Landscape Connectivity: a Conservation Application of Graph
Total Page:16
File Type:pdf, Size:1020Kb
JournalofEnvironmentalManagement(2000)59,265–278 doi:10.1006/jema.2000.0373,availableonlineathttp://www.idealibrary.comon Landscapeconnectivity:Aconservation applicationofgraphtheory A.G.Bunn†§*,D.L.Urban† andT.H.Keitt‡ Weusefocal-speciesanalysistoapplyagraph-theoreticapproachtolandscapeconnectivityintheCoastal PlainofNorthCarolina.Indoingsowedemonstratetheutilityofamathematicalgraphasanecological constructwithrespecttohabitatconnectivity.Graphtheoryisawellestablishedmainstayofinformation technologyandisconcernedwithhighlyefficientnetworkflow.Itemploysfastalgorithmsandcompact datastructuresthatareeasilyadaptedtolandscape-levelfocalspeciesanalysis.Americanmink(Mustela vison)andprothonotarywarblers(Protonotariacitrea)sharethesamehabitatbuthavedifferentdispersal capabilities,andthereforeprovideinterestingcomparisonsonconnectionsinthelandscape.Webuiltgraphs usingGIScoveragestodefinehabitatpatchesanddeterminedthefunctionaldistancebetweenthepatches withleast-costpathmodeling.Usinggraphoperationsconcernedwithedgeandnoderemovalwefound thatthelandscapeisfundamentallyconnectedforminkandfundamentallyunconnectedforprothonotary warblers.Theadvantageofagraph-theoreticapproachoverothermodelingtechniquesisthatitisaheuristic frameworkwhichcanbeappliedwithverylittledataandimprovedfromtheinitialresults.Wedemonstratethe useofgraphtheoryinametapopulationcontext,andsuggestthatgraphtheoryasappliedtoconservation biologycanprovideleverageonapplicationsconcernedwithlandscapeconnectivity. 2000AcademicPress Keywords:landscapeecology,graphtheory,connectivity,modeling,metapopulations,focal species,Americanmink,Mustelavison,prothonotarywarblers,Protonotariacitrea. Introduction anislandmodel.Agraphrepresentsabinary landscapeofhabitatandnon-habitat,where patchesaredescribedasnodesandthecon- Thecurrenttrendinecologicalresearchand nectionsbetweenthemasedges. landmanagementistofocusonlargebiogeo- Graphtheoryisawidelyappliedframe- graphicareas,whichleavestheresearcher workingeography,informationtechnol- andmanagersearchingforlandscape-scale ogyandcomputerscience.Itisprimarily data(Christensenetal.,1996;Noss,1996). concernedwithmaximallyefficientflow Indeed,theinterpretationoflargespatial orconnectivityinnetworks(Grossand data,conceptuallyandtechnologically,can ŁCorrespondingauthor bethelimitingfactorinmakingconservation Yellen,1999).Tothisend,graph-theoretic approachescanprovidepowerfulleverage †NicholasSchoolofthe biologyandecosystemmanagementatan- Environment,Duke giblegoal.Becausetheinternalheterogene- onecologicalprocessesconcernedwithcon- University,Durham,NC ityoflandscapesmakeshabitat-conservation nectivityasdefinedbydispersal.Inpartic- 27708,USA planningaformidablechallenge,modeling ular,graphtheoryhasgreatpotentialfor ‡NationalCenterfor thespatialaspectsoflandscapesisacrit- useinapplicationsinametapopulationcon- EcologicalAnalysisand icalkeytounderstanding.Untilnow,the text.UrbanandKeitt(2000)haveintroduced Synthesis,SantaBarbara, CA93101,USA variedapproachestobuildingthesemod- landscape-levelgraph-theorytoecologists, elshavefocusedprimarilyontwotypesof andherewebuildonthatworkbyexam- §Presentaddress: spatialdata,coveragesofvectors(polygons) ininghabitatconnectivityfortwospecies MountainResearch Center,MontanaState orrastergrids.Wedemonstratetheutility thatsharethesamehabitatbuthavedif- University,Bozeman,MT ofalessfamiliartypeoflattice,thegraph ferentdispersalcapabilities.Specifically,we 59717,USA (Harary,1969),indetermininglandscape askhowAmericanmink(Mustelavison)and Received31May2000; connectivityusingfocal-speciesanalysisin prothonotarywarblers(Protonotariacitrea) accepted30June2000 0301–4797/00/080265C14$35.00/0 2000AcademicPress 266 A. G. Bunn et al. perceive the same landscape. We explore the Focal species sensitivity of landscape connectivity through graph operations concerned with edge defini- Because connectivity occurs at multiple tion. We also examine each habitat patch’s scales and multiple functional levels (Noss, role in maintaining landscape connectiv- 1991), we have chosen two focal species to ity in terms of source strength (Pulliam, apply a graph-theoretic approach to connec- 1988) and long-distance traversability (den tivity. Focal species analysis is an essential Boer, 1968; Levins, 1969) using graph oper- tool for examining connectivity in a real ations concerned with node removal. This landscape, as individual species have differ- type of analysis is done very efficiently ent spatial perceptions (O’Neill et al., 1988). with graph theory. We also present an The American mink and the prothonotary ecologically appealing way to calculate the warbler are appropriate candidates for focal functional distance between habitat patches species analysis as they share very similar using least-cost path modeling. Graph the- habitat but have different ecological require- ory as applied to landscapes represents an ments, and fall into different categories as important advance in spatially explicit mod- focal species. Both species are wetland depen- eling techniques because it is an additive dent and indicators of wetland quality and framework: analysis of a simple, preliminary abundance in a landscape. Both are charis- graph can prioritize further data collection to matic. Furthermore, as meso-predators mink improve the graph model. have small but important roles as a keystone species (Miller et al., 1998/1999). American mink are meso-level, semi- Study area and methods aquatic carnivores that occur in riverine, lacustrine and palustrine environments (Gerell, 1970). In chief, they are nocturnal Study area and their behavior largely depends on prey availability. They have a great deal of vari- Our study focuses on the Alligator River ation in their diet according to habitat type, National Wildlife Refuge (NWR) and sur- season and prey availability (Dunstone and rounding counties on the Coastal Plain of Birks, 1987). Muskrats (Odantra zibethicus) North Carolina (35°500N; 75°550W). It is a are a preferred prey item (Hamilton, 1940; riverine and estuarine ecosystem with an Wilson, 1954), but mink diets in North Car- area of almost 580 000 ha and over 1400 km olina are composed of aquatic and terrestrial of shoreline. The area is rich in wildlife habi- animals, as well as semiaquatic elements tat, dominated by the Alligator River NWR, (e.g. waterfowl; Wilson, 1954). In the south- the Pocosin Lakes NWR, Lake Mattamus- east they have home ranges on the order of keet NWR, Swanquarter NWR, and a variety 1 ha and a dispersal range of roughly 25 km of other federal, state, and private wildlands (Nowak, 1999). (Figure 1). Prothonotary warblers are neotropical mig- The vegetation is characterized by the rants that breed in flooded or swampy mature Southern Mixed Hardwoods forest commu- woodlands. They have two very unusual nity. The area has many diverse vegetation traits in common with wood warblers in that types, including fresh water swamps, pine they are cavity nesters and prefer nest sites woods and coastal vegetation. In the upland over water. They are forest interior birds community, dominant species include many that experience heavy to severe parasitism types of oak (Quercus spp.), American Beech by brown-headed cowbirds (Molothrus ater); (Fagus grandifolia), and evergreen magno- (Petit, 1999). They are primarily insectivo- lia (Magnolia grandiflora). Mature stands rous, occasionally feeding on fruits or seed may have five to nine codominants. The (Curson et al., 1994). Preliminary data indi- wet lowlands are dominated by bald cypress cate that natal dispersal ranges from less (Taxomodium distichum). The pine woods than 1 km to greater than 12 km (Petit, 1999). are dominated by longleaf pine (Pinus palus- Although this is formulated from a small tris), but loblolly pine (P. taeda) and slash sample, it is on the same order as other song pine (P. elliottii) are also important (Vankat, bird dispersal (e.g. Nice, 1933; Sutherland 1979). et al., 2000). Here, we posit warbler dispersal Landscape connectivity 267 Figure 1. Study area in North Carolina with major roads and streams shown along with bottomland hardwood forest (focal species habitat) identified using GIS analysis. to be 5 km and return to the uncertainty of warblers in the Southeast (Price et al., 1995). this statement later. Finer scale spatial information is not readily available. Mink and prothonotary warblers are habi- Geospatial data tat specialists that use the same habitat. To identify habitat patches in the land- To our knowledge there are no current scape we combined data from the US Fish data on the spatial distribution of the focal and Wildlife Service’s National Wetlands species in our study area. The decline in Inventory (http://wetlands.fws.gov) and a trapping of the mink has perversely led to a 1996 land-use coverage from the National decline in good biological information on the Center for Geographic Information and Anal- species. We are unaware of any work done ysis (http://www.negia.ucsb.edu). Both were with mink in the study area since Wilson’s derived from Landsat 7 Thematic Map- (1954) study. The Breeding Bird Survey per imagery with 30-m cells. Cells that indicates that this study area contains one were defined as being bottomland hardwood of the highest concentrations of prothonotary swamp or oak gum cypress swamp, and 268 A. G. Bunn et al. riverine, lacustrine, or palustrine forested as an index to overall traversability of the wetland were selected as habitat. These cells habitat mosaic. were then aggregated into regions using A graph is defined by two data structures: an eight-neighbor