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Fuels Management and Habitat Restoration Activities Benefit Eastern Journal of Herpetology, Vol. 51, No. 4, 468–476, 2017 Copyright 2017 Society for the Study of Amphibians and Reptiles Fuels Management and Habitat Restoration Activities Benefit Eastern Hognose Snakes (Heterodon platirhinos) in a Disturbance-Dependent Ecosystem 1,2 3 1 3 MICHAEL E. AKRESH, DAVID I. KING, BRAD C. TIMM, AND ROBERT T. BROOKS 1Department of Environmental Conservation, University of Massachusetts Amherst, Amherst, Massachusetts USA 3U.S. Forest Service Northern Research Station, Amherst, Massachusetts USA ABSTRACT.—Eastern Hognose Snakes (Heterodon platirhinos) are considered a species of conservation concern in the northeast United States because of their association with rare and declining habitats such as pine barrens and shrublands. These are disturbance- dependent habitats that currently require management to persist. We studied Eastern Hognose Snakes on a pitch pine–scrub oak barren in western Massachusetts from 2008 to 2013 to describe patterns of space use, habitat selection, and survival of this species and to evaluate the effects of habitat restoration and fuels management. We monitored 12 snakes with radio telemetry during the months of May to October 2008–2010. We examined habitat use versus availability using paired logistic regression analyses in which availability was temporally and spatially explicit in relation to radio-tracked snakes’ previous use location and likely movements. We found that radio- tracked snakes significantly avoided closed-canopy forests and power line corridors, and instead primarily used heavily thinned pitch pine and scrub oak barrens. Individuals that used some closed-canopy forested habitat had significantly larger home ranges compared to snakes that used only managed early-successional habitat, congruent with ecological theory that habitat quality can affect home range size. We calculated a probability of 0.61 for adult survival during a 150-d active season (95% confidence interval [CI] = 0.22–0.85), similar to other reports of adult survival for this species. We conclude that fuels reduction and habitat restoration activities, primarily heavy thinning, are increasing the amount of preferred habitat available for this threatened species. Fire-dependent plant communities and the species that snakes often select managed and restored early-successional depend on them are threatened worldwide as the result of fire habitats (Waldron et al., 2008; Bailey et al., 2012); however, the suppression that inhibits the regeneration of fire-adapted effects of thinning and fuels control treatments on habitat species and encourages competition from fire-intolerant species selection, space use, and survival of Eastern Hognose Snakes are (Moreira et al., 2003; Nowacki and Abrams, 2008). In the eastern poorly understood. United States, pitch pine–scrub oak (PPSO; Pinus rigida–Quercus We studied Eastern Hognose Snakes at a PPSO barrens site in ilicifolia) barrens are an imperiled disturbance-dependent western Massachusetts to understand better the effects of community that has declined substantially; in some regions habitat restoration and fuels reduction, and to assist managers <10% of historical barrens remain (Motzkin et al., 1999). in the conservation of threatened snake populations. By radio- Additionally, when fire is suppressed in PPSO barrens, fuels tracking snakes that were able to use adjacent managed and accumulate to hazardous levels that pose extreme fire risk unmanaged areas in our study site, we assessed if snakes were (Duveneck and Patterson, 2007; Bried et al., 2014). Pitch pine– more likely to use managed, early-successional habitat than scrub oak barrens encompass substantial portions of some of the closed-canopy forest. We also examined home range sizes, daily most densely populated regions on the continent (U.S. Census movements, and survival rates of the snake population in our Bureau, 2009). Therefore, residential developments often are managed study area to compare with previously reported adjacent to, or embedded within, the pyrogenic plant commu- parameters of Eastern Hognose Snake populations in other nity (Gifford et al., 2010), such that the risk of human-caused study areas that had relatively more canopy cover and were not ignitions and uncontrollable wildfire is high. To reduce the risk specifically managed for fuels reduction (e.g., Plummer and of wildfire and restore habitat, management efforts have Mills, 2000; LaGory et al., 2009). Lastly, we determined if snakes’ included mowing, controlled burning that kills off fire-intoler- home ranges, movements, and survival in our study site varied ant species, and reducing pitch pine canopy cover (Duveneck among snakes that used, or did not use, closed-canopy forest. and Patterson, 2007). The distinctive fauna of PPSO barrens includes threatened invertebrates (Wagner et al., 2003), birds (King et al., 2011; MATERIALS AND METHODS Akresh and King, 2016), and herpetofauna (Stewart and Rossi, 1981). Eastern Hognose Snakes (Heterodon platirhinos)are Study Site.—We conducted our study between 2008 and 2013 considered a species of conservation concern over much of in the Montague Plains Wildlife Management Area, an ~600-ha 0 0 their range in northeastern United States and are closely site located in Franklin County, Massachusetts (42834 N, 72831 W; associated with disturbance-dependent PPSO barrens within datum WGS 84). Habitats within the study area included pitch this region (Michener and Lazell, 1989; Therres, 1999). Because pine forest, scrub oak barrens, power line corridors, and mixed Eastern Hognose Snakes prefer xeric, sandy soils with open- deciduous forest (King et al., 2011; Akresh, 2012). Pitch pine– canopy habitat and avoid heavily forested areas (Platt, 1969; scrub oak barrens are characterized by xeric, sandy soils, with a Plummer and Mills, 2000; LaGory et al., 2009), they could be plant community dependent on fire or management activities dependent on open-canopy, shrubland habitat created by forest (Motzkin et al., 1999). The Massachusetts Division of Fisheries management. In other disturbance-dependent ecosystems, and Wildlife has been conducting habitat restoration and fuels reduction in the PPSO barrens study site since 2000. Most scrub 2Corresponding Author. E-mail: [email protected] oak stands were mowed or burned once or twice since 2000, DOI: 10.1670/16-049 although stands in the northern part of the study site were not MANAGEMENT ACTIVITIES BENEFIT HETERODON PLATIRHINOS 469 treated. Scrub oak stands had low tree canopy cover (<22%) with and GPSmap 62; Garmin International, Inc., Olathe, Kansas, a dense shrub cover dominated by scrub oak and low-bush USA) at each snake’s capture and relocation sites. blueberry (Vaccinium angustifolium; King et al., 2011). Pitch pine Space Use Analysis.—We examined Eastern Hognose Snake and deciduous forests had a mostly closed canopy with sparse home range sizes with the use of the GPS locations of radio- understory. Since 2004, ~50% of the closed-canopy pitch pine tagged snakes. Statistical analyses were conducted with the use forest had been thinned to 40% canopy cover, creating a of the R Statistical Program version 3.2.1 (R Core Team, 2015) and drastically different habitat type (King et al., 2011; Akresh et al., the ArcGIS program version 10.1 (ESRI, 2012). To compare our 2015). The treated pitch pine stands in our study site had a sparse home range sizes with previously published studies (Plummer pitch pine canopy, with shrub and ground layers of oak and other and Mills, 2000; LaGory et al., 2009), we created 100% minimum tree saplings, low-bush blueberry, Pennsylvania sedge (Carex convex polygons (MCP) with the use of the adehabitatHR pensylvanica), and other ferns and forbs. Power line corridors package in R (Calenge, 2006). We also calculated home range were managed primarily by selective herbicide applications sizes with 95% fixed kernel utilization distributions (UD) with the every 4–5 yr, and were dominated by Spiraea species, regener- use of the ks package (Fieberg, 2014; Duong, 2016) and assessed ating pitch pine, and other shrubs and grasses. Some areas of core use area by determining 50% kernel UDs for each snake. urban development existed on the east side of the study site, Plug-in and reference bandwidths with unconstrained band- including the towns of Lake Pleasant and Millers Falls. Mowed width matrices provide reasonable kernel estimates when there fire breaks and roads existed within and along the borders of the are small numbers of telemetry fixes (Bauder et al., 2015). We study area and two water bodies, Lake Pleasant (21.4 ha) and used plug-in bandwidth estimates to reduce over-smoothing and Green Pond (6.5 ha), were situated on the east side of the study to place emphasis on areas of observed use (Bauder et al., 2015), site. while acknowledging that home range size estimates using Sampling.—We attempted to capture Eastern Hognose Snakes reference bandwidths were highly correlated with the plug-in by using funnel traps, cover boards, constrained searches, and estimates (r = 0.97 using 95% UDs). incidental encounters. In 2008, we installed funnel traps along 10- For our kernel estimates, we excluded telemetry fixes where m silt-fabric drift fences (Corn, 1994) at three points within each the snake was re-located in the same location over multiple days of the untreated pitch pine, treated pitch pine, and scrub oak (1 m of the previous location; LaGory et al., 2009; Timm et al., habitats. We checked traps every other day from May through 2014). Some
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