, or incur lower social and economic costs (Halpern their ecological effects on both ecosystems and the animals
et  al. 2013) or to protect or restore sites that might have that inhabit them (Rudnick et  al. 2012, Olds et  al. 2016).
synergistic benefits for conservation (Thomson et al. 2009, For example, many marine fish move among coral reefs,
Moilanen et al. 2011). seagrass meadows, and mangrove forests throughout their
Conservation and restoration areas, by their very nature, lives, and both fish abundance and diversity are often great-
are positioned within heterogeneous landscapes, which est inside marine reserves that conserve these ecosystems
comprise multiple ecosystems of different sizes and shapes and the pathways that link them across landscapes (Mumby
and with varying degrees of interconnectedness (Forman 2006, Edwards et al. 2009, Olds et al. 2016). Because some
1995, Olds et al. 2018). The landscape context can strongly of these fish perform important ecological functions (e.g.,
influence several key biotic attributes of a site (Holl et  al. herbivory), protecting connections among coral reefs, sea-
2003). Many of these attributes are of direct conservation grasses, and mangroves can also help improve the spatial
and restoration concern (e.g., population dynamics, biodi- of these ecosystems to disturbance (e.g., Magris et al. 2014,
versity, productivity; Ward et al. 1999, Bunn and Arthington Olds et  al. 2014). Furthermore, the benefits of placing
2002), across terrestrial (Hanski and Saccheri 2006), marine reserves into networks to maximize the exchange of indi-
(Micheli and Peterson 1999) and freshwater (Wiens 2002) viduals, matter, and energy among them are widely recog-
realms. Animals respond strongly to landscape configura- nized (e.g., Arturo Sánchez-Azofeifa et  al. 2003, Harrison
tion because most move throughout landscapes, and some et al. 2012, Hermoso et al. 2016). Reserves that are placed
use different habitats throughout their lifecycles. Some sites in areas with greater connectivity among populations,

BioScience 68: 1007–1019. © The Author(s) 2018. Published by Oxford University Press on behalf of the American Institute of Biological Sciences. All
rights reserved. For Permissions, please e-mail: [email protected].
doi:10.1093/biosci/biy126 Advance Access publication 14 November 2018

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among ecosystems, or with other reserves therefore often restoration for the whole ecosystem or landscape (Howe
perform better than reserves with impoverished landscape and Martinez-Garza 2014, Jones and Davidson 2016). For
connections (Stoms et  al. 2005, Ribeiro et  al. 2009, Olds example, restoring to enhance connectivity between habitat
et al. 2016). fragments can help with restoring metapopulation dynam-
Landscape context can have similarly positive ecological ics (Montalvo et  al. 1997, Fischer and Lindenmayer 2007).
effects on the performance of restoration projects (Metzger Increased consideration of metapopulation structure and
and Brancalion 2016). Restored ecosystems with strong connectivity can serve to enhance the available genetic pool
connections to other habitat patches, of either the same or of animal populations, thereby potentially increasing their
different ecosystem types, are more likely to be settled by fitness (Baguette et  al. 2013), reducing population extinc-
animals and to receive larger subsidies of matter and energy tion risk (Reed 2004), and increasing resilience to exog-
from adjacent ecosystems. This landscape context modifies enous disturbances (Etienne 2004). Such considerations are
the distribution, abundance, and diversity of animals and now often considered in the placement of marine reserves,
plants across landscapes (Fahrig 2001, Lees and Peres 2008, especially in cases in which fisheries enhancement is a goal

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Pottier et  al. 2009, Hodgson et  al. 2011) and potentially (Gerber et al. 2003, Puckett and Eggleston 2016). However,
improves the ecological function of restored habitats (da the regularity with which landscape context more broadly is
Silva et al. 2015, Jones and Davidson 2016). considered in restoration site selection and how this affects
Modern algorithms for reserve selection explicitly incor- inhabiting animal populations have, however, not been
porate the importance of positioning within the broader established (Metzger and Brancalion 2016). Consequently,
landscape (Hilty et  al. 2006, Rudnick et  al. 2012, Magris the lines of evidence required for practitioners to properly
et al. 2016, Weeks 2017). These algorithms have been devel- justify the integration of principles from landscape ecology
oped, now, to the point that system- and species-specific into the design of individual restoration projects have not
data are being used in association with commonly used been established.
modeling techniques (e.g., MARXAN, Zonation, and net- In this study, we identified restoration projects from the
work models), broadly across regions, and within individ- global primary literature and assessed the degree to which
ual systems (Kool et al. 2013, Engelhard et al. 2017, Weeks landscape context was used as a criterion to help guide the
2017). Whereas the selection of locations for reserves is selection of sites for restoration when the restoration of asso-
often guided by spatial concepts from landscape ecology, ciated animal populations is also a principle goal and under
including connectivity and landscape context (Margules which scenarios this is most likely to occur. We chose habitat
and Pressey 2000, Sarkar et  al. 2006, Almany et  al. 2009), restoration for animal populations as the focus because ani-
site selection for restoration appears to adopt these princi- mals move throughout a landscape, and so spatial metrics
ples less frequently (Hodgson et al. 2011). However, spatial are likely to be investigated first for these over other aims
prioritization is not widely used in restoration (e.g., Adame for restoration. We conducted literature searches using a
et  al. 2015, Ikin et  al. 2016), so, there have been recent structured literature classification framework. We reviewed
calls for tighter integration of the spatial principles from the literature using multiple terms used for ecological resto-
landscape ecology and conservation biology (Wiens and ration and a suite of ecosystem- and animal-specific phrases.
Hobbs 2015, Audino et al. 2017) to both inform the design We then categorized projects into those in which landscape
of restoration projects, and guide the selection of restora- context was considered in site selection and those in which it
tion sites (Jones and Davidson 2016, McAlpine et al. 2016). was not and further categorized projects into environmental
This is surprising because there have long been calls for the realms, ecosystems, and the animal communities they were
more strategic placement of restoration across landscapes, designed to enhance. We conclude by outlining how greater
including in some important global restoration guidelines uptake of fundamental principles gleaned from landscape
and policies (SER 2008, Keenleyside et al. 2012). ecology can improve restoration success.
The landscape context of restoration sites is likely to be
one of the most important factors influencing the outcomes Methods
of restoration investments, particularly for projects that are Restoration projects were identified using targeted literature
intended to enhance animal populations (figure 1; Miller searches and a structured literature classification framework
and Hobbs 2007, Moreno-Mateos et al. 2012). Because res- (figure 2). All literature searches were conducted using
toration sites usually require colonization by animal popula- the Web of Knowledge database in July 2017. The initial
tions from other habitat patches, the degree of connectivity search was for the term—((“habitat restoration” or “eco-
between restoration sites and other patches of existing habi- logical restoration” or “ecosystem restoration”) and (fauna or
tat is crucial (Scott et al. 2001, Hodgson et al. 2009, Hodgson inhabit* or animal* or biodivers* or wildlife or fish)), result-
et al. 2011). Restoration sites might be placed in areas that ing in 2183 articles returned and 333 articles downloaded
are considered appropriate for the establishment and growth for potential inclusion. On the basis of the composition
of habitat-forming species but that are so isolated or poorly of this initial shortlist, follow up searches included com-
connected that animal populations respond to a lesser ponents of the initial search, as well as wetland*, seagrass,
degree, thereby reducing the overall values of the habitat oyster*, fish*, mangrove*, forest*, grass*, insect*, invert*,

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method of literature searching was not

designed to be exhaustive (i.e., system-
atic, comprehensive) in identifying all
restoration projects in the literature
that also sought to enhance animals. It
was, however, designed to give the best
possible representation of restoration
projects across multiple environmental
realms and ecosystems, and within the
constraints of our search terms.

Inclusion criteria and data extracted. We

followed a set classification framework

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for all studies identified by literature
searches (figure 2). Restoration p ­ rojects
were classified as also targeting the res-
toration of animals to restored habitats
if the study tested for effects of the res-
toration effort for animals or if the arti-
cle states explicitly that a goal of the
restoration effort was also to increase
the abundance or diversity of animals.
We concluded that the landscape
context of restoration sites was consid-
ered in a restoration project if the proj-
ect specifically chose sites or systems
over other areas because of their posi-
tion within the landscape, especially if
the restoration project was intended to
reconnect habitat fragments; included
the position of sites relative to patches
of either similar or different habitat
types; or incorporated an aspect or
understanding of animal movement,
dispersal capacity, or connectivity into
Figure 1. The landscape context of restoration sites has significant consequences
site selection. We did not consider
for the number and types of animals that inhabit restored ecosystems on land
a study to have included landscape
(a), in the sea (b), and where hydrologic connectivity (c) is vital, especially in
context if its sites were chosen sim-
freshwater ecosystems, and wetlands. For example, isolated patches of restored
ply on the basis of existing impacts
grassland (a, left) might perform less effectively for animals than restored
(e.g., restoration of oil spill sites, ero-
grasslands close to nearby alternate habitats (right). In the sea (b), connectivity
sion mitigation); to revegetate mined
between multiple habitats is often an important consideration because fauna
land; using “habitat suitability” models
use multiple habitats throughout their lifecycle. In the present study, restored
for growth of habitat forming species
oyster reefs in the Noosa River, Australia, in two seascape contexts: at a
when the position of sites relative to
distance from nearby mangroves (left) and very close to nearby mangroves
other habitat patches was not also con-
and seagrasses (right). Hydrologic connectivity (c) is limited where weirs or
sidered; or randomly using a number
dams can restrict flow of propagules and animals (left). These challenges can
generator, GIS, or similar method.
be overcome by considering connectivity between water bodies and movement
Restoration projects were grouped
of fauna and implementing restoration interventions such as waterway
according to country, region and habi-
remeandering and fish ways (right). Images courtesy of Cassandra Duncan,
tat restored, and we pooled studies
Matt Lavin (CC BY SA2.0), Nick Carson.
that reported on the same restoration
project to avoid overlap in counting
bird*, mammal*, reptil*, stream*, river*, lake*, and coral*. projects. We also identified the types of animals (in broad
All review articles and meta-analyses identified by lit- groups; e.g., birds, mammals, fish, insects, or any combina-
erature searches were downloaded and scanned by the first tion of these) that each restoration project was intended to
author for additional articles that could be included. This enhance on the basis of their stated outcomes. For projects

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(n = 14, 3%), and the United Kingdom

(n = 14, 3%). The most common eco-
systems restored were forests (n = 134;
28%), followed by streams (n = 87; 18%)
and grasslands (n = 67; 14%). Overall, in
37% of the studies, it was stated explicitly
that the landscape context of restoration
sites was important in determining the
effectiveness of restoration for animals,
irrespective of whether they actually
implemented landscape concepts into
site selection.
Just over half of our data set com-

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prised terrestrial projects (n = 243, 52%;
figure 3). Forests (n = 36, 15% of ter-
restrial projects) and grasslands (n = 34,
14%) in the United States, and forests in
Australia (n = 30, 12%) were the eco-
systems most often targeted for restora-
tion (figure 4). Terrestrial restoration
projects most often sought to enhance
populations of birds (n = 107, 44%),
insects (n = 53, 22%), and mammals
(n = 23; 9%). Freshwater projects rep-
resented roughly one-third of our data
set (n = 151, 32%; figure 3). Streams
and rivers in the United States (n = 47,
31% of freshwater projects) and Finland
(n = 11, 7%) were the ecosystems most
often targeted for restoration (figure 5).
Freshwater restoration projects most
often sought to enhance populations of
Figure 2. Literature classification framework and decision flow chart. Used to fish (n = 89, 59%) and macroinverte-
identify restoration projects from the global primary literature. brates (n = 60, 40%). Marine projects
represented roughly one-sixth of our
data set (n = 78, 17%; figure 3). Oyster
that did incorporate landscape context into their site selec- reefs (n = 30, 44% of marine projects) and saltmarshes
tion, we identified whether the restoration effort resulted (n = 10, 13%) in the United States were the ecosystems most
in the stated aims being achieved for that species (i.e., often targeted for restoration (figure 6). Marine restoration
“positive” effects on fauna). projects most often sought to enhance populations of fish
Studies were classified into environmental realms (terres- (n = 63, 81%), nektonic crustaceans (n = 21, 27%) or macro-
trial, marine, or freshwater) according to their location and invertebrate infauna (n = 20, 26%).
the animal species being restored. For example, wetlands
can be freshwater, marine, or terrestrial; a restored wetland Effects of landscape context in restoration. Across all realms,
in saltwater intended to restore habitat for fish is classified 11% of projects (54 of 472) included landscape context
as marine, whereas a wetland in saltwater enhancing birds or as a criterion in the selection of restoration sites (table 1,
insects is classified as terrestrial. ­figure 3). The integration of landscape context in restoration
projects was remarkably similar across realms: terrestrial
Results (10% of 243 projects), marine (13% of 89), and freshwater
The final database included 472 restoration projects (13% of 164). In terrestrial ecosystems, the projects in which
described in English language journals that sought to landscape context was used most often were those in peat-
enhance animal populations from restoration efforts (for lands (100% of peatland projects, but only one project was
the full list of included articles, see supplemental table S1). identified), wetlands (14% of wetland projects), and forests
We identified restoration projects in 34 countries; most proj- (10%; figure 4b) and those that targeted amphibians (29%)
ects were from the United States (n = 212, 45%), followed and mammals (13%). Freshwater projects that considered
by Australia (n = 44, 9%), Finland (n = 20, 4%), Sweden landscape context most often were those in rivers (40%)

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of the same habitat. or with other

habitats. The projects in which positive
effects of connectivity were reported
on animals were common across ter-
restrial (84% of 25 projects), marine
(60% of 10), and freshwater (68% of
19) landscapes. In the majority of these
(69% of 54 projects), connectivity was
considered between patches of similar
habitats, and in fewer studies were
the effects of connectivity considered
with adjacent alternate habitats (3% of
54 projects, and one in each environ-

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mental realm; table 2). The potential
ecological effects of other landscape
concepts have, however, rarely been
tested with empirical data (26% of 54
projects), with most research limited
to the effects of hydrology in aquatic
ecosystems (table 2).
In most projects (89% of 472 proj-
ects), landscape context was not listed
as a criterion in the selection of resto-
ration sites (table 3, figure 3). When
the spatial attributes of landscapes were
not considered in restoration studies,
the projects were most often at mine
sites (15 projects, 6% of terrestrial proj-
Figure 3. Summary of included studies. Numbers in white circles in the center ects), located to minimize the effects
of each chart denote the number of studies in each category. (a) The total of erosion on freshwater ecosystems (4
number of studies included in the database, divided by environmental realm. projects, 3% of freshwater projects), or
(b) The proportion of studies in which the spatial context of restoration sites placed randomly in landscapes (12 proj-
was considered during ecosystem restoration projects. (c) The proportion of ects, 4% of terrestrial projects and 1% of
studies in which landscape context was considered during the restoration design freshwater projects). For most studies,
process in which the direction of outcomes was reported for animals. however, we were not able to identify
how restoration sites were selected from
the descriptions provided (74% of 472
and streams (7%; figure 5b) and those targeting amphib- projects). This was a common trend across terrestrial
ians (40%) and macroinvertebrate infauna (13%). In marine (177, 73% of all terrestrial projects), marine (58, 75% of all
systems, the consideration of landscape context was most marine projects), and freshwater (113, 75% of all freshwater
prevalent in marine wetlands (30%) and in seagrass (29%; projects) realms.
figure 6b) and when restoration targeted birds (33%) or The first restoration studies in which landscape context
macroinvertebrate infauna (15%). was explicitly considered when selecting restoration sites
There was good evidence to suggest that incorporating were published in 1996 for freshwater ecosystems, 2001
landscape context into the selection of restoration sites for terrestrial ecosystems, and 2004 for marine ecosystems
resulted in positive outcomes for animals, with 56% of the (figure 7a). The proportion of studies in which landscape
projects (30 of 54 projects) in which landscape context was context was considered has remained highly variable, with
considered showing positive outcomes for animals (table 2, no clear trend between years over the past three decades
figure 3). In the remaining 44% (24 of 54) of the projects, (figure 7a). By contrast, there has been a sharp increase in
whether landscape context affected animals directly was the integration of spatial concepts from landscape ecology
not explicitly tested. The projects in which positive effects into the wider fields of biology and ecology during this
of landscape context on animals were reported were more period (figure 7b).
common on land (72% of 25 projects) and in the sea (60% of
10) than in freshwater (32% of 19). Discussion
Most included studies focused on the ecological effects The ecological outcomes of restoration projects can, in many
of connectivity (74% of 54 projects), either between patches cases, be improved by placing sites at locations in landscapes

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(a)

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Coastal dunes (n = 1) Forest (n = 134) Wetland (n = 37) 1 study
9 studies
Marsh (n = 2) Peatlands (n = 1) Grassland (n = 67) 3 studies

(b) Consider spatial ecology Do not consider spatial ecology

100

90
1
80
Percentage of studies

70

60

50

40

30

20 5
13 6
10
0 0
0
Coastal Dunes Forest Grassland Marsh Peatlands Wetland

Habitats

Figure 4. (a) Global distribution of terrestrial restoration projects that aimed to restore animal populations, assemblages,
or diversity. (b) Of the 243 terrestrial restoration projects, only 25 (10%) considered landscape context in their design
phase (table 2). The integration of landscape attributes did, however, differ among terrestrial ecosystems. Numbers above
bars indicate the number of projects that considered landscape context in site selection.

to maximize the recruitment of individuals to newly created considered in restoration projects (12% of studies reviewed).
habitats (Jones and Davidson 2016). Whereas it is widely Furthermore, we found no evidence that the consideration
appreciated that the landscape context of restored habitats of landscape context in restoration has increased over the
can shape the success of restoration projects, the principles past three decades. This is surprising, because spatial con-
of landscape ecology remain rarely considered in restora- cepts have been more widely adopted in conservation during
tion decisions. In this study, we show that a marked dis- the same period, and there are important global restoration
crepancy exists between the stated importance of landscape policy documents that advocate for its inclusion (SER 2008,
context for restoration (37% of the studies reviewed) and Keenleyside et al. 2012). This result could eventuate because
the extent to which the spatial properties of landscapes are of underreporting in the description of restoration sites.

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(a)

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Lake (n = 6) River (n = 35) Spring (n = 1) 1 study
9 studies
Stream (n = 87) Wetland (n = 22) 3 studies

(b) Consider spatial ecology Do not consider spatial ecology

100

90

80
Percentage of studies

70

60

50

40

30 8 5
20
6
10 0
0
0
Lake River Spring Stream Wetland

Habitats

Figure 5. (a) Global distribution of freshwater restoration projects that aimed to restore animal populations, assemblages,
or diversity. (b) Of the 151 freshwater restoration projects, only 19 (13%) considered landscape context in their design
phase (table 2). The integration of landscape attributes did, however, differ among freshwater ecosystems. Numbers above
bars indicate the number of projects that considered landscape context in site selection.

However, restricting our use of only the primary literature principles (e.g., peatlands); however, little can be gleaned
offers some safeguard against substantial under-reporting of from these few examples from habitats with fewer restora-
the design process. tion projects. There were also no clear trends in habitats that
We found no clear reason the uptake of spatial ecological have been studied extensively in terms of landscape context.
principles varied across different habitats, likely because the For example, the study of landscape context on coral reefs
uptake was consistently low across all habitats. The excep- is highly advanced (Pittman and Olds 2015), but no coral
tions were habitats in which few restoration projects were restoration project has yet included it in restoration site
identified but had very high uptake of landscape context selection.

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(a)

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Coral reefs (n = 3) Boulder fields (n = 1) Kelp (n = 1) 1 study
Mangroves (n = 9) Saltmarsh (n = 12) Oysters (n = 34) 9 studies
3 studies
Estuary (n = 1) Seagrass (n = 7) Wetland (n = 10)

(b) Consider spatial ecology Do not consider spatial ecology

100

90
Percentage of studies

80

70

60

50

40
2 3
30

20 1 1 3
10 0
0 0 0
0
Coral reef Boulder fields Macroalgae Mangroves Saltmarsh Oysters Estuary Seagrass Wetland

Habitats

Figure 6. (a) Global distribution of marine restoration projects that aimed to restore animal populations, assemblages,
or diversity. (b) Of the 78 marine restoration projects, only 10 (13%) considered landscape context in their design phase
(table 2). The integration of landscape attributes did, however, differ among marine ecosystems. Numbers above bars
indicate the number of projects that considered landscape context in site selection.

We identified several key types of spatial metrics order of magnitude less than connectivity with similar habi-
that are considered in the selection of restoration sites. tat patches. This relatively low number of studies focused on
Predominantly, researchers assessed connectivity with simi- hydrologic connectivity is surprising because reconnecting
lar patches of habitats (e.g., Angelieri et al. 2016, Derhe et al. water bodies hydrologically is an important focus in wet-
2016), which is unsurprising, given that the role of restora- lands and aquatic ecosystems (Kondolf et al. 2006, Jackson
tion is often to enhance or reconnect threatened habitats, and Pringle 2010). Similarly, concepts regarding the spread
so considering nearby patches of this habitat is usually of propagules by wind or other mechanisms received little
important. Concepts such as hydrologic connectivity and attention in the studies we identified. This means that there
connectivity with alternative habitats (i.e., those other than are several key concepts and metrics within spatial ecol-
the habitat being restored) were, however, considered an ogy that have yet to be properly studied in restoration but

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Table 1. Summary of restoration projects in terrestrial, freshwater and marine realms for all restoration studies.
Terrestrial Freshwater Marine All realms
Percentage of Number of Percentage of Number of Percentage of Number of Percentage of Number of
projects projects projects projects projects projects projects projects
51 243 32 151 17 78 100 472

Table 2. Summary of restoration projects in terrestrial, freshwater and marine realms for restoration studies in which
landscape context was considered in the design phase.
Terrestrial Freshwater Marine All realms
Landscape context Percentage Number of Percentage Number of Percentage Number of Percentage Number of
considered in design of projects projects of projects projects of projects projects of projects projects

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10 25 13 19 13 10 11 54
Positive effects 7 18 4 6 8 6 6 30
No effects reported 4 9 7 11 5 4 5 24
Connectivity with 8 20 8 12 6 5 8 37
patches of similar
habitat
Connectivity with other 1 1 1 1 1 1 1 3
habitats
Hydrology 1 2 2 3 4 3 2 8
Other 1 2 2 3 1 1 1 6
Note: Values in bold indicate total number, and proportion of the percentage of the total number of studies that fall into each category.

Table 3. Summary of restoration projects in terrestrial, freshwater and marine realms for restoration studies in which
landscape context was not considered in the design phase.
Terrestrial Freshwater Marine All realms
Landscape context
NOT considered in Percentage Number of Percentage Number of Percentage Number of Percentage Number of
design of projects projects of projects projects of projects projects of projects projects
46 218 28 132 14 68 88 417
Located on a mine site 7 15 0 0 0 0 4 15
Minimize erosion effects 0 0 3 4 0 0 1 4
Random placement 5 10 2 2 0 0 3 12
Indiscernible 81 177 86 113 85 58 83 348
Other 7 16 10 13 15 10 9 39
Note: Values in bold indicate total number, and proportion of the percentage of the total number of studies that fall into each category.

that are likely to be helpful in placing restoration efforts. likely instructive for the enhancement of animal populations
Broadening the scope of the types of considerations and across landscapes (Montalvo et  al. 1997, McAlpine et  al.
metrics used to place restoration should be a key focus of 2016). Some counterintuitive results were uncovered for
restoration researchers and practitioners alike. oyster reefs. Higher connectivity (in this case, simply prox-
Most projects that incorporated concepts from landscape imity) with adjacent habitats is usually viewed as beneficial
ecology into their design and site selection showed positive for coastal marine organisms (Olds et  al. 2018). However,
outcomes for animals, albeit with some variation among restored reefs in North America contained higher fish abun-
terrestrial (72% positive outcomes), marine (60%), and dance when further from existing marshes because they
freshwater (31%) realms. For example, in freshwater ecosys- provided new complex habitats on previously low-complexity
tems, wetland restorations for amphibians reported marked muddy areas (Grabowski et al. 2005). Conversely, higher con-
increases in adult breeding populations within 3 years of nectivity between extant reefs and restored reefs was viewed
restoration of ponds that were highly connected to each other as a positive influence on inhabiting fauna in other studies
(Petranka et al. 2003), an effect that was inconsistent for other (Gregalis et al. 2009). Although the performance of projects
similar freshwater restoration projects in which context was or sites that did incorporate spatial context into their design
not considered (Shulse et al. 2012). These sorts of consider- and those that did not (e.g., randomly or for some other
ations of adult breeding metapopulations were rare but are ecological reasoning) has not been assessed, these findings

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in a landscape context. Alternatively, res-

toration sites might be placed in areas
to limit the potential for conflict with
other human uses (e.g., fishing, farming,
recreational use, transport) or legislation
(e.g., mooring areas, other forms of con-
servation; Pressey and Bottrill 2008), and
these might not be optimal for the resto-
ration of ecosystems or their inhabiting
animals (Fahrig 2001, Pottier et al. 2009).
Both approaches might result in restora-
tion sites being unintentionally restricted
to locations that provide poor habitat

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values for animals (cf. “residual” conser-
vation areas; Pressey et al. 2000, Pressey
and Bottrill 2008) and that therefore
have the potential to yield suboptimal
restoration returns for animals.
There are several key research areas
that should be promoted to assist in
better prioritizing restoration across
landscapes. We raise in this review a
paucity of restoration projects incorpo-
rating systematic conservation planning
regimens in the selection of restoration
sites. Studies should be conducted to
determine the validity of using system-
atic conservation practices (Margules
Figure 7. Summary of published studies that have considered landscape context and Pressey 2000) for restoration across
in (a) restoration (separated for terrestrial, marine and freshwater realms) environmental realms (research ques-
and (b) across the fields of biology and ecology. Data extracted from the Web tion 1 in box 1). Despite this review
of Knowledge the key words restoration (“habitat restoration” or “ecological uncovering a diversity of ecosystems
restoration” or “ecosystem restoration”), “biology” or “ecology,” and and animals that have been the focus
“landscape ecology” or “spatial ecology.” 2017 is part year only. of restoration, there are several ecosys-
tems (research question 2 in box 1) and
animal groups (research question 3 in
suggest that the spatial properties of landscapes might have box 1) that remain underrepresented within the literature.
broad and largely unrecognized effects on the success of eco- For example, very few studies have assessed the capac-
logical restoration projects. ity for coastal dunes to be restored for animals, especially
Ecological restoration is conducted for many purposes, birds, despite beach ecosystems being regularly restored
and not all types of restoration are designed to benefit ani- for these purposes (Maslo et  al. 2011, Maslo et  al. 2012).
mals. For example, many restoration projects are intended To date, most work on the effects of landscape context on
to restore whole ecosystems (e.g., whole of lake restoration), the outcomes of restoration has been focused specifically
to reverse a particular type of impact (e.g., mine site reha- on how the metric of connectivity (especially proximity
bilitation, oil spill remediation; e.g., Brady and Noske 2010), and isolation) between patches of similar habitat affects the
or to limit the ecological effects of ongoing disturbances, abundance of animals. Restoration sites are always posi-
such as erosion, sedimentation, nutrient enrichment, or tioned within heterogeneous landscapes of multiple habitat
pollution (e.g., restoring oyster reefs to reduce shoreline types, so further research should be conducted on how
retreat or forests to reduce gully erosion; e.g., Piazza et  al. connectivity with nearby patches of different habitat types
2005). Under these circumstances, restoration sites might affects the success of restoration more broadly (research
be placed in areas that are considered appropriate for the question 4 in box 1), as well as the condition of animal
establishment of habitat-forming species but that might be populations themselves, where this is a key restoration
isolated from other remnant patches of the same habitat that goal (research question 5 in box 1). Similarly, there are
serve as sources for the recruitment of animals. Because the several landscape metrics, beyond simple distance-based
locations of these types of projects are often fixed, it might connectivity metrics (especially Euclidean distance), that
not always be possible for restoration decisions to be placed might assist in developing more robust and representative

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Box 1. Priority research questions for integrating landscape principles into restoration ecology.

1. re the principles of systematic conservation planning (Margules and Pressey 2000) directly translatable to all restoration
A
projects and, if not, which principles need to be tailored specifically for restoration?
2. or which habitats does landscape context matter most, and which metrics, and which connections, should be prioritized
F
for individual habitat types?
3. or which animal groups does landscape context matter most, and which metrics, and which connections, should be priori-
F
tized for individual animal groups?
4. o what degree does considering connectivity with other habitats (i.e., interhabitat connectivity) affect the outcomes of
T
restoration projects (e.g., Unsworth et al. 2008)? To date, most research and focus in the literature has been on connectivity
between patches of the same habitat (i.e., intrahabitat connectivity).

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5. o what degree can incorporating metapopulation dynamics and connectivity into restoration planning enhance animal
T
population fitness, persistence, and resilience?
6. re there more thorough, or better, metrics that can be used to optimize the spatial placement of restoration (e.g., McGarigal
A
et  al. 2012)? To date, most work on the spatial metrics that influence restoration has focused on connectivity, especially
proximity (Euclidean) between similar habitat patches.
7. Is there consistency in the efficacy of spatial metrics across environmental realms? (e.g., Pittman et al. 2018)

models for incorporating landscapes ecology into restora- conservation benefits across the entire landscape, and some
tion. Landscape ecologists have developed several multi- potential sites are then eliminated because of other consider-
variate or multimetric variables that can be used to describe ations (e.g., extractive industries, tourism; Watts et al. 2009).
the complexity of landscapes for animals (e.g., McGarigal By contrast, restoration projects often start with a narrower
et  al. 2012). The efficacy of these metrics in restoration perspective focused on the specific ecosystem or habitat to
should be further investigated for different ecosystems be restored. Sites are then selected on the basis of their suit-
and target animals (research question 6 in box 1). Several ability to support the particular ecosystem of interest, with
authors have discussed the validity of landscape metrics and little consideration of the attributes of landscapes beyond
concepts across environmental realms (Pittman et al. 2018). the restoration site. Incorporating the lessons learned from
For example, there is some evidence to suggest that the best systematic conservation planning (i.e., goal setting, data-
metrics to describe spatial patterns in animal abundance based feedback loops, and improvements) and the prin-
might differ between the land and the sea because of the ciples of landscape ecology (i.e., the placement of sites in
movement of water bodies in aquatic ecosystems (Pittman heterogeneous land- and seascapes) into restoration should,
et al. 2018). Establishing the validity of these metrics across therefore, lead to significant improvements in the design,
environmental realms will assist in generalizing restoration placement, and ecological effectiveness of restoration proj-
planning regimens across realms (research question 7 in ects. We advocate a landscape-scale approach to restoration
box 1). Collectively, answering these priority research ques- and suggest that spatial restoration ecology should start with
tions will assist in establishing more effective regimens for the identification of ecosystems and habitats that are in the
systematic landscape restoration across all environmental greatest need of restoration (which can be to be determined
realms, and provide the evidence that managers need to on the basis of the best historical information available).
support their decision-making processes. Sites for restoration should then be selected from all suitable
Systematic conservation planning has made great prog- locations within the landscape of interest to maximize their
ress in using diverse landscape characteristics and sophisti- potential ecological benefits for the ecosystems themselves,
cated algorithms to guide the design of protected areas and the animals and food webs they support, and the ecological
reserve networks (Margules and Pressey 2000, Moilanen functions and ecosystem services they provide. This spatial
et  al. 2009). The principles of landscape ecology and the approach to restoration ecology should, therefore, help to
techniques of conservation planning might be useful in the broaden both the scope and perceived ecological benefits of
design of restoration areas but have not been widely applied many restoration projects and might also improve returns
to introduce a crucial spatial element to restoration planning on investment across restored landscapes.
(Hodgson et  al. 2011). When selecting sites for a network
of reserves, the conservation planning process starts with Acknowledgments
a broad, landscape-scale perspective. Ecosystems, habi- The authors thank Nicholas Ortodossi and Sarah Thackwray
tats, and locations are selected for protection to maximize for assisting with the literature classification. This work was

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funded by Noosa Council and the Noosa Biosphere Reserve Gregalis KC, Johnson MW, Powers SP. 2009. Restored oyster reef location
Foundation. and design affect responses of resident and transient fish, crab, and
shellfish species in Mobile Bay, Alabama. Transactions of the American
Fisheries Society 138: 314–327.
Supplemental material Halpern BS, et al. 2013. Achieving the triple bottom line in the face of inher-
Supplemental data are available at BIOSCI online. ent trade-offs among social equity, economic return and conservation.
Proceedings of the National Academy of Sciences 110: 6229–6234.
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