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Myodes Gapperi) Canadian Journal of Zoology Influence of field technique, density, and sex on home range and overlap of the southern red-backed vole (Myodes gapperi) Journal: Canadian Journal of Zoology Manuscript ID cjz-2018-0338.R1 Manuscript Type: Article Date Submitted by the 14-Jun-2019 Author: Complete List of Authors: Tisell, Honora; University System of New Hampshire, Natural Resources and the Environment Degrassi, Allyson; Shenandoah University, Environment and Society DepartmentDraft Stephens, Ryan; University System of New Hampshire, Natural Resources and the Environment Rowe, Rebecca; University System of New Hampshire, Natural Resources and the Environment Is your manuscript invited for consideration in a Special Not applicable (regular submission) Issue?: core area, mark-recapture, myodes gapperi, radiotelemetry, Keyword: RODENTS;SHREWS < Taxon, southern red-backed vole, space use https://mc06.manuscriptcentral.com//cjz-pubs Page 1 of 35 Canadian Journal of Zoology Influence of field technique, density, and sex on home range and overlap of the southern red-backed vole (Myodes gapperi) Honora B. Tisell1, Allyson L. Degrassi2, Ryan B. Stephens1, Rebecca J. Rowe1 1 University of New Hampshire, Natural Resources and the Environment, 114 James Hall, 56 College Road, Durham, NH 03824, USA. 2 Shenandoah University, Environment and Society Department, 1460 University Drive, Winchester, VA 22601 Draft [email protected] [email protected] [email protected] [email protected] Correspondent: Honora B. Tisell; [email protected]; 707-266-3754 University of New Hampshire, Natural Resources and the Environment, 114 James Hall, 56 College Road, Durham, NH 03824, USA. 1 https://mc06.manuscriptcentral.com//cjz-pubs Canadian Journal of Zoology Page 2 of 35 Influence of field technique, density, and sex on home range and overlap of the southern red- backed vole (Myodes gapperi) Honora B. Tisell, Allyson L. Degrassi, Ryan B. Stephens, Rebecca J. Rowe Abstract Home range is shaped by an individual’s interactions with the environment and conspecifics, and both size and placement may vary in response to population fluctuations. The method used to collect locational data may also affect home range estimates. We examined the effect of density, sex, and field method on home range of southern red-backed voles (Myodes gapperi Vigors, 1830) inhabiting eastern hemlock forests (Tsuga canadensis L. Carrière). Twelve mark-recapture grids were used to census M. gapperi from 2014 to 2017. In 2017, individuals were radio collared. Home rangeDraft size, core area size, and shared space were calculated using kernel density estimators from both mark-recapture and radiotelemetry data. Density effects on home range and core area were analyzed and differences between sex and field method were compared. We found 1) density did not affect home range size, 2) male home range was larger than female home range, 3) females shared space more frequently and to a greater extent with males than other females, and 4) home range estimates were not significantly different between mark-recapture and radiotelemetry. Male home range was, however, larger under radiotelemetry, and may reflect a truncation effect when mark-recapture grid size is smaller than male home range. Keywords: core area, mark-recapture, Myodes gapperi, Northern Forests, radiotelemetry, rodent, Southern red-backed vole, space use 2 https://mc06.manuscriptcentral.com//cjz-pubs Page 3 of 35 Canadian Journal of Zoology Introduction An individual’s home range represents the space they use and reflects a number of important ecological processes and behaviors. The placement and size of the home range is an indicator of a species’ resource requirements (Burt 1943; Powell and Mitchell 2012; Spencer 2012) and is shaped by environmental factors, both abiotic conditions, such as temperature and water availability (Getz 1968), and biotic conditions, such as the distribution of food and habitat resources (Gore 1988; Börger et al. 2008). Intraspecific interactions, such as competition and sociality, can also lead to differences in home range size and placement between sexes or ages (Jorgensen 1968; Bondrup-Nielson and Karlsson 1985; Bondrup-Nielsen and Ims 1986). Changes in population density can affect the strength of intraspecific interactions, particularly for small mammals which often have markedDraft seasonal or inter-annual population fluctuations (Boonstra and Krebs 2012). For example, at high density, intraspecific competition is greater for resources, such as food and space, which often leads to smaller home ranges (Adams 2001; Efford et al. 2016). Because home ranges reflect an animal’s resource needs, movement, and social behavior (Jorgensen 1968) the study of home range is fundamental to understanding population dynamics and responses to changing environments (Spencer 2012). Furthermore, population density and home range (size and overlap) interact to influence ecological processes that are mediated through contact rates, including disease transmission and genetic structure (Kitchen et al. 2005; Sanchez and Hudgens 2015). Identifying an individual’s home range can be challenging because the area is not uniformly used. Areas in which an individual spends disproportionately more time are considered core areas (Samuel et al. 1985; Millspaugh et al. 2012a). These core areas often reflect nesting sites, resting areas, or areas of high resource abundance (Samuel et al. 1985; 3 https://mc06.manuscriptcentral.com//cjz-pubs Canadian Journal of Zoology Page 4 of 35 Millspaugh et al. 2006; Spencer 2012). Overall home range size is limited by the distance an individual can travel and readily return to their core area (Börger et al. 2008). Modelling home range and distinguishing core areas requires multiple spatial or locational observations from the same individual. The ability to differentiate between areas of high and low use may vary depending on the tracking method employed (Samuel et al. 1985). Mark-recapture and radiotelemetry are two common field methods used to gather locational data. Choice of method can depend on the research question and terrain of the study site (Millspaugh et al. 2012b; Fuller and Fuller 2012). An individual’s space use is strongly tied to their body size and the scaling relationships among body size, space use, and energetic requirements (Jetz 2004; Millspaugh et al. 2012a) often dictates the method used to determine home range. In general, because smaller bodied mammals moveDraft shorter distances than larger bodied mammals (Jetz 2004), both mark-recapture and radiotelemetry may be appropriate for measuring their home range. For small-bodied rodents, only a few studies have compared the two field methods for measuring home ranges (e.g. Jones and Sherman 1983, Microtus pennsylvanicus (Ord, 1815); Desy et al. 1989, Microtus ochrogaster (Wagner, 1842); Ribble et al. 2002, Peromyscus boylii (Baird, 1855) and P. truei (Shufeldt, 1885)). However, these studies used minimum convex polygon estimators for home range size and this technique cannot give intensity of use estimates (Harris et al. 1990) and therefore cannot calculate a core area. More recent approaches including utilization distribution (UD) kernel density estimators can calculate the intensity of use of an individual at any given point within their home range (Van Winkle 1975; Powell and Mitchell 2012) and thus can be used to identify core areas (Samuel et al. 1985). The southern red-backed vole (Myodes gapperi Vigors, 1830) is wide spread throughout Canada and the northern United States (Merritt 1981), and is typically associated with montane 4 https://mc06.manuscriptcentral.com//cjz-pubs Page 5 of 35 Canadian Journal of Zoology boreal forests (Miller and Getz 1972). Like many small mammals, it is short-lived, highly fecund and is known to have years of high and low population density (Boonstra and Krebs 2006; Sullivan et al. 2017). Females are thought to be territorial and exclude conspecific females from their defended home range (Perrin 1979b; Bondrup-Nielson and Karlsson 1985). Myodes gapperi is associated with eastern hemlock (Tsuga canadensis L. Carrière) in northern forests (Miller and Getz 1972; Yamasaki et al. 1999). These forests are currently threatened by the introduced hemlock woolly adelgid (Adelges tsugae Annand, 1928) (Orwig et al. 2002; Evans 2004), underscoring the need to document this species’ space use and better understand and manage for response to habitat disturbance (Ellison et al. 2005). In this study, we examined 1) whether M. gapperi density influenced home range or core area size and the amount of shared area, 2) whether there were differences in homeDraft range and core area size and overlap between sexes, and 3) whether home range and core area estimates differed by the field methods of mark-recapture and radiotelemetry. We predict that M. gapperi home range and core area will be density dependent because resources are limited, that there will be sex differences in home range and core area size and overlap because males and females have different reproductive strategies, and that the two field methods will result in different estimates of home range because mark- recapture is spatially constrained while radiotelemetry can follow an animal’s movement freely. Methods Study area Myodes gapperi were captured at the Bartlett Experimental Forest, White Mountain National Forest, New Hampshire (44° 3′ 7.2′′ N, 71° 17′ 25.1′′ W). Twelve mark-recapture grids were established in each of three forest
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