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Habitat composition and availability can affect important life-history traits such as survival, reproduction, and consequently, long-term fitness for ungulate species. For example, management strategies that result in coniferous forest regeneration resulted in small amounts of year-round forage and negatively influenced adult female Sitka Black-tailed Deer (Odocoileus hemionus sitkensis; Farmer and others 2006) survival. Conversely, increasing habitat-management intensity positively influenced Mule Deer (O. hemionus; Bergman and others 2014) fawn survival; fawn survival also increased with summer and fall forage quality (Hurley and others 2017). Female Pronghorn (Antilocapra americana) with fawns also likely minimized fawn predation by selecting for dense cover (Christie and others 2017). Habitat composition and arrangement can even be used to predict movement patterns of White-tailed Deer (O. virginianus; Felix and others 2007). Consequently, understanding habitat availability on the landscape is important, as it influences individual movement and survival.

Although a variety of habitat types may be available to ungulate populations, individuals mostly select habitat types associated with forage and cover (Hobbs and Hanley 1990). Species such as Roe Deer (Capreolus capreolus) and Wild Pigs (Sus scrofa) used habitat types associated with quality nutrition more frequently than expected (Guangshun and others 2006), whereas Elk (Cervus elaphus) selected areas of high-quality cover more often than expected (Beck and others 2006). Elk also selected parturition sites with increased cover habitat (Lehman and others 2016). Therefore, managers should determine if adequate forage and cover can be found on the landscape and, if limited, implement management strategies to increase their availability.

Our goal was to quantify variation in habitat types associated with capture locations of newborn fawns compared to the surrounding area. We assessed these relationships within the 1st month of life of fawns because this is the time period when fawns display a hiding strategy to avoid predation and are dependent upon their mother (Lent 1974; Carl and Robbins 1988); thus, hiding-cover habitats are critical to the fawn. We predicted hiding cover such as grasslands, shrubby areas, and wetlands would be more prevalent at capture locations than in the surrounding area.


Study Area

We assessed whether habitat prevalence at capture locations of free-ranging White-tailed Deer fawns differed from the surrounding area using data collected from 13 counties, comprising 9 study sites across the Northern Great Plains. Counties included: Brookings, Edmunds, and Perkins, South Dakota; Dunn, Grant, Walsh, Grand Forks, Burleigh, Kidder, and Sheridan, North Dakota; and Redwood, Lincoln, and Pipestone, Minnesota (Fig. 1). All counties were located within 5 Level III Ecoregions (Omernik 1987; Bryce and others 1998): Northwestern Glaciated Plains (Burleigh, Kidder, and Sheridan counties, North Dakota), Lake Agassiz Plain (Walsh and Grand Forks counties, North Dakota), Northwestern Great Plains (Grant and Dunn counties, North Dakota and Perkins County, South Dakota), Northern Glaciated Plains (Edmunds and Brookings counties, South Dakota and Lincoln County, Minnesota), and Western Corn Belt Plains (Pipestone and Redwood counties, Minnesota).

Considering the extent of our study sites, weather and vegetation communities varied across this landscape. For example, 30-y (1981-2010) mean annual precipitation ranged from 41.2 (Grant County, North Dakota) to 72.9 cm (Redwood County, Minnesota), and 30-y mean temperatures ranged from a winter low of -15.1[degrees]C (Grant County, North Dakota) to a summer high of 30.3[degrees]C (Perkins County, South Dakota; North Dakota Office of Climatology 2016). Vegetation types were generally classified as Northern Wheatgrass-Needlegrass Plains, Northern Mixed Grass Prairie, and Tallgrass Prairie (Johnson and Larson 2007). Specific vegetation types have been described in detail (Grovenburg and others 2011a; Schaffer 2013; Sternhagen 2015; Moratz and others 2018).

Data Collection

We used postpartum behavior of reproductively active females as an indicator of the presence of fawns (Downing and McGinnes 1969; White and others 1972; Huegel and others 1985) and used vaginal implant transmitters (VITs; Advanced Telemetry Systems, Inc., Isanti, MN, USA) to assist in fawn capture (Swanson and others 2008). We captured fawns by hand or net after locating them. We wore latex gloves and stored all radio-collars and other equipment in natural vegetation to minimize scent transfer. We fitted fawns with expandable breakaway radio-collars (Advanced Telemetry Systems, Isanti, MN, or Telonics Inc., Mesa, AZ) to record mortality events. Fawn-capture methods were generally similar among study sites. For additional information regarding capture methods see Grovenburg and others (2011a), Grovenburg and others (2012a), Schaffer (2013), and Moratz and others (2018). We followed the American Society of Mammalogists guidelines for mammal care and use (Sikes 2016), and the South Dakota State University Institutional Animal Care and Use Committee approved all handling protocols (Approval numbers: 00-A038, 02-A037, 02-A043, 04-A009, 10-006E, 13-091A).

We created 2 circular buffers, one representing habitat associated with the fawn at its capture location, and one representing habitat associated with the surrounding area. We used a 352.3-m circular buffer representing a 39-ha mean core (50%) summer home range around each fawn-capture location (Kie and others 2002) to quantify the habitat found at the capture site. Grovenburg and others (2012a) reported this home range for White-tailed Deer fawns located in north-central South Dakota. We then calculated circular buffers representing maternal 95% summer home ranges reported in previous literature to quantify habitat in the area surrounding the capture location. For example, we used an 1802.3-m circular buffer around a fawn's capture location, which represented a 9.2-[km.sup.2] maternal home range associated with fawns caught in Edmunds County, South Dakota (Grovenburg and others 2009). However, maternal home ranges displayed variation, as they ranged from 2.3 [km.sup.2] in our southwestern Minnesota study area (Lincoln, Pipestone, and Redwood counties, Brinkman and others 2005) to 9.2 [km.sup.2] in Edmunds County, South Dakota (Grovenburg and others 2009; Table 1). We used circular buffers derived from Edmunds County, South Dakota (Grovenburg and others 2009) as a reference and scaled other circular buffers at fawn capture locations accordingly (Table 1). For example, the circular buffer used to quantify habitat in the surrounding area for Perkins County, South Dakota was about 42% smaller than those used for Edmunds County, South Dakota.

We quantified the percent of open water (because of its importance to lactating female ungulates; Jacques and others 2015), developed land (such as farmsteads, roads), forest, shrub, grassland, cropland, wetland, and pasture at each fawn capture location and its surrounding area from the 2011 National Land Cover Database (Homer and others 2015). After quantifying percent habitat type in each circular buffer, we adapted the compositional habitat analysis proposed by Aebischer and others (1993) by defining habitat available in the surrounding area as all habitat available to the fawn. We then defined the habitat found at the capture location as habitat used by the fawn. We conducted our analysis in Program R using the compana function in the adehabitatHS package (R Development Core Team 2015; version 3.3.1; Calenge 2006). We replaced all habitat percentages of 0 in the fawn's capture location with 0.01, an order of magnitude smaller than the smallest amount of habitat found in each habitat type (Aebischer and others 1993). We considered relationships important at [alpha] = 0.05.


We captured 370 fawns from 2001 to 2015. Grassland ([bar.X] = 31.0%, SD = 31.0, n = 370; Table 2) and cropland ([bar.X] = 30.0%, SD = 30.0, n = 370) were the most prevalent habitat types at fawn-capture locations followed by pasture ([bar.X] = 20.3%, SD = 26.4, n = 370). Forested ([bar.X] = 2.2%, SD = 5.6, n = 370) and shrubby areas ([bar.X] = 0.5%, SD = 2.5, n = 370) were least prevalent at fawn capture locations. Conversely, cropland ([bar.X] = 37.8%, SD =23.3, n = 370; Table 2) was the most prevalent habitat type in the surrounding areas, followed by grassland ([bar.X] = 31.4, SD = 26.2, n = 370) and pasture ([bar.X] = 16.0%, SD = 1624, n = 370). As with fawn-capture locations, forested ([bar.X] = 1.9%, SD = 3.9, n = 370) and shrubby areas ([bar.X] = 0.2%, SD = 0.6, n = 370) were least prevalent in the surrounding areas.

Habitat types varied between fawn-capture locations and the surrounding areas ([LAMBDA] = 0.55, P = 0.002). Grasslands were most prevalent at fawn-capture locations, followed by the developed and shrubby habitat types (Table 3). Pasture, forest, and open water were least prevalent at fawn-capture locations, respectively. Although cropland was generally the most abundant cover type at fawn-capture locations, it was only prevalent more than pastures, forests, and open water.


White-tailed Deer display wide variation in their diet (Hewitt 2011) and habitat use (Stewart and others 2011), and are considered generalist species. This is consistent with our results, as most habitat types (approximately 63% of all habitat types used in the analysis) were present at fawn-capture locations. Nevertheless, grasslands, developed areas, and shrubby areas were most prevalent at White-tailed Deer fawn capture locations, whereas pastures, forests, and open water were least prevalent. This suggests that habitat types that are generally associated with increased cover were most prevalent at capture locations. Although wetlands and pastures likely provide hiding cover, prevalence of these habitat types may increase in areas used by fawns after fawns are mobile and may not be as suitable during their 1st month of life if water inhibits a fawn's ability to thermoregulate and pastures are actively grazed. Similarly, forests likely act as quality fawning cover because they represent permanent cover. However, fawn survival may decrease with increasing forested cover (Grovenburg and others 2012b). Given that most forested areas in the Northern Great Plains represent linear shelterbelts, these shelterbelts may improve the search efficiency of predators such as Coyotes (Canis latrans); however, this relationship between patch shape and predator search efficiency needs to be evaluated.

Although open water is important for mothers during lactation (Jacques and others 2015), it was the least prevalent habitat type found at fawn-capture locations. However, simply having water available to a mother, regardless of the quantity, may be adequate to meet her lactation needs. Additionally, annual precipitation likely influences a mother's need for open water (Grovenburg and others 2011b). Cropland was moderately prevalent compared to other habitat types, though it generally comprised the majority of habitat in areas surrounding capture locations. Although cropland also provides fawn cover, it may be inadequate relative to other cover types such as grasslands, likely because of differences in vegetation height, which may be important early in a fawn's life (Grovenburg and others 2010). Regardless, the importance of croplands in providing hiding cover and nutrition to fawns likely increases throughout the summer as crop height increases and agricultural crops increase in availability.

Grasslands and shrubby areas are generally considered high-quality fawning habitat (Grovenburg and others 2010), but farmsteads and roads associated with the developed habitat types are generally considered low quality. Habitat types associated with human presence negatively affect wildlife populations (Benitez-Lopez and others 2010; Torres and others 2016); however, there is some evidence of positive associations. Burr and others (2017) found that nest predation of Sharp-tailed Grouse (Tympanuchus phasianellus) in North Dakota was potentially lower in high-intensity energy-development areas compared to low-intensity energy-development areas because human presence decreased the number of predators in the high-intensity energy-development area. Elk also selected areas closer to houses and trails during the hunting season (Cleveland and others 2012). Areas related to human development may act as refuges from predators during vulnerable time periods (parturition, nesting, human-induced predation events; Berger 2007), though more research is needed to assess whether this pattern is consistent among ungulates. Alternatively, the habitat associated with farmsteads, and not human presence, may be what is influencing its use. Farmsteads tend to have more internal habitat complexity when characterized by a mix of tree rows, low shrubs, and open areas. Habitat complexity could also be associated with indicate increased patch connectivity, which has been related to increased fawn survival (Michel and others 2018).

Additionally, many of these developed areas receive infrquent or no human disturbance, which further complicates interpretation. Using maps of the central North Dakota study areas (ca. 1992; Schaffer 2013), 594 structures outside of incorporated towns were identified as developed areas; 188 (32%) were abandoned farmsteads, rural churches, and cemeteries that were seldom visited by humans (WF Jensen, North Dakota Game and Fish Department, unpubl. data). Since 1992, rural farming operations have become more consolidated, with a subsequent increase in the number of abandoned farmsteads across much of the Northern Great Plains. Therefore, the increased use of developed areas may result from a combination of habitat complexity, configuration, and also a lack of human presence.

Using postpartum behavior of reproductively active females as an indicator of presence of fawns (Downing and McGinnes 1969; White and others 1972; Huegel and others 1985) and using VITs to assist in fawn capture (Swanson and others 2008) may have biased our results. For example, observing postpartum behavior would inherently be more effective in open landscapes where observers can view females at distances at which they would not be detected in heavier cover and therefore, potentially increase the number of fawn-capture locations in open cover types. Conversely, VITs would increase the chances of fawn captures in wooded cover types, given that VITs employ telemetry to find newborn fawns. Regardless, agriculture and grasslands dominated the landscape in our study areas, whereas forested cover types comprised [less than or equal to]3% of land use (Brinkman and others 2005; Grovenburg and others 2010). Indeed, capture locations for fawns found via VITs had only 2.4% of wooded cover types compared to 27.8% of open cover types. Opportunistically captured fawns followed a similar trend, with 0.9% of wooded cover types found at capture locations versus 28.8% of open cover types. Therefore, the disproportional number of fawn captures in open cover types could be related to the availability of open cover types across our study areas.


Our results suggest that grasslands, developed areas, and shrubby areas are most prevalent at White-tailed Deer fawn-capture locations in the Northern Great Plains region. Although grasslands and shrubby areas are generally considered high-quality fawning cover, farmsteads associated with developed areas are not. However, habitat associated with these farmsteads may create important fawning cover, as 40% of grassland and woodland cover has been converted to row-crop agriculture between 2007 and 2013 in the Red River Valley of North Dakota (WF Jensen, North Dakota Game and Fish Department, unpubl. report). This dramatic decline in cover habitat in an agriculturally dominated landscape further emphasizes the importance of cover habitat associated with farmsteads. This is particularly true for abandoned farmsteads that are being targeted for removal at an increasing rate as agricultural commodity prices increase. Regardless, alternative habitat types must be made available in agriculturally dominated landscapes if the objective is to maintain or enhance populations of White-tailed Deer in this region.


We thank the North Dakota Game and Fish Department, South Dakota Department of Game, Fish and Parks, the Minnesota Department of Natural Resources, the Department of Natural Resource Management at South Dakota State University, and numerous private landowners and technicians for their help and cooperation with this project. We thank R. Hoffman and an anonymous reviewer for their helpful comments. We thank BA Schaffer, KL Moratz, BS Gullikson, KM Sternhagen, and TJ Brinkman for their help collecting data used in this analysis. This project was funded by Federal Aid to Wildlife Restoration administered through North Dakota Game and Fish Department (Project W-67-R-57, Study No. C-VIII). Any mention of trade, product, or firm names is for descriptive purposes only, and does not imply endorsement by the US Government.


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Submitted 30 November 2018, accepted 23 January 2019. Corresponding Editor: Robert Hoffman.


Department of Natural Resource Management, South Dakota State University, Box 2140B, Brookings, SD 57007 USA;


North Dakota Game and Fish Department, Bismarck, ND 58501 USA

(1) Current address: Division of Fish and Wildlife, Minnesota Department of Natural Resources, 35365 800th Avenue, Madelia, MN 56062 USA.
TABLE 1. Area-specific 95% summer maternal home ranges, maternal
circular buffers, and associated fawn circular buffers scaled by
maternal home range size of White-tailed Deer in the Northern Great

                                        95% summer       Maternal
                                        home range    circular buffer
               Location                 ([km.sup.2])        (m)

Lincoln, Pipestone, and Redwood             2.3           855.9
counties, MN (1)
Perkins County, SD (2)                      3.4          1037.5
Grant and Dunn counties, ND (3)             3.4          1037.5
Walsh and Grand Forks counties, ND (4)      4.2          1156.5
Burleigh, Kidder, and Sheridan              4.8          1236.4
counties, ND (5)
Edmunds County, SD (6*)                    10.2          1802.3

                                        Percent decrease
               Location                  from reference

Lincoln, Pipestone, and Redwood               52.5
counties, MN (1)
Perkins County, SD (2)                        42.4
Grant and Dunn counties, ND (3)               42.4
Walsh and Grand Forks counties, ND (4)        35.8
Burleigh, Kidder, and Sheridan                31.4
counties, ND (5)
Edmunds County, SD (6*)                        --

                                          Scaled fawn
                                        circular buffers
               Location                       (m)

Lincoln, Pipestone, and Redwood             167.3
counties, MN (1)
Perkins County, SD (2)                      202.9
Grant and Dunn counties, ND (3)             202.9
Walsh and Grand Forks counties, ND (4)      226.2
Burleigh, Kidder, and Sheridan              241.7
counties, ND (5)
Edmunds County, SD (6*)                     352.3

(1) Brinkman and others 2005
(2) Gullikson (unpubl. data)
(3) Gullikson (unpubl. data)
(4) Sternhagen 2015
(5) Schaffer 2013
(6) Grovenburg and others 2009
(*) Used as reference for scaling

TABLE 2. Percent habitat types found within the 95% buffered areas of
370 White-tailed Deer fawn-capture locations and their surrounding
areas in the Northern Great Plains.

Cover type  Capture location  Surrounding area
              Mean   SD        Mean   SD

Cropland      29.70  29.74     37.75  23.35
Developed      6.45   6.56      3.79   2.08
Forest         2.21   5.75      1.86   3.90
Grassland     30.97  31.01     31.41  26.21
Open water     2.56   7.60      3.63   7.21
Pasture       20.32  26.46     15.96  16.20
Shrub          0.45   2.50      0.18   0.62
Wetland        7.29  15.18      5.36  10.01

TABLE 3. Habitat use Rankings of 8 habitat types from White-tailed Deer
from 9 study sites in 3 states comprising the Northern Great Plains.
+++ = habitat type that was used more by fawns compared to other
habitat types;--= less use of that habitat type. Rankings range from
1-8, with 8 being the most used habitat type and 1 the least used.

Cover type  water  Developed  Forest  Shrub  Grassland  Cropland

Open water            --        -       -       --         --
Developed    +++               +++      +        -        +++
Forest        +       --                -       --         -
Shrub         +        -        +               --         -
Grassland    +++       +       +++     +++                +++
Cropland     +++      --        +       +       --
Wetland      +++      --       +++      -       --         +
Pasture      +++      --        +       -       --         +

Cover type  Wetland  Pasture  Ranking

Open water    --       --        1
Developed     +++      +++       7
Forest        --        -        2
Shrub          +        +        6
Grassland     +++      +++       8
Cropland       -        -        4
Wetland                 +        5
Pasture        -                 3
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Article Details
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Author:Michel, Eric S.; Jenks, Jonathan A.; Kaskie, Kyle D.; Jensen, William F.; Hoffman, Robert
Publication:Northwestern Naturalist: A Journal of Vertebrate Biology
Geographic Code:1U4MN
Date:Sep 6, 2019

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