Independent Review of Status of the Bobcat Population in the Northern Lower Peninsula of Michigan Prepared by: Thomas M. Gehring, Timothy S. Preuss, and Bradly A. Potter Department of Biology, Central Michigan University Mount Pleasant, MI 48859 Summary: The number of bobcats harvested by hunters in the northern Lower Peninsula of Michigan (NLP) increased from approximately 140 to 291 during 1985-2002. However, given the current data, it is difficult to determine if this increase was a function of the bobcat population increasing or an increase in the number of hunters or other variables. Statewide, an increase in the number of bobcats harvested by hunters was correlated with the increase in the number of hunters. During the same period, average pelt price for bobcats appeared to decrease. These results suggest that bobcats may now be more valued by hunters as a trophy species rather than an alternate income source and/or bobcat hunting is driven more by the recreational compared to economic incentives. Overall, when the change due to the number of hunters was removed, the bobcat population in Michigan appeared to be relatively stable. However, we could not adequately address the status of the NLP population separately. The proportion of juveniles and yearlings remained relatively high throughout the time period. Based on estimates from opinion surveys, if trapping and hunting occurred annually within in the NLP, an estimated 620 to 3,000 bobcats would be harvested each year assuming 100% filling of bag limits. An estimated 420 to 1,500 bobcats would be harvested annually assuming a 50% success rate. Without accurate population index or other independent data for the entire NLP, it is difficult to evaluate the impact that this harvest figure would have on the NLP bobcat population. Distribution of bobcat harvests in the NLP suggests there are 1 or 2 relatively disjunct regions of Zone 2 that maintain relatively high harvest intensity, whereas the areas between exhibit lower harvest rates. There also appears to be a correlation between harvest intensity and the density of streams and rivers in the NLP. Additionally, we found greater than expected harvest rates on public land compared to private land in the NLP. Investigation into the possible source-sink dynamics and spatial patterning of bobcat harvest in the NLP should be pursued further. These investigations could provide insight into possible new zoning recommendations based on landscape-level features (e.g., hydrologic patterns) and/or spatial patterning of high-quality bobcat habitat (e.g., contiguous patches of lowland forest and non-forested wetland habitat). Introduction: In 1975, bobcats (Lynx rufus) were listed in Appendix II of the Convention on International Trade in Endangered Species (CITES) because of concern for the viability of their populations. This listing required state agencies to provide the U. S. Fish & Wildlife Service with data on the status and viability of bobcat populations. The current geographic range of the bobcat includes all of Michigan although bobcat densities are likely greater in the northern 2/3 of the Lower Peninsula of Michigan. Currently, bobcats are harvested by hunting only in Zone 2 of the northern Lower Peninsula (NLP), with a bag limit of 1 bobcat per hunter. Zone 2 is 1 subdivided into North and South zones with season lengths of 60 days (1 January to 1 March) and 33 days (15 January to 16 February), respectively (Figure 1). Figure 1. Northern Lower Peninsula of Michigan bobcat harvest units depicting Zone 2 North and Zone 2 South units (source: MDNR) Annually, since 1985 the Michigan Department of Natural Resources (MDNR) has collected age, sex, and location data for bobcats harvested in the state. During 1991-1996, MDNR researchers conducted a mark-recapture study and scent-station survey in Crawford, Missaukee, and Roscommon counties in the NLP to evaluate bobcat survival and relative abundance (Earle et al. 2003a). Recently, Central Michigan University (CMU) began a radiotelemetry study of bobcats predominantly in Missaukee and Roscommon counties (Houghton Lake Study Area; Preuss and Gehring, unpublished data). Additionally, in 2003 we began assessing the relative abundance of bobcats using a scent-station survey in Roscommon and Missaukee counties, as well as parts of Clare, Crawford, Gladwin, Kalkaska, and Osceola counties. Collectively, these data allow an examination of bobcat population characteristics in the NLP over an approximately 20-year period. 2 Objectives: 1) 2) 3) Evaluate changes in the relative abundance and population structure of bobcats in the NLP; Evaluate the designation of Zone 2 into a North and South unit; and Provide recommendations for future bobcat management in the NLP. Methods: Relative Abundance We graphically examined the change in the number of registered bobcats over time. Further, we used furbearer harvest survey results (Frawley 2001, 2003, unpublished data) to examine the fur taker population during 1985 to 2002. Because pelt price might influence trapping effort (Royama 1992), we examined this possible influence in the number of bobcats harvested via hunting. We used average bobcat pelt price data from Wisconsin (Dhuey et al. 2003) and regressed (PROC REG, SAS Institute 1994, α = 0.05) this with the number of bobcat hunters. Reliable data were not available for an estimate of the number of bobcat hunters in the NLP only due to high variance. Thus, we pooled Zone 1 and Zone 2 bobcats harvested by hunters and used estimates of the number of bobcat hunters in the State of Michigan (Frawley, unpublished data). We conducted linear regression analysis on the number of registered bobcats hunted (dependent variable) and the number of bobcat hunters. We transformed data using the natural logarithm to gain stationary variance (Royama 1992). We examined the residuals from this regression by plotting residuals in time (i.e., sequence plot) to determine if there was a change in the number of bobcats hunted over time (i.e., a time-related effect, Neter et al. 1996). Additionally, we plotted differenced values [N difference = ln (N t) – ln (N t+1)] for number of bobcats harvested by hunters and the estimated number of hunters. Differencing provided a stationary mean in the values and allowed estimates to be examined as rates of change over time (Royama 1992; Swanson 1998). We conducted a scent-station survey during October – November 2003 in Roscommon and Missaukee counties, as well as parts of Clare, Crawford, Gladwin, Kalkaska, and Osceola counties. The survey was patterned after Linhart and Knowlton (1975), with modifications by Roughton and Sweeny (1982). Recommendations by Sargeant et al. (1998) and Sargeant et al. (2003) also were incorporated. Scent-stations consisted of a 0.9-m diameter circle of sand with a fatty-acid scent tablet placed in the center as an attractant. Seventy transects with 10 stations along each transect were checked for 2 nights. Stations were placed 480-m apart along each transect and transects were located ≥5 km from the nearest transect. The presence of all tracks was recorded to species whenever possible, and visitation rates were calculated for the proportion of stations and the proportion of transects visited by each species. We qualitatively compared various population indices to assess changes in the relative abundance of bobcats over time in Crawford, Missaukee, and Roscommon counties (i.e., the 1991-1996 MDNR study area and our current study site). We only used a qualitative comparison 3 because objectives and methodology for studies were not consistent, and the data record for the total time series (1985-2002) was not complete. These data sources included: 1) number of bobcats harvested during 1985-2002; 2) MDNR scent-station survey data from 1992-1996 (Earle et al. 2003a); 3) CMU scent-station survey data from 2003 (Preuss and Gehring, unpublished data); 4) number of bobcats captured per 1,000 trap nights by the MDNR, 1991-1996; and 5) number of bobcats captured per 1,000 trap nights during 2003 (Preuss and Gehring, unpublished data). We also attempted to estimate the range in the number of bobcats that would be trapped in the NLP if a trapping season was offered. To estimate this number, we used fur harvester opinion survey report data for 2003 (Bull and Peyton 2004), and the number of respondents that indicated that they were ‘very likely’ to trap bobcats in the NLP if it were legal. We estimated a total NLP bobcat harvest (hunting and trapping combined) by adding the mean of the number of bobcats hunted during 1998-2002 in the NLP to our estimated trapper harvest of bobcats in the NLP (i.e., trapping includes additive mortality above the average hunt-related mortality). We used 2 estimates of a trap harvest of bobcats: 1) based on the total proportion of respondents in the Bull and Peyton (2004) survey that indicated they were ‘very likely’ to trap in the NLP (i.e., estimated at 18% of furtaker license holders); and 2) based on the proportion of respondents in the Bull and Peyton (2004) survey that had previously trapped or hunted bobcats and indicated they were ‘very likely’ to trap in the NLP (i.e., 19.5%). We used the mean number of bobcat hunters and trappers during 1998-2002 in our latter estimate (Frawley 2001, 2003). We assumed a bag limit of 1 bobcat per hunter or trapper and a trapping success rate of 1 bobcat per trapper (Frawley 2001, 2003). That is, we assumed that all trappers would fill their bag limit. We also calculated estimates assuming that 50% of trappers would fill their bag limit. We initially attempted to parameterize the Minnesota Furbearer Population Model (e.g., Rolley et al. 2001) with new harvest estimates and existing estimates of other population parameters; however, our simulations of population trends were very sensitive to the value of the initial population size. Furthermore, small changes in the initial population size resulted in vastly different population trends. Rolley et al. (2001) suggested that this model should be calibrated to either an independent source of population trend data or an estimate of absolute population size. We chose not to include these analyses because of concerns about the lack of accurate population indices or absolute counts of the number of bobcats in the NLP over the time period 1985-2002. Population Structure We sorted MDNR databases to isolate the age and sex of registered hunted bobcats. We used the sex identified based on the MDNR’s assessment using lower canine tooth maximal thickness and width of the root (Friedrich et al. 1983) when given. If this lab-determined sex was not provided, we used the field-determined sex. The MDNR estimates age of registered bobcats by counting cementum annuli in cross sections of the lower canine root (Crowe 1975). We classified age as juvenile (0.5 year), yearling (1.5 year) and adult (2.5+ year). Animals that did not have the sex or age identified were excluded from further analyses. Initially, we graphically examined the change in estimated age and sex ratios based on harvested bobcats in 4 Zone 2. Significant changes in ratios over time were determined by linear regression (PROC REG, SAS Institute 1994) with an α-value of 0.05. Preliminary Modeling of Potential Bobcat Population Size and Distribution We obtained a broad estimate of bobcat abundance by determining the number of male and female bobcat home ranges that could theoretically fit in the NLP using ArcView GIS (ESRI, Redlands, California). Average minimum convex polygon (MCP) home ranges were calculated for male and female bobcats radio collared in the Houghton Lake Study Area (HLSA). We captured bobcats during March-June 2003 using No. 3 Victor Soft-Catch padded foot-hold traps (Earle et al. 2003b). We immobilized bobcats via an intramuscular injection of 10 mg/kg ketamine hydrochloride and 1.5 mg/kg xylazine hydrochloride (Kreeger 1999) in order to determine sex, age, reproductive condition, ear tag and radio collar animals. Radio-collared bobcats were located a minimum of 2-3 times weekly from May – December 2003 using a vehicle-mounted 4-element Yagi directional antenna and electronic compass (Lovallo et al. 1994). Triangulations were made from telemetry stations with 3-4 bearings obtained as quickly as possible to reduce telemetry error (White and Garrott 1990). Bobcat locations were estimated using the microcomputer program LOCATE II (Nams 1990). We estimated home-range size using the MCP method (Mohr 1947) using the Animal Movement Analysis Extension to ArcView GIS (Hooge and Eichenlaub 1997). Estimates of home-range size (Table 1) were consistent with those of bobcats in Wisconsin (Lovallo and Anderson 1996) and Minnesota (Fuller et al. 1985). We divided the total area of the NLP by the average home-range size of male ( x = 46.5 km2 = 18 mi2, sd = 27.3 km2 = 10.5 mi2) and female ( x = 10.4 km2 = 4.0 mi2, sd = 5.9 km2 = 2.3 mi2) bobcats to obtain an estimate of the number of male and female bobcats that could potentially reside in the NLP. This model assumed that there was no intrasexual home-range overlap, all bobcats were resident (i.e., bobcats without established home ranges were not accounted for), and bobcats occurred in all landcover types (i.e., bobcats exhibited no selection of habitat). Table 1. Preliminary data on summer home-range size for bobcats monitored during 2003 in the Houghton Lake Study Area as part of a Central Michigan University Study (Preuss and Gehring, unpublished data). Bobcat ID Sex M01 M02 M03 F01 F02 F03 F04 Male Male Male female female female female Minimum Convex Polygon (km2) 26.15 77.44 35.86 9.25 7.01 6.24 19.08 5 Minimum Convex Polygon (mi2) 10.10 29.90 13.85 3.57 2.71 2.41 7.37 We also developed sex-specific models of bobcat abundance relative to preferred landcover types since bobcats exhibit habitat selection (e.g., Lovallo and Anderson 1996; Preuss and Gehring, unpublished data). We used the 2001 landcover GIS theme available from the MDNR. Within ArcView GIS, 2 hexagonal grids were constructed and overlaid on the NLP (Figure 2). The area of each hexagon in the first grid was equivalent to the average home range of male bobcats. The area of each hexagon in the second grid was equivalent to the average home range of female bobcats. We calculated the percentage of streams, lowland conifer forest, and lowland conifer forest/non-forested wetlands within each hexagon of the sex-specific grids. Lowland conifer forest and non-forested wetland cover types were chosen because radio-collared bobcats in the HLSA used these cover types in greater proportion to their availability based on chi-square analysis (χ2 > 29.4, P < 0.001; Neu et al. 1974). We modeled percentage of streams because data from radio-collared bobcats, scent-station surveys, and harvest data suggested a possible relationship between bobcats and streams. We then modeled abundance using 2 approaches. For the first approach, we calculated the mean and minimum percentages of each landcover class occurring within the home ranges of male and female radio-collared bobcats. For each landcover class, we then modeled abundance by assigning 1 bobcat to each hexagon with a percentage greater than or equal to the mean and minimum percentages of that landcover class occurring within a home range. Lovallo and Anderson (1996) found that some home ranges of bobcats in Wisconsin were composed of <5% lowland conifer; however we used 5% as a cut-off and modeled bobcat abundance by assigning 1 bobcat to each hexagon with greater than or equal to 5% lowland conifer composition. For our second approach, we used known bobcat locations obtained from radio telemetry and scent-station surveys to define a sample of hexagons known to be occupied within each grid (n = 42 and 69 hexagons for males and females, respectively). We then calculated the mean proportion of each landcover class within this set of occupied hexagons. For each landcover class, we then modeled abundance by assigning 1 bobcat to each hexagon with a proportion greater than or equal to the mean proportion of that landcover class occurring within known occupied hexagons. This allowed us to obtain potential estimates of the number of male and female bobcats in the NLP. These approaches assume that there is no intrasexual home-range overlap and all bobcats are resident (i.e., bobcats without established home-ranges are not accounted for). 6 a) b) Figure 2. a) Male and b) female hexagonal grids were used to model potential bobcat population size in the northern Lower Peninsula. 7 Distribution of Harvest We qualitatively examined changes in the distribution of harvested bobcats in the NLP using a GIS database and ArcView GIS. We used hydrology, transportation, landcover, land ownership, and township data themes available from the MDNR for analyses. Because of time constraints and the fact that not all harvest locations were mapped to Section, we chose to map locations to the scale of township. We mapped the locations of harvested bobcats as harvestintensity maps with townships mapped into equal intervals of number of bobcats harvested. We mapped the distribution of harvested bobcats in 3-year time periods from 1985-2002. We also created harvest-intensity maps for bobcats harvested in Zone 1 (Upper Peninsula). We overlaid separately the harvest-intensity maps for Zone 2 onto a hydrology and a transportation theme to qualitatively assess any link between harvest and landscape features. Additionally, we examined the distribution of harvested bobcats on public vs. private land to determine any disparity in harvest intensity. We intersected the GAP landownership theme with townships and reclassified townships as public or private land based on the dominant landownership type (i.e., 50% or more of township). We identified commercial forest land as privately owned land. We then overlaid harvest locations for bobcats during the time period 1998-2002 to gain a count for each year. We used contingency analysis (Zar 1996) to determine if bobcats were harvested differentially by year and landownership type. Results: Relative Abundance The number of bobcats harvested in Zone 2 (North and South hunting units combined) increased during 1985-2002; however the linear trend was only slightly positive (Figure 3). When Zone 2-North and Zone 2-South were viewed separately, a similar trend was exhibited. However, more bobcats were generally harvested in the North Unit compared to the South Unit, most likely due to the longer hunting season in the North Unit (Figure 4). 8 Figure 3. Number of bobcats harvested in Zone 2 (hunting units combined) during 1985-2002. Trend line is shown as the dashed line. Figure 4. Number of bobcats harvested in Zone 2 (hunting units separated) during 1985-2002. 9 During 1985-2002, the estimated number of bobcat hunters in Michigan (Zone 1 and Zone 2 combined) appeared to increase (Frawley 2003; Figure 5). This increase corresponded with a decrease in the average pelt price of bobcats, based on Wisconsin pelt prices (Dhuey et al. 2003; Figure 6). We found a significant inverse relationship between the estimated number of bobcat hunters and the average pelt price (F = 20.83; 11,12 df; P = 0.001). Thus, as bobcat fur prices appear to have become reduced and stabilized at approximately $40-50, the estimated number of bobcat hunters in Michigan has increased to nearly 1,900 (Figure 7). This trend suggests that hunter effort may be functioning independent fur prices and bobcats may be Est Number of MI BobcatofHunters valued more as a trophy animal or the recreational aspects of the hunt are more important. Number of Hunters 2500 2000 1500 1000 500 2002 2000 1998 1996 1994 1992 1990 1988 1986 1984 1982 1980 0 Year Figure 5. Estimated number of bobcat hunters in Michigan (Zone 1 and Zone 2) based on WI Avg Bobcat Pelt Price MDNR mail harvest survey (Frawley 2003). 100 Average Pelt Price ($) 90 80 70 60 50 40 30 20 10 2000 1998 1996 1994 1992 1990 1988 1986 1984 1982 1980 0 Year Figure 6. Average pelt prices for bobcats in Wisconsin during 1980-2002 (Dhuey et al. 2003). 10 Average WI Bobcat Pelt Price, $ Relation of Pelt Price to Number MI Bobcat Hunters 100 80 60 40 20 0 0 500 1000 1500 2000 2500 Estimated Number MI Bobcat Hunters Figure 7. Relationship between estimated number of bobcat hunters in Michigan and average price for bobcat pelts. The correlation coefficient was r = -0.81. We found a significant positive linear relationship between the estimated number of bobcat hunters and the number of bobcats harvested in Michigan (F = 14.88; 10,11 df; P = 0.003; Figure 8). The number of bobcat hunters estimated 60% of the variation in the number of bobcats registered (r2 = 0.598). Therefore, most of the changes in the number of bobcats harvested in the entire State of Michigan appear to be due to increases in the estimated number of bobcat hunters in the state. Figure 8. Relationship between estimated number of bobcat hunters and the number of bobcats harvested in Michigan (Zone 1 and Zone 2 combined). 11 We plotted the residuals from the regression model of bobcat hunters and number of bobcats harvested relative to year (i.e., sequence plot of residuals) to assess any time-related patterns. There appears to be no trend in the sequence plot of residuals with relatively random variation around the mean of zero over the time period 1985-2002 (Figure 9). Thus, with the influence of the number of hunters removed, the number of bobcats harvested during 1985-2002 appears to be relatively stable. There does appear to be some annual variation in the number of bobcats registered in Michigan. This variation might be due to changes in the abundance of bobcats although it could reflect annual variation in weather conditions. Variation might also be explained in part by significant declines in pelt price. For example, the apparent decrease in the number of bobcats harvested during 1993 coincided with a bottoming out of fur prices that stabilized at relatively low prices over the next decade (Figure 6 and 9). Determination that similar dynamics are occurring specifically in the NLP bobcat harvest and population are not possible given the lack of goodStd estimates of the number of bobcat Residuals - point plothunters in the NLP. 2 1.5 0.5 2001 1999 1997 1995 1993 1991 1989 -0.5 1987 0 1985 residual 1 -1 -1.5 -2 Year Figure 9. Sequence plot of residuals from the regression model in Figure 8. Plots of differenced values for the number of bobcats harvested and the estimated number of bobcat hunters in Michigan provided similar results. The rate of change in both quantities was closely related, with the change in the number of hunters closely tracking the change in the number of bobcats harvested during 1996-2002 (Figure 10). Prior to 1996, data on the number of bobcat hunters was less complete, making inferences about the rates of change difficult. A regression model of the difference values provided a similar positive linear relationship as outlined above (Figure 11). 12 Figure 10. Time-series plot of difference values (natural-log transformed) for estimated number of hunters and number of bobcats harvested in Michigan (Zone 1 and Zone 2 combined), 19852002. Figure 11. Regression relationship between difference values (natural-log transformed) for estimated number of hunters and number of bobcats harvested in Michigan (Zone 1 and Zone 2 combined), 1985-2002. Additional data on the hunter effort in Zone 2 of the NLP (e.g., estimated number of bobcat hunters and/or number of field-days of hunting effort) would be useful in estimating catch-per-unit-effort (CPUE). These estimates should provide an indication of the current population trend for bobcats in the NLP. With the current analyses, we used estimates of the number of bobcat hunters for Zone 1 and Zone 2, and this may not provide an accurate indication of the trends in the NLP bobcat population in part because of the much greater number of 13 bobcats harvested in Zone 1 compared to Zone 2. There also may be a difference in hunting effort that exists between Zone 1 and Zone 2. We examined several indices of the bobcat population in portions of Crawford, Missaukee, and Roscommon counties of the NLP to assess population fluctuations. We captured and radio-collared 5 bobcats during 2003 with 1,353 trap-nights of effort with foot-hold traps (catch per effort = 3.7 cats per 1,000 trapnights). We monitored a total of 7 bobcats (3 males, 4 females), 1 of which was captured in a box trap and 1 was an incidental capture by federal biologists. Average home-range size (MCP) was 46.5 km2 and 10.4 km2 for males and females, respectively (Preuss and Gehring, unpublished data, Figure 12). Additionally, we recorded 49 bobcat visits across 700 scent stations (7% visitation of stations, 43% of transects with visits) conducted during fall 2003 (Preuss and Gehring, unpublished data, Figure 13). 30 km Figure 12. Location of Central Michigan University bobcat study with home-range polygons of bobcats monitored during 2003-2004. 14 Figure 13. Location of Central Michigan University bobcat study with scent stations and transects identified by rows of circles. Stations with bobcat visits during fall 2003 are identified in red. During 1991-1996, the MDNR bobcat study in approximately the same area we are currently working, reported catch per trapping effort ranging from 3.1 to 6.2 bobcats per 1,000 trapnights ( x = 5.2, sd = 1.2 bobcats per 1,000 trapnights; Earle et al. 2003a). A scent-station survey conducted by the MDNR during 1992-1996 provided estimates of the number of bobcat visits ranging from 27 to 49 bobcats (Earle et al. 2003a). Although the time series are not extensive, nor are they complete, in general the fluctuations in the number of bobcats harvested in the Houghton Lake Study Area appears concordant with the indices used to assess population change during 1992-1996. Scent-station data, but not catch-per-effort data for 2003 also appear to reflect a possible increase in the numbers of bobcats harvested. However, our catch-per-effort estimate was within the confidence intervals for the mean of the MDNR estimate. 15 We suggest caution in interpreting increases in the bobcat population in the north-central LP using harvest data, however, due to the likely strong positive relationship between the number of bobcat hunters and the number of bobcat harvested in Michigan. This caution assumes that this same trend was evident in the NLP. Further, we suggest that comparisons between our study and the 1991-1996 MDNR study are tentative because study methods differed with respect to scent-station protocols and study objectives (e.g., the MDNR study attempted to assess different attractants at stations using a series of experimental treatments that varied over time, Earle et al. 2003a). Furthermore, our study results are all preliminary and based on a relatively small sample of bobcats and 1 scent-station survey to date. Our continued efforts to radio collar bobcats during spring-summer 2004 and our continuation of a scent-station survey during fall 2004 should provide additional insight into the relative abundance and changes in abundance of this local population. In their fur harvester opinion survey report, Bull and Peyton (2004) reported that 19.5% of respondents who had hunted or trapped bobcats before would be ‘very likely’ to trap bobcats in the NLP if it were legal. During 1998-2002, we estimated a mean of 2,324 ± 271 bobcat hunters and trappers in Michigan based on furbearer survey reports (Frawley 2001, 2003). Thus, an estimated 400 to 506 bobcat hunters and trappers in the state would likely trap bobcats in the NLP if it were legal. Additionally, the kill success rate for bobcat trappers averages 1 bobcat per trapper per year. During 1998-2002, an average of 220 bobcats per year was hunted in the NLP. Assuming a bag limit of 1 bobcat per person per year in the NLP, if bobcat trapping and hunting occurred, the annual harvest would be an estimated 620 to 726 bobcats. Assuming a bag limit of 0.5 bobcats per person per year in the NLP (i.e., 50% success rate among trappers), if bobcat trapping and hunting occurred, the annual harvest would be an estimated 420 to 473 bobcats. Additionally, Bull and Peyton (2004) found that 18% of furtakers (approximately 3,000 individuals) indicated they would be ‘very likely’ to trap bobcats in the NLP if it were legal. Thus, an estimated 3,000 and 1,500 bobcats might be harvested annually in the NLP assuming bag limits of 1 bobcat per person per year and 0.5 bobcats per person per year, respectively. Population Structure The proportion of females in the Zone 2 North harvest was approximately 0.40 during 1985-2002 with no significant linear trend (F = 0.21; 16,17 df; P = 0.66; Figure 14a). Conversely, the proportion of females in the Zone 2 South harvest decreased from over 0.50 to near 0.40 during the same time period and this trend was nearly significant (F = 4.00; 16,17 df; P = 0.06; Figure 14b). When harvest data for the Zone 2 units were pooled, the proportion of females remained at nearly 0.40 during 1985-2002 (Figure 14c). The apparent decline in the proportion of females in Zone 2 South may simply be an artifact of the relatively low number of bobcats harvested in this unit prior to 1994 compared to Zone 2 North. Possible fluctuations in the sex ratio also may be related to differential vulnerabilities of animals, differential mortality rates between sexes and/or differential selectivity by hunters. The harvest data do not provide explanations for possible fluctuations in Zone 2 South. 16 North a) South b) Combined c) Figure 14. Proportion of females in the bobcat harvest during 1985-2002 for: a) Zone 2 North; b) Zone 2 South; and c) Zone 2 (North and South units combined). 17 Assuming that the proportion of different age classes in the harvest are representative of age ratios in the bobcat population, it appears that the proportion of adult bobcats is increasing and the proportion of juvenile bobcats is decreasing. This suggests that the population may be aging and recruitment of kittens into the yearling and adult age classes may be decreasing, thus the rate of increase of the NLP bobcat population may be decreasing. During 1985-2002, mean proportions of juveniles, yearlings and adults in the NLP harvest were 23%, 30%, and 47%, respectively (Figure 15). Figure 15. Age ratios for juvenile (0.5 year), yearling (1.5 year) and adult (2.5+ year) bobcats harvested in Zone 2 (North and South units combined) during 1985-2002. Age pyramids suggested that the proportion of juveniles and yearlings remained relatively high throughout the time period. The 2.5 and 3.5-year classes appeared to increase in the harvest during the early and late 1990’s as well as most recently. MDNR analysis of the bobcat population in the north-central NLP suggested that more young bobcats (juveniles and yearlings) were harvested by hunters compared to MDNR trapping efforts (Earle et al. 2003a). Wildlife populations that are exploited by harvest typically have greater numbers of juveniles and yearlings in the harvest record compared to older animals. This may explain the preponderance of younger animals in the NLP harvest (Fritts and Sealander 1978; Rolley 1985) or it may simply be a function of hunter selectivity for younger animals, whereby any bobcat that is treed is harvested. Younger bobcats are presumably more naïve and easier to tree compared to older individuals. Interpretation of these population structure data should be cautious, however. Caughley (1974) cautioned about using age ratio data to infer dynamics of populations. Instead, information on the rate of increase for populations would more accurately reflect population dynamics compared to age ratios (Caughley 1974). 18 GIS Methods for Modeling Potential Bobcat Population Size and Distribution Our GIS modeling efforts produced theoretical estimates of bobcat population size in the NLP with assumptions outlined above. Estimates, however, varied widely from a total of 843 to 3,543 bobcats in the NLP (Table 2). Table 2. Population size estimates for resident bobcats in the northern Lower Peninsula of Michigan assuming habitat preference relationships and that intra-sexually there are nonoverlapping home ranges. Estimated Estimated Estimated Model # Males # Females Total Figure No selection for habitat (complete saturation) 1,044 4,665 5,709 --Mean Lowland Forest 255 908 1,163 16 Mean Lowland Forest & Non-Forest Wetland 185 658 843 17 Minimum Lowland Forest 94 922 1,016 18 >5% Lowland Forest (Lovallo) 699 2,844 3,543 19 Models depicted in figures 16, 17, and 18 highlight similar regions of the NLP that may be core habitat areas for bobcats. These same general areas also correspond with areas that typically have the highest intensity of bobcat harvest (see Figure 20). Figures 16-18 also might offer some insight into the disjunct nature of the North and South hunting units in Zone 2. That is, 2 relatively disjunct habitat patches are distributed in the northeastern and southwestern quarters of Zone 2. A possible 3rd habitat patch might exist in the east-central region of Zone 2. Figure 19 probably provides a maximum population size since most of the NLP is depicted as potential bobcat habitat using a lower proportion of lowland forest in bobcat home ranges. We caution, however, that these estimates are based on preliminary data and small sample size of the number of bobcats monitored via radio telemetry as well as only 1 location where scent stations have been conducted in the NLP to provide landcover type associations in these models. For example, our sample sizes for figures 16-18 were n = 42 and 69 hexagons for males and females, respectively. These models will need to be refined and validated before they might offer greater insight into potential bobcat habitat and better approximations of the size of the bobcat population in the NLP. 19 a) b) Figure 16. a) Male and b) female hexagonal grids predicted to potentially contain male (blue) or female (red) resident, adult bobcats in the NLP. Predictions are based on the mean proportion of lowland forest in bobcat home ranges as determined by a sample of radio-collared bobcats and bobcats detected using a scent-station survey. a) b) Figure 17. a) Male and b) female hexagonal grids predicted to potentially contain male (blue) or female (red) resident, adult bobcats in the NLP. Predictions are based on the mean proportion of lowland forest and non-forested wetland habitat in bobcat home ranges as determined by a sample of radio-collared bobcats and bobcats detected using a scent-station survey. 20 a) b) Figure 18. a) Male and b) female hexagonal grids predicted to potentially contain male (blue) or female (red) resident, adult bobcats in the NLP. Predictions are based on the minimum proportion of lowland forested habitat found in bobcat home ranges (22% for males, 18% for females) as determined by radio-collared bobcats in the CMU bobcat study. a) b) Figure 19. a) Male and b) female hexagonal grids predicted to potentially contain male (blue) or female (red) resident, adult bobcats in the NLP. Predictions are based on the proportion of lowland forest in bobcat home ranges (>5%) as determined by Lovallo and Anderson (1996) in northwestern Wisconsin. 21 Distribution of Harvest The distribution of bobcats harvested during 1985-2002 is relatively widespread throughout Zone 2; however, harvest locations do appear to concentrate into 1 large patch in the northeastern portion of the NLP with a secondary linear patch in the central NLP (Figure 20). A portion of this distribution pattern may be related to the different season lengths between the North and South hunting units in Zone 2, although these same regions are identified in our habitat analysis above. Figure 20. Distribution of harvested bobcats across the NLP during 1985-2002. Harvest intensity of bobcats is mapped to the scale of township. Cooler colors indicate lower harvest intensity, whereas warmer colors represent greater numbers of bobcats harvested. We mapped the distribution of harvested bobcats across 3-year intervals to examine changes in the pattern of harvest intensity. We maintained the same scale of harvest intensity in Figure 21. In general, harvest intensity remained relatively constant and localized in townships of the northeastern portion of the NLP, with 8-18 bobcats harvested per township during 19852002. The central portion of the NLP exhibited periods of greater harvest intensity during 19851990, reduced harvest pressure during 1991-1996, and an apparent increasing harvest intensity 22 since 1996 (Figure 21). Conversely, the distribution of harvest intensity for hunted bobcats in Zone 1 appeared relatively stable despite the increases in bag limit in 1994 and 1996 (Figure 22). These patterns may suggest that there are 1 or 2 relatively disjunct regions of Zone 2 that maintain relatively high harvest intensity, whereas the areas between exhibit lower harvest rates. As such, these patterns provide some support to the idea that the NLP bobcat population may exhibit source-sink dynamics and may need to be managed as 2 separate units. However additional demographic data on rate of population increase and dispersal would be needed before characterizing it as a true source-sink population (e.g., Pulliam 1988). We also mapped harvest intensity of bobcats during 1985-2002 with respect to landscape features (hydrology and road systems) to identify potential patterns. In the NLP, bobcat harvest pressure appears to be concentrated predominantly along streams and rivers, including headwater tributaries (Figure 23). This pattern is likely linked to the lowland forest and non-forested wetland habitats (including beaver sloughs) located along water courses. These habitats appear to be preferred habitat for bobcats in the NLP (Preuss and Gehring, unpublished data). A more ambiguous pattern was noted for the relation between bobcat harvest intensity and road density in the NLP, perhaps a function of the extensive road network that exists throughout much of the NLP (Figure 23). 23 a) 1985-1987 b) 1988-1990 Figure 21. Distribution of harvest intensity of bobcats, mapped to the scale of township, over the NLP during 1985-2002. Lighter colors indicate lower harvest intensity, whereas darker red colors represent greater numbers of bobcats harvested. 24 c) 1991-1993 d) 1994-1996 Figure 21. Distribution of harvest intensity of bobcats, mapped to the scale of township, over the NLP during 1985-2002. Lighter colors indicate lower harvest intensity, whereas darker red colors represent greater numbers of bobcats harvested. 25 e) 1997-1999 f) 2000-2002 Figure 21. Distribution of harvest intensity of bobcats, mapped to the scale of township, over the NLP during 1985-2002. Lighter colors indicate lower harvest intensity, whereas darker red colors represent greater numbers of bobcats harvested. 26 a) 1985-1987 b) 1988-1990 Figure 22. Distribution of harvest intensity of bobcats, mapped to the scale of township, over the UP during 1985-2002. Lighter colors indicate lower harvest intensity, whereas darker red colors represent greater numbers of bobcats harvested. Bag limit was increased to 2 bobcats per hunter and 3 bobcats per hunter in 1994 and 1996, respectively. 27 c) 1991-1993 d) 1994-1996 Figure 22. Distribution of harvest intensity of bobcats, mapped to the scale of township, over the UP during 1985-2002. Lighter colors indicate lower harvest intensity, whereas darker red colors represent greater numbers of bobcats harvested. Bag limit was increased to 2 bobcats per hunter and 3 bobcats per hunter in 1994 and 1996, respectively. 28 e) 1997-1999 f) 2000-2002 Figure 22. Distribution of harvest intensity of bobcats, mapped to the scale of township, over the UP during 1985-2002. Lighter colors indicate lower harvest intensity, whereas darker red colors represent greater numbers of bobcats harvested. Bag limit was increased to 2 bobcats per hunter and 3 bobcats per hunter in 1994 and 1996, respective 29 a) b) Figure 23. Distribution of harvested intensity of bobcats across the NLP during 1985-2002 mapped to the scale of township. Lighter colors indicate lower harvest intensity, whereas darker red colors represent greater numbers of bobcats harvested. a) Overlay of harvest intensity and hydrology illustrating the general pattern of greater number of bobcats harvested along streams and rivers and associated tributaries; b) Overlay of harvest intensity and primary and secondary roads illustrating no clear pattern of higher harvest in more or less roaded areas. 30 During 1998-2002, a greater number of bobcats was harvested on private land compared to public land in both the NLP ( x = 64 bobcats on public and x = 155 bobcats on private land, P<0.001) and UP ( x = 278 bobcats on public and x = 416 bobcats on private land, P = 0.02). However, harvest was not allocated proportional to the amount of public or private land area in either area. Contingency analysis indicated that a greater than expected proportion of bobcats was harvested on public land and a lower than expected proportion was harvested on private land in the NLP during 1998-2002 (χ2 = 78.21, 4df, P < 0.001). Similarly, a greater than expected proportion of bobcats was harvested on public land and a lower than expected proportion was harvested on private land in the UP during 1998-2002 (χ2 = 69.28, 4df, P < 0.001). These are likely underestimates because we classified public vs. private land to the scale of township and harvest locations were mapped to this same scale (Figure 24). Figure 24. Distribution of harvested bobcats across public (red) and private (white) townships in the NLP during 1998-2002. 31 Summary: Effective management of wildlife populations requires ecological data pertaining, in part, to abundance, distribution, and habitat use of species. Furthermore, effective management of a harvested species requires additional scientific information. Rolley et al. (2001) identified information needs for harvested bobcat populations to include: mandatory harvest registration, population indices, population models, user statistics, market-value surveys, and periodic habitat inventories. In part due to the difficulty in adequately surveying for low-density carnivores such as bobcats, Wisconsin has used a relatively conservative approach using restrictive harvest strategies (Rolley et al. 2001). Rolley (1987) recommended that at least 2 population-assessment methods be incorporated into a monitoring program in order to provide independent data on population trends. For example, scent-station surveys or winter-track surveys and sighting data (e.g., Archer’s index or sighting reports) are currently used by some states (Bluett et al. 2001). The Archer’s index or other sighting data from hunters could provide long-term trend data, but may not adequately assess annual population fluctuations (Bluett et al. 2001). Recently, the increased use of GIS modeling to identify and predict areas of quality habitat has allowed wildlife managers to focus their management and conservation efforts more efficiently. Models developed by Lovallo et al. (2001) resulted in the first harvest of bobcats in Pennsylvania in 30 years. Nielson and Woolf (2002) developed models that linked habitat and relative abundance to evaluate distribution and abundance of bobcats in Illinois. These models were used to assess bobcat status and contributed to the delisting of bobcats as a threatened species in Illinois (Woolf et al. 2002). Integrating demographic data (e.g., data determined via telemetry methods) and GIS spatial models can provide a powerful tool to focus management efforts. These tools are rare for solitary carnivores and region-specific models should be developed to more effectively direct bobcat management (Lovallo et al. 2001). Additional modeling efforts based on our CMU study of bobcats in Michigan should aid in making management recommendations. The number of bobcats harvested in the NLP increased from approximately 140 to 291 during 1985-2002. However, given the current data, it is difficult to determine if this increase was a function of the bobcat population increasing or an increase in the number of hunters. Statewide (Zone 1 and Zone 2), the increase in the number of bobcats harvested by hunters was correlated with the increase in the number of hunters. As such, the use of bobcat harvest numbers without an assessment of hunter effort does not appear to provide great insight into bobcat population trends. In particular, good estimates of the number of hunters and/or hunter effort for the NLP are needed to fully assess changes in that bobcat population. Estimates of the number of bobcat hunters in the NLP might be obtained via an increased sampling of this stratum. Additionally, a more extensive (and independent) assessment of the relative abundance of bobcats across the NLP is needed. Based on more localized data for the north-central counties of Crawford, Missaukee, and Roscommon counties, the bobcat population there appeared to be fairly stable during 1992-1996 (Earle et al. 2003a). However, harvest intensity has fluctuated in this region with relatively high harvest rates during 1988-1990 and then again during 1997-1999. Presumably, the number of hunters did not decrease in between these time periods and this 32 decrease in harvest may reflect a decline in the population. Additional data from other sites throughout the NLP would be needed to more fully assess the dynamics of the Zone 2 bobcat population. Additional population indices such as winter track counts, an Archer’s index, an estimate of the number of bobcats sighted by MDNR personnel, and/or the number of bobcats treed per day by hunters would provide valuable information on population trends at least over a longer time period. Based on estimates from surveys, if trapping and hunting occurred annually in the NLP, an estimated 620 to 3,000 bobcats would be harvested each year assuming a current bag limit of 1 bobcat per individual. An estimated 420 to 1,500 bobcats would be harvested each year assuming a current bag limit of 0.5 bobcats per individual. Without an accurate population index or other independent data for the entire NLP, it is difficult to evaluate the impact that this harvest figure would have on the NLP bobcat population. However, the high estimates likely exceed the total bobcat population in the NLP. The lower estimates might approximate 50% of the total NLP bobcat population assuming that our preliminary GIS modeling of potential size of the NLP bobcat population. Investigation into the possible source-sink dynamics and spatial patterning of bobcat harvest in the NLP should be pursued further because harvest intensity appears to be clustered into 2 distinct regions of the NLP and bobcat habitat distribution appears to demonstrate a similar patchiness. These investigations could provide insight into possible new zoning recommendations based on landscape-level features (e.g., hydrologic patterns) and/or spatial patterning of high-quality bobcat habitat (e.g., contiguous patches of lowland forest and non-forested wetland habitat). A possible opening of a trapping season in the NLP as a whole would not be warranted until additional population indices and hunter effort measures are developed and implemented to assess dynamics of the population before a change in harvest regulations. In particular, there remains a need to assess the impacts of predicted higher harvest levels if a trapping season would be added. A possible adaptive resource management model could be applied to bobcat management in a portion of the NLP, whereby trapping would be allowed for a 3 or 4-year period. However, at the end of such a trial period, an assessment would need to be conducted to determine the effects of a trapping season. In order to conduct an adequate assessment, a sound monitoring program should include: 1) harvest effort data; 2) bobcat population demographic data such as abundance, fecundity, survivorship, and dispersal; 3) multiple population indices to detect changes in bobcat abundance; and 4) habitat assessments. Ideally, the monitoring program would be established for several years: 1) before a change in harvest regulations; and 2) during the period of changes to the regulations. These monitoring efforts would be conducted at a site (or sites) where trapping was allowed and control sites where no trapping was allowed. Further, if warranted, these data could be useful in setting bobcat harvest quotas within the NLP. The current CMU bobcat study site might serve as a useful control site, assuming that trapping pressure elsewhere does not influence the study area (e.g., bobcat dispersal rates are not impacted and/or hunting pressure is not displaced away from a trapping zone and into the control study area). 33 Acknowledgements: We are grateful to Central Michigan University (CMU) and the Department of Biology for providing time and access to facilities to complete this review. We also thank CMU, Defenders of Wildlife, Wildlife Forever, and CITGO Petroleum Corporation for funding of our bobcat research. We thank the MDNR for providing trapping equipment, housing facilities and assistance with logistics for our research. 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