The workshop will produce a report listing dozens of potential linkages. Conservationists have only enough money, planning capacity, and attention spans to attack the most important ones.
There are two ways to think about importance. One is the biological value of the linkage: If the linkage is lost, which species would become extinct or at significantly greater risk of extinction? Which species might persist, but in such small numbers that they would be ecologically irrelevant? How much degradation would occur in ecosystem processes such as top-down control by large carnivores, gene flow, recolonization after disturbance, seasonal migration, interspecific competition, and evolution?
A second way to think about importance is threat and opportunity. A potential linkage can also be more important because it is at greater risk of being irreversibly lost if we do not conserve it immediately. Because conservationists must be opportunistic, we also want to give higher priority to a linkage if there is an active conservation effort already underway.
We recommend considering the two types of importance separately, such that each potential linkage can be scored in two dimensions as indicated in the graph below. Potential linkages in the upper right quadrant would be the top priorities.
How do you get those scores for biological value, and for threat and opportunity? You guessed it—another workshop involving all interested stakeholders. Most participants will come to the meeting wanting to ensure that their pet linkage is a high priority, or that linkages serving their pet wildland are conserved. This is natural. Conservationists are motivated more by love of place than love of abstract ideas like biodiversity and ecosystem function. Because “it's all important” and “it's all about love,” some participants may resist attempts at quantification. But you can't prioritize by comparing one participant's love for linkage A with another person's love for linkage B.
Before the workshop, set up a spreadsheet with columns for at least 10 criteria related to biological value and at least 6 criteria related to threat and opportunity, and one row for each linkage. Above the header row, have a row in which the weight of each criterion can be set and changed. Set up a column that multiplies row entries by weights and sums the weighted scores to produce overall biological value score and an overall threat and opportunity score for each linkage. Link these two columns to an x-y graph, so that participants can see where each linkage falls compared to others.
Fill in as many columns as possible before the workshop begins. For instance, you can calculate size—or at least size class—of each wildland to be connected by a potential linkage. Some columns (e.g., habitat quality in the smaller wildland block) may require information from participants, or may be derivable from a GIS (if for example, you are willing to use road density as a surrogate for habitat quality). You want to spend most workshop time arguing about values (weights), not about mere facts.
We have participated in enough of these workshops to know that it is pointless for us to propose weights for criteria, or even an exhaustive list of potential criteria. However, in our experience, the following criteria will be viewed as important by all participants, and will have relatively high weights:
Threat relates to the risk that roads, canals, urbanization, border security operations, or other problems will sever the linkage if we do not act now. Participants can decide whether they want to consider current threat or anticipated future threat. Most workshops ignore threats such as off-road vehicle use or agricultural conversion, because these are more reversible than urbanization and roads. Some workshops started with separate scores for each threat, but used only the maximum threat score, reasoning that a corridor at dire risk of being closed by urbanization and highways is not twice as threatened as a corridor threatened by only one of these factors.
Opportunity typically relates to active conservation efforts. If several local groups and funders are working to conserve connectivity in the area, a linkage design would be more useful than it would be in area where local planners are openly hostile to conservation and no conservation groups are ready to push the plan forward. A potential linkage can also be given high priority if the state transportation agency anticipates a major new project in the area. The rationale is that the linkage design would provide timely input into the transportation planning process.
Adding threat and opportunity scores is like adding apples and oranges. Participants at every workshop commented on the incongruity. But participants have always agreed that it produces rankings that better reflect the non-biological value of a potential linkage. No participant has argued for a third dimension to the prioritization scheme.
Participants will use scientific evidence to argue for a relatively large or small weight for a criterion. But is size of the wildland blocks 50%, 100%, or 200% more important than presence of an endangered species in the linkage area? The principles of conservation biology, ecology, and related sciences cannot answer this question because it is a matter of values. The prioritization process is not about finding 'the correct weights' but rather about consistently applying a consensus set of weights to all of the potential linkages.
Sometimes a participant, upset that their pet linkage is in the upper left quadrant, will propose a new biological value criterion that might push their linkage to the upper right quadrant. Or a participant might suggest a weighting scheme that strikes you as just plain silly. The beauty of the workshop format is that you do not have to argue about values. Instead, you try the new scheme, and use the spreadsheet to instantly show participants how the new scheme rearranged linkages in the prioritization graph space. If the participant sees a silly collection of potential linkages in the upper right quadrant, he or she will withdraw their selection. Alternately, you may be surprised to learn that the suggestion improved the prioritization!
Determining the criteria and scoring system is an iterative process. Participants gradually reach consensus on the conceptual underpinnings of the gestalt ratings that each person held at the start of the workshop. The process does not pretend to seek “truth.” Instead the process forces every participant to be consistent, and to discuss their conservation values in a respectful way. One could even argue that values are formed by this sort of public discussion. By the end of every workshop, almost every participant will agree that the consensus scheme is superior to their own initial guess, proving once again that “none of us is as smart as all of us.”
The process of designing corridors is frustrating, because there are many decisions to make throughout. After spending way too many meetings arguing over the details related to designing corridors, and getting a few too many bruises on our foreheads from repeatedly banging our heads against our desks, we thought maybe we could save you some of the agony.
For a printable version of this material, download the Conceptual Steps PDF on our downloads page!
There are many potential ways to design wildlife corridors, ranging from a hand drawn map, to acquiring any possible land between two blocks of habitat, to the GIS-intensive modeling discussed throughout the site. Neither of these techniques are necessarily better, and all are probably optimal in some situations. If we could stress one thing, it would be to use modeling as an aid to think about how to best design coridors for a landscape, not as a replacement for thinking! Don't model because you think you have to. Don't expect to get the truth. Expect to get frustrated, and come up with more questions than answers.
Do you want to design a corridor within one habitat type? Then our tools and methods probably won't help you too much. How about a 30-m urban greenway? Again, GIS modeling might not be the best approach. Our methods, together with the CorridorDesigner ArcGIS tools, are best suited for designing corridors in a heterogenous landscape at a regional (e.g. 2 - 500 km long) scale. We have found GIS-based corridor modeling to be a useful tool which helps us design the best corridors we can, but it's certainly not the only game in town!
While CorridorDesigner provides one method of modeling wildlife corridors with ArcGIS, it's not the only game in town. Here are some other free tools you may find handy for modeling wildlife or ecological corridors, connectivity, or habitat. Please note: we're just providing the links, and can probably not offer help with any problems you may have with the following tools.
Linkage Mapper is a GIS tool designed to support regional wildlife habitat connectivity analyses. Linkage Mapper uses GIS maps of core habitat areas and resistances to identify and map linkages between core areas. Each cell in a resistance map is attributed with a value reflecting the energetic cost, difficulty, or mortality risk of moving across that cell. Resistance values are typically determined by cell characteristics, such as land cover or housing density, combined with species-specific landscape resistance models. As animals move away from specific core areas, cost-weighted distance analyses produce maps of total movement resistance accumulated.
Circuitscape is a stand-alone Python program which borrows algorithms from circuit theory to predict patterns of movement, gene flow, and genetic differentiation among populations in heterogeneous landscapes. It uses raster habitat maps as input, and predicts connectivity and movement patterns between user-defined points on the landscape. Circuitscape is distributed by Brad McRae of the National Center for Ecological Analysis and Synthesis (NCEAS) at University of California, Santa Barbara.
The Connectivity Analysis Toolkit provides conservation planners with newly-developed tools for both linkage mapping and landscape-level 'centrality' analysis. Centrality refers to a group of landscape metrics that rank the importance of sites as gatekeepers for flow across a landscape network. The Toolkit allows users to develop and compare three contrasting centrality metrics based on input data representing habitat suitability or permeability, in order to determine which areas, across the landscape as a whole, would be priorities for conservation measures that might facilitate connectivity and dispersal. The Toolkit also allows application of these approaches to the more common question of mapping the best habitat linkages between a source and a target patch.
FunConn is a functional connectivity modeling toolbox for ArcGIS, distributed by Dave Theobald and the Natural Resource Ecology Laboratory at Colorado State University. The goal of the functional connectivity model is to allow landscape connectivity to be examined from a functional perspective. Functional connectivity recognizes that individuals, species or processes respond functionally (or behaviorally) to the physical structure of the landscape. From this perspective, landscape connectivity is specific to a landscape and species/individual/process under investigation.
Formerly called ArcRstats, the HabMod/ConnMod toolboxes produce multivariate habitat prediction rasters using ArcGIS and the open-source R statistical package for implementing classification and regression (CART), generalized linear models (GLM) and generalized additive models (GAM), and includes a connectivity modeling toolbox. The toolbox is distributed by Pat Halpin and the Marine Geospatial Ecology Laboratory at Duke University
Conefor Sensinode quantifies the importance of habitat areas for the maintenance or improvement of landscape connectivity. It is conceived as a tool for decision-making support in landscape planning and habitat conservation, through the identification and prioritization of critical sites for ecological connectivity.
Path Matrix is a tool for ArcView 3 to compute matrices of effective geographic distances among samples, based on a least-cost path algorithm. It was developed by Nicholas Ray of the Computational and Molecular Population Genetics Lab, University of Bern, Switzerland.
openModeller aims to provide a flexible, user friendly, cross-platform environment where the entire process of conducting a fundamental niche modeling experiment can be carried out. The software includes facilities for reading species occurrence and environmental data, selection of environmental layers on which the model should be based, creating a fundamental niche model and projecting the model into an environmental scenario.
StatMod is an extension for ArcView 3.3 that helps users create logistic regression and CART models using the statistical packages SAS and S-PLUS. It is distributed by Chris Garard of Utah State University.
Maxent provides a maximum-entropy approach for species habitat modeling. The software takes as input a set of layers or environmental variables (such as elevation, precipitation, etc.), as well as a set of georeferenced occurrence locations, and produces a model of the range of the given species. It is distributed by Princeton University.
GRASP (Generalized Regression and Spatial Prediction) is a package for the statistical softwares R and S-PLUS for creating predictive models of species distributions using Generalized Additive Models (GAMs). It is distributed by A. Lehmann, J.R. Leathwick and J.McC. Overton at the Landcare Research Institute, New Zealand.
BioMapper is a stand-alone package for developing ecological niche and habitat suitability models using only a species presence data. It is distributed by Alexandre Hirzel of the Laboratory For Conservation Biology, University of Lausanne, Switzerland.
DesktopGarp is a software package for biodiversity and ecologic research that allows the user to predict and analyze species distributions using presence data. The package is distributed by the University of Kansas Natural History Museum.
Spatial Data Modeler is a collection of geoprocessing tools for adding categorical maps with interval, ordinal, or ratio scale maps to produce a predictive map of where something of interest is likely to occur. The tools include the data-driven methods of Weights of Evidence, Logistic Regression, and two supervised and one unsupervised neural network methods, and a knowledge-driven method Fuzzy Logic.
PatchMorph is an extension for ArcMap which delineates habitat patches from a habitat suitability or land cover map.
Marxan delivers decision support for reserve system design. Marxan finds reasonably efficient solutions to the problem of selecting a system of spatially cohesive sites that meet a suite of biodiversity targets.
PANDA is a stand-alone application developed to provide a user friendly framework for systematic protected areas network design to ArcGIS users. Through the use of P.A.N.D.A. the designer can explore different hypothetical configurations of a system of protected areas in the planning area.
CLUZ is an ArcView GIS interface that allows users to design protected area networks and conservation landscapes. It can be used for on-screen planning and also acts as a link for the MARXAN conservation planning software. It is currently being developed at DICE and is funded by the British Government through their Darwin Initiative for the Survival of Species.
LINK is a set of ArcGIS tools designed to analyze habitat patterns across a landscape. LINK uses species habitat matrices to model potential species habitat and habitat diversity. What sets LINK apart from its predecessors is that it uses raster data sources—raster data sources allow LINK to model habitat over a much larger area than its vector based ancestors.
Jenness Enterprises has created many extensions for ArcView 3.3 useful to ecological applications. Jeff Jenness created the CorridorDesigner ArcMap extension.
Hawth's Analysis Tools is an extension for ArcGIS (specifically ArcMap). It is designed to perform spatial analysis and functions that cannot be conveniently accomplished with out-of-the-box ArcGIS. Most of these analysis tools have been written within the context of ecological applications such as movement analysis, resource selection, predator prey interactions and trophic cascades.
Although roads and urban areas usually occupy only a small fraction of a linkage design, their impacts threaten to block animal movement between the habitat blocks. In this section, we review the potential impacts of these features on ecological processes, identify specific barriers in the linkage design, and suggest appropriate mitigations.
While roads and fences impede animal movement, and the crossing structures we recommend are important, we remind the reader that crossing structures are only part of the overall linkage design. To restore and maintain connectivity between any two wildland blocks, it is essential to consider the entire linkage design, including conserving the land in the linkage. Indeed, investment in a crossing structure would be futile if habitat between the crossing structure and either wildland block is lost.
While the physical footprint of the nearly 4 million miles of roads in the United States is relatively small, the ecological footprint of the road network extends much farther. Direct effects of roads include road mortality, habitat fragmentation and loss, and reduced connectivity, and the severity of these effects depends on the ecological characteristics of a given species. Direct roadkill affects most species, with severe documented impacts on wide-ranging predators such as the cougar in southern California, the Florida panther, the ocelot, the wolf, and the Iberian lynx (Forman et al. 2003).
In a 4-year study of 15,000 km of road observations in Organ Pipe Cactus National Monument, Rosen and Lowe (1994) found an average of at least 22.5 snakes per km per year killed due to vehicle collisions. Although we may not often think of roads as causing habitat loss, a single freeway (typical width = 50 m, including median and shoulder) crossing diagonally across a 1-mile section of land results in the loss of 4.4% of habitat area for any species that cannot live in the right-of-way. Roads cause habitat fragmentation because they break large habitat areas into small, isolated habit patches which support few individuals; these small populations lose genetic diversity and are at risk of local extinction.
In addition to these obvious effects, roads create noise and vibration that interfere with ability of reptiles, birds, and mammals to communicate, detect prey, or avoid predators. Roads also increase the spread of exotic plants, promote erosion, create barriers to fish, and pollute water sources with roadway chemicals (Forman et al. 2003). Highway lighting also has important impacts on animals (Rich and Longcore 2006).
Wildlife crossing structures that have been used in North America and Europe to facilitate movement through landscapes fragmented by roads include wildlife overpasses & green bridges, bridges, culverts, and pipes. While many of these structures were not originally constructed with ecological connectivity in mind, many species benefit from them (Clevenger et al. 2001; Forman et al. 2003). No single crossing structure will allow all species to cross a road. For example rodents prefer to use pipes and small culverts, while bighorn prefer vegetated overpasses or open terrain below high bridges. A concrete box culvert may be readily accepted by a mountain lion or bear, but not by a deer or bighorn sheep. Small mammals, such as deer mice and voles, prefer small culverts to wildlife overpasses (McDonald & St Clair 2004).
Wildlife overpasses are most often designed to improve opportunities for large mammals to cross busy highways. Approximately 50 overpasses have been built in the world, with only 6 of these occurring in North America (Forman et al. 2003). Overpasses are typically 30 to 50 m wide, but can be as large as 200 m wide. In Banff National Park, Alberta, grizzly bears, wolves, and all ungulates (including bighorn sheep, deer, elk, and moose) prefer overpasses to underpasses, while species such as mountain lions prefer underpasses (Clevenger & Waltho 2005).
Wildlife underpasses include viaducts, bridges, culverts, and pipes, and are often designed to ensure adequate drainage beneath highways. For ungulates such as deer that prefer open crossing structures, tall, wide bridges are best. Mule deer in southern California only used underpasses below large spanning bridges (Ng et al. 2004), and the average size of underpasses used by white-tailed deer in Pennsylvania was 15 ft wide by 8 ft high (Brudin 2003). Because most small mammals, amphibians, reptiles, and insects need vegetative cover for security, bridged undercrossings should extend to uplands beyond the scour zone of the stream, and should be high enough to allow enough light for vegetation to grow underneath. In the Netherlands, rows of stumps or branches under crossing structures have increased connectivity for smaller species crossing bridges on floodplains (Forman et al. 2003).
Drainage culverts can mitigate the effects of busy roads for small and medium sized mammals (Clevenger et al. 2001; McDonald & St Clair 2004). Culverts and concrete box structures are used by many species, including mice, shrews, foxes, rabbits, armadillos, river otters, opossums, raccoons, ground squirrels, skunks, coyotes, bobcats, mountain lions, black bear, great blue heron, long-tailed weasel, amphibians, lizards, snakes, and southern leopard frogs (Yanes et al. 1995; Brudin III 2003; Dodd et al. 2004; Ng et al. 2004). Black bear and mountain lion prefer less-open structures (Clevenger & Waltho 2005). In south Texas, bobcats most often used 1.85 m x 1.85 m box culverts to cross highways, preferred structures near suitable scrub habitat, and sometimes used culverts to rest and avoid high temperatures (Cain et al. 2003). Culvert usage can be enhanced by providing a natural substrate bottom, and in locations where the floor of a culvert is persistently covered with water, a concrete ledge established above water level can provide terrestrial species with a dry path through the structure (Cain et al. 2003). It is important for the lower end of the culvert to be flush with the surrounding terrain. Many culverts are built with a concrete pour-off of 8-12 inches, and others develop a pour-off lip due to scouring action of water. A sheer pour-off of several inches makes it unlikely that many small mammals, snakes, and amphibians will find or use the culvert.
| Effect of roads | |||
|---|---|---|---|
| Characteristics making a species vulnerable to road effects | Road mortality | Habitat loss | Reduced connectivity |
| Attraction to road habitat | x | ||
| High intrinsic mobility | x | ||
| Habitat generalist | x | ||
| Multiple resource needs | x | x | |
| Large area requirements/low density | x | x | x |
| Low reproductive rate | x | x | x |
| Behavioral avoidance of roads | x | ||
Based on the small but increasing number of scientific studies on wildlife use of highway crossing structures, we offer these standards and guidelines for all existing and future crossing structures intended to facilitate wildlife passage.
The previous previous describe how to construct one single-species corridor. The union of single-species corridors constitutes a preliminary linkage design. This is morphed into a final linkage design after you tweak the design to accommodate focal species for which no corridor was modeled, buffer against edge effects, and remove areas from the design that likely increase financial costs of acquisition or management without substantially improving ecological utility.
The previous procedures produce a corridor model for a single species. You will repeat these procedures for several focal species. Now you are ready to create a comprehensive linkage design. There is more than one way to do this, but we recommend that this process include the following steps:
After creating corridor models for all focal species, we recommend joining all single-species corridors into a preliminary linkage design. This union of corridors is the most obvious way to fulfill our goal of “no species left behind.”
By overlaying the linkage design on a map of modeled habitat patches for each focal species, you will usually find that most breeding patches for each species were already captured by the union of corridors. Sometimes one or two species-specific habitat patches can be added if the addition would reduce the need for individuals to move distances longer than the estimated dispersal capability of the focal species. Because dispersal distances are only known for few species, and probably biased low due to difficulties associated with collecting dispersal data, we recommend using the longest known dispersal distance of the species as an estimate of dispersal capability. If there are no data for a species, data for a closely related species can be used, or another species of similar body size, mobility, and natural history.
Least-cost procedures will always produce a least cost corridor or path—even if the best is entirely inadequate for the focal species. This overlay procedure is a good way to assess how well the linkage design serves each species. Evaluating corridors and linkages describes additional descriptors you can use to assess how well a linkage design or corridor serves each species.
In Who to connect: selecting focal species, we mentioned that it may not be possible or appropriate to develop a corridor model for some focal species. In our efforts, we address needs of these species in one of the following ways:
If there are enough data to model potential habitat, map potential population and breeding patches. Overlay this patch map on the preliminary linkage design. Ask a species expert if the preliminary linkage design captured enough of these habitat patches to conserve the species. If not, add any additional patches near the preliminary linkage design that would improve the species' prognosis.
Overlay a map of known occurrences of the species on the preliminary linkage design. Ask a species expert if it would be helpful to expand the linkage design to include more of these occurrences, and if so, whether the occurrences should be included as disjunct steppingstones or by widening part of the linkage design.
In the arid southwest, adding perennial streams to the linkage design is a simple way to meet the needs of fishes and other riparian obligates. Buffering each stream 100m from the edge of the riparian zone is useful to reduce pollutants, sedimentation, and other edge effects. While many studies document edge effects on streams only out to about 50m, there are already many irreversible human alterations within 50m of most streams. Attempting to have 100m buffers in other areas may compensate for these impacts. Finally, the uplands adjacent to the stream are important movement areas for many terrestrial species. A 100m buffer helps reduce human disturbance and edge effects in this upland zone.
At this point, your preliminary linkage will likely have multiple strands, each serving the needs of one or more focal species. You may have some deeply looped strands. The thin strands winding through the linkage design in the map below capture the only continuous strands of perennial waters between the two wildland blocks.
A linkage design like this is clearly not analogous to the linear corridors—such as hallways in our office buildings, or interstate highways—we construct to facilitate human movement. This is not surprising. We want the linkage to fit the species needs. We do not try to make the species fit the corridor.
Next, you will evaluate the preliminary linkage design as modified so far, and widen one or more strands to provide the following benefits of wide linkage strands:
Although edge effects and home range widths of focal species are relevant to linkage width, we recommend asking not “how narrow a linkage strand might possibly be useful to focal species?” but rather “what is the narrowest width that is not likely to be regretted after the adjacent area is converted to human uses?”
Conservation dollars are limited, so you do not want to propose a linkage design that includes large areas that do not substantially improve connectivity for a species.
In our linkage designs, we examined the various slices of single species corridors to identify the “balloon” areas that emerged when we selected a slice that met our target minimum width (e.g., areas in the eastern ends of Slice 3 and Slice 4 in the linkage below).
To determine if the balloon areas are important, we overlay the selected slice on a map of population patches, breeding patches, and habitat quality. We look for opportunities to delete areas so long as the deletion will not significantly increase the travel cost for that species, or any other focal species.
This entire section describes procedures that are only weakly quantitative. Many procedures, such as the “no-regret” standard for width, are quite subjective. Does it really matter? Is it worth it?
Modeling is a tool to help you examine your landscape using all of your brain-power and the best available scientific knowledge. Modeling is not a substitute for such hard thinking!
In practice, all this tweaking has rarely caused big changes in our linkage designs. But we still go through the checklist. Mulling over the linkage design in this way is better than rushing a plan out the door and later wishing we had addressed these issues.
In a few cases, these considerations have improved the linkage design, and we sleep better at night for that. In many cases, it helps us more fully discuss the likely benefits (or lack thereof) for particular species and other conservation goals.
Least cost methods always provide a “best” solution, even when the best is not very good. You need tools that can effectively describe how well your proposed linkage design serves each focal species. These same tools can be used to compare the linkage design to alternative designs that may be proposed to meet cost or political constraints.
Three of the most useful tools are:
As conservation investors try to implement the linkage design you helped produce, they will face some tough choices. For example, they may have an opportunity to buy two huge parcels in the linkage area from conservation-friendly landowners. These two parcels form a continuous swath that overlaps about half of your linkage design, plus an even larger amount of land outside the linkage design. This opportunity represents an alternative linkage design. The investors want to know: How will it compare to the optimal linkage design? Is it almost as good, half as good, or markedly inferior? In another linkage area, a large development company owns most of the land, and wants to develop 3 new cities there. They propose an alternative linkage design that allows them to proceed with their development plans. The developer produces a glossy booklet touting the virtues of their alternative. The County Planning & Zoning Department needs answers to the same questions: How will this alternative compare to the optimal linkage design? Is it almost as good, half as good, or markedly inferior?
There are an endless number of such scenarios. As the analyst, you cannot simply say “We've presented the optimum design. Now please go away.” You want to provide some useful descriptors that allow decision makers to make choices with their eyes open.
Please avoid the following ways of summarizing utility of alternative linkage designs:
There is no silver bullet. You will have to provide several useful descriptors of utility and help summarize them in a way decision-makers can understand.
In the illustration, the alternative is clearly inferior. Notice that a frequency distribution is much more informative than the mean habitat suitability. You may also want to provide frequency distributions for other GIS layers, such as land cover types, elevation, and topographic positions. All of these can help decision makers appreciate the differences between alternatives.
In the illustration, the inset graph shows width of a corridor at each point along the corridor midline (the purple line on the map). By placing your ruler horizontally at any y-axis value, you can see the number and length of bottlenecks that are narrower than any threshold of interest. For instance, in this case, there are no bottlenecks as severe as 250 m. There are six bottlenecks narrower than 400m, the longest of which is 1.2 km long (from 6.8km to 8.0 km along the midline). By producing a similar graph for an alternative, you provide a useful way to compare them.
The linkage design below has two strands. For Species A, six potential population patches are fully or completely overlapped by the linkage design. In addition to presenting this map, you can provide a list of interpatch distances that a dispersing animal would have to cross to move between wildland blocks. These distances are represented by the green lines connecting 5 of the patches to the wildland blocks. Notice that the patch in the lower strand is not part of this movement path, and is not included, and that the green line is confined to a single linkage strand. Thus the green line represents our best estimates of interpatch distances that would need to occur after the remaining matrix has been converted to uses incompatible with wildlife movement.
You can produce this same output for any alternative linkage design. The results could be displayed in the format of the following table. The modeled distances are most useful if you compare them to the species' estimated dispersal ability. In the example below, for instance, three interpatch distances in the Alternative exceed the dispersal ability of the species, compared to none in the proposed linkage design.
Cost distance of each pixel is the lowest possible cumulative resistance from that pixel to terminuses in each habitat block. A map of cost-distance always produces continuous swaths of permeable pixels; these swaths have been used as the basis for all published corridor designs. The most difficult issue is selecting a corridor width ('slice') wide enough to facilitate movement, but narrow enough to minimize monetary costs of conservation. We recommend a corridor slice wider than the home range width for corridor-dwellers, and various ad-hoc procedures for passage species.
As illustrated in the figure below, low resistance pixels may not form a continuous swath. To identify well-connected low-resistance pixels, all published corridor designs calculated each pixel's accumulative cost-distance as the lowest possible cumulative resistance from that pixel to terminuses in each habitat block.
These cost-distance values do form continuous swaths, as illustrated in the next figure. In this figure, the pixels with the lowest cost-distance are shown in black, and progressively lighter colors represent progressively higher thresholds of cost-distance.
As the maximum cost-distance increases, the pixels with lowest cost-distance define a nested set of increasingly broad “slices” of the landscape. As the cost threshold increases from slice 1 to slice 3 to achieve minimum width, w, the modeled corridor becomes very wide in areas outside the bottleneck. As the cost threshold increases even more (slice 4) the modeled corridor gains additional strands.
Note that resistance or travel cost is an attribute of a pixel resulting from the pixel's internal characteristics, and is simply the inverse of the pixel's permeability or habitat suitability. In contrast, cost-distance (sometimes called effective distance or cost-weighted distance) is a pixel attribute resulting from the pixel's resistance plus the resistance of a chain of pixels reaching to each terminus. Thus in moving from resistance to cost-distance, you are moving from a single pixel's content to its landscape context.
The previous analytic steps produce a map of increasingly wide corridors displayed as nested polygons, each defined by the largest cost allowed in the polygon. Although a wider corridor is better in the sense that broad swaths include narrower ones, financial constraints favor smaller corridors. In multiple-species linkage designs, partial overlap among species corridors will tend to widen each single-species corridor, so the width needs only to be “about right” at this state. But still, you need a stopping rule to map each single-species corridor.
For a corridor dweller, one suggestion is that the corridor should be as wide as the species' typical home range. However, if the focal species is strongly territorial, this could result in corridors fully occupied by home ranges where social interactions impede movement through the corridor. Such social interactions probably led to the disturbing case of an occupied but non-functional corridor previously described. Overview of habitat modeling. Therefore we suggest that minimum width for a corridor dweller should be at least two home range widths along all or most of the length of the corridor.
Species able to move through a corridor in one or several days can doubtless pass through areas narrower than their typical home range. Like other corridor designers, we consider the biology of the focal species to recommend a reasonable minimum width. But a cost threshold that achieves a minimum width in a bottlenecked area may cause impractically broad swaths elsewhere (e.g. swath 4 in the figure above). Reasonable procedures include:
No matter what approach you adopt, choosing the right corridor slice is an iterative process of examining a series of slices and evaluating the advantages and disadvantages of moving to the next larger or smaller slice. An objective set of decision rules, and an automated way to run them, would be significant advances. However, given the myriad of possible landscape configurations and reasonable differences of opinion about when a corridor is “big enough,” this may be an impossible goal.
We see no excuse for using least cost paths instead of corridor swaths to define wildlife corridors. A least-cost path is only one pixel wide. Because it is easy to identify in GIS software, it is popular. But a pixel-wide path surrounded by otherwise inappropriate habitat is unlikely to be used, and would be biologically irrelevant. Furthermore, the location of a least cost path is highly sensitive to pixel size and errors in classifying single pixels. Finally, you would never recommend conservation of a pixel-wide path.
The starting and ending points of a corridor can greatly impact the location of the modeled corridor. A terminus is a part of a wildland block that forms one end of a modeled corridor. You can define it as a point (or pixel), a linear edge (e.g., the wildland boundary), or a patch (population patch or breeding patch). Often there is more than one potential terminus in each wildland block.
Some linkage designs use each wildland block in its entirety as a terminus for each single-species analysis. However, in our experience this procedure sometimes produces a corridor that connects to a part of the wildland block far from any potential habitat for the focal species. You can avoid this result by allowing only patches of potential breeding habitat within each wildland block serve as a potential terminus. Better yet, if you have a map or known breeding patches (Arizona has such data for pronghorn, for example), you should use each known breeding patch as a terminus.
Wildland blocks vary in how much known or potential breeding habitat occurs in the block. A flexible strategy is to select start/end points in the following order of priority:
If you have no pixels of breeding habitat in either wildland block, you probably should not build a corridor model for that species. You can modify the linkage design to serve this species later.
An important component of cost-distance is Euclidean distance. If your two wildland blocks nearly touch at one or more locations, least-cost modeling will tend to identify the narrowest gap as the best corridor, even if the modeled corridor is low in habitat value and does not effectively connect the most important habitat patches between wildland blocks. To reduce this problem, we often give the GIS model “room to run” by defining the terminuses well inside of the wildland blocks. The easiest way to do this is to draw roughly parallel lines a few kilometers apart, with most matrix land centered between the two lines, and then selecting population patches behind these lines. Even if this modification does not change the modeled corridor for the focal species, it will demonstrate that the modeled corridor is not merely an artifact of a point where two wildland blocks nearly touch.
To get from a habitat suitability map to a corridor map, we follow three steps:
We have been operating under the assumption that habitat suitability and habitat permeability are synonyms. In Overview of habitat modeling, we admitted that we don't know this for sure. The big leap of faith is assuming that habitat suitability is the same as habitat permeability. At this point, we simply define resistance or travel cost as the inverse of suitability or permeability, such that
Resistance (cost of travel through a pixel) = Maximum suitability minus pixel suitability.
In this tutorial, we have defined habitat suitability and permeability on a scale of 0-100. Thus,
Resistance (travel cost) = 100-pixel suitability.
Resistance or travel cost reflects the ecological cost of travel through a pixel. In general resistance increases with the risk of dying in the pixel and the energetic cost of travel through the pixel. Resistance decreases with how much food, water, and cover an individual might find in the pixel, or the probability of finding a mate and breeding in an area that includes the pixel. It is not necessarily related to the speed of travel through the pixel. In fact, animals may tend to move quickly through costly pixels, to minimize risks and more quickly get to better habitat.
The starting and ending points of a corridor can greatly impact the location of the modeled corridor. A terminus is a part of a wildland block that forms one end of a modeled corridor. You can define it as a point (or pixel), a linear edge (e.g., the wildland boundary), or a patch (population patch or breeding patch). Often there is more than one potential terminus in each wildland block.
We discuss this step further in Defining end points for corridors.
Cost distance of each pixel is the lowest possible cumulative resistance from that pixel to terminuses in each habitat block. A map of cost-distance always produces continuous swaths of permeable pixels; these swaths have been used as the basis for all published corridor designs. The most difficult issue is selecting a corridor width ('slice') wide enough to facilitate movement, but narrow enough to minimize monetary costs of conservation.
We discuss this step further in Cost distance and single-species corridors.
We use the term “corridor” for a swath of land that is best expected to serve movement needs of an individual species after the remaining matrix has been converted to other uses. We use “preliminary linkage” to refer to the union of all single species corridors. We use “linkage design” to refer to this map after it has been modified to (a) serve focal species for which you could not build a corridor model, (b) remove redundant strands, and (c) meet other conservation goals. The linkage design includes not only this map, but also recommendations for mitigating barriers and managing lands in and adjacent to the mapped area.
Sometimes a habitat suitability model needs to be modified to better reflect what we know about a species or study area. We give three scenarios:
Habitat factors such as land cover are often developed using remotely-sensed data that is several years old. When used to create a habitat suitability model, this can result in a model depicting newly developed land as optimal habitat, or recently restored land as unsuitable.
To create more realistic habitat and corridor models, we recommend modifying habitat suitability models to account for previously unmapped influences. One simple way to do this in GIS is to simply digitize the unmapped influence, then reassign all pixels falling within the digitized feature a new score which better reflects your understanding of the new habitat suitability.
We suggest caution if tweaking a habitat map to account for possible new developments. When this modified habitat map is used for corridor analyses, you will still get a corridor, and the corridor may run right through the pixels you just re-mapped as developed. A naïve comparison of the 2 maps would suggest “The development won't affect the corridor.” But even if a change does not affect the location of the corridor, it may affect its quality. To assess the impact on corridor quality, use the metrics outlined in Evaluating corridors.
The typical corridor model is for a species that can move from one wildland block to another in a single movement event of a few days or weeks. Some species—corridor dwellers—take more than one generation to move between wildland blocks. CorridorDesigner lets you build more realistic corridors for such species.
Most corridor models assume that an individual animal can move between wildland blocks in a single movement event of a few hours to a few weeks. These animals can be called passage species, in contrast to corridor dwellers, which require more than one generation to move their genes between wildland blocks. The distinction is based on an interaction between the species and the landscape. Thus a species could be a passage species if the wildland blocks are within dispersal distance, but a corridor dweller where wildland blocks are farther apart. Corridor dwellers must find suitable breeding opportunities within the linkage.
One way to accommodate corridor dwellers is to assign the highest suitability value to patches of potential breeding habitat. This tends to produce a corridor that links those patches in steppingstone fashion. By dispersing from patch to patch, one interpatch movement per generation, these animals can gradually recolonize a linkage and wildland block after a local extinction event, or move their genes between wildland blocks.
In our experience, this procedure makes sense only when modeling a species with a few habitat patches imbedded in a matrix dominated by poor habitat. Do not use this procedure if most of the matrix is breeding habitat. In such a case, the procedure creates a highly linear corridor that often fails to include the highest-quality habitat. Similarly, unless you know the threshold between breeding and non-breeding habitat precisely, don't use this procedure when a large fraction of the matrix is near the estimated threshold. In that situation, a tiny error in the threshold can drastically affect modeled patches and the modeled corridor.
Using a standard habitat suitability model for a species that is dependent on proximity to a critical resource can greatly overpredict the amount of suitable habitat in an analysis area. Neither geometric mean nor arithmetic mean habitat models adequately account for a situation where a species is absolutely dependent on close proximity to one specific resource. For example:
One way to create a more realistic model is to reclassify a habitat suitability model to better reflect declining suitability with increasing distance from a particular critical factor. Using GIS, the basic steps are:
A habitat patch is a cluster of pixels that are good enough, big enough, and close enough together to support breeding by a particular species. In corridor modeling, patches are useful as start and end points for corridors, as steppingstones in the matrix, and as descriptors to evaluate utility of a linkage design for each focal species. In a GIS context, modeling patches requires you to set:
A habitat patch is a cluster of pixels that are good enough, big enough, and close enough together to support breeding by a particular species. “Good enough” means that they have sufficient resources for the animal. “Big enough” reflects the fact that there needs to be enough area to support at least one breeding unit, typically considered a mating pair of animals with overlapping home ranges. “Close enough together” means that the pixels must be clustered, rather than divided into a checkerboard by too much interspersion with pixels of bad habitat. “By a particular species” emphasizes the fact that one species' breeding patch may be another species' worst nightmare.
You can design a linkage without delineating habitat patches. In fact, most corridor designs do not incorporate patch models. But in our experience, modeling patches of breeding habitat is useful in three ways:
To delineate habitat patches, you must specify the threshold habitat quality for breeding, and the minimum area of suitable habitat necessary to sustain a breeding pair or population. This is easily done in a GIS, and in CorridorDesigner, by counting pixels that exceed the threshold value and that can touch at an edge or corner. However, two problems sometimes arise with this procedure in this simple form:
The procedure could fail to recognize some patches usable by an animal with a large home range. Such animals would probably “ignore” a narrow ribbon of non-habitat imbedded in otherwise suitable pixels. However, if that narrow ribbon divides the suitable pixels into two clusters, each slightly below the minimum size, this procedure would not recognize the habitat patch.
Conversely, this procedure could recognize some habitat patches that an animal would probably not use. The extreme example would be a diagonal string of pixels touching only at their corners, surrounded by pixels of very low habitat suitability. In this case, edge effects such as predators, nest parasites, or exotic species might make this area unsuitable for breeding, despite its being identified as a “patch.”
CorridorDesigner gives you an option to address both of these problems by computing the average habitat suitability score of all pixels within a moving window around the focal pixel and using this “neighborhood habitat suitability score” to define patches. In corridor designer we use the neighborhood score only to define patches; each pixel retains its raw score in all other procedures.
Because appropriate, species-specific data are usually lacking, it is difficult to determine the optimal neighborhood size for a species. Estimates of home range size, daily spatial requirements, and the relationship between body mass and spatial requirements may all be useful in determining an ecological neighborhood. In our modeling, we used one of three moving window sizes, namely a 200-m radius, a 3x3-pixel square, and none, depending on our understanding of the biology of the species.
Whether you use raw habitat suitability scores or the habitat suitability in a moving window, you must specify the cutoff between breeding and non-breeding habitat. In some of our early designs, we used an arbitrary habitat suitability scale, and we designated the top 40% of the pixels as potential breeding habitat. This was obviously unsatisfactory: a species might find 100% of one landscape is suitable for breeding, and 0% of another landscape. An arbitrary 40% does not make biological sense. So we switched to a scheme in which habitat suitability scores had a biological meaning, as illustrated below.
Assigning meaning to the scores made it much easier to assign a threshold (60 in this example). Of course, for this to work, you must parameterize habitat models explicitly keeping this framework in mind, instead of applying this framework after having already parameterized a model.
It is useful to map at least one patch size: the area sufficiently large enough to support a breeding event (usually a home range). We recommend also defining a larger habitat patch size capable of supporting a larger population of individuals. We mapped patches in two size classes, namely:
To combine multiple habitat factors into an overall habitat suitability score for each pixel, you must:
The weighted arithmetic mean is the most commonly used algorithm to combine weights, but the weighted geometric mean better reflects a situation in which one habitat factor limits suitability in a way that cannot be compensated by other factors.
To combine multiple habitat factors into one aggregate habitat suitability model, you must first assign weights to each factor that reflect their relative importance. In our linkage designs, we found it intuitive to assign each factor a percentage weight, such that the sum of the weights is 100%. For example, land cover could be assigned a weight of 60%, topographic position a weight of 20%, and distance-to-roads a weight of 20%, making land cover three times more important than the other factors. If a habitat factor is not important for a species, it is assigned a weight of 0%.
Weighting is one of the weakest parts of our models, lacking any underlying theory or hard data. One theoretical issue, for example is this: When the scores are combined across factors, does the overall pixel score still have the same biological interpretation we established when scoring suitability for each factor? Quite honestly, we don't know, but we suspect that the biological meanings have been altered, at least a little bit. The lack of hard data is obvious: We have never built or seen a corridor model that used weights based on empirical data—100% of them are based on expert opinion.
While there are many potential ways to combine the relative influence of multiple factors, we focus on two: weighted arithmetic mean and weighted geometric mean. Under many circumstances, these algorithms will produce a similar habitat suitability model.
The practical difference between the two algorithms is this: weighted arithmetic mean allows a deficiency in one factor to be compensated by other factors, while weighted geometric mean better reflects a situation in which one habitat factor limits suitability in a way that cannot be compensated by other factors.
What are some examples of this difference?
Most linkage designers have used a weighted arithmetic mean algorithm to combine multiple habitat factors. The weighted arithmetic mean is calculated by multiplying the class score times the percentage weight assigned to its factor, then adding across factors. It is equivalent to the weighted overlay function in ArcGIS, and maintains the same range of suitability scores—if you used a 0-100 scale to score classes within each factor, your weighted sum will also be scaled 0-100.
While the weighted geometric mean algorithm is not used as often to build habitat models, we find the approach intuitively appealing.
Weighted geometric mean better models a situation in which a deficit in one factor cannot be compensated by high scores for other factors. For instance, if urban areas are poor habitat under all circumstances, you'd want to combine factors in a way that a pixel of urban habitat doesn't get a high score because it has ideal elevation, topography, and distance to road.
This reflects the limiting factor concept, one of the earliest ideas in ecology. As originally expressed, the idea was that a species population, or an ecosystem flow like primary productivity, is limited by whichever essential factor was most scarce relative to needs. Early ecologists called it Leibig's law of the minimum. It's the same concept you learned when you balanced the equation of a chemical reaction. The amount of chemical product produced depends on the reagent in limited supply. Because we think that habitat suitability probably is limited by the worst factor, we now use this routinely.
The bottom line is that the weighted geometric mean gives more influence to suitability scores near 0 and absolute influence to suitability scores of 0.
Assign a suitability score to each of the different classes within every factor (such as desert scrub or grassland within landcover). Set biologically meaningful thresholds to divide habitat suitability scores into categories, paying particular attention to the suitability threshold required to support breeding habitat. Assign a score of zero only when the species absolutely won't use a particular class.
Whenever possible, recruit an expert biologist knowledgeable of the focal species to parameterize the model for each species. When this is not possible, we recommend recruiting several non-expert biologists to review all relevant literature for the species, parameterize the model independently, then compare and discuss differences and assumptions in parameterizations before averaging them into the final model.
A fundamental assumption is that habitat suitability and permeability are synonyms, and that both are the inverse of ecological cost of travel. Estimating suitability values (this section) and factor weights is the link between the behavior of the focal species as it moves through the landscape and non-ecological GIS data. Virtually all the relevant literature concerns habitat use, not animal movement, so we find it easier to estimate habitat suitability rather than habitat permeability to movement.
In least-cost modeling, habitat suitability and permeability are synonyms. Big numbers indicate good habitat suitability and high permeability, while small numbers indicate poor suitability. In the literature on corridor modeling, the term travel cost is used more commonly than suitability or permeability, and least-cost modeling is the generic term for the most common corridor models. Throughout this tutorial, just remember that cost and suitability are just opposite sides of the same coin, such that cost plus suitability = 100 (or other maximum value). When we get into corridor modeling, we will shift into cost terminology. But for now, it is easier to write and read this discussion using suitability terminology.
We recommend (and CorridorDesigner requires) using a scale with fixed end points (such as 0 to 100) rather than a scale with no upper limit. We also recommend that you use verbal descriptions of threshold values, such as those given below. As you will see in 2.5 Modeling habitat patches, providing biological interpretations to habitat suitability scores provide a rational basis for modeling habitat patches. An arbitrary scale does not have this virtue.
A score of zero should only be assigned to a class when the animal would not use the class, even if the other factors were optimal. For instance, a score of zero for elevations above 7,000 feet means that “the animal won't use this, even if the vegetation, topography, and road density are otherwise ideal.” This would be appropriate if 7,000 feet is the upper elevation limit of the species distribution.
Sometimes you will have to parameterize models yourself. But whenever possible, we recommend recruiting a biologist who is an expert on the species, especially if he or she has worked in or near the linkage analysis area. Even if they have published papers on habitat use, experts have reams of unpublished data and field experience that you can't get by reading papers.
In addition to scoring habitat preferences based on GIS variables, you should also ask the species expert to provide estimates of uncertainty, estimates of factor weights, and estimates of the sizes of areas needed to support a single breeding event and a breeding population.
It has been demonstrated that a species expert will do a better job parameterizing the model if they refer to the scientific literature while they do so. Even if the expert published most of those papers, he or she has forgotten a lot of it, and filling out the form without referencing the literature will result in a poorer model.
Sometimes you will not be able to find a species expert. When this happened to us, we assigned the task to 3 persons on our team. We provided each scorer with copies of the relevant literature, and we each independently filled out the spreadsheet. If our scores differed by ‹20 (on the 100-point scale described above), we used the average. We discussed each score that differed by 20 or more until we reached a consensus score.
Land cover is a factor in every corridor model we've seen, and it tends to come in distinct flavors, so that it makes sense to assign a permeability score to each class . But elevation and distance to road are inherently continuous variables. How do we estimate permeability as a function of a continuous variable?
We don't. Instead we define a few classes of elevation, and a few classes of distance to road, and estimate suitability for each class. But you need not be constrained by our lack of imagination, and there is nothing inherently wrong with developing a function that relates permeability to a continuous variable. All the issues discussed above still apply.
Assuming you are covering this material in order, you now have several sets of permeability scores that look something like this:
| Land cover | Topographic position | Distance to roads | Elevation | ||||
|---|---|---|---|---|---|---|---|
| Class | Score | Class | Score | Classs | Score | Class | Score |
| Pine forest | 60 | Canyon bottom | 80 | 0-50 m | 30 | 0-300 m | 0 |
| Grassland | 30 | Ridgetop | 20 | 50-200 m | 50 | 300-500 m | 40 |
| Urban | 00 | Steep slope | 20 | ›200 m | 90 | 500-1000 m | 100 |
| Agriculture | 30 | Gentle slopes | 50 | 1000-1200 m | 40 | ||
| Riparian | 100 | › 1200 m | 0 | ||||
As you can see, each pixel will have 4 suitability scores—one for each of the four factors. To estimate the overall permeability of a pixel, you must combine these four scores. To do this, you must assign a weight for each factor, and choose an arithmetic operation to apply these weights.
Many habitat models are based on factors such as land cover, topography, and human disturbance, not because they fully describe habitat, but because these are the only relevant factors available as GIS layers. Given the low accuracy of land cover layers and incompleteness of the set of available factors, we guess most models are no more than 70% successful. We discuss how to deal with the crudeness of habitat models, and we recommend metrics for some factors.
We recommend categorical metrics over continuous ones, and use of few rather than many categories within each categorical factor. For some species, steepness or ruggedness are important factors, and easy to model. Topographic position can be useful, but requires us to guess how topographic position was defined in the habitat-use studies we rely on. We recommend using distance to roads rather than road density as a measure of human disturbance. If appropriate soil maps are available in your linkage analysis area, we encourage using soil properties as factors in habitat models for some species.
Metrics for habitat factors can be categorical (land-cover types, topographic classes) or continuous (percent slope, distance from a cover type or road). When we have the choice between the two, we usually prefer categorical metrics. For example, if habitat suitability is a function of steepness, we find it easier to characterize the suitability of 2 or 3 steepness classes than to estimate intercept, slope, quadratic terms, or inflection points that would be needed for a linear, curvilinear, or step function of a continuous variable.
When using a categorical variable, we usually limit the number of classes based on biological understanding. For example, suppose we are using distance-to-road classes in a habitat model for a snake. We know snakes get killed on roads. The average daily movement of this snake has a width of about 200 m, so snakes up to 200 m away might be affected by increased risk of mortality. We also know that snakes hear through their jaws, and a study has shown that these reptiles can perceive vibrations from cars passing 50 m away. These vibrations may confuse the snake, or may cause it to avoid the area within 50 m of a road. This suggests that 3 classes (0-50 m, 50-200 m, and >200 m from a road) are all I need. I could create 10 classes, but how would I estimate habitat suitability for each of them? The complex model would be no better than the simple one. Let's face it—our model is crude, and making it more complex is just polishing a turd.
GIS layers commonly used in habitat suitability models include land cover, topographic variables, distance to streams, human disturbance, and soils.
Land cover is often the most important factor in the habitat models of many species. The importance of land cover reflects the fact that land cover is related to food, hiding cover, thermal cover, and (for classes like urban land use) human disturbance. The term “vegetation type” is sometimes used for this factor, because most land cover classes are names of vegetation communities. However, land cover also includes mines, farms, urban areas, open water, and other classes that make “vegetation type” an inappropriate term.
Land-cover data are usually treated categorically. Examples of continuous metrics would be tree-canopy closure or distance from forest. Land cover data may be available in a GIS layer with 20-30 coarse classes (National Land Cover Database in the USA) or 70-100 classes (GAP data layers in the USA). However, we have found it useful to lump the 70-100 GAP classes into 25-50 classes for two reasons. First, the scientific literature we use to parameterize our models does not distinguish among habitat suitabilities of several closely related land-cover types—we'd end up scoring them all the same anyway. Second, the tables of cross-classification accuracy for GAP data layers show that many errors involve confusion between closely-related land covers. Pooling these closely-related types thus likely increases the classification accuracy of the map.
Most wildlife habitat studies using land cover layers present the data as if they represent reality, although classification accuracy is typically 60% to 80%. Digital maps developed from different remotely-sensed images can produce markedly different depictions of vegetation. The GIS analyst should always report the resolution and source for land cover data. Typically the developers of land-cover data layers also report classification accuracy; you should pass this information on to the users of your models. It is depressing to report that the land cover map—the most important factor in the model—is also more error-ridden than digital elevation models, census layers, or road layers. But transparency is a hallmark of science, and we gotta tell it like it is.
Elevation is a determinant of land cover. It also affects the thermal environment of an animal, the amount of precipitation, and the form (rain, snow) of precipitation. Fortunately, digital elevation models (DEM) are available for every land area on Earth.
In our models, we typically use elevation as a factor when we have literature stating that the species occurs within a certain range of elevation. Depending on our interpretation of the literature, we often recognize 3 classes (below, within, and above the elevation limits) or 5 classes (if we suspect the literature was a crude generalization and we want to assign intermediate suitability to elevation classes near the reported limits).
DEMs are also the basis for several derived variables, including aspect, slope, and topographic position.
Topographic position is correlated with moisture, heat, cover, and vegetation. It also is relevant to cost of movement, and is therefore an attractive factor to include in a habitat model that will be used as a travel cost model. In scientific papers, some animals are reported to be associated with canyon bottoms, steep slopes, or other topographic positions.
Topographic position can be estimated by classifying pixels into any number of classes such as steep slope, ridgetop, or valley bottom. Topographic position algorithms analyze pattern within a moving window, the size of which must be specified by the analyst. While it is tempting to scale the moving window size to reflect the way each focal species may perceive the landscape, we caution against this. There have been virtually no studies on how any non-human organism assesses topography. More important, all published habitat-selection studies refer to the topographic position as perceived by the human researcher, not the animal! This still leaves the non-trivial issue of estimating the moving window size human researchers use to characterize topographic position. Unable to find any scientific papers on this topic, we have found a moving window size of 200-300 m to yield reasonable results.
Slope are ruggedness are correlated with protection from predators and cost of movement. Two of the best documented examples are the close association between bighorn sheep and steep terrain they require to escape predators, and the strong association between pronghorn and gentle slopes.
In temperate zones, aspect is a determinant of solar radiation, and consequently temperature, soil moisture, and vegetation. Few habitat models use aspect, however, because few habitat studies suggest that aspect is directly associated with habitat suitability for animals.
Distance to water is correlated with water, movement, and food for some species. The scientific literature occasionally includes statements that a certain species is usually found within a certain distance of water. In the arid southwest we have unfortunately found that GIS layers often depict springs or artificial waters (earthen tanks) that do not exist on the ground, and do not accurately portray perennial versus ephemeral sections of mapped watercourses. A site visit and conversations with local land or wildlife managers can greatly increase accuracy of any water map.
Most habitat models contain a factor related to human disturbance. All of our models in California and Arizona used either road density or distance to roads.
Many linkage designs use road density within a moving window around the focal pixel. Unfortunately, despite the seeming scale-invariance of length per length-squared, the calculated value of road density changes erratically and non-intuitively with the size of the moving window. For example, in the image below, if a straight-line road runs through the focal pixel (yellow box). The road density is 6.4 km/km2 within the 100-m radius moving window, 1.3 km/km2 in the 500-m window and 0.6 km/km2 in the 1000-m window!
Thus, it is difficult to reliably estimate resistance for road density classes, and published estimates of animal occurrence with respect to road density cannot be translated to a different moving window size. Because distance to nearest road avoids this problem, and because scientific reports using this metric can be directly imported into a model, we now use it in preference to road density.
Some models assign pixels containing a road a resistance value so high that the pixel is impermeable, or nearly so. However, we advise against this practice because the raster representation of curves in a road will always have spuriously thicker and thinner areas. The “thin” areas will be spuriously modeled as areas of lower resistance. Such distortion can seriously affect the modeled corridor. For example, this would cause a modeled corridor to completely avoid a road that runs partway through the width of the matrix, even if all other habitat characteristics near that road are far superior for the animal. Following “The Cinderella Principle,” we prefer to make the road fit the animal (e.g., by adding underpasses) rather than making the animal's movement fit the road (conserving inferior habitat as a linkage and lengthening the linkage because the large resistance value blinded us to the otherwise optimal route).
Some corridor models use human density, but census blocks are often polygons within which humans are not uniformly distributed. Allocating the mean population density to every pixel in a census block will create errors, especially when the mean density does not occur anywhere in the block! Census data can be useful for corridors at a continental scale, or assessing potential release sites for reintroducing a wide-ranging animal, but are not helpful for most linkage designs.
Soil texture is important for burrowing species such as kit foxes, badgers, and some toads. Many lizards, rattlesnakes, and pikas are closely associated with rockpiles. However, know of no linkage design which has included soil as a factor in a habitat model.
There are several problems with most soil maps. First, it is often difficult to find a seamless soil map for any linkage area. Even at the county level, there are several maps, each compiled by a different protocol and each providing an idiosyncratic set of soil classes. Second, metadata are sometimes lacking, leaving the user to guess the meaning of soil attributes. In other cases, each polygon had many attributes, none of which (as near as we could tell) were highly correlated with presence of rockpiles, soil suitable for burrows, or the factors we are interested.
Perhaps more useful maps exist in areas where agriculture is more important than where we work (Arizona and southern California). Bottom line: we would like to include soil as a factor, but so far we haven't been able to do so.
Habitat use is driven by availability of food, nest sites, and other resources, safety from predators and other hazards, presence of competitors or facilitating species, and other factors. However these factors are rarely included in GIS models for linkage design! Instead these models are typically based on one to five factors, including land cover, one or two factors related to human disturbance, and one or two topographic factors. The model is built on these factors for a simple reason: they are the only relevant factors for which georeferenced data are available for the entire planning area.
As we described above, each of these GIS layers is related to some aspect of food, cover, and other important components of habitat. But, these GIS layers don't correspond exactly with habitat factors. Statisticians tell us that any statistical or GIS model that fails to cover all aspects of the problem can give misleading results. We simply do not know how strongly the GIS layers we use are correlated with habitat use or movement by most focal species. We'd be delighted with 90% explanatory power and disappointed with 10%. Given the low accuracy of land use layers and incompleteness of the set of available factors, we guess most models are no more than 70% successful. Much better than letting a monkey with a crayon create a habitat map, but far short of the certainty we'd like to provide conservation investors who are risking scarce resources to conserve a linkage.
What can be done about the incompleteness of our models? We propose three responses:
Statisticians tell us that a model can give unreliable results when there is redundancy among the factors. If two factors in a model were elevation in feet, and elevation in meters, these two factors are perfectly redundant. Although you are too smart to build a habitat model that silly, your model will include factors that are correlated with each other (land cover is related to elevation, for instance). However, the problem is only serious when a variable in the model is over 90% correlated with the other variables. Furthermore, the main impact is on ecological interpretation of the model, not on accuracy of predictions. In general, predictions improve as variables (even highly redundant ones) are added to the model.
In this section, we review the concept of habitat and habitat modeling approaches, and call attention to a critical, untested assumption about the relationship between habitat modeling and corridor modeling—namely, that animals make decisions about how to move across the landscape using the same rules they use to select habitat.
Habitat is “where an animal lives” or “the living and non-living characteristics of a landscape that an animal uses.” Although habitat is fundamentally a description of what animals use and where animals are found, most ecologists assume that habitat also is what animals need to survive and reproduce. Technically, only experiments can determine what animals need, and wildlife ecologists regularly engage in soul-searching about this slippery concept and whether our habitat studies are properly designed and interpreted. We will not get bogged down in this important and valuable debate, however. We will try to keep the focus on habitat as a description of what animals use, but at times we will slip into the assumption that habitat is what animals need to survive and reproduce—and to move across the matrix between wildland blocks.
Habitat is often broken down into several components, depending on what the animal is doing in a particular area or with a particular element of the landscape. Five components are usually listed as food, water, hiding cover (prey) or ambush cover (predators), thermal cover (against heat or cold or both), and nest sites (or other special needs for reproduction). Some ecologists add a 6th component, namely the minimum amounts and spatial arrangement of the first 5 components. Survival and reproduction require that an animal has enough of each habitat component within the range of its daily, seasonal, or annual activities.
Habitat models allow you to assess the quality of habitat for a species within the study area or a modeled corridor, and serve as the required cost layer for least-cost path and corridor analyses. In GIS, habitat suitability models relate suitability to raster-based layers such as land use/land cover, elevation, topographic position, human disturbance (e.g. distance from roads, road density, housing density, etc), or other important factor available as a GIS layer. We refer to these raster layers as factors. Within each factor, there are several to many classes. For instance, the factor land cover may include classes such as juniper woodland, desert scrub, and urban land. There are two common ways to build these models:
The most common habitat suitability modeling technique—and that used by CorridorDesigner—is based on literature review and expert opinion, and generally follows the ideas found in the 1981 U.S. Fish and Wildlife Service publication Habitat Evaluation Procedures Handbook. While literature-based models are subject to uncertainty and errors when translating literature-based habitat studies to a habitat suitability score, they are relatively easy to create, do not require new collection of detailed field data for all species in the linkage zone, and can be applied to multiple study areas, allowing for rapid analyses and linkage designs.
The procedure requires a biologist to assign a weight to each factor (section 2.4) and a habitat suitability score to each class within a factor (section 2.3). Suitability scores for all habitat factors are then combined to form a single habitat suitability map with a suitability score for each pixel. The two most common methods of combining factors are arithmetic (or additive) mean and geometric mean models. We elaborate on the differences between these algorithms in 2.4 Combining habitat factors. Further details on these models can be found in the Standards for Development of HSI Models section of the Habitat Evaluation Procedures Handbook.
If presence-absence data or abundance is available for the species in the study area, then empirical statistical models can be created by relating the species occurrence data to habitat factors. Statistical techniques such as generalized linear or generalized additive models (e.g. logistic or Poisson regression), artificial neural networks, classification and regression trees (CARTs), and genetic algorithms can all be used to create a map of a species probability of occurrence at any pixel in the landscape.
With these models, data is typically extracted from the GIS layers, assembled into a site by occurrence matrix, analysed with a statistics package such as R, S-Plus, or SAS, then fed back into the GIS software to create a map depicting probability of occurrence. Stand-alone modeling packages such as Biomapper, openModeller, Maxent, or DesktopGARP can also be used.
While empirical models are probably more accurate than rule-based or literature-review based models, they require gathering a good set of field observations for every species in the linkage area, which can take a considerable amount of time.
Our approach has a fundamental, untested assumption—we assume that animals make decisions about how to move across the landscape using the same rules they use to select habitat. It is reasonable to assume that an animal prefers to move through areas that provide food, water, cover, and reproductive opportunities. But it is important to admit that we don't know this for sure. And in one study conducted by Horskins, Mather, and Wilson (Landscape Ecology 21: 641-655) we know this eminently reasonable assumption was false!
Horskins, Mather, and Wilson studied two small mammals which occurred in an 85-year old woodland corridor in Australia and in the woodland blocks it connected, but did not occur in the matrix of grassland and pasture surrounding the corridor. Reproductive individuals were trapped in the corridor, suggesting that the animals bred there, but there was apparently no gene flow between the two woodland blocks for either species! Their genetic divergence was just as extreme as populations in isolated woodland patches.
Given even one counter-example as demoralizing as this one, why do we make the assumption that animals make decisions about how to move across the landscape using the same rules they use to select habitat.? We have no choice. Over 95% of the ecological literature we use to parameterize our habitat models are papers on habitat use. For any single species there will be at most 2 papers on animal movement; typically there are none. And only a small fraction of papers on movement describe the type of movement we are most interested in—namely how animals move between patches of suitable habitat.
Above all, remember that your goal is to conserve or restore a functioning wildland network that maintains ecological processes and provides for the movement of all native species between wildland blocks. Your goal is not to use a particular GIS tool. Work with biologists who know the analysis area to select 10 or more focal species that collectively will serve as an umbrella for all native species and ecological processes.
Examples of focal species include
While large carnivores are excellent focal species for linkage designs, a linkage should never be designed solely to serve large carnivores. For species for which a corridor model cannot be created, we give recommendations in From corridors to linkages.
We encourage the selection of focal species likely to collectively serve as an umbrella for all native species and ecological processes. In linkage designs we created in California and Arizona, we often had 10-20 focal species, including reptiles, fish, amphibians, plants, and invertebrates.
Important types of focal species include:
Relying solely on large carnivores to design a linkage will likely harm more than help a linkage design. Because large carnivores like bears and wolves live at low density and are among the first to be harmed by loss of connectivity, they are excellent focal species for linkage design. They also make popular flagships to increase stakeholder support for a linkage. Large carnivores were the only focal species in almost half of the linkage designs published to date.
But we argue against designing a linkage solely for large carnivores—or any single species. Many species besides large carnivores need linkages to maintain genetic diversity and metapopulation stability. Furthermore most large carnivores are habitat generalists that can move through marginal and degraded habitats, and a corridor designed for them does not serve most habitat specialists with limited mobility. Worst of all, successful implementation of a single-species corridor for large carnivores could have a “negative umbrella effect” if land use planners and conservation investors become less receptive to subsequent proposals for less charismatic species. The umbrella effect of large carnivores best serves biodiversity if these species are part of a linkage designed for a broad array of native species.
Biologists familiar with the study area should be invited to identify focal species. Even the foremost ecologist in the linkage area cannot provide a comprehensive list of all focal species. As an analyst or planner, you would hate to publish your plan and then discover that you had failed to include an important focal species. To avoid this, contact biologists working for agencies, NGOs, academic institutions, consulting companies, and major landowners in the area to develop a comprehensive list of focal species.
If stakeholders are concerned that a linkage may increase the spread of invasive species into wildlands, then one or more invasive species could be included in the suite of focal species. Any expected invasion via the linkage should be compared to invasion expected from edges and matrix land regardless of the conserved linkage.
As an analyst, you will groan when someone proposes a focal species about which little is known. How can you possibly design a linkage to serve such a species? Shouldn't we just forget about it? In general, our answer is “no.”
Recall that our motto is “no species left behind.” We can't just say, “Sorry, butterfly, we don't know how you move, so you are out of luck.” We may not be able to model movement, but as conservationists, we must do what we can. We return to this issue below.
There is one circumstance in which it can be OK to exclude a poorly-known species. This can occur when there is another, better-understood focal species that plausibly captures the needs of this focal species. For instance, if a species was proposed because it is suspected to prefer steep slopes, but little else is known about the species, you can identify another focal species whose close affinity to steep slopes is more susceptible to modeling.
Even though species may not occur in one or both wildland blocks, they may still be important to the functioning of an ecosystem linkage. Managing for species endemic to the linkage can help us ensure that we are managing the linkage as a semblance of a fully-functioning ecosystem, rather than a narrow gauntlet that lets focal species pass between wildland blocks.
One example of this scenario that we have had to face was with the plant Rainbow manzanita in California. Rainbow manzanita does not occur in either the Santa Ana or Palomar Mountains protected wildland blocks to be connected, but is widely distributed in the matrix between them. The plant's geographic range is nearly 20 miles long, and contained almost entirely in the matrix between wildland blocks. Since part of our underlying goal is to conserve evolutionary processes, including the crucial processes of evolution, range shifts, and response to climate change, Rainbow manzanita was a most appropriate focal species.
We now routinely include such species, and refer to them as “species the corridor needs” (to ensure its ecological integrity) in contrast to “species that need the corridor” (to get from one wildland block to the other). This leaves the question of what exactly we do with such species, because we cannot handle them in the same framework as species that occur in both wildland blocks.
Just because we cannot build a corridor model for some species, we do not just remove these focal species from consideration!
In the last two sections we mentioned two types of focal species that don't fit well in our standard framework of designing corridors between wildland blocks. The first group consists of species for which we cannot model movement as function of GIS layers. For example, this would include animals that can fly (birds, many insects) and plants or insects whose propagules are wind-dispersed. We may be able to model suitable patches of habitat, but we don't know how they move from patch to patch.
The second group consists of species that occur in the matrix, but not in the wildland blocks. Even if we can model their movement, they differ from all the other focal species in that they are not moving between wildland blocks—we have no logical start and end points for the corridor. More precisely, we have an impractically large number of patches within the matrix, any of which could be considered start and end points. You will doubtless find other types of species that are legitimate focal species, but for which a modeled corridor would not pass the “laugh test.” We describe methods to accommodate species that don't fit well in our framework of designing corridors in From corridors to linkages
The analysis area for a linkage design typically includes:
The analyst and stakeholders in the linkage design should agree on meaningful boundaries for the habitat blocks to be connected.
Every corridor or linkage design must connect blocks of habitat for species. We will call these habitat blocks wildland blocks. Between wildland blocks is a mosaic of wildlands and developed lands in which the wildland blocks are imbedded; we call these lands the matrix. The linkage design typically recommends that portions of the matrix be managed for connectivity.
Besides being composed of potential habitat for focal species, an important characteristic of wildland blocks is that they should be likely to remain wild for at least several decades. Conserving a linkage is an expensive endeavor, and there is no point designing a linkage that connects to a wildland that will soon be converted to urban uses. Although the US Congress may occasionally fund a “bridge to nowhere,” conservationists should not emulate this practice.
The stakeholders and the analyst should agree on how the wildland blocks are defined, because the decision will affect the map of the modeled linkage. Wildland blocks may be restricted to lands with the strongest conservation mandate, such as designated wilderness areas or strict nature reserves. But some wildland blocks have no areas in such status, and consist entirely of multiple use natural lands. Stakeholders must judge whether the conservation mandate of Forest Service, BLM, military lands, tribal lands, state trust lands or private conservation easements justify including these lands in wildland blocks, or whether they should be part of the matrix for which conservation recommendations will be made. As long as the areas to be connected are likely to remain wild, these blocks can be delineated on the basis of what conservation investors have an interest in conserving.
Within a wildland block, habitat for each focal species may be limited in quality and amount, an issue we return to in Modeling habitat patches. GIS procedures will require the analyst to specify a start-end point within each habitat block. A terminus is usually not the same as the entire wildland block. We discuss this in Corridor end points.
The analysis area for a linkage design is typically a rectangle that includes
Potential linkage areas must be defined in terms of the wildland blocks they connect. It makes no sense to conserve or restore a corridor without an explicit idea of what you want to connect.
A potential linkage is an area where connectivity between wildland areas is at risk. Some potential linkages allow free movement of plants and animals, others have been severely compromised, but all have some potential to maintain or restore connectivity.
Before you can prioritize a list of potential linkages, you must first identify the potential linkages in the landscape. Typically a region has many potential linkages at risk. Wouldn't it be great to immediately develop and implement conservation plans for all such areas? Sadly, resources are limited, and conservationists must prioritize, meaning we must select a few linkages as the first to be conserved.
Each stakeholder tends to feel that the wildland he or she knows and loves best should be the highest priority for a linkage design. Because conserving a linkage requires coordinated action by transportation agencies, owners of conservation lands, donors, and others, somehow the stakeholders must agree on a prioritized list. A rational and transparent prioritization helps all stakeholders work together.
California, Arizona, New Mexico, and Colorado have undertaken statewide efforts to map and prioritize potential linkages. In our first California effort, when we proposed a “top twelve” list to a stakeholder group, we were bombarded with questions on why each stakeholder's pet area was not at the top of the list, and why some areas were not on the list at all. Big mistake, but we recovered from it.
Who develops the list of potential linkages, and how do they do it? We recommend allowing any interested party to put a linkage on the list at a workshop where they can talk face-to-face. Invitees to the workshop should include land management agencies (Forest Service, state and national parks, BLM, etc.), state and federal wildlife management agencies, conservation NGO's, transportation agencies, county and municipal planners, local land trusts and conservancies, Native American tribes, military bases, utility districts, developers, ranchers, universities and other research entities (like USGS), and biological consulting firms. Some states have held smaller regional meetings instead of or in addition to the statewide workshop. If any person or group asks to attend, invite them, but make the purpose of the meeting clear so they do not waste their time.
Invitees are more likely to attend if the invitation has the logo of major organizations (including their own). These same logos will appear on the cover of the report, so take time to assemble a diverse list of inviters. Of course, someone must first invite the inviters. If you are that someone, avoid the temptation to give top billing to your organization. Conservation success will be greatly enhanced if all the inviters are given equal prominence.
The workshop goal is to develop a comprehensive list and map of all potential linkages. At this point, do not exclude any potential linkage, even if the linkage area has been totally destroyed by urbanization and would link only to a small, degraded wildland. You want to honor everyone's participation. In the next step, the less-important or unrestorable linkages will fall to the bottom of the list, but there is no reason to exclude them entirely from the start. The only requirements are that nominators must explicitly state:
We have contributed to over 30 linkage designs in California and Arizona. We failed at this task when we tried to tell managers what to do. We succeeded when we asked management agencies and conservation organizations how we could help them identify wildlife linkages at risk and develop plans to conserve them. We share four lessons.
I (Paul Beier) had barely heard of corridors when I started a 5-year study of mountain lions in southern California in 1988. But I soon learned that mountain lions were on the road to extinction in every southern California mountain range. As the encirclement of each mountain range became complete, each mountain lion population would wink out, one by one.
It is more exciting and rewarding to work for connectivity than against fragmentation.
But it doesn't have to end that way. In 1990, mountain lions were still moving between mountain ranges. If they could continue to do so, they could survive in every linked mountain range. More important, by radio-tagging cougar cubs, I learned that these animals would find and use narrow, highly disturbed corridors through urban areas. Imagine how successful a corridor would be if we designed them to facilitate movement by animals. Not just mountain lions, but also badgers, jackrabbits, bighorn sheep, arroyo toads, steelhead trout, and even plants and invertebrates at risk.
As recipient of these scientific insights, I felt obliged to bring them to the attention of managers. I published scientific papers on my findings, but I knew managers wouldn't read them. So for the next 7 years, I did the only thing I could think to do. I fought against proposed projects that would sever the two potential corridors linking the Santa Ana Mountains (my study area) to other areas. I read environmental impact reports and wrote scathing critiques of them. I testified at hearings on proposed projects. I wrote letters to the editor, and helped reporters write news stories. But mostly I fought proposed housing developments. Typically I'd end up with a few token mitigations that left the corridor worse off, but perhaps not as bad off as it could have been.
Your goal is not slowing down the rate at which things get worse; your goal is to make the landscape more permeable than it is now!
This work had to be done. I am glad I did it. But fighting development proposals is not a strategy for victory. A victory may stop one bad project, but next year there will be another proposal, just as bad, on the same piece of ground. It took me 7 years to figure out that we could only win if we moved beyond reacting to bad proposals and put forward a positive proposal—a linkage design.
Having learned to work for connectivity, I worked on an effort that produced the “South Coast Regional Report”—basically a map of a connected wildland network in California. But the South Coast Regional Report had a fatal flaw: It was a plan written by 15 PhDs who wanted to help the befuddled management agencies see the need for connectivity. While I'm sure we did help some managers think about a positive vision for a connected wild system, many managers saw that our map failed to connect some important wildlands under their jurisdiction. If they had been part of the process, they might have agreed with our priorities, but instead they were handed a map and told to “make it happen.” Worse yet, most managers, already forced to read the mountain of paperwork from their own agency, didn't even have time to pick it up the Regional Report. The Report gathered dust. The press ignored it.
This time we learned faster. If you want agencies to read a document, it really helps if it is their document! And a year later, when 5 big agencies invited managers to a workshop to create a map identifying wildlife corridors at risk, 200 of them showed up and enthusiastically contributed to the Missing Linkages report. When we asked how we could help, they gladly said “Please take all our input and write up the report and put our logo on it.” Ironically, when we tried to lead (I'll write a report for you to follow) nobody followed, and when we served (How can I help all you agencies tackle this difficult problem?) we were given the very sort of power we had earlier wrongly assumed was our natural right as scientists!
Lead by serving. Leadership is not “getting others to follow.” Leadership is engaging diverse people to develop fair, sound, and comprehensive solutions to difficult problems.
When the report came out, managers read it. They had to—it bore their logo. And they liked it. Quite honestly, the report written by 200 people, mostly non-scientists, was better than the report by 15 PhDs. These 200 people knew more than we did about what was important. They loved the land as much as we did. They were just as passionate about creating a landscape more than the sum of its parts, because they owned the parts.
None of us is as smart as all of us. There are a lot of great people who will do great things when they work as part of a team rather than as gophers for scientists who fancy themselves as leaders.
This lesson has permeated every aspect of the linkage designs that are now being successfully implemented in southern California. The Missing Linkages report (http://scwildlands.org) was a map of potential linkage areas at risk. The next step was to identify the top priorities for detailed plans and immediate action. Instead of relying solely on scientists to prioritize linkages, we invited every interested party to another workshop to select criteria. After participants saw the priorities resulting from the first weighting scheme, they argued to change the weights. It took forever, but at the end of the day, each participant agreed that the final criteria were better than the scheme each of us had advocated at the start of the day. And everybody owned the final priorities.
At virtually every juncture in the linkage design process, we had another workshop. As a scientist, I took a while to embrace the idea of inviting non-scientists to participate in scientific issues. But science is nothing more than a way of knowing that is transparent, evidence-based, logical, and open to correction. No assumption or logical chain in ecology is so esoteric that a manager can't understand it. A scientist who wants to be a conservationist simply must invite managers to participate in the science. The product is improved by having managers challenge our assumptions and offer alternative evidence and alternative interpretations of the evidence.
Who should be invited to participate? Land management agencies, state and federal wildlife management agencies, conservation NGO's, transportation agencies, county and municipal planners, local land trusts and conservancies, first nations (Native American tribes, etc.), military bases, utility districts, developers, ranchers, universities and other research entities, and biological consulting firms.
Because large carnivores like bears and wolves live at low density and are among the first to be harmed by loss of connectivity, they are appropriate focal species for linkage design. And people love them, so they are popular flagships to increase public support for a linkage. In fact, large carnivores are the only focal species in about half of all published linkage designs based on focal species. But please don't design a linkage solely for large carnivores—or any single species! Many other species need linkages to maintain genetic diversity and population stability. Furthermore most large carnivores are habitat generalists that can move through marginal and degraded habitats, and a corridor designed for them does not serve most habitat specialists with limited mobility.
Develop linkage designs to accommodate all species that move between wildland blocks; not just large carnivores
Finally, implementation of a single-species corridor for large carnivores will have a “negative umbrella effect” for the other species. We simply cannot ask land use planners and conservation investors to create a mountain lion corridor this year, and then come back and ask them to add a bighorn sheep corridor next year, and a desert tortoise corridor the year after that. If the mountain lion is going to be an umbrella for biodiversity, it must be part of a linkage designed for a broad array of native species.
Conserving land will not create a functional linkage if major barriers are not mitigated, an excellent crossing structure will not create a functional linkage if the adjacent land is urbanized, and an integrated land acquisition-highway mitigation project could be jeopardized by inappropriate practices (e.g., predator control, fencing, artificial night lighting). An adequate linkage design will recommend crossing structures and management practices to restore native vegetation and minimize the impact of exotic species, fences, pets, livestock, and artificial night lighting. An emerging issue is how to mitigate the impact of fences, mowed strips, and stadium lighting designed to discourage human traffic on international borders.
The linkage design also must address how landowners living in or adjacent to the linkage area will become stewards of the linkage. Of course, in keeping with the philosophy that “None of us is as smart as all of us,” landowners will have been invited to participate from the outset, and some of them will already be on board. Many homeowners may initially decline the invitation to work on the plan. But once the plan is on the street, it may be necessary to ask all homeowners to help, either through individual voluntary actions, through a homeowners association, or in other ways.
A wildlife linkage is “all edge” and will require active management forever. The linkage design may ban off-road vehicles and eradicate major invasive plants, but in another decade there will be another recreational threat and a new invasive plant. Your plan cannot address all of these. But if it is to have any hope of being more than a pretty map, it must comprehensively address land conservation and roads and management practices and involve landowners as stewards.
Before you pop open ArcGIS...before you start running models...before you start making maps...take a long hard look at your landscape, and ask yourself some questions:
Designing corridors is not about pushing buttons!
If you can't first answer these questions, you probably shouldn't be designing corridors. To design a successful corridor, remember that the problem is greater than knowing what buttons to push in GIS, and the solution requires a lot more than GIS, computers, and maps.
In the following sections, we address these questions you must tackle before designing corridors. We urge the reader and analyst to always keep these questions in mind throughout the corridor design process. While the details are enticing, without a clear focus on the big picture, you're probably just wasting your time.