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Journal ArticleDOI

Conceptual domain of the matrix in fragmented landscapes

01 Oct 2013-Trends in Ecology and Evolution (Elsevier)-Vol. 28, Iss: 10, pp 605-613
TL;DR: The conceptual domain of the matrix, defined as three core effects and their interaction with these five dimensions, provides a much-needed framework to underpin management of fragmented landscapes and highlights new research priorities.
Abstract: In extensively modified landscapes, how the matrix is managed determines many conservation outcomes. Recent publications revise popular conceptions of a homogeneous and static matrix, yet we still lack an adequate conceptual model of the matrix. Here, we identify three core effects that influence patch-dependent species, through impacts associated with movement and dispersal, resource availability, and the abiotic environment. These core effects are modified by five ‘dimensions’: spatial and temporal variation in matrix quality; spatial scale; temporal scale of matrix variation; and adaptation. The conceptual domain of the matrix, defined as three core effects and their interaction with these five dimensions, provides a much-needed framework to underpin management of fragmented landscapes and highlights new research priorities.

Summary (3 min read)

A matrix focus is now both important and possible

  • The patch-matrix model of landscapes [20] includes patches that are useful for conservation and the matrix in which the patches are embedded [21] (see Glossary).
  • Assumptions underpinning the patch-matrix model are reasonable in many situations, particularly in fragmented and relictual landscapes where there are patch-dependent species [22] [23] [24] .
  • The growth in knowledge about the matrix means it is now possible to develop a detailed synthesis of the mechanisms by which the matrix directly, or indirectly drives the distribution of patch-dependent species in space and time.
  • Habitat loss and fragmentation are the biggest threat to biodiversity globally [31] .
  • There is now a pressing need for a comprehensive theoretical framework of the matrix to guide the way scientists and land managers think about matrix ecology.

Movement and Dispersal. Matrix quality influences the outcome of movement into the matrix

  • Recent reviews report that movement between patches is enhanced as the matrix becomes structurally more similar to the remnant patches [40, 41] .
  • The matrix can influence immigration and emigration in other ways.
  • If a species does venture into the matrix, rapid movement through unfavourable habitat could enhance connectivity between separated habitat patches [42] .
  • Red squirrel Tamiasciurus hudsonicus populations thrived on pine-seeds in Canadian pine plantations.
  • With the possible exception of bees that can forage outside of the nesting patch [e.g. 14], evidence that patch-dependent species gather resources outside of the patch to support higher population densities inside the patch is limited [e.g. 46].

Abiotic environment. The matrix influences microclimate and disturbance regimes of patches.

  • The physical structure of the matrix is often different from habitat patches and can alter the environmental conditions within patches [19, 37] , particularly when treed landscapes are cleared [25] .
  • Changes to disturbance regimes in the matrix can also affect patch-dependent species.
  • Larger and more frequent fires can occur if there are more ignitions in the matrix [11] , or when the fuel structure in the matrix is changed by forest logging [11, 49] or by invasive grasses [17] .
  • Altered microclimate and disturbance regimes can advantage some species, often invasive exotic species [6, 17] , but disadvantage others, often species that depend on remnant vegetation [8] .
  • Increased disturbance associated with urban or mining landscapes can also drive local extinctions in patches [9, 10] .

Spatial scale. The extent of the matrix influences its impacts on patch-dependent species

  • The spatial scale of the matrix, including geographic extent and distance between patches (see Glossary), has an important effect on patch-dependent species.
  • If patch-dependent species exploit resources in the matrix [34] , a proportionally greater area of matrix to patch could increase the relative abundance of such resources.
  • Most edge studies disregard the scale of the adjacent matrix and so understanding of such effects is rudimentary.
  • By examining the extent to which changes 9 in population size were synchronous, Powney et al. [58] found that matrix permeability to dispersal had the strongest effect on movement between patches at intermediate distances.
  • Abiotic effects are highly dynamic [7] and change over time as a consequence of succession, seasonality, and changes in species composition, management and disturbance regimes.

Temporal scale. Demographic and dispersal rates influence responses to changes in the matrix

  • In poorly dispersing lichen species, forest succession through plantation harvest cycles can be too rapid for colonisation, particularly when the matrix is extensive [70] .
  • In contrast, where resources change gradually, dietary specialists can replace generalists as succession advances [73] .
  • A species response to the matrix can change over time.
  • As a research planning tool, it stimulates new ways of framing hypotheses about the matrix, including drawing attention to novel interactions among the dimensions and core effects (Box 5).

Figure 1. Matrix core effects

  • The matrix can influence species abundance, community composition and ecological processes within patches of native vegetation through three core effects associated with (i) movement and dispersal, (ii) resources provided within the matrix, and (iii) the abiotic environment of patches.
  • The matrix can also alter dispersal by acting as a barrier to emigration, or can promote dispersal leading to increased immigration.
  • The matrix can provide resources that allow non-patch species to breed and subsequently spill over into patches.
  • Each of these effects can have consequences for individual species, and subsequently for community composition (see Box 2 for a more detailed description of some pathways and Box 1 for consideration of species interactions).
  • Numbers indicate studies listed in the references that support parts of each pathway.

Figure 2. Five dimensions modify matrix core effects

  • The conceptual model of the matrix consists of the three core effects whereby the matrix influences patch-dependent species through effects associated with movement and dispersal, resource availability, and the abiotic environment.
  • This does not imply any priortity of effects (although difficult to draw, these could also be imagined as overlapping spheres encompassing the core effects, like electrons around an atom's nucleus).the authors.
  • The importance of patch characteristics and species interactions are well established (Boxes 1, 4).the authors.
  • In another example, increasing resources in the matrix (seeds in wet years) enabled seed-eating rodents to forage widely throughout the landscape [84] .
  • Competition-colonisation trade-offs or predator-prey patch dynamics [85] might also drive feedbacks between pathways in Figure 1 .

Box 2. New species colonise patches by multiple pathways

  • By defining three core effects , their conceptual model puts colonisation of patches into a mechanistic context.
  • Coffee plantations have received widespread attention as a matrix capable of supporting forest species [89] , but these plantations also provide resources for pest species.
  • Such spill-over edge-effects could be more widespread than is currently recognised in the literature [56, 90] .
  • Invasion changed the microclimate which reduced amphibian abundance and diversity [48] , along with effects on the invertebrate fauna [92] .
  • Key responses to the changing matrix include: (i) new species were recruited to the landscape because the pine matrix provided breeding habitat ; (ii) a illremnants , and; (iii) a habitatwith measured temporal changes in vegetation attributes as the patches responded to the changed abiotic conditions and management regime.

Box 4. The patch still matters

  • The matrix affects local populations through core effects associated with dispersal, the resource base and the abiotic environment, but patch dynamics are also strongly influenced by characteristics of the habitat patch itself.
  • The rate of change of habitat quality within patches could allow, for example, long-lived species to readily survive short-term changes in habitat quality [99] .
  • Patch size is often important, but spatial scale issues are more relevant when considering a matrix with multiple embedded patches.

Matrix

  • The matrix is an extensive land-cover with different types of landcover embedded within it .
  • The matrix does not provide for self-sustaining populations of some species, which are dependent upon the patches.
  • The matrix therefore, includes the extensive landcover types that patch-dependent species cannot sustainably live in.
  • This definition means that what is the matrix for some species, or was the matrix at one time, might not be at other times [15] or for other species [16] .

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Content maybe subject to copyright    Report

1
This is the accepted version of the article. For the final version visit 1
http://www.cell.com/trends/ecology-evolution/ 2
3
4
Conceptual domain of the matrix in fragmented landscapes 5
6
Don A. Driscoll, Sam C. Banks, Philip S. Barton, David B. Lindenmayer, Annabel L. Smith 7
8
ARC Centre of Excellence for Environmental Decisions, the National Environmental Research 9
Program Environmental Decisions Hub, Fenner School of Environment and Society, The 10
Australian National University, Canberra ACT 0200, Australia. 11
12
In extensively modified landscapes, how the matrix is managed determines many 13
conservation outcomes. Recent publications revise popular conceptions of a homogeneous 14
and static matrix, yet we still lack an adequate conceptual model of the matrix. Here, we 15
identify three core effects that influence patch-dependent species, through impacts 16
associated with movement and dispersal, resource availability and the abiotic environment. 17
These core effects are modified by five 'dimensions': (i) spatial and (ii) temporal variation 18
in matrix quality, (iii) spatial scale, (iv) temporal scale of matrix variation, and (v) 19
adaptation. The conceptual domain of the matrix, defined as three core effects and their 20
interaction with the five dimensions, provides a much-needed framework to underpin 21
management of fragmented landscapes and highlights new research priorities. 22
23

2
A matrix focus is now both important and possible 24
Biodiversity conservation often focusses on patches of native vegetation in a surrounding matrix 25
that is highly modified by agriculture or urbanisation [18, 19]. The patch-matrix model of 26
landscapes [20] includes patches that are useful for conservation and the matrix in which the 27
patches are embedded [21] (see Glossary). Assumptions underpinning the patch-matrix model 28
are reasonable in many situations, particularly in fragmented and relictual landscapes where 29
there are patch-dependent species [22-24]. However, the matrix surrounding remnant vegetation 30
can have a strong influence on species occurrence and spatial dynamics [25, 26] and can be more 31
important than the size and spatial arrangement of remnant patches [2, 27, 28]. The growth in 32
knowledge about the matrix means it is now possible to develop a detailed synthesis of the 33
mechanisms by which the matrix directly, or indirectly drives the distribution of patch-dependent 34
species in space and time. 35
36
Not only is such a synthesis possible, it is also urgent. The nature of the matrix has profound 37
implications for conserving biodiversity [28, 29]. Management of the matrix can limit or 38
exacerbate the impacts of habitat loss and fragmentation [30]. Habitat loss and fragmentation are 39
the biggest threat to biodiversity globally [31]. In highly modified landscapes, further loss of 40
remnant vegetation is limited because most of it is already gone, or because what remains is 41
legally protected [32, 33]. Where this is the case, modifying the matrix will be the major form of 42
landscape change in the future, and will therefore likely be the main process influencing 43
biodiversity conservation. There is now a pressing need for a comprehensive theoretical 44
framework of the matrix to guide the way scientists and land managers think about matrix 45
ecology. 46

3
47
While there has been much conceptual development in the habitat fragmentation literature [22, 48
26, 34], the concepts related to how the matrix influences patch-dependent species have not been 49
thoroughly synthesised. In this review, we build on progress made within ecological sub-50
disciplines [25, 35, 36], and on research into edge-effects [37] and habitat fragmentation [26, 51
34], to describe the conceptual domain of the matrix in fragmented landscapes. 52
53
Our approach to understanding the conceptual domain of the matrix is to synthesise ideas from 54
the empirical literature. However, instead of providing a list of matrix effects [e.g. 25, 35, 36, 55
38, 39], we illustrate relationships among mechanisms in a conceptual model. We demonstrate 56
through the conceptual model that what previously were considered primary effects of the matrix 57
are actually secondary outcomes of three 'core effects' (see Boxes 1 and 2). In the second part of 58
our review we identify five influential 'dimensions' and show how these modify the way that core 59
effects play out. The resulting conceptual model of the matrix can help to improve 60
communication of matrix ideas, and guide future research, including research that addresses new 61
questions about interactions between core effects and dimensions associated with time, space and 62
adaptation. 63
64
Core effects of the matrix 65
After considering the range of effects that the matrix can have on patch-dependent species [using 66
empirical literature, also canvased in numerous reviews: 19, 25, 34-36], we identified three 67
fundamental ways that the matrix influences the spatial dynamics of populations and species 68
occurrence in fragmented landscapes. The matrix can influence population persistence in 69

4
fragmented systems through effects associated with (i) movement and dispersal; (ii) resource 70
availability, and; (iii) the abiotic environment (Figure 1). 71
72
Movement and Dispersal. Matrix quality influences the outcome of movement into the matrix 73
Recent reviews report that movement between patches is enhanced as the matrix becomes 74
structurally more similar to the remnant patches [40, 41]. For example, when pastures are 75
replaced by tree plantations, colonisation of forest patches by forest specialists can increase [4]. 76
However, the matrix can influence immigration and emigration in other ways. Sharp ecotonal 77
boundaries between a patch and the matrix can cause individuals to cluster inside remnants 78
('fence effects') [1]. If a species does venture into the matrix, rapid movement through 79
unfavourable habitat could enhance connectivity between separated habitat patches [42]. On the 80
other hand, dispersal or movement between disjunct habitat patches might decline due to altered 81
behaviour, or increased mortality [2, 5, 26, 43]. The influence of the matrix as a demographic 82
sink has received little research attention, although in theory, density-independent emigration can 83
increase the risk of local extinctions [44]. 84
85
Resource availability. Matrix resources could aid patch-dependent species or support matrix 86
specialists. 87
The role of the matrix as a resource base for species that invade remnant patches has long been 88
understood [19] (Box 3). For example, red squirrel Tamiasciurus hudsonicus populations thrived 89
on pine-seeds in Canadian pine plantations. Squirrels subsequently invaded remnant broad-leaf 90
forest and ate Brown Creeper Certhia americana eggs, increasing the rate of nest failure of this 91
patch-dependent bird [16]. On the other hand, if the right resources are provided, the matrix can 92

5
be converted to habitat and desirable native species can live throughout the landscape [e.g. 45]. 93
However, if species remain patch-dependent, they might nevertheless use resources within the 94
matrix as a food subsidy [34]. With the possible exception of bees that can forage outside of the 95
nesting patch [e.g. 14], evidence that patch-dependent species gather resources outside of the 96
patch to support higher population densities inside the patch is limited [e.g. 46]. 97
98
Abiotic environment. The matrix influences microclimate and disturbance regimes of patches. 99
The physical structure of the matrix is often different from habitat patches and can alter the 100
environmental conditions within patches [19, 37], particularly when treed landscapes are cleared 101
[25]. Microclimatic changes associated with increased light and wind penetration can have far-102
reaching effects on patch-dependent species, increasing the risk of local extinction [7, 47]. In 103
addition, species that prosper under the altered microclimate can colonise remnant vegetation 104
and drive edge-sensitive species into the remnant core [37, 48]. 105
106
Changes to disturbance regimes in the matrix can also affect patch-dependent species. Larger 107
and more frequent fires can occur if there are more ignitions in the matrix [11], or when the fuel 108
structure in the matrix is changed by forest logging [11, 49] or by invasive grasses [17]. 109
Conversely, active fire suppression in matrix environments can reduce rates of natural 110
disturbance in patches [3]. Altered microclimate and disturbance regimes can advantage some 111
species, often invasive exotic species [6, 17], but disadvantage others, often species that depend 112
on remnant vegetation [8]. Increased disturbance associated with urban or mining landscapes can 113
also drive local extinctions in patches [9, 10]. 114
115

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TL;DR: In this paper, the authors focus on the biogeograpbic consequences of the creation of habitat islands of different sizes and have provided little of practical value to managers in the field of landscape management.
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Abstract: . Abstract Research on fragmented ecosystems has focused mostly on the biogeograpbic consequences of the creation of habitat “islands” of different sizes and has provided little of practical value to managers. However, ecosystem fragmentation causes large changes in the physical environment as well as biogeograpbic changes. Fragmentation generally results in a landscape that consists of remnant areas of native vegetation surrounded by a matrix of agricultural or other developed land. As a result fluxes of radiation, momentum (La, wind), water, and nutrients across the landscape are altered significantly. These in turn can have important influences on biota within remnant areas, especially at or near the edge between the remnant and the surrounding matrix. The isolation of remnant areas by clearing also has important consequences for the biota. These consequences vary with the time since isolation distance from other remnants, and degree of connectivity with other remnants. The influences of physical and biogeographic changes are modified by the size, shape, and position in the landscape of individual remnant, with larger remnants being less adversely affected by the fragmentation process. The Dynamics of remnant areas are predominantly driven by factors arising in the surrounding landscape. Management of, and research on, fragmented ecosystems should be directed at understanding and controlling these external influences as much as at the biota of the remnants themselves. There is a strong need to develop an integrated approach to landscape management that places conservation reserves in the context of the overall landscape

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Frequently Asked Questions (10)
Q1. What is the key trait determining the ability of species to exploit changes in the matrix?

Demographic and dispersal rates influence responses to changes in the matrix 216 Dispersal rate is a key trait determining the ability of species to exploit changes in the matrix 217 [69]. 

Research addressing these questions has the 272 potential to generate novel conservation strategies and improved understanding of ecological 273 phenomena in fragmented landscapes. 

Changes in the amount of tree 310 cover, the prevalence of exotic plant and animal species, fire regimes and land-use intensity 311 (among others) all contribute to making the matrix more or less hostile for patch-dependent 312 species. 

Sharp ecotonal 77 boundaries between a patch and the matrix can cause individuals to cluster inside remnants 78 ('fence effects') [1]. 

lists of possible 290 approaches have been proposed, such as maintaining a certain proportion of forest cover of 291 particular size [30], maintaining hedge-rows or reducing insecticide use [83]. 

245 Plastic and evolutionary responses of species to the matrix are rarely considered, but have the 246 potential to influence response pathways. 

These changes could make the conservation outlook more bleak as land use intensifies, 313 for example, but matrix changes also provide opportunities to support species in patches. 

387 388 Competition-colonisation trade-offs or predator-prey patch dynamics [85] might also drive 389 feedbacks between pathways in Figure 1. 

If patch-dependent species exploit 167 resources in the matrix [34], a proportionally greater area of matrix to patch could increase the 168 relative abundance of such resources. 

188 However, the effects of matrix heterogeneity are most likely to be apparent on the spatial scale of 189 individual movement behaviour [59] or the scale over which population synchrony occurs [58]. 

Trending Questions (1)
What is matrix resistence in landiscape context?

Matrix resistance refers to the ability of the matrix (non-habitat areas) in a landscape to impede or facilitate movement and dispersal of species.