Habitat-Relative Abundance Relationship for Bobcats in Southern Illinois Clayton K. Nielsen; Alan Woolf Wildlife Society Bulletin, Vol.

30, No. 1. (Spring, 2002), pp. 222-230.

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Carnivore Research and Management

222
HABITAT-RELATIVE A B U N D A N C E RELATIONSHIP F O R BOBCATS

Habitat-relative abundance relationship

for bobcats in southern Illinois

CZaj^t~ii K. .\i'el.seti aitcl Alan Wtoolf

Abstract Large-scale multivariate
models of habitat suitability have been developed for several carnivore species. However, rarely have habitat models been combined with demographic information to create habitat-relative abundance relationships. We used remotely sensed land-cover information, bobcat habitat use data, multivariate distance statistics, and a geographic information system to provide wildlife managers a spatially explicit depiction of bobcat (Lynx rufus) relative abundance in a 12,512-kmL portion of southern Illinois. We used the Penrose distance statistic to model regional habitat similarity to areas within core areas of 52 radiocollared bobcats captured during 1995-1 999. Bobcat core areas were comprised primarily of forest cover (61%). Conversely, the southern Illinois region consisted of a more even mix of agricultural (36"/0), forest (29%), and grass cover (22%). Mean patch size of forest cover and proportion of forest cover were most correlated ( ~ 0 . 3 9 to ) Penrose distance. The Penrose distance model was validated using an independent data set of bobcat sighting locations (n= 248). Thirty-one percent and 81 OO / of independent bobcat sightings occurred in the top 10% and 25Y0 of distributions of Penrose distances, respectively. We then modeled relative abundance for the region based on Penrose distance, abundance information from areas occupied by radiocollared bobcats, and bobcat sighting locations. Given the collection of radiotelemetry data, this modeling technique can provide a better alternative than more traditional methods (e.g., scent station surveys) to assess large-scale distribution and abundance of solitary carnivores.

Key words bobcat, habitat-relative abundance relationship, habitat modeling, Lynx rufus, multivariate distance statistics, southern Illinois

Large-scale multivariate models of habitat suitability have been developed for several carnivore species (e.g.. black bears [Ursus anzericnnus], Clark et al. 1993;Florida panthers [Putnu concolor coryi], Maehr and Cox 1995; and gray wolves [Canis Ez~pus], Mladenoff et al. 1995). Two primary components of these models are habitat use data from several animals and remotely sensed landcover information. These models are then often used to describe or predict habitat needs for conservation purposes.

In a few instances, habitat models have been combined with demographic information to create habitat-population density relationships (Roseberry and Woolf 1998). Roloff and Haufler (1997) demonstrated this approach, combining habitat suitability maps and species home range illformation using published data from Canada lynx (Lynx canadensis) studies. Although such approaches could pro\-ide managers with desirable methods to estimate regional abundance or as part of a population viability analysis (Roloff and Haufler 199'),

Authors' address: Cooperative LVildlife Research Laboratory and Department o i Zoolog)i versitv, Carbondale, IL 62901, USA; e-mail for Nielsen: kez0920siu.edu.

mailc code 6504, Southern Illinois Uni-

Wildlife Society Bulletin 2002, 30(1):222-230

Peer refereed

Habitat-relative abundance relationship for bobcats they are rare for solitary carnivores. Wildlife managers in Illinois needed such information to focus management of bobcats (L. rufus). Based on increased sightings and a statewide assessment of relative abundance and habitat suitability (Woolf et al. 2002), bobcats were taken off the state's threatened species list in 1999 (Woolf et al. 2000). We used remotely sensed land-cover information, bobcat habitat use data, multivariate distance statistics, and a geographic information system to provide wildlife managers a spatially explicit depiction of bobcat relative abundance in southern Illinois.

Nielsen and Woolf

223

Next, we modeled habitat similarity between bobcat core areas and the rest of the region. Finally,we modeled relative abundance for the region based on habitat similarity, abundance information from areas occupied by radiocollared bobcats, and bobcat sighting locations.

Trapping and handling

We captured bobcats during November- march 1995-1999 with either cage-type traps constructed of galvanized wire mesh (38 x 38 x 90 cm) or padded number 3 Soft-catchB (Woodstream Co.. Lititz, Pa., USA) foothold traps. All traps were baited with whole or partial carcasses and chunks of meat from a variety of animals (e.g., roadkilled white-tailed deer [Odocoileus z~irginianus]). Visual attractants (e.g., bird wings) and commercial lures also were used at most trap sites. Captured bobcats were chemically immobilized for handling with a combination of ketamine hydrochloride (HCl) and xylazine HCl (both in 100 mg/mL concentration solution). We sexed,weighed, measured, and classified bobcats as adults Q 2 yr) or juveniles based on mass (bobcats <5 kg were considered juveniles) and condition of dentition. Capture and handling procedures were conducted in accordance with Southern Illinois University at Carbondale IACUC Animal Assurance #A-3078-01 and under provisions of Illinois Endangered Species Permit #95-14s issued to the second author.

Study area
We trapped and radiocollared bobcats on 2 study areas (eastern study area, 1,000 km2;western study area, 791 km2) in the 13 southernmost counties of Illinois (12,512 km2;Woolf and Nielsen 1999). This region included the Shawnee Hills, Ozark, Lower Mississippi River Bottomlands, and Coastal Plain natural divisions (Neely and Heister 1987) and was comprised of 39% cropland, 25% hardwood forest, and 24% rural grassland (Luman et al. 1996). Streams and roads were abundant on the landscape (stream density = 1.1 km/km2, road density = 1.4 km/km2). Elevation ranged from 92 to 316 m, with an average slope of l.dO. Human population density was 21.5persons/km2. Land cover of the eastern study area consisted primarily of closed-canopy mixed hardwood forests (55%) dominated by white oak (Quercus alba), black oak (Q. rubra) and hickory (Ca y a spp.), rural grasslands (26%),and cropland (11%) characterized by corn and soybeans (Luman et al. 1996). Land cover of the western study area consisted of a lower proportion of forests (46%) and rural grassland (8%)and a higher proportion of cropland (28%). Bobcats have been protected from legal harvest in Illinois since 1971 and were considered threatened in the state until 1999.

Radiotelemetry
We fitted bobcats withTelonics (Mesa,Ariz.,USA) model 315-S6A and Wildlife Materials (Carbondale, Ill., USA) model HLPM-2 140M radiocollars equipped with mortality sensors. We used standard ground and aerial radiotelemetry techniques to monitor bobcats (White and Garrott 1990). We used a TS-1 scanner (Telonics), hand-held 2- or 3-element yagi antenna, and a compass to ground-track. We used 2-element yagi antennas mounted on the wing struts of a Cessna 172 aircraft or on the skid of a Bell Long Ranger I1 helicopter for aerial telemetry. We determined point locations of bobcats from radiotelemetry, capture, and visual locations of radiocollared individuals. We obtained most (91%) locations by taking 22 bearings from bearing stations <2 km from bobcats. Bearing stations were located from 7.5-minute series (1:24,000) United States Geological Survey (USGS) topographic maps at road intersections and other landmarks. Less

Methods
Approach
Our goal was to create a habitat-relative abundance relationship for adult bobcats in the southern Illinois region. We calculated core areas of radiocollared bobcats and used core area size to determine an appropriate scale for habitat modeling. We then determined a set of regional habitat variables based on remotely sensed land-cover data.

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Wildlife Society Bulletin 2002,30(1):222-230

than 20 minutes elapsed between first and last bearings for 94% of all locations. We used the program LOCATE I1 (Nams 1990) to estimate locations according to the maximum likelihood estimator (Lenth 1981) and to calculate bearing error (n= 200, .t= 4.2f0.2) and error polygons (72 = 200,9= 1.6k0.1 ha, Springer 1979). Homing, capture, al, and aerial locations comprised 9% of total locations and were plotted on USGS topographic maps. We radiotracked most (87%) bobcats for less than 1 j7ear (2radiodaysbobcat = 477i2G). We collected an average of 8 3 f 12 annual locations/bobcat, >50 locations were obtained for 81% of bobcats, and >29 locations were collected for all bobcats. We obtained most (97%) locations of individuals 220 hours apart.

Figure 1. Hexagons used to clip land co\,er for habitat-relative abundance modeling for bobcats i n southern Illinois, 1995-1 999.

Core area estimation

We used the program RANGES V (Kenward and Hodder 1996) to estimate 50% core area boundaries of a d d t bobcats using the minimum convex polygon (MCP) estimator (Mohr 1947). We estimated core areas based on all locations obtained for each individual. Although other methods exist to depict spatial use (White and Garrott l990), we used the ,MCP estimator because it provided 1 area of use/individual, which was suitable for our analysis. We used core areas rather than home ranges because core area overlap was minor (Nielsen and Woolf 2001) and core areas are estimated more reliably than home ranges (Seatnan et al. 1999). We estimated core areas for 52 individuals (22 M, 30 F; Nielsen and Woolf 2001).

Habitat variable selection

%re calculated habitat variables based on landscape metrics for the 13 southertltnost counties of Illinois to represent the entire region. We created a continuous coverage of 2,'04 non-overlapping hexagons of 4.5 km2 (i.e.,2 size of MCP core areas: Nielsen 2000) that covered southern Illinois but did not overlap the regional boundary (Figure 1). We used FRAGSTATS (McGarigal and Marks 1995) to calculate landscape metrics associated with land cover within each hexagon. We log-transformed cover-type proportions and used a value of 0.001 for n d l proportions (Aebischer et al. 1993). We chose representative variables from each of 8 major metric groups, resulting in 153 potential variables for analysis (Table 1). We retained a smaller group of variables for habitat modeling based on presumed importance to bobcats and univariate statistics. First, for each cover type, we retained its log-transformed proportion. Second, within each co~ier type, we calculated nonparametric Spearman rank correlations for variables within each metric group and determined the number of nonsignificant correlations per variable. Third, we eliminated 1 of all pairs of correlated (1'50.0 5) variables within each metric group depending on number of nonsignificant correlations with other variables in the group. Values of significant correlations varied (r= 0.088-0.920) because occasionally variables were not present in hexagons, resulting in different sample sizes. We retained variables most correlated to others within each group (i.e., most representative of the group). In cases of ties, we chose the variable with suspected greater biological importance to bobcats (e.g., patch density of woods was selected

Land-cover data
We performed all geographic information systems operations using ARCANFO and ARCVIEW (Environmental Systems Research Institute, Redlands, Calif,. USA) and all statistical analyses (a= 0.05) using STATISTIX c44nalytical Software 1996). We derived land-cover information from the Illinois Critical Trends Assessment Project Land Cover Database (Luman et al. 1996) and reclassified from the original 23 cover classes into the follom7ing 8 aggregations: urban. transportation (i.e., roads and railroads), agriculture, grass. forest, open water, stream, and marsh. Because some roads and streams were not included in the original landcover classification,we converted road and stream data (Illinois Department of Natural Resources 1996) from vector to raster format and merged with the land-cover classification.

Habitat-relative abundance relationship for bobcats

Table 1. Potential habitat variables used to model habitat use by bobcats in southern Illinois, 1995-1 999. Variables were calculated using FRAGSTATS software (McGarigal and Marks 1995: 14-1 5). Class metrics were calculated for all 8 cover classes except when noted otherwise. Cover classes included urban (URB), transportation (TRAN), agriculture (AG), grass (GRS), forest (FOR), open water (WATI, stream (STR), and marsh (MAR).
-

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225

Calculation

Acronym

Metric (unit) Percentage of landscape (ha) ' 1 0 ) Largest patch index ( Patch density (no.il00 ha) Mean patch size (ha) Patch size coeificient of variation ('10) Edge density (mlha) Mean shape index Area-weighted mean shape index Landscape shape index Double log fractal dimension Core area percentage of landscape Core area density (no.1100 ha) Total core area index ('10) Mean core area (Oh) Mean core areafpatch (ha) Mean areafdisjunct core Patch core area coeificient of variation (%) Disjunct core area coefiicient of variation (%) Shannon's diversity index Modified Simpson's diversity index Patch richness density (no./100 ha) Relative patch richness (%i Shannon's evenness index Modified Sim~son'sevenness index Mean nearest neighbor distance (m) Nearest neighbor coefficient of variation (%i Mean proximity index Interspersion and juxtaposition index (%) Contagion index (%I

Area metrics PCTa Class LPI Class-landscapeb Patch metricsC Class-landscape PD MPS~ Class-landscape PSCV Class-landscape Edge metricsC EDe Class-landscape Shape metricsC MSI Class-landscape AWMSl Class-landscape LSI Class-landscape DLFD Class-landscape Core area metricsci CO/oLAND Class Class-landscape CAD TCAlg Class-landscape MCAl Class-landscape MCAl Class-landscape MCA2 Class-landscape CACVl Class-landscape CACV2 Class-landscape Diversity metricsC Landscape SHDl MSlDl Landscape Landscape PRD Landscape RPR Landscape SHE1 Landsca~e MSlEl Nearest neighbor metricsbc Class-landscape MNK Class-landscape NNCV Class-landscape MPI Contagion metricsC Class-landscape 111 Landscape CONTAG
a

reduced this data set to 15 variables by correlating physiognomic variables (e.g.,cover-type diversity) within each cover class and retaining the variable most correlated to others. Because urban land cover existed within only 2 of 52 (4%) bobcat core areas, we excluded the 2 variables associated with urban land cover and excluded 2 additional variables (edge density of water and marsh cover types) because they existed in only 25 and 35 of 52 (148%) bobcat core areas, respectively. This resulted in 11 final variables for habitat-relative abundance modeling. Further analysis required data normality; therefore, we transformed 2 variables (mean patch size of forest and grass cover) to normal distributions (Wilk- Shapiro statistic = 0.989-0.992) using square-root transformations. The other variables were distributed normally and not transformed (Wilk-Shapiro statistic = 0.898-0.989).

Habitat- relative abundance relationship

Modeling habitat sirizilarity to areas occupied bll radiocollared bobcats. We used the Penrose distance statistic to measure similarity between the mean habitat vector calculated from core areas of radiocollared adult bobcats and the rest of southern Illinois. We calculated Penrose distance as

Retained for analysis for all cover classes.

Landscape refers to each cell used to clip land cover. Not calculated for TRAN or STR cover classes. C ' Retained for analysis for the GRS and FOR cover classes only Retained for analysis for the landscape only. Not calculated for the WAT cover class. 6 Retained for analysis for the AG cover class only.

over patch density of water). This approach resulted in 50 potential habitat variables. We further

where populations i and j represented bobcat core areas and study area hexagons, respective1y;pwas the number of habitat variables evaluated, y was the variable value, k was each observation,

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WlEdltfe Society BuUetin 2002,30(1):222-230

represented only 1 bobcat core area. We assumed that areas occupied by Correlation radiocollared bobcats repbetween resented areas of greatest Mean Study study area Variable" vectorb area and PDC relative abundance (1.0 bobcat/hexagon) and dePercentage of transportation cover Percentage of agricultural cover termined mean Penrose Percentage of grass cover distance for those hexaPercentage o i forest cover gons containing bobcat Percentage of water cover core areas (Figure I). Percentage of stream cover Mean Penrose distance Percentage of marsh cover was 0.91; hence. we Edge density of the landscape assumed that hexagons Total core area index of agricultural cover with Penrose distances Mean patch size of grass cover < 0.91 to contain 1.0 bobMean patch slze of forest cover cat. "over type values are shown as raw percentages and not their log-ratio transformed We estimated zero and equivalents. intermediate relative abunCalculated from 50% minimum convex polygon core areas of 52 bobcats. dance based on the aforeSignificant ( P < 0.05) correlat~ons are denoted as is). mentioned sighting location data. We overlaid and Vwas variance (Manly 1986). This technique is sighting locations on the Penrose distance map and similar to the LMahalanobis distance statistic (Manly calculated frequency distributions. Because no 1986) used previously by several researchers for bobcats were sighted in hexagons with Penrose habitat modeling (Clark et al. 1993, Knick and Dyer distance >6.7, we considered hexagons having 1997, Corsi et al. 1999). We calculated the mean these values to contain no bobcats. We created a habitat vector as the mean values of the 11 habitat mathematical relationship between Penrose disvariables within bobcat core areas. We evaluated tance versus frequency of bobcat sightings to estieach hexagon on the study area relative to the mean mate intermediate abundance (i.e., between 0 and habitat vector, where a Penrose distance of 0 was 1.0 bobcat/hexagon). A logarithmic curve yielded closest to the mean vector. We made all calculations the greatest R2 (0.84); hence, the relationship J J = in a spreadsheet and appended the output database -0.51 ln[x] +0.95, where . Y is Penrose distance and to the hexagon coverage to create a GIS map of -2' is bobcat abundance. regional Penrose distance. We then correlated Penrose distance to each habitat variable to determine Results relative importance of each variable in calculating Penrose distance over the entire study area. Bobcat core area hexagons were comprised priModel validation. We validated the Penrose dis- marily of forest cover (61%) with agricultural and tance model using an independent data set of bob- grass cover combining for an additional 30% (Table cat sighting locations collected during 1995-1999 2). Conversely,the region consisted of a more even ( n= 248;Woolf and Nielsen 1999). Following Knick mix of agricultural (36%). forest (29%), and grass and Dyer (1997) and Corsi et al. (1999), we overlaid cover (22%). Although represented in relatively sighting locations onto the habitat map and deter- low proportions, percentage water cover differed mined the cumulative frequency of bobcat sight- most between core areas and the entire region; pering~ relative to Penrose distance. centage agricultural and forest cover also differed Modeling regional relative abundance. We markedly. Mean patch size of grass cover and permodeled bobcat relative abundance for the region centage stream cover were nearly the same based on Penrose distance associated with each between core areas and the entire region. Mean hexagon, abundance information from areas occu- patch size of forest cover and proportion of forest pied by radiocollared bobcats, and bobcat sighting cover were most correlated to Penrose distance data. For simplicity,we assumed that each hexagon (Table 2).
Table 2. Mean values (+ SE) of 11 habitat variables used to model bobcat habitat in southern Illinois and correlations between each variable and Penrose distance (PDI, 1995-1999.

Habitat-relative abundance relationship for bobcats

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Figure 2. Penrose distance map depicting habitat similarity between bobcat core areas and the southern Illinois region, 1995-1 999. Lesser Penrose distances indicate greater habitat similarity to bobcat core areas.

Figure 3. Bobcat relative abundance in southern Illinois based on a habitat-relative abundance relationship, 1995-1 999.

Mean Penrose distances for bobcat core areas and the entire region were 0.91f 0.80, range= 0.10-5.32) and 1.41 0.92 (range = 0.14-9.13, respectively. Regionally 1,018 of 2,704 (38%)hexagons contained Penrose distances 10.9 (Figure 2). Least average Penrose distance (i.e., with most similarity to bobcat core areas) occurred in east-southeast and west-central portions of southern Illinois. Penrose distance was greatest in the extreme northeast and Mississippi River bottomland areas. These areas consisted of proportionately more agricultural cover than the rest of the region. Thirty-one percent and 81% of independent bobcat sightings occurred in the top 10%and 25%of distributions of Penrose distances, respectively. Predicted relative abundance was relatively uniform across the study area, except for the western portions comprising Mississippi River bottomland and the extreme northeast (Figure 3).

Discussion
Roloff and Haufler (1997) used existing literature from Canada lynx studies to illustrate the utility of a novel approach to assess population viability. Their goal was not to actually conduct a population viability assessment, but to illustrate the use of their technique (Roloff and Haufler 1997). W e created a habitat-relative abundance relationship for bobcats in southern Illinois using a similar approach; however, ours was based on empirical data and was intended directly for use by wildlife managers in Illinois. Biologists frequently use habitat suitability information to focus wildlife management. These efforts

usually rely on habitat use data from relatively few animals,where inferences within a small study area are extrapolated to regions animals are known to inhabit but that were not monitored explicitly (Maehr and Cox 1995, Mladenoff et al. 1995). We used a multivariate distance statistic (i.e., Penrose distance) to achieve this goal for bobcats in southern Illinois. Other studies have used distance statistics to model habitat suitability (Clark et al. 1993, Knick and Dyer 1997, Corsi et al. 1999). The primary advantage this has over other techniques (e.g., logistic regression) is that there is no need to assume used and unused habitats are classified without error. Other studies used the Mahalanobis distance statistic, which accounts for correlations among variables (Ttlanly 1986). Although Penrose distance accounts for variance of each variable, it does not account for correlations; in our model, high correlations of habitat variables were removed during variable reduction procedures. Other researchers have used primarily compositional variables and a set of vector grids coded 0 or 1 for nonuse and use, respectively, whereas we performed calculations for each core area hexagon in a spreadsheet using raw values. Because the Mahalanobis distance equation does not contain squared terms (Manly 1986), the negative values provided by physiognomic variables disallowed use of the Mahalanobis distance statistic. Regardless, Manly (1986) demonstrated that Mahalanobis distance and Penrose distance statistics provide similar results. W e determined hexagon size for modeling based on core areas of radiocollared bobcats and modeled Penrose distance relative to areas occupied by

these bobcats. Bobcat core areas represented a general measure of areas used by bobcats and represented a useful scale for modeling. Most core areas were estimated based on individuals with >50 locations. However, we obtained fewer locations for some individuals, which may have somewhat biased estimates of core area size. Number of locations was not correlated to core area size (r=0.22, P<0.05); therefore, varying location sample sizes had little effect on core area size for bobcats in our study. There also was concern that location sample sizes were insufficient to delineate core areas; such is especially true given that we obtained relatively few locations for some bobcats. However, average location sample sizes for bobcat spatial use analysis rarely exceed those obtained during our study especially for those involving >50 individuals. Increasing the number of locations/bobcat likely would have resulted in larger MCP core area estimates Uennrich and Turner 1969). However, we believe this would not have changed modeling results, because habitat use did not differ between bobcat home ranges and core areas in southern Illinois (C. K. Nielsen, unpublished data). Thus, habitat within slightly larger core areas would not have differed from habitat within core areas as reported in this study. The Penrose distance model was very accurate when validated with an independent data set of bobcat sighting locations. Although validity and location accuracy of sighting data can obscure wildlife-habitat associations (Agee et al. 1989, Stoms et al. 1993, Palma et al. 1999), sighting data was the only set of independent data over the entire region that was useful for model validation. Further discussion of limitations of our sighting data are described in Woolf et al. (2000) and Woolf et al. (2002). Of additional concern were potential compounding of errors from multiple models and statistical techniques, whose aggregate effect on the final relative abundance model was unknown. However, we had no other means by which to assess this error or validate the model (e.g., via scent-station indices). Mean patch size of forest and percentage forest cover were the 2 variables most correlated to Penrose distance, which demonstrated their importance in determining bobcat habitat suitability. These variables were correlated negatively to Penrose distance, such that when mean patch size of forest cover was large and forest cover plentiful, Penrose distances were low, indicating suitable

habitat. These results agree with our previous analyses (Woolf et al. 2002), which found that forest compositional and physiognomic variables were primary predictors of bobcat abundance and distribution and habitat suitability statewide in Illinois. Forest cover is important to bobcats over large scales (Conner et al. 2001, Lovallo et al. 2001) for numerous reasons, including providing denning cover and prey resources (Anderson 1987, Rolley 1987). Positive correlations of percentage transportation, water, marsh, and agricultural cover and total core area index of agriculture to Penrose distance indicated poorer habitat suitability These cover types likely provided relatively little habitat value for bobcats when compared to forest cover types. Wildlife managers often require information on population abundance and distribution, which is especially challenging to determine for carnivores. Census techniques such as scent-station surveys are of limited value over larger scales due to poor precision of population estimates and low visitation rates (Diefenbach et al. 1994, Woolf and Nielsen 1999). Therefore, we used habitat suitability information in conjunction with calculated abundance in trapped areas and extrapolated abundance estimates to other areas using a bobcat sighting-habitat relationship. Van Horne (1983) warned about using habitat suitability information to assess abundance without knowledge of survival, reproduction, and physical condition of the species in question. However, because bobcats in southern Illinois have relatively high natural survival and recruitment rates (Nielsen 2000), a habitat-relative abundance relationship was appropriate.

Management implications
Because of the rarity and secretive behavior of medium-sized and large carnivores, quantifying their relative abundance over large scales is logistically difficult. We used data commonly collected in radiotelemetry studies (e.g., spatial use estimates) that provided a more desirable alternative than other methods (e.g.,scent-station surveys) to assess distribution and abundance of bobcats in southern Illinois. These techniques can be used by wildlife managers to estimate relative abundance and delineate management unit boundaries of other species, given that similar data is collected from several radiocollared animals. Managers also can use these techniques to estimate regional population size for management,

Habitat-relativeabundance relationship for bobcats

especially to initiate a harvest season. It is impossible to determine precision of population estimates based on this technique, and the technique is not useful to track short-term changes in population size. However, a habitat-relative abundance relationship can serve as a foundation for a population model because appropriate data is frequently rare for bobcats over large scales. Given the quantification of demographic (e.g., survival, reproduction) and spatial characteristics (e.g.,juvenile dispersal). such a model could be used to predict population response to harvest and potential gene flow among subpopulations.

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Acknou~ledgments. This study was funded through the Federal Aid in Wildlife Restoration program (Project W-126-R, Status of the Bobcat in Illinois), and the Cooperative Wildlife Research Laboratory at Southern Illinois University at Carbondale and the Division ofwildlife Resources of the Illinois Department of Natural Resources cooperated. We thank B. Bluett of the Illinois Department of Natural Resources for support and encouragement. C. Greene, D. Kennedy,J. Kolowski,M. Krecja,C. Schieler, C.Vincent,J.Waddell, and M. Woodruff provided field assistance with bobcat trapping and radiotelemetry. We thank all cooperating trappers and landowners for their assistance and access to property. M. Chamberlain, D. Diefenbach, and an anonymous reviewer provided helpful comments on an earlier draft of this manuscript.

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Clayton K. Nielsen received his B.S. in natural resources trom the University of Nebraska-Lincoln, his M.S. in en\,~ronmental and forest biology iron? State Univers~ty of New York College of Environmental Science ancl Foi-estry, and his Ph.D. in zoology from Southern Illinois University at Carbondale (SlliCl. He cur1 post-doctoral research tello\lj at the Cooperative rently is ' LZiildlite Rcsearch Laboratory (CWRL) at SIUC and operates 2 wildlite consulting tirms, Holtcrra Wildliie Management and Kinetic Conservation Group. Clay's professional interests include research and managenient of ungulate and carnivore populations in both urban antl rural landscapes. Alan Woolf is the director of the CWRL and a professor of zoologv at SIUC. He receiveti his B.S. from Cornell University, his M.S. irom Colorado State University, ancl returned to Cornell for his Ph.D. Alan's research interests are varied, but his professional emphasis centers around graduate programming, research, ant1 education. He has served the CWRL for the past 2 2 vears and has heen director since 1987.

Associate editor: Chamberidin