2012 SOUTHEASTERN NATURALIST 11(2):337–348 Influence of a Large-scale Removal of an Invasive Plant (Melaleuca quinquenervia) on Home-range Size and Habitat Selection by Female Florida Panthers (Puma concolor coryi) within Big Cypress National Preserve, Florida Paul Julian II1,2,*, Edwin M. Everham III1, and Martin B. Main3 Abstract - The control of invasive exotic plants is often deemed important for managing native wildlife, but surprisingly little research exists that evaluates benefits to wildlife, including species of conservation concern. Melaleuca quinquenervia (Melaleuca) is an invasive, non-native, broad-leaved tree that aggressively displaces native plant communities in south Florida. We used land-cover maps to document changes in plant communities and radio-telemetry data to compare habitat selection and mean home-range size of the endangered Puma concolor coryi (Florida Panther) within Big Cypress National Preserve (BCNP) during a Melaleuca removal project’s removal phase (1991–1997) and the ensuing maintenance and habitat recovery phase (1998– 2006). During the removal phase, Panthers incorporated areas infested by Melaleuca as components of their home range. Following >99.9% removal of Melaleuca, we documented pronounced increases in total cover of native upland forest (227%), wetland forest (211%), and prairie (54%) communities. During the habitat recovery phase, Panther habitat selection in the study area included significantly more upland forest within home-range core areas, and mean home-range size contracted by 16%. However, similar reductions in mean home-range size were not observed during the same time period for the regional population of radio-collared Panthers occupying contiguous conservation lands in south Florida. Although our findings are correlational and do not demonstrate cause and effect, the increase in native plant community cover, the increased use of native plant communities by Panthers, and the reduction in mean home-range size following the removal of Melaleuca are consistent with what would be expected if improvements in habitat quality reduced Panther home-range size requirements. Restoration of native plant communities and particularly native forest types, therefore, may have improved habitat quality for the Florida Panther and resulted in smaller home-range sizes and a potential increase in carrying capacity within BCNP. Introduction Puma concolor coryi Bangs (Florida Panther) is listed as endangered under the United States Endangered Species Act, and population declines have been attributed to a combination of factors (US Fish and Wildlife Service 2008). Loss 1Florida Gulf Coast University, Department of Marine and Ecological Sciences, 10501 FGCU Boulevard South, Ft. Myers, fl33965. 2Florida Department of Environmental Protection, Office of Ecosystem Projects, 3900 Commonwealth Boulevard, Tallahassee, fl32399. 3University of Florida, Department of Wildlife Ecology and Conservation, 2685 SR 29 Nth, Immokalee, fl34142. *Corresponding author - paul.julian@dep.state. fl.us , pjulian@eagle.fgcu.edu. 338 Southeastern Naturalist Vol. 11, No. 2 and fragmentation of habitat and unregulated killing over the past two centuries have reduced and isolated populations to extreme south Florida, where the population was estimated at 30–50 adults in 1990. More recent estimates indicate an increase in the Panther population to approximately 117 adults and semi-adults (McBride et al. 2008). In addition to habitat loss resulting from anthropogenic land-use changes, invasive non-native plants may also degrade habitat and reduce habitat availability (Rayamajhi et al. 2007, Turner et al. 1998, Wilcove et al. 1998). In south Florida, Melaleuca quinquenervia Blake (Melaleuca) is an invasive, non-native tree of concern due to dense stand formation, its ability to outcompete native plants, and its hypothesized impacts upon wildlife habitat (Rayamajhi et al. 2007). The effect of invasive plants, including Melaleuca, on Panther habitat use is unknown (US Fish and Wildlife Service 2008), but may be important because poor habitat quality has been associated with larger home-range sizes and reduced carrying capacity (Swihart et al. 1988). Additionally, the Florida Panther is a wide-ranging and territorial predator whose major limitations to recovery include the availability of suitable habitat (US Fish and Wildlife Service 2008). Studies documenting use of Melaleuca-infested areas by wildlife in Florida report that Melaleuca forests create dense canopies with an absent or sparse herbaceous understudy that provides limited food and habitat value for native wildlife, and have the potential to reduce indices of native species in Florida wetlands by as much as 80% (Bodle et al. 1994, Dray et al. 2006, O’Hare and Dalrymple 1997, Porazinska et al. 2007). Even where Melaleuca invasion of herbaceous wetlands increased structural diversity and species richness of avifauna, abundance of many species was lower in Melaleuca than in native wetland forest communities (O’Hare and Dalrymple 1997). Although the effects of Melaleuca displacing native vegetation is well documented, information documenting the response of native wildlife to Melaleuca infestation is scarce, and documentation of the response of Melaleuca removal and recovery of native plant communities is non-existent (Craven 2010). During 1991–2006, the Big Cypress National Preserve undertook a landscape- scale Melaleuca-removal project that included an active removal phase (1991–1997) followed by a period of maintenance removal and habitat recovery (1998–2006) at a cost of approximately $4.5 million (J. Burch, Big Cypress National Preserve, Ochopee, FL, unpubl. data). The objectives of this study were to document recovery of native plant communities and subsequent changes in habitat use patterns and mean home-range size of the Florida Panther following removal of Melaleuca at BCNP. Study Area The BCNP (Fig. 1) is located in extreme south Florida and covers 2940 km2 of flat, swampy areas that merge into coastal marsh and mangrove swamps. The ecology of this region is water dependent and is rich in biota (Klein et al. 1970). The BCNP encompasses almost half of a unique water-dependent ecosystem 2012 P. Julian II, E.M. Everham III, and M.B. Main 339 called the Big Cypress Swamp. Unlike the Everglades, it is still a relatively pristine wetland system, with little hydrologic modification. Nearly 80% of the rain normally falls during the six-month wet season of May through October and averages 135 cm per year (Schneider et al. 1996). Melaleuca was introduced into BCNP in the mid-1940s and spread unchecked, primarily by wind dispersal, until the establishment of the invasive plant control program within the preserve in 1984 (Laroche 1999). According to land-use maps produced by the South Florida Water Management District (SFWMD) in 1990, approximately 57.6% (1700.7 km2) of the preserve was infested with stands of Melaleuca. BCNP has since instituted an intensive invasive plant removal program (J. Burch, unpubl. data) targeting priority exotics such as Melaleuca, Lygodium spp. (old-world climbing fern), Casuarina equisetefolia L. (Australian Pine) , and Schinus terebinthifolius Raddi (Brazilian Pepper). Melaleuca removal peaked during the mid- to late- 1990s, and the species was reported to be eradicated from BCNP in 2006. Methods Habitat classification We grouped landscape cover into five habitat classifications based on landcover/ land-use data from the Florida Land Use and Cover Classification System (FLUCCS) for 1990, 1995, 2000, and 2004 (SFWMD 2010). The five landcover classifications were upland forest, wetland forest, Melaleuca, prairie, and anthropogenic land use. Reclassification of similar land-cover types (e.g., Taxodium spp. [cypress species] and cypress-mixed hardwoods) into five land-cover Figure 1. The study area at Big Cypress National Preserve, which is located in southwestern Florida, covers 2940 km2, and encompasses the core of the Florida Panther’s primary range. Primary range indicated by dark shaded grey, secondary range shown as light shaded grey (Kautz et al. 2006). 340 Southeastern Naturalist Vol. 11, No. 2 categories was based on function and form of these communities and deemed more ecologically meaningful for a wide-ranging species such as the Florida Panther. The upland forested habitat classification includes forests dominated by a mixture of hardwood trees, Pinus elliottii Engelm (Slash Pine), and Sabal palmetto Lodd. ex Schult. & Schult. f. (Cabbage Palm) with a Serenoa repens Small (Saw Palmetto) understory. Wetland forest habitat typically included species such as cypress and other hardwood species adapted to prolonged hydroperiods. Prairie habitat was dominated by a diverse herbaceous community of grasses and sedges, with Cladium jamaicense Cranz (Sawgrass) and Typha spp. (cattail species) dominating in areas with longer hydroperiods. Anthropogenic habitat delineated as pastures in private inholdings represented a minor component (less than 1.0% cover) of the landscape and were grouped as prairie habitat due to similar habitat structure. All prairie habitat within BCNP is seasonally influenced by hydrology, particularly during the dry season. Melaleuca cover was specifically classified in FLUCCS (FLUCCS Code 4240) and included Melaleuca infestation in both upland and wetland areas. Water classifications contained lakes, natural drainages, and canals, but for the purpose of analysis were combined with prairie due to proximity and low percent cover (less than 0.5% cover) in the study area. Anthropogenic land-use classification included roads, human-built structures, and non-pasture agriculture such as row crops and citrus, but this classification was excluded from analysis because this land-use type only occurred on the periphery of BCNP, was included in the home ranges of only two Panthers during two separate years (2000 and 2005), and was calculated to represent less than 1% of those Panther home ranges (0.9% and 0.7%, respectively). We evaluated changes in percent habitat cover following Melaleuca removal with a change analysis using the ArcGIS Proximity Analysis Union Tool. To quantify changes in percent habitat cover we used available land-cover data that bracketed the study (SFWMD 2010) and were from 1990 (intensive removal phase, defined as time point one) and 2004 (maintenance removal and habitat recovery phase, defined as time point two). Florida Panther habitat use and home range size We documented Panther habitat use and home-range size during 1991–2006 using radio telemetry data obtained from the Florida Fish and Wildlife Conservation Commission (FWC) in conjunction with land-use maps reclassified as described for the purpose of habitat analysis. Telemetry locations used in analyses were collected in a standardized manner from fixed-wing aircraft by FWC and National Park staff between the hours of 0600–1000, and telemetry error associated with Panther locations has been reported to range from 77–230 m (Belden et al. 1988, Dees et al. 2001, Janis and Clark 2002). We used telemetry locations from 13 individuals that included 12 female Florida Panthers, and one intentionally introduced female Puma concolor stanleyana Goldman (Texas Cougar; Johnson et al. 2010), which we also refer to as a Panther, during the period 1991–2006 to evaluate changes in habitat use and home-range size for Panthers within BCNP. We restricted our comparisons of 2012 P. Julian II, E.M. Everham III, and M.B. Main 341 changes in Panther home-range size during the Melaleuca removal and recovery phases to female Panthers living exclusively within the BCNP study area and did not include males because of small sample size (n = 1 during removal phase, n = 2 during recovery phase) and the lack of multiyear data on collared males during the recovery phase. Our comparison of mean home-range size for the regional Panther population during Melaleuca removal and recovery included both males and females and was used to evaluate whether the mean Panther home-range for the regional population changed over time in response to increasing Panther population density in south Florida (McBride et al. 2008). We included males in this analysis because they are an integral part of the home-range processes, with male home ranges often overlapping multiple female home ranges, and the minimum Panther count did not distinguish between sexes. Only Panthers (male and female) with a minimum of 69 locations were included in analyses. Minimum telemetry data requirements were established due to the natural breaks in the data and the sufficiency of the lower limit to adequately estimate Panther home ranges (i.e., greater than one telemetry point per week for 52 weeks.). Annual home ranges were determined by the 95% fixed kernel (FK) and 50% FK methods (Seaman and Powell 1996, Worton 1987) using ArcGIS and Hawth’s Tools (Beyer 2004, Environmental Science Research Institute 2009). The kernel-smoothing parameter was selected by using the reference band-width (href) method (Seaman and Powell 1996). Analyses We evaluated habitat use versus habitat availability with a Manly selection index (Manly et al. 2002). The Manly selection index produces a resource selection function (RSF) proportional to the probability of use and provides a maximum likelihood estimate based on the proportion of utilized to available habitat. Although the utility of RSFs in habitat-selection studies has been questioned (Keating and Cherry 2004), the response by Johnson et al. (2006) demonstrated that RSFs can provide valid information on habitat selection and that use versus availability studies are often the most appropriate design for understanding wildlife-habitat relationships, particularly for mobile species. For analysis, the chronological time line was divided into two periods. These included the intensive Melaleuca-removal phase (1991–1997), during which the majority of Melaleuca was removed from BCNP, and the maintenance removal and habitat recovery phase (1998–2006) (J. Burch, unpubl. data.). We used analysis of variance (ANOVA) to compare Manly selection indices for the four habitat classifications (upland forest, wetland forest, Melaleuca, and prairie) within the two defined time periods. A square root transformation was applied as Manly selection indices were non-normally distributed for individual habitat selection within home ranges (Zar 2008). To detect differences among means, we used Tukey-Kramer honestly significant difference (Tukey HSD). We compared mean home-range size between the Melaleuca removal and habitat recovery periods for 95% and 50% FK home range areas with the 342 Southeastern Naturalist Vol. 11, No. 2 non-parametric Wilcoxon/Kruskal-Wallis test. We evaluated the effect of population density on home-range size with linear regression using mean regional annual home-range size and annual minimum count data (McBride et al. 2008). Linear regression was also utilized to assess both regional and BCNP mean annual home ranges between 1991 and 2006. All statistical operations were performed with JMP® (Ver 7.0.4, SAS, Cary, NC). The critical level of significance was set at α = 0.05. Results Changes in habitat and Florida Panther habitat selection During the 15-year period of Melaleuca removal and control on BCNP, 1701 km2 of Melaleuca were cleared and >99% of Melaleuca was reported removed from the preserve (J. Burch, unpubl. data). The removal of Melaleuca enabled native plant communities to recover, and total cover of native upland forest, wetland forest, and prairie plant communities increased by nearly 500% between 1990 and 2004. Wetland forest and upland forest types had the greatest increases (211% and 227%, respectively), and total native prairie cover increased by 54%. Absolute changes in total cover were greatest for native wetland forest, which increased by 1295 km2 (41% of BCNP), followed by prairie 380 km2 (13% of BCNP), and upland forest increased cover by 123 km2 (3% of BCNP). Anthropogenic land cover represented <1% of total cover on BCNP during 1990 (26 km2), and that cover declined to less than half that amount by 2004 (11 km2). Habitat use by Panthers within their 95% FK home-range areas differed among the different habitat types during both the Melaleuca removal (F = 8.84, df = 3, P < 0.01) and maintenance removal and habitat recovery period (F = 121.52, df = 2, P < 0.01). During the period of intensive Melaleuca removal, Panther use of upland forest, wetland forest, and areas infested by Melaleuca did not differ statistically. Prairie was used significantly less than either upland Table 1. Habitat selection by female Florida Panthers (n = 13) for 95% and 50% fixed-kernal (FK) home-range areas as measured by mean Manly selection indices during intensive (1991–1997) and maintenance (1998–2006) Melaleuca removal periods. Statistical comparisons evaluated using Tukey-Kramer HSD tests. Intensive removal (1991–1997) Maintenance removal (1998–2006) Habitat Mean SE Tukey HSD Mean SE Tukey HSD 95% FK habitat selection Melaleuca 0.37 0.06 A B 0.00 0.00 - Prairie 0.20 0.04 B 0.15 0.01 C Upland forest 0.52 0.04 A 0.54 0.02 A Wetland forest 0.36 0.04 A 0.32 0.02 B 50% FK habitat selection Melaleuca 0.37 0.05 AB 0.00 0.00 - Prairie 0.26 0.04 B 0.26 0.03 B Upland forest 0.47 0.03 A 0.54 0.04 A Wetland forest 0.39 0.04 A 0.48 0.04 A 2012 P. Julian II, E.M. Everham III, and M.B. Main 343 or wetland forest, but was similar to the use of Melaleuca (Table 1). During the maintenance removal and habitat recovery period, habitat selection within 95% FK home-range areas differed significantly among habitat types, with upland forest being preferred over wetland forest, which was preferred over prairie habitat. Melaleuca, which was greatly reduced, was not selected by Panthers during the maintenance removal and habitat recovery phase (Table 1). We found similar patterns of habitat use within each time period for the 50% FK home-range core areas, except Melaleuca was a least preferred habitat type within Panther home-range core areas. Habitat use by Panthers within 50% FK core areas differed among the different habitat types during both the intensive removal period (F = 5.24, df = 3, P < 0.01) and the maintenance removal and habitat recovery period (F = 18.06, df = 2, P < 0.01). During the intensive removal period, habitat selection of upland forest and wetland forest were similar and significantly greater than prairie, and selection for Melaleuca was similar to prairie (Table 1). During the maintenance removal and habitat recovery period, habitat selection of upland forest and wetland forest were similar and signifi- cantly greater than prairie, and Melaleuca was not selected (Table 1). Florida Panther home-range size Annual mean home-range (95% FK) size for female Panthers living in the BCNP study area declined by 16% between the 1991–1997 period of intensive Melaleuca removal and the 1998–2006 maintenance removal and habitat recovery period, and this change was statistically significant (χ2 = 17.37, df = 1, P < 0.01). In absolute terms, the 95% FK home-range size for Panthers averaged 115.5 km2 ± 6.6 (n = 13) during the intensive Melaleuca-removal period and 99.5 km2 ± 4.3 (n = 37) during the maintenance removal and habitat recovery period. Mean core home-range size (50% FK) for female Panthers on BCNP did not differ significantly (χ2 = 3.09, df = 1, P = 0.08) between the intensive removal (26.1 km2 ± 2.2, n = 13) and maintenance removal and habitat recovery (24.7 km2 ± 1.5, n = 37) phases. Mean annual home-range size for the regional Panther population did not undergo the same reductions in size over time as were observed for the female Panthers on BCNP (BCNP: F=119.5, df = 14, P < 0.01, R2 = 0.46; Regional: F=1.57, df = 14, P = 0.23, R2 = 0.10; Fig. 2). Instead, the mean annual homerange size for the regional Panther population changed little during the study time frame. Although the regional Panther population increased during the time frame of this study (McBride et al. 2008), we found no significant relationship between annual Panther population size and mean annual home-range size for the regional population (F = 3.13, df = 17, P = 0.10, R2 = 0.16). Discussion The removal of Melaleuca infesting approximately 1701 km2 (57.6% of the land area) on BCNP resulted in the recovery of native plant communities on a landscape scale, with native plant community cover increasing by approximately 500%. The greatest increase in cover was observed for wetland forest, but large 344 Southeastern Naturalist Vol. 11, No. 2 increases in upland forest and prairie cover were also documented, all of which are used by native wildlife including important prey species for the Florida Panther such as Odocoileus virginianus Zimmermann (White-tailed Deer) and Sus scrofa L. (Feral Hog) (Maehr et al. 1990). Although there is no published data on the value of Melaleuca as foraging habitat to White-tailed Deer and Feral Hogs, herbaceous understory vegetation is typically sparse in stands of Melaleuca, and Melaleuca is reported to be herbivore adapted and to provide limited food and habitat value for native wildlife (Craven 2010). Consequently, Melaleuca control efforts on BCNP were successful in restoring native plant communities and, presumably, improving habitat for native wildlife, including the Florida Panther, for which these cover types have been described as critical habitat (Belden et al. 1988, Benson et al. 2008, Cox et al. 2006, Kautz et al. 2006). Following the intensive Melaleuca-removal phase and during the maintenance removal and habitat recovery phase on BCNP, changes were documented both in patterns of habitat use and mean home-range size among radio-collared female Panthers in the study area. Following Melaleuca removal, Panthers selected upland forest and wetland forest equally above all other cover types. Melaleuca was not selected for during the maintenance removal and habitat recovery phase although it still remained in the system. Interestingly, the changes observed in habitat use by Panthers on BCNP were only documented in the 95% FK homerange areas, not in the 50% FK core home-range areas. This result was because Figure 2. Florida Panther mean annual home-range size (95% fixed kernel) for radio collared female Florida Panthers on the Big Cypress National Preserve (BCNP) and for the entire south Florida regional Panther population (males and females) during the Melaleuca removal phase (1991–1997) and habitat recovery phase (1998–2006). The average number of radio collared Panthers per year for the BCNP and regional Panther populations during 1991–1997 and 1998–2006 were 1.9 and 23.9, and 4.1 and 33.4, respectively. 2012 P. Julian II, E.M. Everham III, and M.B. Main 345 upland forest and wetland forest were preferred habitat types in Panther core home-range areas during both the intensive Melaleuca-removal phase and the habitat recovery phase, whereas Melaleuca was a least preferred habitat type during both phases. Consequently, Panthers used native habitats to a greater extent following Melaleuca removal, which is both consistent with the habitat preferences demonstrated for core home-range areas and intuitive because native habitats were more available on the landscape and are thought to provide better habitat for important prey species. The reduction in mean home-range size observed for Panthers on BCNP following the intensive Melaleuca-removal period is a correlational result and does not demonstrate cause and effect as might be provided by an experimental design. Consequently, it is not possible to rule out other potential causes for the observed changes in Panther home-range size following Melaleuca removal on BCNP. Having acknowledged this possibility, circumstantial evidence suggests that improvements in habitat quality may have resulted in reduced home-range requirements for the female Panthers living in the study area. First, the 16% reduction observed in mean home-range size following the Melaleuca- removal phase was only evident for the 95% FK home-range areas, not the 50% FK core home-range areas. This finding seems logical because core homerange areas are assumed to represent the highest quality habitat within a home range and, consistent with this explanation, Panthers selected upland and wetland forest types significantly more than Melaleuca in core areas during the Melaleuca-removal phase. Consequently, there would be little reason to anticipate significant changes in core home-range sizes because Melaleuca removal had little effect on habitat selection within the Panther core home-range areas. Secondly, similar reductions in mean home-range size were not observed for the regional population of radio collared Panthers over the same time period (Fig. 2). This is an important finding because Panther populations have steadily increased since the genetic introgression of female pumas from Texas in 1995, which resulted in a steady increase in the regional Panther population during the study time frame (Hostetler et al 2010, McBride et al. 2008, Pimm et al. 2006). Some studies have suggested that home-range dynamics may also be a densitydependent phenomenon (Clutton-Brock et al. 1987, Saether 1997), which might explain the reduction in home-range size for the Panthers on BCNP. However, if increasing Panther density caused home-range size to decrease on BCNP, it seems reasonable to expect reductions in annual mean home-range size would also have occurred among Panthers in the regional population, but no reduction in home-range size in the regional population occurred (Fig. 2). Additionally, Florida Panther home-range expansion or contraction is not believed to be a density- dependent process at the current population size (D. Onorato, Florida Fish and Wildlife Conservation Commission, Naples, FL, upubl. data). Whereas mean annual home-range size for the regional Panther population changed little over the study time frame, a distinct reduction in mean annual home-range size for female Panthers in the study area occurred following the Melaleuca-removal phase. These findings are correlational but are consistent with what would be expected if 346 Southeastern Naturalist Vol. 11, No. 2 improvements in habitat quality reduced home-range size requirements (Fig. 2). Note that mean annual home-range size for both Panthers on BCNP and in the regional Panther population decreased dramatically during 1995. Although again correlational, changes in home-range size during 1995 may have been influenced by restricted Panther movements in response to extreme flood conditions in BCNP and the greater Everglades ecosystem that began with Tropical Storm Gordon (November 1994) and was exacerbated by the highest annual rainfall on record during 1995 (for the period 1970–2011; R. Sobczak, Big Cypress National Preserve, Ochopee, FL, unpubl. data.). Our results quantified an increase in native plant communities in response to removal of Melaleuca, including wetland forest and upland forest types reported as critical habitats for the Florida Panther. These habitat types are also important for native wildlife including White-tailed Deer, which has been reported to constitute 40–50% of the Panther’s diet (Maehr et al. 1990). Reduced homerange size has been reported as a response to increased habitat quality for other species (Braunisch and Suschant 2008, Whitaker et al. 2007), and is often correlated to time and energy costs involved in securing requisite resources (Powell 2000, Whitaker et al. 2007). Our results, although correlational, are consistent with these findings and suggest that the recovery of native plant communities on BCNP following removal of Melaleuca may have improved habitat quality for the Florida Panther, which resulted in smaller home-range requirements and increased carrying capacity. These results also illustrate the need to establish wildlife-monitoring programs at the initiation of exotic-plant control programs to more effectively document wildlife response to management actions. Acknowledgments We thank Big Cypress National Preserve for assistance with this project, including providing Melaleuca removal data. 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