2009 Reports Re Connectivity

Journal of Applied Ecology 2009, 46, 964969
doi: 10.1111/j.1365-2664.2009.01695.x
FORUM
Climate change, connectivity and conservation decision making: back to basics

Jenny A. Hodgson*1, Chris D. Thomas2, Brendan A. Wintle3 and Atte Moilanen4
Institute of Integrative & Comparative Biology, Miall Building, University of Leeds, Leeds LS2 9JT, UK; 2Department of Biology, University of York, PO Box 373, York YO10 5YW, UK; 3School of Botany, University of Melbourne, Victoria 3010, Australia; and 4Metapopulation Research Group, Department of Biological and Environmental Sciences, PO Box 65 (Viikinkaari 1), University of Helsinki, Helsinki FI-00014, Finland
1
Summary 1. The challenge of climate change forces us to re-examine the assumptions underlying conservation planning. 2. Increasing connectivity has emerged as the most favoured option for conservation in the face of climate change. 3. We argue that the importance of connectivity is being overemphasized: quantifying the benets of connectivity per se is plagued with uncertainty, and connectivity can be co-incidentally improved by targeting more concrete metrics: habitat area and habitat quality. 4. Synthesis and applications. Before investing in connectivity projects, conservation practitioners should analyse the benets expected to arise from increasing connectivity and compare them with alternative investments, to ensure as much biodiversity conservation and resilience to climate change as possible within their budget. Strategies that we expect to remain robust in the face of climate change include maintaining and increasing the area of high quality habitats, prioritizing areas that have high environmental heterogeneity and controlling other anthropogenic threatening processes. Key-words: adaptation, biodiversity, conservation prioritization, habitat quality, landscape planning, spatial ecology, speciesarea relationship, uncertainty
cic dispersal. We argue that uncertainty associated with connectivity is generally higher than uncertainty about habitat area and quality, and threatening processes such habitat destruction. We aim for a more balanced approach to developing climate change conservation strategies where connectivity is treated as a potentially useful tool, but not as an end in itself.
Introduction
How should we adapt our conservation strategies in the face of climate change? Of the multitude of suggested answers, the single most repeated suggestion is to increase connectivity (Heller & Zavaleta 2009). Connectivity conservation (Crooks & Sanjayan 2006) is gathering pace and political support (e.g. Australian Government 2004; IUCN WCPA 2006; Kettunen et al. 2007). The idea is to maintain and build connected environments that will enable species to move with the climate. Whilst laudable, our concern is that this strategy could cause more harm than good if it inadvertently redirects resources and attention away from more certain and eective conservation actions. In this study, we revisit the principles of spatial ecology and conservation planning. We summarize how connectivity emerges as a complicated function of habitat area, habitat quality, the spatial arrangement of habitat and species-spe-
Spatial conservation planning: the basics

We start from a consideration of individual species; the population theory we discuss applies to any species, but we apply this primarily to spatial planning for terrestrial landscapes. The regional carrying capacity, and hence the population size of a species depends principally on the area of suitable habitat, the quality of that habitat and on the spatial arrangement of suitable habitat (Fig. 1; Andrewartha & Birch 1954; MacArthur & Wilson 1967). Habitat arrangement has multiple dimensions, but we consider the main one to be aggregation (Fig. 1); how habitat is concentrated in space. We consider habitat quality to be a measure of potential population
*Correspondence author. E-mail: J.Hodgson@leeds.ac.uk
2009 The Authors. Journal compilation 2009 British Ecological Society
Climate, connectivity and conservation 965

(a) (b)
Connectivity and uncertainty

Connectivity is seen as something that should be accounted for in conservation planning (Margules & Pressey 2000; McCarthy, Thompson & Possingham 2005; Moilanen et al. 2005), but there are numerous overlapping denitions (Tischendorf & Fahrig 2000; Moilanen & Hanski 2001; Goodwin 2003; Fagan & Calabrese 2006). Broadly, functional connectivity estimates the actual or potential rate of immigration into a point, cell, or patch on the landscape (Hanski 1998; Tischendorf & Fahrig 2000), and thus depends on several attributes of the species, as well as the interaction between the species and the landscape (Fig 2). As a result, most connectivity measures subsume inuences of habitat area, quality and spatial aggregation, and some also include information about heterogeneities in the non-habitat (Fig. 2). We argue that uncertainties in the estimation and eects of connectivity make it potentially inecient as a primary conservation metric. Conservation planning is plagued with uncertainties (Regan, Colyvan & Burgman 2001). Uncertainty about area and quality derive from uncertainty about which environmental and biotic factors explain the local carrying capacity for a given species (Elith, Burgman & Regan 2002). Uncertainty about the functional connectivity of the species automatically includes uncertainties relating to area, quality and habitat pattern because functional connectivity depends partly on the distribution and quality of habitats in the landscape (Fig. 2). Additional uncertainties about the measurement of connectivity include: species-specic inuences of distance on dispersal; tails of dispersal distributions, which are notoriously dicult to estimate; eects of source and target habitat quality on emigration and immigration; how dispersing individuals search for habitat; how movement behaviour is aected by the spatial structure of non-habitat, and the inuence of spatially correlated environmental stochasticity on population-dynamical processes (Moilanen & Nieminen 2002). By combining all of these, uncertainty about measuring and predicting connectivity is always higher than uncertainty about the constituent factors that contribute to connectivity. Not only is the measurement of connectivity uncertain, but so are its eects on long-term expected population size. Total carrying capacity always steadily increases with increasing habitat area and quality, but does not do so with increasing habitat aggregation (Fig. 1bd) or increasing dispersal, the other components of connectivity. There is only a relatively narrow window at intermediate levels of habitat aggregation where increased aggregation makes a major dierence to expected population size (because once habitats are close enough to be colonized, further benets of aggregating habitat or increasing dispersal are diminished). Thus, increasing area and quality are more certain to increase population size than are increases in aggregation and dispersal, unless isolation is already known to be the main constraint for a particular species and landscape. As uncertainties about connectivity tend to be high, and increases in habitat quantity and quality coincidentally improve connectivity, we conclude one should generally
(c)
(d)
Fig. 1. Fundamental variables of spatial population biology. (a) Habitat area, habitat quality and habitat aggregation (see text for denitions) are independent axes that all aect regional population size (they also aect functional connectivity, see Fig. 2). (bd) For each factor, there is a threshold below which the regional population is unable to persist. Solid lines indicate potential carrying capacity, whilst dashed lines denote the long-term expected population size.
growth and or density, and area to be the total area with a positive quality. Importantly, each of these three quantities has a threshold below which the regional population of the species will not persist (Fig. 1) too little habitat area, too low habitat quality or excessive dispersion of habitat will all lead to regional extinction of the focal species (Fig. 1; With & King 1999; Hanski & Ovaskainen 2000). Even though area, habitat quality and aggregation are abstractions and their exact denitions might be debated, there is very good evidence for their eects on single species (e.g. Thomas, Thomas & Warren 1992), and they are used as the basis of much conservation planning (Margules & Pressey 2000; Moilanen & Wintle 2006). Dening area, quality and aggregation for multiple species simultaneously is not straightforward because the requirements of species vary. The relationship between species and area is most strongly established (MacArthur & Wilson 1967; Simberlo 1976a,b). More area generally means more individuals, more resources and more environmental variation, giving opportunities for niche specialization. Quality is a dicult notion when considering multiple species, but for practical purposes can be described in terms of freedom from anthropogenic degradation, disturbance, pollution, etc. Inuences of habitat aggregation and isolation on diversity are seen for many communities (e.g. MacArthur & Wilson 1967; Simberlo 1976b; Hanski 1998), but dierences among species in their habitat requirements and dispersal characteristics mean that there is no universal relationship.
2009 The Authors. Journal compilation 2009 British Ecological Society, Journal of Applied Ecology, 46, 964969
966 J. A. Hodgson et al.

What the organism needs (niche) defines Habitat Potential for barriers or conduits in non habitat Aggregation (or other pattern)
Dispersal mechanism, etc.
is quantified in terms of Area Quality
Connectivity (proxy for immigration rates)

Fig. 2. A schematic of the place of (functional) connectivity in spatial ecology and conservation. Functional connectivity is a quantity that always incorporates some aspect of spatial pattern, but it usually also includes information about the amount and quality of habitats and factors inuencing dispersal behaviour and success.
Vital rates and carrying capacity Population size, spatial distribution and persistence probability
provide higher weight in decision-making to actions that increase area and quality. Theoretically, we know that populations will sometimes benet more from a small, well-connected piece of habitat than a larger, more isolated one, but the relative uncertainties and the probability of worse-than-expected outcomes should also aect our decision making. From a decision theoretic perspective, when faced with two options that convey similar expected (mean) returns, one should choose the option with the lowest variance of expected outcome to maximize the probability of achieving ones conservation goal (e.g. Ben-Haim 2001). We are particularly concerned that in a planning process, people who want to release areas from conservation, e.g. for development, could actually exploit the profusion of connectivity measures (Kindlmann & Burel 2008) to choose one that works for them (Walker et al. 2009). For example, restored corridors, stepping stones or softening of the anthropogenic matrix will cause increases in some connectivity measures. Depending on the connectivity metric used, a large percentage increase in connectivity could be used to argue that a large percentage decrease in habitat area is acceptable allowing a development to proceed. Loss of habitat implies certain and immediate decreases in population sizes, whereas compensating long-term benets of additional connectivity might be largely unknown and possibly small (Falcy & Estades 2007). Notwithstanding these misgivings, the functional connectivity of landscapes applied to single species is a very useful concept in appropriate circumstances, when the constraints on a particular species are known. But, ultimately, conservation is a multi-species enterprise. In this context, various measures of structural connectivity have been proposed, that generalize
the connectivity of vegetation types without reference to particular species. Combining species responses in this way magnies uncertainty because multi-species responses are not a simple function of each individual species response. There is an attractive simplicity to increasing structural connectivity for multi-species conservation, but the scientic basis for such a strategy is largely absent and the applicability of this concept under climate change is also highly uncertain. The trade-o between increased structural connectivity per se and increased protection for existing natural or semi-natural habitats are always very dicult to calculate. However, maintaining and increasing the area of natural or semi-natural habitats will add useful habitat area for many species and, as described above, will usually coincidentally increase connectivity.
The new world order

So far, our discussion has been most relevant to situations where the regional distributions of species can be assumed to be relatively stationary. Given climate change and the lagged responses of species (Menendez et al. 2006), dynamic change will be the norm for the foreseeable future. At large scales there are shifts to higher latitudes and elevations (Hickling et al. 2006; Parmesan 2006; IPCC 2007) and movements along moisture gradients, and at smaller scales there are shifts in preferred microclimates and changes to the nature of the vegetation that constitutes habitat (Thomas et al. 2001; Davies et al. 2006). These changes undermine three common presumptions in conservation planning. First, we often presume that vegetation type can be used as a reasonable proxy for habitat availability for one or many species. Quaternary evidence shows that
Climate, connectivity and conservation 967 species exhibit individualistic responses to climate change, and vegetation types change with the climate (Williams, Shuman & Webb 2001), so that the community classication schemes we now use to describe vegetation types will become redundant in the long term. Secondly, we assume that structural connectivity, measured on the basis of land cover types, is a reasonable proxy for functional connectivity of multiple species. As with the rst assumption, this will become less reasonable the more climate change advances and changes communities. Thirdly, we commonly assume that protecting locations with the most populations of a species will maximize the chances of persistence in both the short- and long-term survival. Under climate change, prioritizing only existing core populations carries the danger of promoting short-term persistence in current strongholds at the expense of long-term survival; but prioritizing only marginal populations that are predicted to expand is risky because of massive uncertainties about the true consequences of climate change. In essence we are required to deal with trade-os through time, as well as continuing to pay attention to trade-os in space and trade-os between species. We have to address the additional question How much short-term conservation success should we forgo in order in increase the long-term probability of achieving our targets? Perhaps not much, as benets that are to materialize a long time in the future may have a tendency of disappearing on the way (Walker et al. 2009). Such considerations need to be incorporated within population viability analyses and decision frameworks, which can no longer assume long-term stasis in environmental conditions. retain a particular community composition may be expensive and ultimately doomed to failure. A second generalization is that environmental heterogeneity provides opportunities for populations to survive dierent extremes by shifting between dierent types of vegetation, soils, aspects or elevations (Thomas et al. 2001; Davies et al. 2006). Species diversity and endemism are also increased in regions with high topographic and habitat diversity (e.g. Simberlo 1976a), especially where there are steep elevation and climatic gradients (Ohlemuller et al. 2008). Our second mes sage is that focussing eorts on regions with high existing environmental heterogeneity is likely to be a robust strategy. In a sense, we are identifying the importance of a dierent kind of connectivity that between cooler and warmer (and drier and moister), habitats rather than between currently similar habitats. Further research is needed to quantify the benet of habitat diversity, especially when there might be a trade-o between this and the aggregation of existing habitat for many species. Thirdly, the majority of small-range terrestrial species are clustered into a small percentage of the land surface (centres of endemism areas of high irreplaceability, cf. Wilson, Carwardine & Possingham 2009), many of which are mountain ranges. A high percentage of the species threatened with extinction from climate change are found in such locations: they are expected to show range retractions within the regions where they currently occur, and are unlikely to achieve long distance colonization of other parts of the world (Midgley et al. 2002; Williams, Bolitho, & Fox 2003; Thomas et al. 2004; Malcolm et al. 2006; Ohlemuller et al. 2008). So, our third message is that concentration of conservation eort in centres of endemism remains a valid strategy. Fourthly, almost all threatened species are negatively impacted by multiple factors. In some instances, mitigating known threats other than climate change may be sucient to permit a population to persist, even if the local climate has deteriorated. When this strategy cannot ensure persistence in its own right, mitigating known threats should be regarded as an essential rst step in making populations robust to climate change. We recommend dealing with known (stoppable) threats for which there are known solutions before addressing uncertain and or unstoppable threats with less certain or less feasible solutions (Pressey et al. 2007; Wilson et al. 2007). These four principles, increasing protected area, maintaining and in some cases increasing environmental heterogeneity, concentrating eorts in centres of endemism, and reducing other pressures are likely to be benecial and robust, with or without climate change. However, these are rules of thumb, and there is great potential for improved planning at regional scales with improved adaptive decision-making methods. Importantly, decision-making tools need to weight strategies according to their relative costs, expected benets and the uncertainty in achieving that benet (Burgman, Lindenmayer & Elith 2005; McDonald-Madden, Baxter & Possingham 2008). Ecological research needs to contribute by quantifying benets (including the benets of connectivity and the benets of habitat heterogeneity) in terms of a common currency, e.g. long-term species persistence, and by quantifying
Reasons to be cheerful
It is easy to be overwhelmed by the complexity and uncertainty involved in conservation planning for a world with climate change. There is a huge desire to do something but what exactly is it that we should do? A vast number of suggestions have been made, and there is limited direct evidence to assess which of these is likely to be most eective (Heller & Zavaleta 2009). So, can we step back and ask if any principles hold true with or without climate change, and thereby which conservation strategies are most likely to be robust? One such principle is that increasing numbers of species are associated with increasing area (Connor & McCoy 1979; Guilhaumon et al. 2008). But how much area is enough? Eective conservation requires sucient habitat where a species currently occurs and additional locations that will support populations whilst the distribution is changing, until it reaches a new equilibrium (assuming the climate does; Araujo et al. 2004). Any previously used target [e.g. the 10% terrestrial protected area target (IUCN 2004)] will deliver lower conservation outcomes under climate change than originally hoped. Therefore, renewed eort and additional funding to conserve extra land is warranted. Locations that have low human impacts should remain good for many species, even if the identities of those species change. Maintaining sites of high value to biodiversity should be feasible, but management that attempts to

Table 1. Suggested re-assessment of basic ideas in conservation planning, assuming climate change Ideas to keep More total area is benecial; including expanding present conservation areas Biodiversity hotspots centres of endemism should be prioritized Environmental habitat heterogeneity facilitates diversity and persistence Human activities that diminish diversity should be minimized or reversed Ideas to modify and develop The time frame for conservation planning (the trade-o between current and future benet; and incorporation of environmental change in population viability assessment) The role of connectivity, including trade-os between connecting landscapes and other conservation actions Ideas that hold us back Attempting to maintain existing or past community composition Using permanently xed conservation targets (e.g. 10% terrestrial area target)
uncertainty. Research is required in the development of decision frameworks (Hoegh-Guldberg et al. 2008) to integrate and visualize the costs and benets, and to ensure that methods are easy to adapt and update as new information becomes available.
Conclusion
In summary, we think that the political and ecological reality of climate change should prompt us to reassess which ideas to keep, which to modify and which to abandon (Table 1). Connectivity usefully reminds people that excessive isolation is a threat to populations, but, as increased attention is paid to the spatial arrangement of habitats and dispersal, more fundamental issues may be overlooked (Fig. 1; Table 1). Whilst climate change adds extra challenges, potential damage can still be alleviated by removing other sources of threat. Land conversion and land-use change leading to habitat loss is still the most cited threat to currently endangered species, and the most straightforward way to tackle this is to maintain and restore larger areas of natural habitat. Species will not be able to survive where they are or shift their distributions to new climatically suitable areas unless there are sucient habitats for them, and it should be remembered that increasing habitat area is an eective way of increasing connectivity. Furthermore, conserving habitats will be benecial even if the particular species found in an area are gradually replaced by others as the climate changes. The conservation of high quality existing habitats should therefore remain the primary focus of conservation eorts to maintain biodiversity.
Acknowledgements
Thanks to two anonymous reviewers whose comments helped to improve the manuscript. J.H. and C.D.T. were supported by Natural England. B.W. was supported by the Australian Research Council (DP0774288) and the Australian Governments Commonwealth Environment Research Facility program. A.M. was supported by the Academy of Finland, Finnish Center of Excellence Programme 2006-2011, grant 213457.
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doi: 10.1111/j.1365-2664.2010.01899.x
FORUM
Connectivity, dispersal behaviour and conservation under climate change: a response to Hodgson et al.
Veronica A. J. Doerr1,2*, Tom Barrett3 and Erik D. Doerr1,2
CSIRO Ecosystem Sciences, GPO Box 284, Canberra ACT 2601, Australia; 2Research School of Biology, Australian National University, Canberra ACT 0200, Australia; and 3New South Wales Department of Environment, Climate Change & Water, PO Box 494, Armidale NSW 2350, Australia
1
Summary 1. Hodgson et al. [Journal of Applied Ecology 46 (2009) 964] argue that connectivity is complex and uncertain, that it can be improved incidentally by increasing habitat extent, and that connectivity conservation is unlikely to be eective under climate change. 2. We believe that they have overlooked recent research on dispersal behaviour and structural connectivity, which has improved our understanding of functional connectivity and revealed that it will not necessarily increase with habitat extent. 3. New modelling techniques including least-cost path models incorporate this more detailed understanding of connectivity into conservation planning, facilitating the true aim of connectivity conservation to ensure appropriate interactions between habitat extent, quality and connectivity. 4. Synthesis and applications. Advances in behavioural research and modelling techniques allow us to manage structural connectivity with as much certainty as we manage extent and quality of habitat. Successful landscape conservation to address both current threats and future climate change must manage these three elements in concert. Key-words: aggregation, behavioural ecology, connectivity conservation, corridor, fragmentation, gap-crossing, metapopulation, population viability, range shift, stepping stone
Introduction
For most of the worlds ecosystems, human-induced habitat loss, degradation and fragmentation are primary causes of declines in biodiversity (Fahrig 2003; Lindenmayer & Fischer 2006). Furthermore, climate change is predicted to interact with and intensify the eects of these problems. Connectivity conservation has emerged as an overarching solution with considerable political and popular support (Crooks & Sanjayan 2006). However, Hodgson et al. (2009) highlight the dangers of investing in connectivity per se, and argue that other strategies may provide better protection for species in a changing climate. We wholeheartedly agree with Hodgson et al. that connectivity should not be the sole focus of conservation actions, and that conservation investments should be based on analysis of their likely benets. Yet Hodgson et al. suggest that connectivity conservation is never likely to be a robust strategy, and here we disagree. Specically, Hodgson et al. argue that there is too
*Correspondence author. E-mail: veronica.doerr@csiro.au
much uncertainty surrounding connectivity, that connectivity is primarily a result of habitat aggregation, and it can coincidentally be improved by increasing habitat extent. They also suggest that connectivity conservation sacrices long-term conservation success under a changing climate in favour of short-term gains. In this response, we suggest that Hodgson et al. have overlooked recent advances in our understanding of connectivity, particularly arising from research on dispersal behaviour. These advances provide a clearer distinction between structural and functional connectivity and greater certainty regarding the eects of structural connectivity. They also bring a new awareness that increases in habitat extent alone will not necessarily increase functional connectivity. In addition, we believe that Hodgson et al. have misinterpreted connectivity conservation, which carries a specic meaning that involves more than just conserving structural connectivity, and can provide long-term solutions to many of the threats associated with climate change including those highlighted by Hodgson et al. Finally, we suggest that the dierences in our perspectives may partly result from dierences in the scales at which empirical research and conservation planning are conducted. Fortunately, new modelling techniques are allowing us
2010 The Authors. Journal of Applied Ecology 2010 British Ecological Society
144 V. A. J. Doerr, T. Barrett & E. D. Doerr to move beyond simple measures of aggregation to incorporate a more detailed behavioural understanding of connectivity into conservation planning, despite dierences in scale. individuals use a foray-based search strategy, they may have a maximum search distance beyond which they will not travel, regardless of how much structural connectivity is present in the landscape (Conradt, Roper & Thomas 2001; Doerr & Doerr 2005; Doerr, Doerr & Davies 2010). The species-specic nature of behavioural research may be viewed as an impediment to its usefulness for ecosystem conservation, but patterns are emerging which suggest that responses to structural connectivity may not be as species-specic as was once thought (Haddad et al. 2003; Doerr, Doerr & Davies 2010; Gilbert-Norton et al. 2010). Instead, movement behaviour may be shaped by the structure of environments experienced over evolutionary time, and species in any given ecological community with broadly similar life-histories may have evolved similar movement behaviours as responses to their shared environments (Fahrig 2007). For example, Doerr, Doerr & Davies (in press) found that use of scattered trees, foray distances, and gap distances crossed were similar among ve Australian woodland birds despite substantial dierences in their ecology. Belisle (2005) proposed that travel costs may provide one mechanism through which landscapes can exert similar evolutionary pressures across species. Thus, theories from behavioural ecology such as the marginal value theorem could provide the basis for general theories of connectivity, allowing us to predict the eects of dierent types of structural connectivity for large suites of species at once (Belisle 2005).
Dispersal behaviour and structural connectivity

The intent of connectivity is to facilitate dispersal of individuals. Thus, an empirical understanding of connectivity depends on understanding animal behaviour, particularly movement and dispersal behaviour, to reveal what parts of landscapes individuals are willing to move through and why (Lima & Zollner 1996; Chetkiewicz, Clair & Boyce 2006). The value of behavioural research for conservation has been debated (Blumstein & Fernandez-Juricic 2004; Caro 2007), and thus it is unsurprising if conservation biologists are not familiar with the movement behaviour literature, very little of which existed during early discussions about connectivity. Yet movement behaviour is a rapidly growing eld (Nathan 2008), with extensive empirical analyses and emerging theories that can provide a strong foundation for modelling and conserving connectivity. When connectivity began to be viewed from a behaviourbased perspective, it became a characteristic of the matrix between subpopulations (Taylor, Fahrig & With 2006), rather than a characteristic of patches or landscapes. Movement could be dependent not just on distances between subpopulations (i.e. aggregation) but on the physical characteristics of the matrix itself, particularly the presence of habitat elements too small for settlement but which might nonetheless facilitate movement. As a result, a much clearer distinction emerged between structural and functional connectivity. Structural connectivity refers to physical characteristics of the landscape between patches of occupied habitat. Functional connectivity refers to the degree to which movement of individuals and or their genetic material actually occurs, and is inuenced by both movement potential due to structural connectivity and by local subpopulation dynamics (Hilty, Lidicker & Merenlender 2006). Empirical research on movement behaviour has concentrated on revealing what types of structural connectivity provide the potential for dispersal movements and thus contribute to functional connectivity. A number of studies have shown that various species use corridors to move through fragmented landscapes (Haddad et al. 2003; Haddad & Tewksbury 2005), and some have demonstrated the use of simpler landscape elements such as scattered trees (Fischer & Lindenmayer 2002; Doerr, Doerr & Davies 2010). Research on gap-crossing behaviour has been particularly critical, identifying gap distances either within corridors or among scattered trees that may prevent movements, thus revealing details of structural connectivity that contribute to movement potential (St. Clair et al. 1998; Grubb & Doherty 1999; Robertson & Radford 2009). In addition, research on overall movement strategies such as foray search is illustrating that distances between subpopulations (i.e. aggregation) may have a threshold eect rather than a linear eect on movement potential. When
Conservation certainty
All of these advances are making structural connectivity a much more measurable and manageable concept than Hodgson et al. suggest. Functional connectivity remains complex because it integrates movement potential with the dynamics of subpopulations (which is why Hodgson et al. deem it too intractable for conservation planning). Yet structural connectivity contributes signicantly to functional connectivity by determining movement potential. The resulting eects on population persistence are also increasingly predictable thanks to controlled research in experimental landscapes which is demonstrating that connected patches experience fewer local extinctions than isolated patches (Damschen et al. 2006; Brudvig et al. 2009). Thus, structural connectivity can be directly quantied in the landscape, has predictable eects on movement potential, and is known to contribute to population persistence, making it a worthwhile focus for management. Unfortunately, Hodgson et al. omit structural connectivity from their schematic of the place of connectivity in conservation (Hodgson et al., Fig. 2). We have revised their diagram to distinguish between structural and functional connectivity and depict the relationships between them, as well as relationships with the area and quality of habitat suitable for settlement (Fig. 1). Structural connectivity is independent of habitat area and quality and is what denes habitat for dispersal, just as area and quality are what dene habitat for settlement. Structural connectivity, habitat area and quality interact to determine functional connectivity, but they also interact to determine subpopulation dynamics and thus the eective
2010 The Authors. Journal of Applied Ecology 2010 British Ecological Society, Journal of Applied Ecology, 48, 143147
Connectivity and dispersal behaviour 145

What the organism needs for survival and reproduction Defines Habitat for settlement
What the organism needs for dispersal Defines Habitat for dispersal Is quantified in terms of
Is quantified in terms of Area Quality Structural connectivity
Which is the interaction between Landscape characteristics between areas of habitat for settlement Distance between areas of habitat for settlement
Which determine
Potential rates of dispersal between subpopulations
Which interact to determine Functional connectivity (actual rates of dispersal) and subpopulation size and dynamics Which are integrated over multiple subpopulations to determine Effective population size, spatial distribution and persistence probability
provide for many more of a species needs than habitat for dispersal (Haddad & Tewksbury 2005; Doerr, Doerr & Davies 2010). Restoring habitat for settlement, either for a single species or particularly for an entire community, may thus be more complex and uncertain than restoring habitat for dispersal (i.e. increasing structural connectivity). Finally, Hodgson et al. argue that increasing habitat extent will coincidentally improve connectivity by increasing aggregation and thus reducing distances between patches. However, behavioural research suggests that reducing inter-patch distances without providing structural connectivity will only be benecial once patches become close enough to allow gap-crossing between them. That distance may be as little as 60100 m (unlikely to be achieved by most eorts to increase habitat extent), as many species are unwilling to cross gaps any larger (Desrochers & Hannon 1997; Robertson & Radford 2009; Doerr, Doerr & Davies 2010). Individuals can traverse much greater distances between habitat patches if structural connectivity is present, but the existence of foray-based search means that increases in aggregation may only be benecial if distances between patches can be reduced below a critical foray distance threshold, which may only be 12 km (Doerr, Doerr & Davies 2010, in press). Thus, the benets of reducing aggregation per se (as opposed to managing it in concert with structural connectivity) are risky because they are not commensurate with eort.
Fig. 1. A schematic illustrating the role of both structural and functional connectivity in spatial ecology and conservation (revised from Fig. 2 in Hodgson et al. (2009)). Functional connectivity results from interactions between the amount and quality of habitat suitable for settlement as well as the inuence of the rest of the landscape (i.e. structural connectivity) on the potential for dispersal. Structural connectivity is therefore a vital component of functional connectivity that is tractable to model and manage.
Connectivity conservation is more than just conserving connectivity

Hodgson et al. interpret connectivity conservation as the eort to increase structural connectivity with the primary purpose of enabling species range shifts due to climate change. However, as highlighted above, structural connectivity interacts with other aspects of the landscape and thus is not necessarily the sole or most important aspect to improve in every landscape. Connectivity conservation acknowledges this, and has a very specic meaning in the literature (IUCN WCPA 2006; Worboys 2010), much like systematic conservation planning has a specic meaning and doesnt merely refer to taking a systematic approach to planning conservation actions (Margules & Pressey 2000). As a result, connectivity conservation is broader than Hodgson et al.s interpretation. Connectivity conservation can be dened as coordinated eorts to achieve metapopulation viability across a range of spatial scales, which involves evaluating and improving the interactions between habitat area, habitat quality and structural connectivity (Crooks & Sanjayan 2006; Worboys 2010). There is no overarching rule about which action is always more eective this depends on the existing conditions in a given landscape. Connectivity conservation thus aims to develop exible solutions, tailored to the dierent needs of dierent landscapes. This may involve protecting large continuous areas of existing habitat, but may also involve protecting or increasing connections between multiple small discontinuous areas of habitat where that is all that remains. The preference of Hodgson et al. to focus on habitat area and habitat quality can thus be
population size of the population as a whole. None has a direct inuence on populations completely independent of the others all provide the same degree of conservation certainty because their benets depend on the interactions between them. Hodgson et al. also argue we can be relatively certain about the positive eects of increasing habitat extent and habitat quality often assumed to be accomplished through increasing the size and or number of protected areas. Yet in highly disturbed ecosystems, there may be little habitat left outside of already existing protected areas. Thus, increasing habitat extent and improving habitat quality involves restoring habitat in areas where it has been lost to other land uses. Unfortunately, there are substantial limitations and uncertainties in our ability to restore ecosystems. For example, nitrogen enrichment via fertilisation reduces plant diversity as well as the stability of ecosystems worldwide (McIntyre 2008; Bai et al. 2010). These eects can last long after fertilisation has ceased, inhibiting full recovery of the ecosystem despite restoration attempts (Munro et al. 2009). It is also reasonable to argue that habitat quality will often be more species-specic than structural connectivity, as habitat for settlement must
146 V. A. J. Doerr, T. Barrett & E. D. Doerr encompassed by connectivity conservation wherever these actions are deemed to provide the greatest benets. Finally, the ultimate purpose of connectivity conservation is not simply to facilitate range shifts, but to increase the resilience of populations to the variety of threats caused by or intensied by climate change. Under connectivity conservation, structural connectivity is desired where it links multiple subpopulations via dispersal, allowing subpopulations to function collectively as one larger, more resilient population. These principles can be applied at any scale, not just scales that might be relevant for possible range shifts under climate change (Opdam & Wascher 2004). Thus, connectivity conservation can be used to reduce pressures other than climate change, can be applied to increase viability of populations in centres of endemism, and can concentrate on areas of high environmental heterogeneity all of which are principles that Hodgson et al. suggest will underlie robust conservation strategies under climate change. data on foray-based search behaviour and foray distances (Doerr & Doerr 2005; Doerr, Doerr & Davies 2010) were used to dene the maximum distance individuals will move through structural connectivity, modelled as suboptimal habitat. Using modern satellite imagery, suboptimal habitat could be mapped at a ne scale of resolution to detect very small elements of structural connectivity known to support dispersal movements, such as single trees. Fine-scale habitat mapping and simplied behavioural rules were then modelled together using the least-cost path approach (Drielsma, Manion & Ferrier 2007), with habitat quality as a surrogate for movement cost. This modelling approach simultaneously evaluates habitat area, quality and structural connectivity, identifying where these elements currently exist in concert in the landscape versus where they are unable to interact due to deciencies in one or more elements (Barrett et al. 2010). This gives conservation planners the ability to make practical recommendations that maximise the likelihood that actions at a local scale will contribute to population viability and resilience at large scales (Barrett et al. 2010). These models are currently being used to guide conservation planning decisions in several regions of New South Wales, Australia.
Moving beyond aggregation in large-scale conservation planning models

Conservation modellers may still be unsure how to incorporate advances in our understanding of connectivity and connectivity conservation due to the dierent scales at which behavioural research and conservation planning are usually conducted. Conservation planning often occurs at very large scales regions to global scales. Yet a behavioural understanding of connectivity is shaped at scales relevant to movement of individuals local to landscape scales. The diculty is that incorporating small-scale detail in large-scale models is often deemed computationally intractable. Fortunately, there are promising new advances that can model connectivity over large spatial scales in ways that align more closely with a behaviourbased view of connectivity. First, we have already noted that the types of structural connectivity that facilitate dispersal movements are not necessarily species-specic. Thus, models may only need to incorporate general principles (such as threshold distances between habitat patches) rather than behavioural detail specic to many dierent species. Another way in which conservation planning models can incorporate a tractable amount of behavioural detail is through the use of least-cost path modelling and state-space modelling. These new types of models simultaneously explore behavioural and landscape parameters to identify which landscape details most need to be incorporated into large-scale models (Chetkiewicz, Clair & Boyce 2006; Kadoya 2009), which can then be kept relatively simple by modelling only the few most relevant small-scale parameters. Remaining computational challenges can often be overcome by decreasing the sizes of grid cells only to a relevant scale. Further behavioural detail can then be incorporated by modelling resistance of grid cells that have dierent compositions (McRae & Beier 2007). One example of the success of these new approaches comes from our own work, in which data on gap-crossing distances (Doerr, Doerr & Davies 2010) were used to dene the maximum distance individuals will move through non-habitat, and
Reasons to be cheerful indeed!

As Hodgson et al. suggest, it is easy to be overwhelmed by the challenges of conservation under climate change. It is worthwhile returning to basic principles, focusing on actions that will be cost-eective and that will address current threats as well as those anticipated due to climate change. Fortunately, connectivity conservation provides such an approach by focusing on habitat area, quality, and structural connectivity as independent attributes that must all work together to support viable, resilient populations. Thanks to a growing body of behavioural research, we can reliably identify situations in which fostering structural connectivity in the matrix is likely to yield positive benets for populations. We can also use behavioural information to model connectivity alongside habitat extent and quality and thus tailor management to specic landscapes. Ultimately, these new techniques ensure that large-scale conservation planning can take advantage of up-to-date connectivity knowledge to truly provide evidence-based conservation guidance.
Acknowledgements
Thanks to members of the Great Eastern Ranges Initiative, David Westcott, Paul Sunnucks, Sasha Pavlova, and Colleen Cassady St. Clair for discussions that shaped these ideas. The manuscript was greatly improved by the comments of Richard Fuller, Sue McIntyre, Dan Lunney, Vicki Logan, and ve anonymous reviewers.
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doi: 10.1111/j.1365-2664.2010.01919.x
FORUM
Habitat area, quality and connectivity: striking the balance for efcient conservation
Jenny A. Hodgson1*, Atte Moilanen2, Brendan A. Wintle3 and Chris D. Thomas1
Department of Biology Area 18, University of York, York YO10 5DD, UK; 2Metapopulation Research Group, Department of Biological and Environmental Sciences, PO Box 65 (Viikinkaari 1), FI-00014, Finland; and 3School of Botany, University of Melbourne, Vic., 3010, Australia
1
Summary 1. Population viability can depend on habitat area, habitat quality, the spatial arrangement of habitats (aggregations and connections) and the properties of the intervening non-breeding (matrix) land. Hodgson et al. [Journal of Applied Ecology 46 (2009) 964] and Doerr, Barrett & Doerr (Journal of Applied Ecology, 2011) disagree on the relative importance of these landscape attributes in enabling species to persist and change their distributions in response to climate change. 2. A brief review of published evidence suggests that variations in habitat area and quality have bigger eects than variations in spatial arrangement of habitats or properties of the intervening land. Even if structural features in the matrix have a measurable eect on dispersal rates, this does not necessarily lead to signicant increases in population viability. 3. Large and high-quality habitats provide source populations and locations for colonisation, so they are the main determinants of the capacity of species to shift their distributions in response to climate change because populations must be established successively in each new region. 4. Synthesis and applications. Retaining as much high quality natural and semi-natural habitat as possible should remain the key focus for conservation, especially during a period of climate change. Key-words: aggregation, climate change, conservation planning, corridor, landscape, matrix, uncertainty
limiting extinctions and maintaining functioning ecosystems. Because of the complexities of ecology and human impacts in dierent regions, no single prescription for conservation will work everywhere. So, we agree with Doerr, Barrett & Doerr (2011) that conservation strategies need to be tailored to the landscape. We also fundamentally agree about which factors aect the persistence of populations and metapopulations (Doerr, Barrett & Doerrs Fig. 1 essentially expands our boxes for dispersal mechanism and potential for barriers or conduits in non habitat). However, we disagree about the relative importance of the various factors: habitat area, habitat quality, the spatial arrangement of habitats (aggregations and connections) and the properties of the intervening non-breeding (matrix) land. Hence, we provide here evidence in support of our assessment of the relative importance of these landscape attributes.
Introduction
Hodgson et al. (2009) noted that habitat area, quality, and aggregation were key components of landscape-scale conservation, and that prioritising habitat area and quality would be a robust way to facilitate connectivity and the persistence of biodiversity in the face of climate change. Doerr, Barrett & Doerr (2011) suggest that structural features to facilitate dispersal in non-breeding habitat are valuable targets in themselves, and can be measured with certainty equal to that of area, quality, and aggregation. They further suggest that we misunderstood the term connectivity conservation. In this reply to the Forum by Doerr, Barrett & Doerr in this issue, we begin by summarising where we are in agreement. We then present our case that Doerr, Barrett & Doerr, in focussing narrowly on non-habitat structural features, have ignored the bigger picture of relative priorities for conservation under climate change. Hodgson et al. (2009) and Doerr, Barrett & Doerr (2011) present dierent ways of approaching the same overall goal;
Relative importance of landscape attributes

HABITAT AREA
*Correspondence author. E-mail: jenny.hodgson@york.ac.uk
Successful reproduction is conned to natural and semi-natural habitats for the majority of terrestrial species.
2010 The Authors. Journal of Applied Ecology 2010 British Ecological Society
Area, quality and connectivity 149 In the context of climate change, multi-generational range shifts are facilitated initially by large habitat areas that support large source populations, by substantial intervening habitat areas to support breeding and generate propagules en route, and by high habitat availability within target regions to ensure eventual persistence (Table 1). Maximising area is also likely to increase diversity via increased habitat heterogeneity. Contrary to Doerr, Barrett & Doerr (2011), we contend that there is generally a large amount of natural or semi-natural habitat with insucient protection. Tropical forest, for example, is converted to other land uses at around 05% annually (FAO 2005). It is usually much more eective and cheaper to retain what is still present than to attempt to recreate it.
NON-BREEDING HABITAT (OR THE MATRIX)
HABITAT QUALITY
Quality improves persistence by increasing population growth, resulting in larger propagule numbers, increased likelihood of colonisation, and higher population growth rates following colonisation. Many aspects of habitat quality are, as Doerr, Barrett & Doerr (2011) suggest, speciesspecic and dicult to control. However, some aspects of quality aect many species similarly, such as nutrient pollution, the spread of invasive species or major habitat disturbances. Much of the worlds biomes are now partly or substantially altered by, for example, selective logging, partial fertilisation of grasslands, drainage of wetlands, and elimination of re or large mammals. Preventing further degradation and increasing the quality of already-degraded areas can generate extremely large dierences in population densities (e.g. Table 1) and, therefore, colonisation and population growth, including range expansion.
Interventions in the matrix could contribute to population viability by overcoming behavioural barriers to crossing certain boundaries or land-cover types, or by reducing dispersal mortality. Managing the matrix can result in modest increases in dispersal between nearby habitat patches (by ca. 25%, Table 1). However, we could nd no robust evidence that matrix condition alters long-distance, multi-generational range changes. The likelihood of leaving an individuals natal habitat patch may increase if the intervening matrix is favourable, but movements typically become faster and straighter when an individual is in a hostile matrix environment, and this leads to much longer realised dispersal distances (Ovaskainen et al. 2008; Zheng, Pennanen & Ovaskainen 2009). The longest dispersal distances are the most important for maintaining genetic diversity and for range expansions under climate change (Neubert & Caswell 2000; Trakhtenbrot et al. 2005). We also note that a whole dierent set of considerations apply to wind- and water-dispersed species, and to the four kingdoms other than animals. The suggestion of Doerr, Barrett & Doerr (2011) that dispersal behaviour could be universal is centred on an example that involved a small number of well-studied and related species with high cognitive abilities.
Uncertainty and robustness

Doerr, Barrett & Doerr (2011) argue that structural connectivity can be measured with reasonable certainty. The issue to us is not the development of a repeatable metric, but whether variation in such a metric is a good predictor of multi-generational range expansions for a wide range of dierent taxa. The eect of the matrix on long-distance dispersal and colonisation is virtually unknown; and we already know that short-distance matrix eects are species specic (Eycott et al. 2009; Prevedello & Vieira 2010). Doerr, Barrett & Doerr (2011) reasonably counter that measurement of habitat quality is an issue, and also species-specic, but it is much easier to investigate, for example by relating the population densities of individual species or broader measures of diversity to environmental and biotic variables. We strongly disagree with the Doerr, Barrett & Doerr (2011) view of area, quality and connectivity that all provide the same degree of conservation certainty because their benets depend on the interactions between them. It is clear to us (e.g. Table 1) that dierent variables that interact in a given model do not all have the same eect sizes or associated uncertainties. Uncertainty about functional connectivity is higher than any of the above-mentioned factors because it adds uncertainty about dispersal distances and behaviour on top of uncertainty about what constitutes habitat quality. Crucially, though, this uncertainty does not negate the large eects of habitat area and quality over large numbers of species and landscapes, because only metapopulations close to their extinction threshold are substantially limited by connectivity (Hodgson et al. 2009: Fig. 1).
SPATIAL ARRANGEMENT OF HABITATS
The locations of remaining habitat fragments in landscapes are known to be important to long-term population persistence (Hanski & Ovaskainen 2000), but the size of this eect is smaller than the eects of quantity and quality (Table 1), principally because the production of new individuals takes place within habitats, regardless of their location (Ovaskainen 2002). Habitat aggregation generally increases the chance of a propagule landing in suitable habitat, and therefore of a patch of habitat being colonised occupied, but it can only compensate a little for deciencies in quantity and quality (Table 1). Under climate change, the benet of aggregation may be less certain because aggregating remaining habitat within a few regions may leave dispersal barriers that will eventually need to be bridged. When suitable habitat is a very low proportion of the landscape, there may be a tradeo between maximising aggregation and reducing the largest dispersal barriers. There is evidence that corridors may increase dispersal rates between patches to some extent (Table 1), but an increase in dispersal per se is not direct evidence of an increase in population viability.

Table 1. Brief overview of evidence for the importance of dierent landscape-scale factors in conservation. Priority has been given to review papers and meta-analyses, papers that compare more than one factor, and papers that relate factors specically to range expansion Landscape attribute Habitat area Summary of evidence Eastern US trees range expansions dependent on source populations Patch area has consistently bigger eect (up to 100 times greater F ratio in ANOVA) than connectivity measures on species richness and abundance of plants and butteries in fragmented grasslands in southern Germany Plant and buttery species richness of calcareous grasslands in northern Germany is aected by patch area, and not signicantly by isolation or surrounding landscape heterogeneity Simulated and observed expansion rates of buttery Pararge aegeria >40% slower in landscape containing 24% less woodland cover Hesperia comma buttery range expansion rates in UK strongly related to habitat quantity Habitat quality* Positive eects of quality on occupancy for plants, butteries, moths, other insects, amphibians, birds, small mammals, primates and carnivores Correct vegetation management can increase buttery densities 10- to 100-fold Macaw (Ara and Orthopsittaca spp) density in Amazon varies >10-fold with respect to indicators of human disturbance Sea otters (Enhydra lutris nereis) range expansion dependent on population growth rate Quality the dominant factor aecting buttery and moth abundance and diversity in a Finnish landscape Spatial pattern 1: aggregation (aka spatial autocorrelation) Isolation negatively aects colonisation rate and occupancy Negative eect of isolation on patch occupancy in three English butteries (eect of habitat quality is 23 times larger). Positive eects of aggregation on ower visitation and seed set in pan-European study of 10 plant species (although patch area eect is bigger) Aggregation measures have positive eects on species richness and abundance of butteries (and to lesser extent plants) in fragmented grasslands, though not as large as the eects of area Spatial pattern 2: habitat connections (corridors and stepping stones) Average 16-fold increases in exchange rates between patches, based on systematic review in which the corridor is the same type of habitat as the connected patches Modest positive eect of corridors on dispersal rate: standardised eect size <05 if study controlled for distance between patches. Smaller eect for created corridors than natural corridors. Substantial evidence that hedgerows are used by animals as corridors between woods, but very little on the quantitative eect this has on population viability Corridors of habitat relatively unimportant for spread of an invasive plant Sources Iverson, Schwartz & Prasad (2004) Bruckmann, Krauss & Stean-Dewenter (2010)
Krauss, Stean-Dewenter & Tscharntke (2003);Krauss et al. (2004) Hill et al. (2001)
Wilson, Davies & Thomas (2010) Reviewed by Mortelliti, Amori & Boitani (2010) Thomas, Simcox & Hovestadt (2010) Karubian et al. (2005) Tinker, Doak & Estes (2008) Poyry et al. (2009) Reviewed in Hanski (1999: Chapter 9) Thomas et al. (2001)
Dauber et al. (2010)
Bruckmann, Krauss & Stean-Dewenter (2010)
Eycott et al. (2009)
Gilbert-Norton et al. (2010)
Davies & Pullin (2007)
Andrew & Ustin (2010)
Area, quality and connectivity 151

Table 1. (Continued) Landscape attribute Matrix (non-breeding habitat) quality Summary of evidence On average, 13-fold increases in exchange rates between patches, based on systematic review of movements between patches separated by more versus less benign matrix Meta-analysis found an eect of the type of matrix surrounding habitat patches in most studies, but such eects were smaller than patch size or isolation eects, and strongly species-specic (whenever species were compared). Ranking of alternative landscape congurations was the same with a simple habitat-patch model as one with complex matrix-dependent movement behaviour Sources Eycott et al. (2009)
Prevedello & Vieira (2010)
Jepsen et al. (2005)
This leads to the issue of robustness. Understanding and modelling the detailed behavioural responses of individuals to multiple landscape elements is an interesting area of research. However, if such details are to be incorporated within multigenerational distribution models, we would argue that other details (e.g. habitat changes, local adaptation, species interactions, evolution of dispersal) are equally relevant, and that they all have associated uncertainties. The robustness of such complicated modelling will almost necessarily be low. The intention of the original Hodgson et al. (2009) argument was to identify robust, easy to understand and easy to implement conservation strategies.
What role for connectivity conservation?

It is possible that everybody understands connectivity conservation to mean a holistic, landscape-to-continental scale conservation method that accounts for all important factors (including habitat quality, quantity, spatial habitat arrangements, the character of the intervening landscape), and the consequences of these for persistence and resilience of multiple species, in a location-specic manner (Doerr, Barrett & Doerr 2011). If so, our fears about an over-emphasis on connectivity may be unfounded. But if connectivity conservation is all this, then the term is actually an unneeded alias for biodiversity conservation. Such a re-branding may be useful to stimulate investment in conservation, but we should still be aware of the potential for confusion it creates. Our original concern was not that there were problems with every activity labelled connectivity conservation, but that the review of Heller & Zavaleta (2009) found that increasing connectivity was the predominant proposed solution for conservation under climate change. We think that considerations of connectivity have an important place in conservation planning (e.g. Moilanen et al. 2005; Moilanen & Wintle 2007; Moilanen, Wilson & Possingham 2009; Carroll, Dunk & Moilanen 2010). Recent awareness of connectivity has undoubtedly been helpful to conservation: it has freed conservationists from focusing too narrowly on individual protected areas, and has brought landscape-scale and spatial considerations into conservation. But our concern was how connectivity is often seen as the solution when the funda-
mental problem is an inadequate quantity of high-quality habitats. It is fortunate that many of the existing connectivity conservation programmes such as the Great Eastern Ranges (GER) initiative are heavily focused on increasing and consolidating natural habitat area (the rst three of ve GER goals; Mackey, Watson & Worboys 2010). This serves to reinforce the impression that connectivity conservation may be a new name for the business of conservation that has long recognised the role of habitat quantity, quality, and connectivity (e.g. Diamond 1975). Based on the weight of current evidence (Table 1) and our original arguments (Hodgson et al. 2009), we remain of the opinion that maintaining (and where feasible restoring) large areas of environmentally diverse, high-quality (low human impact) breeding habitats should be the primary focus of conservation when resources limit the range of conservation actions that can take place. Compared to this, structural features of the matrix which are not breeding habitat for many or any species are a minor consideration.
Acknowledgements
We thank Amy Eycott, Jeremy Thomas and Kevin Watts for providing material permission to cite work in press.
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2009 Reports Re Connectivity

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Journal of Applied Ecology 2009, 46, 964969

Climate change, connectivity and conservation decision making: back to basics

Spatial conservation planning: the basics

*Correspondence author. E-mail: J.Hodgson@leeds.ac.uk

2009 The Authors. Journal compilation 2009 British Ecological Society

Climate, connectivity and conservation 965

Connectivity and uncertainty

966 J. A. Hodgson et al.

Dispersal mechanism, etc.

is quantified in terms of Area Quality

Connectivity (proxy for immigration rates)

The new world order

968 J. A. Hodgson et al.

Climate, connectivity and conservation 969

Journal of Applied Ecology 2011, 48, 143147

*Correspondence author. E-mail: veronica.doerr@csiro.au

Dispersal behaviour and structural connectivity

Connectivity and dispersal behaviour 145

Is quantified in terms of Area Quality Structural connectivity

Potential rates of dispersal between subpopulations

Connectivity conservation is more than just conserving connectivity

Moving beyond aggregation in large-scale conservation planning models

Reasons to be cheerful indeed!

Connectivity and dispersal behaviour 147

Journal of Applied Ecology 2011, 48, 148152

Relative importance of landscape attributes

*Correspondence author. E-mail: jenny.hodgson@york.ac.uk

NON-BREEDING HABITAT (OR THE MATRIX)

Uncertainty and robustness

SPATIAL ARRANGEMENT OF HABITATS

150 J. A. Hodgson et al.

Dauber et al. (2010)

Bruckmann, Krauss & Stean-Dewenter (2010)

Eycott et al. (2009)

Gilbert-Norton et al. (2010)

Davies & Pullin (2007)

Andrew & Ustin (2010)

Area, quality and connectivity 151

Prevedello & Vieira (2010)

Jepsen et al. (2005)

What role for connectivity conservation?

152 J. A. Hodgson et al.

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