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Of Habitat Suitability for the Delmarva

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Of Habitat Suitability for the Delmarva Abstract.-Discriminant function analysis corn par- Habitat Structure, Forest ina 36 occu~iedand 18 unoccu~iedsites revealed Composition and Landscape thGt structural variables discriminked between sample groups better than compositional variables, Dimensions as Com~onents and the latter discriminated better than landscape variables. These results are encouraging that habitat of Habitat suitability for the structure will provide a reliable basis for a predictive classification model of habitat suitability. Such a Delmarva Fox model would be useful both for pre-screeningthe biological suitability of potential release sites and for planning, implementing and monitoring prescriptive Raymond D. Dueser,' James 1. Dooley, JL,~ habitat management. and Gary J. Taylor" The Delmarva fox squirrel (Sciurus of habitat requirements will be essen- Methods niger cinereus) was placed on the fed- tial for both initiatives (Dueser and eral endangered species list in 1967 Terwilliger 1988). Data Base (32 FR 4001; US. Department of Inte- Habitat requirements might be rior 1970). Remnant populations expressed through any of three sepa- During a 12-mo search for remnant were restricted to four counties in rate but related components of habi- populations of the Delmarva fox eastern Maryland (Taylor and Flyger tat suitability: forest habitat struc- squirrel on the Maryland Eastern 19731, representing less than 10% of ture, forest tree species composition, Shore, Taylor (1976) located 36 "fox the historic range of the subspecies and surrounding landscape struc- squirrel present" (Present) sites with on the Delmarva Peninsula. Forest ture. Both habitat structure and for- extant populations and 18 "fox squir- clearing and habitat fragmentation est composition have been shown to rel absent" (Absent) sites. The gray throughout the range undoubtedly influence the distribution and abun- squirrel (Sciurus carolinensis) was contributed significantly to the pres- dance of fox squirrels in heterogene- present on all 54 sites. Taylor then ent endangerment (Taylor 1973). ous landscapes (Nixon and Hansen sampled the forest habitat of each The U.S. Fish and Wildlife Service 1987). site, to compare Present forest stands Recovery Plan for the restoration of Recent research has demonstrated with Absent stands. He established a the Delmarva fox squirrel to secure the potential influence of landscape representative 4 m x 200 m belt status emphasizes both the reintro- composition and structure on popu- transect on each site, on which he re- duction of this subspecies to suitable lations of woodland mammals occu- corded the number of trees by spe- habitats throughout the former range pying farmland mosaics (Wegner cies per diameter-breast-height and prescriptive habitat management and Merriam 1979, Middleton and (DBH) size class (5-20 cm, 20.1-30 for established populations (Taylor et Merriam 1983, Fahrig and Merriam cm, 30.1-50 cm, and 50.1+ cm), per- al. 1983). A thorough understanding 1985). Furthermore, changes in the cent crown cover, percent understory landscape of the Delmarva Peninsula cover, understory density, and 'Paper presented at symposium, Man- almost certainly played a major role understory species composition. All agement of Amphibians, Reptiles and Small on the decline of the fox squirrel habitat measurements were taken Mammals in North America. (Flagstaff.AZ, (Taylor 1973). from June through September 1972 July 19-2 1, 1988.) Given this background, the objec- and 1973. These data formed the ini- 2RaymondD. Dueser is an Associate tive of this study was to compare the tial data base for this study. Professor in the Department of Environ- mental Sciences, University of Virginia, apparent effects of habitat structure, Taylor (1976) reported the number Charlottesville,VA 22903. forest composition and landscape di- of trees measured in each of two size JJamesI. Dooley, Jr., is a graduate re- mensions on the presence and ab- classes: "small" trees (5-30 cm DBH) search assistant in the Department of Envi- sence of the Delmarva fox squirrel on and "large" trees (> 30 cm DBH). We ronmental Sciences, University of Virginia. 54 study sites in eastern Maryland. assigned each tree to one of five taxo- Charlo~esville,VA 22903. This analysis is the first step in the nomic groups: loblolly pine (Pinus 4GaryJ. Taylor is Associate Director of Wildlife. Department of Natural Resources, development of a predictive classifi- taeda), combined oak species (Quercus Maryland Forest, Park and Wildlife Service, cation model of habitat suitability for spp.), American beech (Fagus gran- Tawes State Office Building, Annapolis, MD this subspecies (cf. Houston et al. difolia), combined hickory species 27401. 1986). (Ca ya spp.), and combined mixed hardwoods. We estimated the ap- planimetry of 1:I000 black-and-white ground?"), (2) tree species composi- proximate total basal area for each photographs (dated 1978) obtained tion ('Which tree species predomi- size-taxonomic class by assuming an from Eastern Shore offices of @e nate in this forest and give it its char- average DBH of 17.5 cm for small USDA Agricultural Stabilization and acter?"), and (3) landscape dimen- trees and 40.0 cm for large trees. We Conservation Service. We initially sions ("What are the land use and then estimated total basal area for all measured each landscape variable cover dimensions of the landscape trees 2 5 cm DBH and the fraction of for a 2-km2 circular sample unit cen- mosaic in which this forest is embed- that total basal area represented by tered on the sample woodland. This ded?"). Conceptually, these compo- each taxonomic group. These basal unit was chosen as a first-approxima- nents represent a gradient of scales area estimates provide a basis for tion of "minimal population area" on from "microscopic" habitat structure comparing forest "composition" in- the basis of home range size and ac- to "macroscopic" composition to dependently of forest "structure" as tivity (Flyger and Smith 1980). Based "rnegasc~pic'~ con text. reflected, for example, in the raw on the results of analyses for the 2- Two-group discriminant function percentage of trees counted in each km2unit, both smaller (1-km2)and analysis was used to compare the taxonomic group. larger (4-km2) sample units subse- Present and Absent forest stands Original data were collected on quently were described in the same identified by Taylor (1976). Each land use and cover composition of way. analysis (1) computed the univariate the landscape surrounding a random F-ratio comparing -Presentand Ab- subset of Taylor's (1976) study areas sent sites for each habitat variable, (fig. 1). Landscape variables included Statistical Analyses (2) tested the centroids of Present area of open fields, percentage of and Absent sites for equality on the area forested, internal forest perime- This comparison of habitat compo- basis of a linear combination of the ter ("edge") within the sample unit, nents is based on multivariate statis- habitat variables (i.e., a linear dis- forest shape (Blouin and Connor tical analyses of three separate but criminant function), using multivari- 1985), and distance to next nearest related components of forest habitat ate analysis of variance (MANOVA), woodland. These variables are re- suitability: (I) habitat structure (3) indicated the relative contribution ferred to below as landscape "dimen- ('What does the forest 'look like' to of each habitat variable to any ob- sions." They were measured by an observer passing through on the served difference between centroids, based on the correlation between the variable and the discriminant func- tion, (4) tested the sample variance- covariance matrices of Present and Absent sites for homogeneity using a Box's M test statistic, and (5) indi- cated the percentage of the variation in group membership (Present or Absent) explained by the discrimi- nant function, based on the correla- tion between the membership vari- able and the discriminant function. (Dueser and Shugart 1978). All analy- ses were computed both with and without arcsin-square root transfor- mations of percentage variables. Re- sults of the parallel analyses were qualitatively similar in each case. For purposes of interpretability, only the results for untransformed variables are presented here. All analyses used the MANOVA and DISCRIMINANT routines of the Statistical Package for the Social Sciences (SPSS, Nie et al. 1975). Figure 1 .-Schematic diagram of 1 -, 2- and 4-km2 sample units for measuring landscape dimensions of "fox squirrel present" and "fox squirrel absent" study areas on the Eastern As an unbiased test of the ability Shore of Maryland..Eachsample unit was centered on one of Taylor's (1976) study areas. of each set of habitat variables to classify the group membership of the centage shrub-ground cover (0.564) Forest Tree Species Composition study sites (i.e., Present or Absent), a are particularly important in dis- jackknife procedure (Efron 1979) was criminating between sites. Conceptu- All 54 study areas supported a mix used to classify each of Taylor's ally, Present sites have larger trees, of hard- and soft-mast tree species. study areas. Each site was deleted in less shrub-ground cover vegetation, Although Present sites had some- sequence, DISCRIMINANT was run and less understory than Absent sites what greater basal areas for Ameri- for data from the remaining 53 sites, (fig. 2). Present sites were correctly can beech (p > 0.07) and mixed hard- and a classification function was classified 79% of the time
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