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Home Range and Movement of Blanding’s Turtles (Emydoidea blandingii) in New Hampshire
Robin J. Innes, Kimberly J. Babbitt, and John J. Kanter

Northeastern Naturalist, Volume 15, Issue 3 (2008): 431–444

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2008 NORTHEASTERN NATURALIST 15(3):431–444 Home Range and Movement of Blanding’s Turtles (Emydoidea blandingii) in New Hampshire Robin J. Innes1,3, Kimberly J. Babbitt1,*, and John J. Kanter2 Abstract - Emydoidea blandingii (Blanding’s Turtle) is a Species of Special Concern in New Hampshire, yet it has received little research attention. As part of a broader study to establish conservation and management guidelines for this species, we radiotagged 18 Blanding’s Turtles to determine home range and movement patterns within two study areas of southeastern and central New Hampshire from 2000–2002. Mean daily movement of female turtles peaked in June coincident with nesting, whereas movement of males peaked in August and September coincident with an increase in mating activity. Median adaptive kernel home range (HR) and core range (CR) estimates for turtles in central New Hampshire (HR = 12.5 ha, CR = 3.1 ha) were larger as compared to southeastern populations (HR = 3.7 ha, CR = 1.6 ha). This difference could not be readily explained by characteristics of the turtles, such as age, size, or sex. New Hampshire populations of Blanding’s Turtles exhibit intermediate home range sizes compared to other parts of the species distribution. Forty-five percent of turtles exhibited multimodal location distributions in core range estimates. Mating and estivating turtles were always found within their core range area, whereas turtles occasionally traveled outside of core ranges to seek suitable overwintering sites. Thirty-eight percent of turtles used the same overwintering habitat for 2 consecutive winters. Location-specific information regarding key nesting and overwintering areas may be important to the conservation of the species. Introduction Emydoidea blandingii (Holbrook) (Blanding’s Turtle) is a semi-aquatic species whose primary range is in the Great Lakes region and midwestern United States, with disjunct populations occurring in portions of Massachusetts, New Hampshire, Maine, New York, and Nova Scotia, Canada (Ernst et al. 1994). Blanding’s Turtle populations in New Hampshire have received little research attention. There is concern that Blanding’s Turtle populations in New Hampshire may exist in very low densities and that the species is declining due to a variety of factors including habitat loss and fragmentation, and increased road building and consequent increases in road kill. The range of the Blanding’s Turtle overlaps with the four counties in New Hampshire having the fastest growing human population (Society for the Protection of New Hampshire Forests 2005). The status of Blanding’s Turtles in New Hampshire is best described as poorly known. Its classification as a Species of Special Concern may 1Department of Natural Resources, University of New Hampshire, Durham, NH 03824. 2New Hampshire Fish and Game Department, Concord, NH 03301. 3Current address - Wildlife, Fish, and Conservation Biology Department, University of California, Davis, CA 95616. *Corresponding author - kbabbitt@cisunix.unh.edu. 432 Northeastern Naturalist Vol. 15, No. 3 underestimate the conservation and management needs of this species, as the Blanding’s Turtle is classified as Threatened (Massachusetts) or Endangered (Maine) in adjacent states where it has received greater research attention (e.g., Grgurovic and Sievert 2005, Joyal et al. 2000). Further, several lifehistory features, including delayed sexually maturity (Congdon and van Loben Sels 1991; Congdon et al. 1983, 1993; Gibbons 1968; Graham and Doyle 1977; MacCulloch and Weller 1988, Petokas 1986; Ross 1989), low recruitment caused by low fecundity, high juvenile and egg mortality, and high levels of egg and nest failure (Congdon et al. 1983, Herman et al. 1995, Linck and Moriarty 1997, Petokas 1986, Power 1989, Ross 1989, Ross and Anderson 1990, Standing et al. 2000) enhance the vulnerability of this species to the effects of habitat loss and degradation, and limit their ability to recover from high levels of additive mortality caused by anthropogenic factors. Further, cumulative evidence suggests that factors that increase adult mortality are the most significant threat to this species (Congdon and van Loben Sels 1991; Congdon et al. 1983, 1993; Doak et al. 1994; Heppell et al. 1996; Herman 1997; Iverson 1991). More than 358,000 people are projected to move into New Hampshire between 2000 and 2025 (Society for the Protection of New Hampshire Forests 2005), increasing the amount of suburban development, roads, and traffic, all of which threaten the future of this species. Understanding the home range and habitat use patterns are fundamental to guiding appropriate land management and conservation approaches. As part of a broader effort to determine the best conservation management strategies for the Blanding’s Turtle in New Hampshire, we conducted a trapping and radiotelemetric study to determine the home range, movement patterns, and habitat use of Blanding’s Turtles. We used the information to address the following objectives: 1) determine space use (i.e., home range) of Blanding’s Turtles throughout the year, and 2) evaluate how space use is influenced by turtle movements and activity (i.e., nesting, mating, estivation, and overwintering). Methods We studied Blanding’s Turtles during their active season (April–October) from 2000 to 2002. We examined the home range and movement patterns of Blanding’s Turtles from six sites (wetland-upland complexes) in two areas of New Hampshire separated by 60 km (Fig. 1). Three sites were located in the southeastern corner of New Hampshire in the towns of Newmarket and Lee within Strafford and Rockingham counties (hereafter Strafford). Three additional sites were located in central New Hampshire, in the towns of Weare and Dunbarton within Merrimack and Hillsborough counties (hereafter Hillsborough). Wetland habitat types were defined by geographic information system data layers provided by the New Hampshire GRANIT database. Wetland habitat types within the study sites included freshwater marsh, wet meadows, scrub-shrub and forested wetlands, permanent ponds, vernal pools, and riparian and lacustrine habitats. The upland matrix included mixed deciduous-coniferous forests, primarily deciduous woodlands, and 2008 R.J. Innes, K.J. Babbitt, and J.J. Kanter 433 scattered open lands, including agricultural fields and active and abandoned sand and gravel pits. Sites were located in both rural and suburban areas. We captured Blanding’s Turtles with baited, nylon hoop traps (2 m long x 0.8 m dia; 5-cm mesh), opportunistically by hand, and during road surveys to and from study sites. Hoop traps were set in wetlands from April to October and checked every 24 to 48 hours. The number of hoop traps placed in each wetland ranged from one to six depending on wetland size. We uniquely marked all captured Blanding’s Turtles by notching the marginal scutes of the carapace (Cagle 1939). We weighed turtles to the nearest 5 g, measured straight-line carapace and plastron width and length with calipers to the nearest 1 mm, and estimated the age of turtles by counting plastral humeral laminae (Condgon and van Loben Sels 1991, Germano and Bury 1998, Graham 1979, Innes et al. 2005). Sex was determined in adults by plastral concavity and distance of the cloacal opening from the carapacial margin (Ernst et al. 1994, Graham and Doyle 1977). Figure 1. Map of New Hampshire indicating the current known distribution of Blanding’s Turtles (Emydoidea blandingii) based on records from the Reptile and Amphibian Reporting Program (RAARP) at the Nongame and Endangered Species Program, New Hampshire Fish and Game Department (light gray shaded areas). Areas shaded in dark gray indicate study areas (Hillsborough County and Strafford County). 434 Northeastern Naturalist Vol. 15, No. 3 We fitted 18 adult Blanding’s Turtles with 11-g radio transmitters with whip antennas (Advanced Telemetry Systems, Model LTC-7PN), by attaching the transmitter package to the anterior portion of the carapace using epoxy putty as a base and clear waterproof epoxy as a coating. The transmitter package weighed about 66 g (n = 18, 5.1% of body weight, mean = 1294 g, range = 825 to 1725). We released turtles near the point of capture within 24 hours of capture. To minimize temporal autocorrelation, we did not locate individuals more than once in any 48-hour period; we located individuals approximately three times per week between 0700 and 1900 hours. Locations were determined by homing (Samuel and Fuller 1996), using a hand-held receiver (Communications Specialists, Inc., Orange, CA; Model: R-1000) and Yagi antenna. We recorded locations using a hand-held geographical positioning system unit (Model: Garmin 12, Garmin International, Inc., Olathe, KS) accurate to within 10 m. We conducted home range analyses on radio-tagged individuals when the following conditions were met: 1) the individual was determined to have the minimum number of sequential radio locations required to reach an asymptote in home range size (Kenward 2001), and 2) the radiotelemetric duration for the individual encompassed at least 5 months of activity, thus representing the active season of Blanding’s Turtle in New Hampshire. Several Blanding’s Turtle home ranges examined in preliminary analyses had more than one area of intensive use (i.e., multimodal). Therefore, we chose the adaptive kernel method (ADK) to estimate home range because it relies on probability densities and is a better technique for accurately determining home range size in animals exhibiting bimodal distributions (Aebischer et al. 1993, Worton 1989). We also used the minimum convex polygon method (MCP) to facilitate comparison with previous studies (Mohr 1947). Extended trips, such as those associated with nesting females, were excluded from home range estimates and evaluated separately (Rowe and Moll 1991). We calculated home range and average daily distance traveled using the CALHOME Home Range Analysis Program (Kie et al. 1996). For individuals tracked multiple years, we selected one year randomly for each individual for analyses. We estimated home ranges and core ranges (a.k.a., activity centers) using 95% and 70% of all radio locations, respectively, by omitting outlying locations (Anderson 1982; White and Garrott 1990; Worton 1989, 1995). The number of polygons comprising the core range was calculated in CALHOME using ADK. In addition, ADK utilized a grid of 50 x 50 cells overlaid onto the dataset and an optimum bandwidth (smoothing parameter), following the methods of Piepgras and Lang (2000). A grid of 50 x 50 cells is the maximum allowable by CALHOME, and results in the smoothest fit to the data (Kie et al. 1996). For radio-tagged turtles having a small home range (less than 3.5 ha), it was not possible to use a grid of 50 x 50 cells. In these cases, CALHOME selected the optimum grid cell size and bandwidth to achieve the lowest least-squares cross-validation score (Kie et al. 1996). To determine if movement habits changed over the tracking 2008 R.J. Innes, K.J. Babbitt, and J.J. Kanter 435 period, we computed distances between successive locations and estimated mean distance traveled per day during a month, assuming a constant rate of movement in a straight line between locations. We analyzed turtle movements in the context of monthly activities based on: 1) change in overall activity (i.e., decrease in basking, nesting; increase in estivating, mating); 2) change in average movement distance; and 3) apparent shift in habitat-type used (i.e., to overwintering habitat). Gross habitat shifts within hydrologically connected wetlands (inter-habitat shifts) and between isolated wetlands (inter-wetland shifts requiring overland movement) were examined. We identified shifts by a change in the most dominant habitat used. Shifts were generally easy to recognize, as daily movements within a habitat type were generally small, whereas inter-habitat shifts and inter-wetland shifts were often associated with larger and often sudden movements (i.e., large short-term movement preceded and followed by little movement). Small movements alongside transitional zones between wetland habitat types were sometimes apparent. Movements associated with nesting, estivation, mating, and overwintering were examined for differences between these activity sites and core range areas. Turtles were assumed to be inactive and potentially estivating if 3 or more consecutive radio-locations (5 or more days) occurred within the same location. Estivation as defined here is a period of inactivity coinciding with decreasing water availability in the late summer and fall, and is a behavior that has been observed to occur terrestrially (e.g., Rowe and Moll 1991) and aquatically (e.g., Ross and Anderson 1990) in Blanding’s Turtles. Overwintering movements were associated with a lull in activity at the end of the active season in October and November. Results We captured 31 Blanding’s Turtles in Strafford (17 males, 11 females, 3 juveniles), and 9 Blanding’s Turtles (6 females, 3 males) in Hillsborough throughout the course of the study. We radio-located 18 turtles during the active period from 2000 to 2002: 1 juvenile (PL = 158 mm, 10 years old), 6 adult males, and 3 adult females in Strafford, and 3 adult males and 5 adult females in Hillsborough. Of these adults, 11 radio-tagged turtles were included in analyses. Three radio-tagged turtles were located at Hillsborough at one study site, and 8 radio-tagged turtles were located at Strafford at 3 study sites. The juvenile radio-tagged turtle in Strafford was retained for comparative purposes since the home range ecology of Blanding’s Turtle in this age class is generally lacking (Piepgras and Lang 2000). The number of locations obtained for radio-tagged animals used in analyses averaged 52 (range = 37–65 locations) and spanned 5 to 7 months of activity (April to November). At Strafford, median home range and core range estimates of females were generally smaller and less variable than those of males (Table 1). The home range estimate (11.6 ha ADK) and core range estimate (4.6 ha ADK) of the juvenile was intermediate to that of other radio-tagged turtles in Strafford. The median home range of female turtles in Hillsborough was nearly 5 times larger 436 Northeastern Naturalist Vol. 15, No. 3 than that of female turtles in Strafford. Similarly, median core range size of females tended to be larger in Hillsborough compared to Strafford. Patterns of location distribution and concentration also varied between study areas and between sexes. For example, of 3 turtle core ranges examined in Hillsborough, all had more than one area of intense use (i.e., bimodal). However, of 8 Blanding’s Turtle core ranges examined in Strafford, 2 were bimodal and 6 were unimodal; both males and females exhibited multimodal location distributions. We compared the average distance traveled per day during each month to characterize movement patterns. Overall, the average distance traveled per day for females peaked in June followed by a slow decline in average movement distance until hibernation (Table 2). Male average movement distance slowly increased to a peak in August. High average movement distances for males was maintained into September, but rapidly decreased thereafter. Average movement distance patterns were similar for females at Strafford and Hillsborough. Patterns in average movement distance may or may not be explained by home range size. For example, a male Blanding’s Turtle at Strafford with consistently large average movement distances (range = 41.5–44.6 m/d, Jul–Sep) also had the largest home range area (24.8 ha Table 1. Median (interquartile range) home range (95% of radio locations) and core range (70% of radio locations) area (ha) estimated by minimum convex polygon (MCP) and adaptive kernel (ADK) methods for male, female, and juvenile Blanding’s Turtles in southeastern (Strafford) and central (Hillsborough) New Hampshire during 2000–2002. Combined estimate includes adult male and female turtles only. Study area MCP ADK and turtle class n Home range Core range Home range Core range Strafford Female 3 1.5 (0.2–3.0) 0.7 (0.09–1.6) 2.7 (0.4–4.7) 1.0 (0.2–2.7) Male 4 3.7 (2.0–25.6) 2.8 (1.1–7.1) 8.7 (3.4–22.0) 2.7 (1.3–9.2) Juvenile 1 3.2 2.0 11.6 4.6 Hillsborough Female 3 6.8 (3.7–8.7) 2.4 (2.1–3.2) 12.5 (5.0–14.6) 3.1 (2.1–3.2) Combined 10 3.3 (1.9–7.3) 1.9 (0.9–3.5) 4.9 (3.2–14.0) 2.4 (1.2–3.4) Table 2. Mean rate of travel (m/day/month), assuming a constant straight-line movement between successive locations for male (M) and female (F) Blanding’s Turtles in southeastern (Strafford) and central (Hillsborough) New Hampshire during 2000–2002. Strafford Hillsborough F M F Month m/day SD n m/day SD n m/day SD n May 22.5 24.1 2 15.3 17.5 2 21.7 3.1 3 June 49.8 35.8 3 28.1 7.3 3 142.9 72.0 3 July 26.4 25.3 3 28.5 15.0 5 74.0 52.8 3 August 21.6 12.8 3 38.3 15.3 4 30.0 11.4 3 September 11.4 6.6 3 32.6 23.3 4 16.3 3.7 3 October 10.3 3.9 3 15.9 12.3 4 8.2 7.0 3 November 4.2 2.1 3 11.3 14.6 4 2008 R.J. Innes, K.J. Babbitt, and J.J. Kanter 437 ADK) throughout the active season in 2001. Conversely, a male turtle with a relatively small home range area (3.7 ha ADK) had equally large average movement distances during that time (range: 41.9–58.1 m/d, Jul–Sep). Radio-tagged turtles were present within overwintering habitat into April. Turtles began transitioning into surrounding habitats in early May. From early May to early July, turtles moved frequently among habitat types, often exhibiting large, dramatic inter-habitat movements and exploratory forays. Where vernal pools were available, radio-tagged turtles often traveled from hibernation sites overland to vernal pools upon emergence in the spring. The movements of one male turtle suggest that he traveled through the upland during inter-wetland movements on at least 7 occasions in the spring of 2001 and 2002. Inter-habitat or inter-wetland movements were generally followed by longer periods of residency before an animal attempted another inter-habitat or inter-wetland movement, although periods of residency varied widely and periods of residency were not apparently correlated with early or late-season activities. Average minimum residency period within a habitat was approximately 7 weeks (range = 2–20 weeks). Nesting forays, defined here as movements to and from nesting areas, occurred between June 6 and July 3 and lasted a minimum of 4 to 16 days (n = 3). Movement distances from residential wetlands (i.e., wetlands occupied previous to initiation of nesting activity) to nesting areas ranged from 1.0 to 1.2 km. Vernal pools acted as important staging areas for nesting turtles, as vernal pools were used during all documented nesting forays in Stafford and Hillsborough. Nesting areas were generally adjacent to or within 0.2 km of staging areas. Nesting females always returned to residential wetlands upon completion of nesting activity. For a nesting female in Hillsborough, the same staging and nesting areas were used in 2000 and 2001. In addition, in 2000, this female shared these areas with another nesting female. Following nesting, activity of female turtles decreased and movements became less dramatic, as suggested by the decrease in average movement distance. This shift coincided with a reduction in the amount and variety of wetland habitat types available due to low water levels. Nonetheless, male movements continued to increase. Also at this time, estivation and mating were observed more frequently. We recorded 10 mating observations during the course of the study; 70% of these occurred during August through October (Jenkins and Babbitt 2003). Male and female turtles always mated within their core range. Likewise, radio-tagged turtles always estivated within their core range. Some radio-tagged turtles did not estivate; however, other individuals estivated on several occasions. Thirty-nine percent of radio-tagged turtles (4 radio-tagged turtles in Strafford [3 F, 1 M], in addition to the juvenile, and 2 in Hillsborough [1 F, 1 M]) estivated during July to October. Periods of inactivity or estivation lasted 5 to 48 days. Although inactivity or estivation occurred in July and August, it was more likely to occur just prior to overwintering in September and October. In these cases, estivation was likely to occur within or near to overwintering habitat. For example, a female in Strafford made infrequent 438 Northeastern Naturalist Vol. 15, No. 3 movements within a 30-m diameter pond on several occasions in September and in November prior to hibernating there. Turtles mostly estivated in permanent ponds, except on one occasion when a turtle estivated in upland habitat. Starting in August, and more frequently in September, turtles began to transition into overwintering habitat, and movements became exclusive to one habitat or two adjacent habitats. Turtles ceased movements and entered hibernation from late October to mid-November. The radio-tagged juvenile began overwintering earlier (26 October 2001) than the adult radio-tagged turtles in Strafford (n = 7). Males and females entered dormancy at similar times. Generally, radio-tagged turtles overwintered within their core range (84%); however, a male and a female in Strafford overwintered outside of their core range in 2001 and 2000, respectively, and two females in Hillsborough overwintered outside of their core range in 2001. Fidelity to overwintering habitat between years was frequently observed. Thirty-eight percent of turtles used the same overwintering habitat for 2 consecutive winters. Fidelity to hibernacula was recorded for one male Blanding’s Turtle in 2000–2001 and 2001–2002. Two female turtles in Hillsborough were found to enter hibernation on or before 1 November and 11 October, 2000, and both were found within their hibernacula until 18 April the following spring, although we did not monitor these turtles throughout the winter. Discussion Accurate home range and movement pattern information is critical for determining Blanding’s Turtle habitat requirements and for guiding land management and conservation approaches that protect critical resources, such as key nesting and foraging areas, estivation refugia, and hibernacula. Home range estimates for Blanding’s Turtles reported in previous studies vary widely (0.56– 63.0 ha MCP; reviewed in Grgurovic and Sievert 2005). Although home ranges in this study were larger in Hillsborough than in Strafford, home range sizes in these two areas were more similar to each other than to those recorded from most other areas of the species’ distribution. Home range sizes recorded in this study are larger than those recorded in many other populations (Maine [Joyal 1996], Wisconsin [Ross and Anderson 1990], Illinois [Rowe and Moll 1991]), but smaller than those reported in Massachusetts (Grgurovic and Sievert 2005) and Minnesota (Hamernick 2000, Piepgras and Lang 2000). Overall, comparison of home range estimates among studies suggests that New Hampshire Blanding’s Turtles have intermediate home range sizes. There are many factors that may contribute to the large variation in home range estimates among studies. In addition to potential methodological differences, factors such as age, size, sex, population density, and year-to-year fluctuations in climatic conditions can influence home range estimations (Brown and Brooks 1993, Brown et al. 1994, Burke and Gibbons 1995, Morreale et al. 1984, Schubauer et al. 1990, Stickel 1989, Tuberville et al. 1996). In addition, locality differences related to habitat composition and availability, and resource distribution, particularly refugia and hibernacula, 2008 R.J. Innes, K.J. Babbitt, and J.J. Kanter 439 also likely affect home range estimates (Carter et al. 1999, Hamernick 2000, Pettit et al. 1995, Piepgras and Lang 2000). In our study, differences in home range size between study areas could not be readily explained by differences in age, body mass, or body size between study areas (Innes et al. 2005). Further study is needed to elucidate the mechanisms underlying home range size and movement patterns of this species in New Hampshire. We found differences between the sexes in activity and movement patterns, with female activity peaking in June coincident with nesting activity, and male activity peaking in August and September. Turtles moved less in October and November as they shifted to overwintering sites. Increased activity of males in August and September in this study may be explained by an increase in mating activity observed at this time (Jenkins and Babbitt 2003), a pattern also observed in Nova Scotia, Canada, populations (Herman et al. 1995). Kofron and Schreiber (1985) found that feeding patterns of Blanding’s Turtles peaked in June and August, but did not detect a difference in feeding patterns between males and females. In general, studies concerned with this species’ movement patterns have found that male Blanding’s Turtles make more long-distance movements than females in early to late spring, whereas during the nesting season, females make more long-distance movements than males (Herman et al. 1995, Ross and Anderson 1990, Rowe and Moll 1991). Average movement distance was generally not a good predictor of home range area, with many turtles exhibiting relatively large average movement distances (>20 m/d) and small home range areas (<5 ha ADK). This result suggests that, for these turtles, movements of greater distance are of short duration, likely between core areas, with more frequent smaller movements occurring within core areas. We found that 45% of Blanding’s Turtle core ranges had bimodal location distributions. It is not uncommon for Blanding’s Turtles to exhibit disproportionate use of parts of home range areas during certain times of the year (Piepgras and Lang 2000, Rowe and Moll 1991). Rowe and Moll (1991) suggest that location of core range area and duration of core range occupancy may be indicators of increased habitat quality in these areas; however, support for this correlation was lacking, and turtles in this study may have been responding to the other factors. The importance of examining the distribution and placement of core range areas is further emphasized in our study by the frequent occurrence of overwintering, mating, and estivation activities within core ranges. Nonetheless, on several occasions, we found that hibernacula were located outside of a turtle’s core range. Other studies have found that Blanding’s Turtles overwinter in summer activity centers (Ross and Anderson 1990), but many report that Blanding’s Turtles travel great distances to arrive at overwintering sites (Hall and Cuthbert 2000, Piepgras 1998, Ross and Anderson 1990, Rowe and Moll 1991). These findings emphasize the importance of including late season activities in home range and movement estimates in this species. Overwintering and estivation sites are critical components of Blanding’s Turtle habitat requirements, as prolonged periods of inactivity may be spent 440 Northeastern Naturalist Vol. 15, No. 3 in a single location. In our study, Blanding’s Turtles reduced activity or estivated at any one location for periods of 5–48 days, and overwintered in any one location for as long as 6 months. We found that Blanding’s Turtles shifted into overwintering habitat beginning in mid-July, with the majority arriving from mid-August to mid-September, and entered winter dormancy from late October to mid- to late November. We also found that estivation frequently graded into hibernation on several occasions, a behavior frequently observed in turtles (Carr 1952). Other studies found that Blanding’s Turtles arrived at overwintering areas and entered dormancy at similar times (Hall and Cuthbert 2000, Herman et al. 1995, Rowe and Moll 1991). However, turtles may be active throughout the winter in some locations (Conant 1938, Kofron and Schreiber 1985). Herman et al. (1995) found that males generally entered hibernation later than females. This pattern was not apparent in our study. Patterns in estivation activity showed greater variation among studies than overwintering activities. We observed estivation on 9 occasions. Although unusual, Piepgras and Lang (2000) did not observe any periods of estivation in 47 radio-tagged turtles throughout the duration of a 16-month study. Ross and Anderson (1990) and Rowe and Moll (1991) reported terrestrial estivation on several occasions, with most estivation activity occurring between late July and August. Mechanisms underlying estivation activity are not generally understood. Terrestrial estivation has been attributed to cool water temperatures (Rowe and Moll 1991). Rowe (1987, as referenced in Ross and Anderson 1990) did not find a correlation between aquatic estivation and water temperature. Future studies should examine specific habitat features associated with overwintering and estivation activity since availability and location of refugia and hibernacula also likely affect movement and home range sizes. In addition to estivation and overwintering sites, particular emphasis should be placed on the location and habitat features associated with nesting areas. Blanding’s Turtle nesting behavior and ecology, as well as clutch and hatchling demography, are well documented. However, location-specific information regarding nesting activity and key nesting areas is important to the conservation of the species because nest-site fidelity is common (Congdon et al 1983; Joyal et al. 2000; Standing et al. 1999, 2000), and nests are generally found closer to bodies of water other than the maternal female’s residential wetland (Congdon and Rosen 1983, Piepgras and Lang 2000, Rowe and Moll 1991). In our study, one female was found to use the same upland pathway for nesting for 2 consecutive years. In addition, another female was found to use a similar pathway as this radio-tagged turtle. Nesting females spent long periods of time outside of home range areas (1–3 weeks) and used a variety of habitats not available within home range areas (R.J. Innes and K.J. Babbitt, unpubl. data). For example, all 3 nesting females in this study were found in small wetlands (i.e., staging areas) near to or adjacent to nesting areas both prior to and immediately after nesting. Use of staging areas by females during nesting forays has been reported elsewhere as well (Congdon et al. 2000). Females in that study always returned to their 2008 R.J. Innes, K.J. Babbitt, and J.J. Kanter 441 areas of residence immediately following a nesting excursion. Joyal et al. (2000) is the only published account that found a female Blanding’s Turtle who did not return to her residential wetland after nesting. Knowledge of all aspects of reproduction and life history will be critical in determining the best conservation and monitoring strategies for this species (Dodd 1997, Pedrono et al. 2001, Tinkle et al. 1981). Information on a larger sample of Blanding’s Turtles, especially nesting females, would be beneficial for conservation efforts aimed at all aspects of Blanding’s Turtle ecology. Identifying important movement corridors, for example, where more than one turtle share the same pathway for nesting year after year, may be important when considering conservation approaches in areas subject to fragmentation. In addition, identifying wetland habitats that are important overwintering and estivating areas is an important component of effective habitat management for the Blanding’s Turtle. Acknowledgments We thank E. Snyder of UNH, Cooperative Extension for her continued, enthusiastic support. We thank D. Carroll for providing data and sharing his expertise and vast knowledge of Blanding’s Turtles in New Hampshire. We recognize the New Hampshire GRANIT database for providing the data layers used in the analyses. The following people made notable contributions to this study by assisting with data collection and turtle captures: M. Baber, A. Beaulieu, S. Callaghan, D. DeGraaf, M. Hinderliter, M. Libby, A. Shutt, E. Snyder and T. Tarr of the University of New Hampshire; C. 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