Integrating Socioeconomic Factors into Gap Analysis Products Applications of Urban Growth Models and Wildlife Habitat Models to Assess Biodiversity Losses FINAL REPORT 30 October 2001 Christopher B. Cogan, Researcher Ph.D. candidate, Environmental Studies, University of California-Santa Cruz Frank W. Davis, PI Donald Bren School of Environmental Science and Management University of California-Santa Barbara Keith C. Clarke, Co-PI Dept. of Geography, University of California-Santa Barbara Contract Administration Through: University of California-Santa Barbara Institute for Computational Earth System Science Submitted by: Christopher B. Cogan Research Performed Under: Cooperative Agreement No. 00HQAG0009 as part of the GAP Socioeconomic Seed Grant Program, managed by Dr. Gary Machlis U.S. Department of the Interior U.S. Geological Survey Biological Resources Division Gap Analysis Program Applications of Urban Growth Models and Wildlife Habitat Models to Assess Biodiversity Losses ABSTRACT Habitat loss and subsequent fragmentation due to urban development is part of a larger suite of anthropogenic impacts on biodiversity, but it now ranks among the principle causes of species endangerment in the United States. Several types of urban growth simulation models have been developed which can supply useful information for biodiversity planning. In many cases however, the data required for biodiversity planning may not be compatible with the urban models, leading to analytical inaccuracies and misleading conclusions. Here, we examine several lines of logic likely to be employed in biodiversity assessment and show how assumptions built into the data influence model outcome. Introduction Biodiversity can be described conceptually as a collection of indices, ranging from diversity of compositional, structural, and functional biotic elements (Noss 1990), to ecosystem, species, and genetic diversity (Soulé and Mills 1998). Representing such a broad ecological concept requires the combination of several models (Cogan in press), and central amongst these are predictions of species habitat quality and quantity. Habitat loss or fragmentation due to urban development is only one of many anthropogenic impacts on biodiversity today (Tilman and Lehman 2001), but it now ranks among the principle causes of species endangerment in the United States (Dobson et al. 1997, Vitousek et al. 1997, Wilcove et al. 1998). Urban growth models developed recently, incorporate a broad range of variables and spatial dependencies (Makse et al. 1995, Couclelis 1997, White et al. 1997, Batty 1998, Clarke and Gaydos 1998, Landis and Zhang 1998, Makse et al. 1998). These models do not themselves attempt to determine the environmental consequences of future urbanization. However, by providing a localized abstraction of the direction and magnitude of land use change, they offer a starting point for assessing future impacts on biodiversity. Planning efforts are not currently making full use of basic biodiversity information (Press et al. 1996, Crist et al. 2000), and improved linkages between urban growth models, biodiversity models, and land use planning is urgently needed. By using predictive models of urbanization and its effects on biodiversity, county planners and other stakeholders will be able to visualize and evaluate different future growth scenarios as an effective way to lessen the impact of urbanization on biodiversity (e.g., Landis 2000) Biodiversity assessments often rely on ecological models that simplify and abstract the biophysical world based on a series of assumptions about ecosystem functions. The set of assumptions may not be clearly understood by the model user and this can lead to inappropriate applications of the models. Problems of generalization and appropriate model selection are further compounded as more models are used in combination. When planning for both urban growth and biodiversity conservation today, a planner may be faced with the task of using two potentially complex types of models: one of urbanization processes and another of ecological processes. Many urban growth models use a simple binary classification of land use into urban or non-urban, although in fact urbanization includes a wide range of settlement patterns and human densities. Perhaps most significant for biodiversity conservation is the phenomenon of urban and suburban sprawl at the margins of existing metropolitan areas. Because the sprawl is incremental, the loss of habitats in these “front line” areas can be difficult to control. The boundary between urban and non-urban is fuzzy at best. Similar challenges are involved in predicting the impact of urbanization on biological species, involving simple habitat classification schemes and crude habitat suitability rating systems to predict whether a given land use or land cover class is or is not suitable habitat for a species. As a first step, urban growth models sensitive to increasing human population density and urban expansion in rural regions can be usefully combined with biodiversity models. Since even relatively sparse development on a parcel-by-parcel basis can dramatically affect biodiversity, the growth models should be able to detect change over fairly small spatial areas (e.g. 100 meter grids). The biodiversity models should likewise be sensitive to land use change at a similar spatial grain. Even with the constraints of generalization and spatial grain compatibility, information gained from such a union of predictive urban and biodiversity models will be valuable in helping to anticipate and avoid biodiversity erosion caused by habitat loss and fragmentation. In this report, we examine several lines of logic likely to be employed in the combination of models used to plan for urbanization and biodiversity conservation. We also show how assumptions inherent to the data can influence model outcome. Examples are presented from a case study in Santa Cruz County, California, where multiple urban growth scenarios are combined with land cover data, and wildlife habitat relationship (WHR) models. Methods Habitat quality and quantity aspects of biodiversity were examined using three principle inputs: urbanization scenarios, wildlife habitat maps, and species habitat models. Output from the analyses is reported as loss of habitat area, or in some cases, in terms of impact to the vertebrate species associated with degraded habitats. Limited data availability did not permit an impact analysis for invertebrate species. A flow chart of the models and analyses provides an overview of the biodiversity sensitivity analysis (Figure 1). Three different previously developed models for predicting patterns of urban expansion were tested. The three models included the “urban buffer” (see below), “Landis” (Landis and Zhang 1998) and “Clarke” (Clarke and Gaydos 1998) scenarios. Outputs from the different growth models were then used in conjunction with coarse grain (100 ha minimum mapping unit) land cover maps from the California Gap Analysis Project (GAP, Davis et al. 1998). The Landis and Clarke models were also used with a finer grain (1 ha) land cover data set. This map layer was commissioned by the Association of Monterey Bay Area Governments (AMBAG) based on 30-meter Landsat Thematic Mapper (TM) imagery. Spatial distributions of 21 individual vertebrate species predicted to occur in the study area were made possible by applying wildlife habitat relationship (WHR) models (Airola 1988) to the courser grained GAP land cover data. Potential impacts of urban growth to these species were explored by intersecting scenarios of future urban growth from each of the three models with the WHR-based predicted distributions of the species. The onehectare finer grained land cover data was intersected with output from the three urban growth models to generate impact assessments for generic habitat types, such as “coastal oak woodland”, without evaluating potential to individual vertebrate species from WHR (Fig. 1). urban buffer growth scenario Landis growth scenario GAP 100 ha land cover Clarke growth scenario County 1 ha land cover WHR models vertebrate species impacts habitat impacts Figure 1. Flow chart for biodiversity sensitivity analysis. Three urban growth scenarios and two land cover models combine to evaluate vertebrate and habitat impacts in Santa Cruz County, California. Urban Growth Models: As a case study, we have employed three different urban growth scenarios, each portraying a possible future urbanization pattern in Santa Cruz County, California, USA. All three scenarios are based on the same initial urbanization pattern from the California Gap Analysis Project, derived largely from Landsat TM satellite imagery (Davis et al. 1998). The Gap urban data have a coarse (100 hectare) spatial grain, which can potentially cause problems when analyzing some land cover effects (e.g. absolute measures of fragmentation). In this study, we use the Gap urban data only for urban model starting points and relative area comparisons. The first growth scenario is based on a simple spatial buffer, which is generated by expanding current urban land use areas outwards by a distance of 500 meters. The 500meter forecasted growth area appears as a narrow red band around the current urban areas (Figure 2). A second growth scenario is based on a model of urbanization developed by Landis and Zhang (1998), which incorporates socioeconomic and physical data to predict areas of future urbanization. Using logit (natural log of the odds ratio) models of historical land use change, 100-meter grid cells are predicted to be urban or non-urban in a future time period. Probability and magnitude of land-use change is predicted from projected population growth individually modeled for each city and county in the study area. With population growth specifying the demand levels for urbanization, a series of six independent variables is used to forecast which areas will be developed. The variables are: 1) initial site use; 2) demand factors, i.e. employment data; 3) accessibility, i.e. commute distance; 4) cost constraints, i.e. development costs for the site, availability of services; 5) policy constraints, i.e. zoning categories; 6) adjacent-use effects and proximities, i.e. industrial neighbors or nearby shopping centers. Current Urban Landuse Forecast Urban Landuse Other Landuse Areas in Santa Cruz County, California KILOMETERS 0 10 10 Data Source: CA GAP 5 0 5 MILES 10 20 15 Figure 2. Urban growth forecast using an urban buffer model. Santa Cruz County, California. The combination of population growth models with site-specific spatial allocation models uses large amounts of data to characterize the spatial, political, economic, and historic circumstances of the local study area. Output from the model can be in table form or in map form. The Landis model results for Santa Cruz County were transformed into a geographic information system (GIS) map (Figure 3), depicting the pattern of urbanization used in this case study. The Landis model can generate alternative outcomes depending on adjustments to the input parameters, though variations are not temporally explicit. For a detailed discussion of the model, see Landis and Zhang (1998). The third urban growth scenario used a cellular automata approach developed by Clarke (1998) to predict a year-by-year sequence of growth based upon physical landform and land use data. Model inputs included road locations, presence of existing urban areas, slope, and protected exclusion areas. The Clarke model uses a series of precalibrated control values, which are self modified with each theoretical year of growth. Initial calibration is based on historic trends. Output of the model is an image of forecast growth areas. The digital image was converted to GIS grid map format (Figure 4). As with the Landis model, a 100-meter grid template was used to map the presence or absence of urban land cover. In contrast to the Landis model, the physical data inputs were largely derived from available remote sensing products, using a combination of aerial photography and satellite imagery. The Clarke model is spatially and temporally explicit, scaleable to any particular spatial grain or extent, and does not utilize socioeconomic data. Current Urban Landuse Forecast Urban Landuse - Landis Model Other Landuse Areas in Santa Cruz County, California KILOMETERS 0 10 10 Data Source: CA GAP 5 0 5 MILES 10 20 15 Figure 3. Urban growth forecast using the Landis model. Santa Cruz County, California. Current Urban Landuse Forecast Urban Landuse - Clarke Model Santa Cruz County, California KILOMETERS 0 10 10 Data Source: CA GAP 5 0 5 MILES 10 20 15 Figure 4. Urban growth forecast based on the Clarke model. Santa Cruz County, California. Landover Data: Two different land cover datasets provide an opportunity to assess changes in model output resulting from differences in the land cover parameters. The coarser of the two land cover datasets is the GAP land cover GIS map layer (Figure 5). This map layer was originally created by human photo interpretation of Landsat TM imagery with a resampled pixel size of 100 by 100 meters (1 ha). The GAP interpreters also used a variety of ancillary data, including historic vegetation maps dating from the 1930’s. GAP land use and land cover polygons were delineated with a 100-hectare minimum size generally referred to as the minimum map unit (MMU). The 100 ha MMU is coarse compared to many other map products, however it is the most detailed statewide land cover product for California. For more information on the development and appropriate use of the California GAP data, see Davis et al. (1998). One aspect of the GAP land cover data is particularly useful for biodiversity analysis. Each land cover polygon is linked to a list of vertebrate species, based on the California Wildlife Habitats Relationship (WHR) system (Airola 1988, Davis et al. 1998). With this linkage, each land cover polygon in the GAP database can be associated with habitats for particular species. Using standard GIS techniques, we used urban growth model outputs in conjunction with GAP land cover to determine which habitats and which vertebrate species may be impacted in the near future. KILOMETERS 0 10 10 Data Source: CA GAP 5 0 5 MILES 10 20 15 Figure 5. California Gap Analysis Project (GAP) land cover polygons in Santa Cruz County, California. Another land cover map is also available for Santa Cruz County. This GIS map layer was commissioned by the Association of Monterey Bay Area Governments (AMBAG) based on the same 30 meter TM imagery the GAP project used. In this case however, the land cover partitions were machine classified, resulting in a product with a 30 meter MMU (1/9 ha), and approximately 40 land cover classes (Figure 6). The AMBAG cover classes are not directly compatible with the WHR system. In parallel to the analysis using the GAP products, the AMBAG data was also combined with the urban growth models, for investigation of impacted habitats following urbanization. The combination of urban growth data and land cover data was modeled in two ways. As a measure of vertebrate species impact, when any portion of a land cover polygon was predicted to become urbanized, the entire polygon was assumed to be compromised for that species. As a measure of habitat impact, the exact proportion of the land cover polygon predicted to become urbanized was calculated for the analysis of habitat loss. AMBAG landcover polygons Santa Cruz County, California KILOMETERS 0 10 10 Data Source: AMBAG 5 0 5 MILES 10 Figure 6. AMBAG land cover polygons in Santa Cruz County, California. 20 15 Results Urban Growth Models: From the broad county perspective, the Landis and Clarke urban growth model scenarios had some similarities in their predictions of future urbanization in Santa Cruz County, California. Both models avoided the steep rural areas in the northwest and northern ridge areas, and typically selected sites closer to preexisting urban land use. Several agricultural areas in the south were selected by both models. There were also several key differences between the Landis and Clarke model results. The Clarke cellular automata approach (Figure 4) usually forecast urbanization to occur in areas within 500 meters of current urban land use, as well as other regions near major roads. Very little development was forecast for remote coastal areas, in spite of the presence of roads and flat terrain. The Landis model did not predict correspondingly high levels of roadside expansion, but did forecast large amounts of isolated new development in areas not contiguous with existing urban land use. The buffer model may not appear to be realistic as a growth scenario, however the land areas identified by this approach were consistently targeted by the other two models. Thus, the buffer model represents a simplified starting point for predicted urbanization, without using the model parameters of more sophisticated approaches. Land Cover Models: The two different models of land cover; one from GAP and a second from AMBAG produced useful data for comparison. A few simple statistics on these data models reveal important differences that affect the biodiversity analysis. The GAP land cover data are spatially coarser, with a minimum map unit of 100 ha, and a maximum polygon size of approximately 4,000 ha. The GAP data describe Santa Cruz County using 208 polygons in 14 primary classes. Secondary and tertiary classes describe areas of mixed land cover smaller than the MMU. In contrast, the AMBAG data uses a much finer spatial grain, derived directly from 30 meter Landsat Thematic Mapper Satellite imagery. These data have a minimum map unit of 1/9 ha, and a maximum polygon size of 49,000 ha. The AMBAG data use approximately 11,000 polygons to map Santa Cruz County in 40 classes, with no secondary classification. Habitat Impacts: Habitat impacts were assessed in terms of land area lost to urbanization using the three separate growth models. This comparison is based on the AMBAG 30 meter MMU land cover data. For each of the 30 impacted habitat types, the sum of area lost is shown as a bar graph (Figure 7). Maximum impact is predicted using the Clarke model, resulting in over 10,000 ha of Redwood / Douglas-fir habitat converted to urban land cover. Minimum impact occurred with the 500 meter buffer model, though for a few habitats this trend was inconsistent. Also shown in the graph (Figure 7), are the large differences in land area and the range of inconsistencies among the three growth models. The large range of variability necessitated the use of a log scale on the y-axis. 10000 Clarke Model Landis Model Area Urbanized (ha) (log scale) 1000 Buffer Model 100 10 Re dw Gr ood Co as / D s & ou as t al Co glas Oa as F k W Co Co tal O ir D a o oo sta a dl l O stal ak W min an ak O a d & W ak ood nt Re Re ood Wo land dw dw la od oo oo nd lan d/ d d/ & Do Do G ug ug rass las Spa las Fi rse Oth F Co r & ly V er A G ir as Co eg g ra tal as et ric ss O ta at u Or ak l O ed/ lture ch W ak Fa ar oo W llo d d Kn s/O lan oodl w ob utd d & and co oo ne r N Shr u Pa Pine urse b Sh lu Vi rub str Do ries ne m & i n Eu ya e E ina ca rds Coa m nt ly pt /Bu stal Str erg us sh O aw en & es ak b t Co (eg W erri as . R oo es tal a dla Oa spb nd lo lo w Euc k W errie w de aly oo s) de ns ns ity ptus dlan ity ur & d ur Po ba G ba nd n & ras n & er Gr s Sh C osa as ru oa Pi s b s n W t Re & al e D a dw Re Oa om ter oo dw k W in d/ oo Do d ood ant ug /Do lan Eu las F ugla d Co ca i r s F as tal Gr lypt & S ir as us hru Oa s& & b kW Eu Shr oo ca ub dl l an yp d& tu s Eu Shr ca ub l y Go pt lf u s C Eu ou ca rse ly pt us 1 Figure 7. Comparison of three growth scenarios in Santa Cruz County California: 500 meter urban buffer, Landis growth model, and Clarke growth model. Habitat data are from AMBAG 30 meter land use / land cover maps. Habitat classes are rank ordered based on the results from the Clarke model. Vertebrate Impacts: Native vertebrate impacts were assessed using the GAP 100 ha MMU land cover data. As with the habitat assessment, the maximum impact case is predicted when using the Clarke model (Figure 8). Percent habitat loss is based on recent (1993), not historic species habitat in the ecoregion. All three growth models predict vertebrate species impacts in similar rank order, with Vaux’s swift (Chaetura vauxi) most heavily impacted (38% under the Clarke model) followed by hermit warbler (Dendroica occidentalis) and golden-crowned kinglet (Regulus satrapa). The Landis and Clarke models also have overall similar magnitudes of predicted impact, while the 500 meter buffer model predicts up to 30% less impact and does not clearly distinguish among the top 12 species. 40 35 30 Habitat Loss (%) Clarke Model Landis Model 500 m buffer 25 20 15 10 5 go Va u lde her x's s n-c mit wi row wa ft ne rble dk r pin ingle M t e ac gil shr siski e l n oli ivr wmo a ve -si y's le Tr ded warb ow fly le bri ca r Ca tc d lif orn blac ge's her k sh i no a gia salam rew rth nt ern sal and e a sh allig man r arp at d -sh or l er inn iza r he ed h d a r mi w wh tt k ite -cr ru hrus ow bbe h ne rb d o so spar a ch l r i est tar ow nu t-b wi y vir eo a c nt Am ked er w r c e e Ca ric hic n lif an kad orn g ia bro old ee f sle w nd n c inch er r sal eep am er an de r 0 Figure 8. Comparison of three growth scenarios in Santa Cruz County California: 500 meter urban buffer, Landis growth model, and Clarke growth model. Species and habitat data are from the California Gap Analysis Project (GAP). Habitat classes are rank ordered based on the results from the Landis model. Discussion The species habitat analysis presented here is a close examination of one major factor in the assessment of biodiversity. Other biodiversity elements such as ecoregional analysis, restoration potential, special features, and habitat shape are also important, though these are not specifically addressed in this study. We used three urban growth models and two land cover maps in a relatively straightforward combination of data (Figure 1) to compare measures of habitat and vertebrate impacts. Here, habitat impacts are considered to be actual habitat areas converted to urban land use. For example, if a 1000 ha forest is reduced to 900 ha after urbanization the habitat loss is 10%. If the same forest is reassessed in terms of native vertebrate habitat, it may be more important to consider buffer distances from impacts, non-linear predation effects, and other complex landscape metrics. These more specific approaches can be valuable in some instances, however, when applied to a regional study with many species the results can be misleading. Stated differently, it is challenging to model disturbance effects as realistically as possible, while working with a group of dissimilar species over a broad area. The approach to vertebrate habitat assessment presented here assumes that if a highly intrusive land use such as urbanization enters a habitat patch, then the entire patch is likely to be compromised in terms of vertebrate species habitat quality. In some instances, this assumption may overemphasize the impact of urbanization. On the other hand, it is also likely that urbanization effects are underemphasized in cases where urban expansion approaches (but not actually enters) a habitat area. An alternate model could employ spatial buffers to model the neighborhood effects of urbanization, however this approach introduces additional complexities such as splitting map polygons, and imposes the need for species-specific analysis. Both the habitat and species types of impacts are important, however it is necessary to clarify the conceptual differences between habitat and vertebrate impacts when evaluating or discussing urban growth impacts. A noteworthy example of this distinction was shown in a study of mesopredators and avian prey in Southern California (Crooks and Soulé 1999). The methods used in this analysis are based upon an underlying logical sequence most simply presented as a flow chart (Figure 9). A central assumption here is that different urban growth patterns should have measurably different biodiversity impacts. As with any metamodel, it is also important to ensure that the data and various component models are compatible for integrated analysis. It is often illuminating to investigate where the logic of a scientific investigation might become unsound, as well as where it is strong. The logical flowchart outlines key junctions where this type of biodiversity assessment might face impediments and offers explanations and recommendations for each situation. Biodiversity Analysis Variations in urban growth patterns are not critical in biodiversity analysis. Explanation: Particular species will always be impacted – perhaps due to their rarity in the county vs. the ecoregion. Action: Treat these species and habitats as special cases; use the biodiversity model to evaluate the remaining biodiversity elements. Variations in urban growth patterns do impact biodiversity. Model error prevents variation in growth pattern from producing a measurable biodiversity response. Urban growth scenarios are constrained into similar patterns. Explanation: Urban models lack sufficient realistic variation. Action: test with different or random growth scenarios. Variation in growth is measurable in terms of biodiversity. Biodiversity data are too coarse to respond to fine urbanization differences. Explanation: Habitat models are too coarse grained for measurable response to urban change. Action: use as is for coarse grain analysis, but use finer grain habitat model and new WHR models for fine biodiversity analysis. Explanation: model is working with available data. Action: use urban growth scenarios and existing species habitat data to evaluate biodiversity impacts. Figure 9. Logical flow chart for the evaluation of biodiversity analysis with urban growth models. Given perfectly accurate biodiversity and urban growth models, lack of biodiversity response will still occur if the two models are not spatially or thematically compatible. One indicator of this type of incompatibility can be seen in the comparison of vertebrate habitat losses following different urbanization scenarios (Figure 8). One interpretation of this result suggests that vertebrate impacts are much the same following either the Clarke or the Landis models. Indeed, it seems remarkable that the rank order of species and even habitat impacts is so similar under two independent and seemingly different growth models. It would seem to require a radically different growth model like the simplistic 500-meter buffer to produce a significantly different outcome. Another, perhaps more likely interpretation is also possible. If the GAP data on wildlife habitat relationships is spatially coarser that the growth models, our ability to differentiate between the Landis and Clarke models will be diminished. In support of this hypothesis, the appearance of the map products, and (most importantly) the habitat impacts (Figure 7) indicate substantial differences between each of the three urban models. The balance of spatial grain and thematic detail is an important consideration when producing and using maps of land cover for use in biodiversity analysis. Using the AMBAG 30 meter MMU land cover map (Figure 6), the fine map grain results in relatively large areas (up to 49,000 ha) to be mapped as contiguous albeit marginally connected patches. At slightly coarser map grains, many of the corridors of connecting habitat would merge into other classes resulting in a very different dataset for the habitat modeler. This example illustrates how fine grain maps with coarse thematic detail can overemphasize habitat connectivity. In this case, the assumption that urban disturbance on the edge of a habitat patch impacts the entire patch becomes tenuous when using fine spatial grain, coarse thematic grain data such as the AMBAG 30 meter land cover map. As 100 meter or finer grain urban growth models gain acceptance as a reasonable spatial scale to model the biodiversity land use complex, more research is needed to ascertain the appropriate levels of thematic resolution in land use and land cover mapping. There are several difficulties associated with measuring regional urban impacts on vertebrate species. The model presented here uses polygons of habitat to represent potential distributions of vertebrate species, and assumes that analysis of divided polygons is not a valid application of the data. Detailed study of a specific divided habitat polygon is possible given appropriate species-specific data, however this local approach will not be effective regionally. Urban development is sometimes seen as a continuous creeping of small steps whereby each development project in isolation is difficult to assess for regional biodiversity impact. The species assessment method presented here uses habitat polygons to model impacts, effectively dealing with the “urban creep” issue while maintaining biologically meaningful area units. The complementary combination of a discrete species metric along with a continuous habitat model is a powerful and much needed approach. As biodiversity models such as those discussed here evolve and build in complexity, our land cover maps and wildlife habitat relationship models will be pressed to deliver more information with higher quality standards. Some of our data sources have already evolved from simple maps of predicted species location to become temporally dynamic models of predicted species connectivity and spatial pattern. Unfortunately, most of our current maps are not up to this advanced standard. Like most modelers, cartographers have long known that the design constraints of producing the best habitat maps will depend on the specific questions being asked of the data. This fundamental principal is sometimes obscured or overlooked when we allow technological capabilities such as satellite sensor resolution and radiometric spectral response to overly influence our understanding of habitat classification and vertebrate distribution. This paper has outlined a method for assessing habitat and species degradation given different development scenarios. The inherent assumptions and complexities of this type of analysis were also explored. Examples have been presented which show how assessment of biodiversity following urbanization may be misleading. We have also shown how seemingly significant differences in urban growth pattern may be obscured by incompatible habitat data. These findings are presented to facilitate an improved understanding of habitat and species impact models, and to provide direction for future land use and land cover mapping. The specific models discussed here are important elements of more generalized biodiversity assessments, which are continually improving our understanding of biodiversity and promise to provide additional guidance to minimize the disruptive impacts of urbanization and development. Acknowledgments: We thank Bob Johnston, Mike Jennings, Mary Anne Van Zuyle, and Uta Passow for reviewing earlier versions of this paper and providing constructive comments. Their insights were greatly appreciated. This work was partially funded by the United States Geological Survey, Cooperative Agreement No. 00HQAG0009 as part of the GAP Socioeconomic Seed Grant Program, managed by Dr. Gary Machlis. References: Airola, D. A. 1988. Guide to the California wildlife habitat relationships system. Jones and Stokes Associates, Sacramento, CA. Batty, M. 1998. Urban evolution on the desktop: simulation with the use of extended cellular automata. Environment and Planning A 30: 1943-1967. Clarke, K. C., and L. J. Gaydos. 1998. Loose-coupling a cellular automaton model and GIS: long-term urban growth prediction for San Francisco and Washington/Baltimore. International Journal of Geographical Information Science 12: 699-714. Cogan. in press. Biodiversity conflict analysis at multiple spatial scales. in J. M. Scott, ed. Predicting Species Occurrences: Issues of Accuracy and Scale. Island Press, Washington, D. C. Couclelis, H. 1997. From cellular automata to urban models: New principles for model development and implementation. Environment and Planning B-Planning & Design 24: 165-174. Crist, P. J., T. W. Kohley, and J. Oakleaf. 2000. Assessing land-use impacts on biodiversity using an expert systems tool. Landscape Ecology 15: 47-62. Crooks, K. R., and M. E. Soulé. 1999. Mesopredator release and avifaunal extinctions in a fragmented system. Nature (London) 400: 563-566. Davis, F. W., D. M. Stoms, A. D. Hollander, K. A. Thomas, P. A. Stine, D. Odion, M. I. Borchert, J. H. Thorne, M. V. Gray, R. E. Walker, K. Warner, and J. Graae. 1998. The California gap analysis project: Final report. University of California, Santa Barbara, CA. Dobson, A. P., A. D. Bradshaw, and A. J. M. Baker. 1997. Hopes for the future: Restoration ecology and conservation biology. Science 277: 515-522. Landis, J. 2000. CUF, CUF II, and CURBA: A family of spatially explicit urban growth and land-use policy simulation models. Pages 157-200 in R. K. Brail and R. E. Klosterman, eds. Planning Support Systems. ESRI Press, Redlands, CA. Landis, J., and M. Zhang. 1998. The second generation of the California urban futures model. Part 1: Model logic and theory. Environment and Planning B-Planning & Design 25: 657-666. Makse, H. A., J. S. Andrade, M. Batty, S. Havlin, and H. E. Stanley. 1998. Modeling urban growth patterns with correlated percolation. Physical Review E 58: 70547062. Makse, H. A., S. Havlin, and H. E. Stanley. 1995. Modelling urban growth patterns. Nature 377: 608-612. Noss, R. F. 1990. Indicators for monitoring biodiversity - a hierarchical approach. Conservation Biology 4: 355-364. Press, D., D. F. Doak, and P. Steinberg. 1996. The role of local government in the conservation of rare species. Conservation Biology 10: 1538-1548. Soulé, M. E., and L. S. Mills. 1998. No need to isolate genetics. Science 282: 1658-1659. Tilman, D., and C. Lehman. 2001. Human-caused environmental change: Impacts on plant diversity and evolution. Proceedings of the National Academy of Sciences of the United States of America 98: 5433-5440. Vitousek, P. M., H. A. Mooney, J. Lubchenco, and J. M. Melillo. 1997. Human domination of earth's ecosystems. Science 277: 494-499. White, R., G. Engelen, and I. Uljee. 1997. The use of constrained cellular automata for high-resolution modelling of urban land-use dynamics. Environment and Planning B-Planning & Design 24: 323-343. Wilcove, D. S., D. Rothstein, J. Dubow, A. Phillips, and E. Losos. 1998. Quantifying threats to imperiled species in the United States. Bioscience 48: 607-615.