Home Range and Habitat Use of Coyotes in an Area of Native Prairie, Farmland and CRP Fields

By Kamler, Jan F.; Ballard, Warren B. et al. | The American Midland Naturalist, April 2005 | Go to article overview

Home Range and Habitat Use of Coyotes in an Area of Native Prairie, Farmland and CRP Fields


Kamler, Jan F., Ballard, Warren B., Lemons, Patrick R., Gilliland, Rickey L., Mote, Kevin, The American Midland Naturalist


ABSTRACT.-

From 1999 to 2001 we monitored 12 coyotes (Canis latrans) in northwestern Texas to determine their home ranges and habitat use in a landscape interspersed with native prairie, farmland and Conservation Reserve Program (CRP) fields. Annual home range size was 10.1 km^sup 2^ for residents and 84.5 km^sup 2^ for transients. We determined habitat use at two spatial scales: within home ranges and within study area. Habitat use patterns were similar at both scales, as residents selected for native prairie and transients selected for CRP fields. Habitat use between residents and transients differed in both seasons, with residents selecting more native prairie, less farmland and less CRP (summer only) than transients. Habitat at natal den sites also differed from expected for residents, us most dens (8 of 10) were located in CRP fields. The CRP fields contained the only tall permanent vegetation on our study sites and appeared to provide important foraging habitat for transient coyotes, and denning habitat for resident coyotes.

INTRODUCTION

Despite substantial losses of native habitat during the past 200 y, coyotes (Canis latrans) increased their numbers and have thrived in human-altered habitats, including urban and agricultural environments (Bekoff, 1982; Voigt and Berg, 1987). Coyotes, which primarily occupied the prairie biome prior to European settlement, have since expanded their range into nearly all available habitats in North America (Nowak, 1978; Voigt and Berg, 1987; Moore and Parker, 1992). Despite extensive research on coyotes during the past 30 y, there is little published information concerning habitat selection of prairie compared to nonnative habitats. Information concerning habitat use of coyotes in landscapes interspersed with native prairie and human-altered habitats may help elucidate why this species has been so successful in human-dominated environments. Coyotes have been classified in their social organization as residents and transients (Andelt, 1985; Gese et al., 1988; Kamler and Gipson, 2000). Because transients and residents can exhibit different habitat use patterns (Kamler and Gipson, 2000), effects of heterogeneous landscapes should be investigated for both types of coyotes.

With passage of the 1985 Farm Bill, the Conservation Reserve Program (CRP) was created with the goal of retiring highly erodible land from agricultural production and converting it to permanent cover. Since 1985, CRP has removed approximately 14.7 billion ha nationwide from agricultural production (Weber et al., 2002). Most (87%) CRP acres are planted to grasses, with most (66%) being nonnative species (Osborn and Heimlich, 1994). By law, CRP fields are not grazed, unless restrictions are lifted in times of extreme drought. Thus, the vegetation structure and diversity of most CRP fields differ considerably from native grasslands, which are often grazed by cattle. Although originally designed to reduce soil erosion, CRP fields are now widely acknowledged to have positive benefits for many wildlife species (Delisle and Savidge, 1997). For example, previous research showed that permanent cover provided by CRP fields is important to many bird species (Johnson and Schwartz, 1993; Best et al., 1998; McCoy et al, 1999). The CRP also can be beneficial to larger species such as white-tailed (Odocoileus virginianus) and mule (O. hemionus) deer (Could andjenkins, 1993; Kamler et al., 2001). However, use of CRP by other large mammal species, especially carnivores, has received little attention. A better understanding of CRP use by coyotes is needed, as this information would help determine if this relatively recent, but widespread, habitat type is beneficial to this species.

The purpose of this paper was to compare home ranges and seasonal habitat selection of resident and transient coyotes in a heterogeneous landscape. We determined habitat selection at two spatial scales: within home ranges and within the study area. We also compared habitat selection of natal den sites for resident coyotes. Our study site in northwestern Texas occurred within the historical range of coyotes and consisted of native prairie, farmland and CRP fields.

METHODS

Study area.-Research was conducted on a 110-km2 area on the border of Dallam and Sherman counties in northwestern Texas (36°24'N, 102°19'W). The center of the study site was located on a private ranch surrounded by other ranches, farmland and CRP fields. Vegetation on ranches consisted of shortgrass-prairie species (hereafter, native prairie) that were dominated by blue grama (Bouteloua gracilis) and buffalograss (Buchloe dactyloides) and were moderately to intensively grazed by cattle (Bos taurus). Farmland consisted of irrigated and nonirrigated agricultural fields. Primary crops on our study site were corn and winter wheat planted on a rotational basis. Most CRP fields in our study were enrolled in 1985, and planted to warm-season grasses, dominated by old world bluestem (Andropogon spp.) and sideoats grama (Bouteloua curtipendula). The CRP fields had vegetation that was taller and thicker than the shortgrass prairie that historically dominated this region (Barbour and Billings, 1988), especially because they were not grazed by cattle (Fig. 1).

Capture and radiotekmetry.-From January 1999 to September 2000 we radio-collared and monitored 17 coyotes (10 males, 7 females). Due to early deaths (n = 2) or dispersal (n = 3), 5 coyotes (4 males, 1 female) were not included in the analyses. Coyotes were captured in padded leg-hold traps (Kamler et al., 2002) placed throughout the study area. Most traps (>95%) were set along fences that separated different fields. All fence-lines within the study area were trapped and all habitat types were represented when trapping. Captured coyotes were ear-tagged and fitted with radio-collars (Advanced Telemetry Systems, Inc., Isanti, Minnesota) and aged by tooth wear, body size and reproductive condition (Gier, 1968). Coyotes were classified as residents if they maintained relatively small home ranges that overlapped little with adjacent family groups (Andelt, 1985; Gese et al., 1988; Kamler and Gipson, 2000). Coyotes were classified as transients if maintained relatively large home ranges that overlapped those of other coyotes (Andelt, 1985; Gese et al., 1988; Kamler and Gipson, 2000).

We recorded independent telemetry locations (White and Garrott, 1990) for study animals 1-2 times per week and >12 h apart. We radio-tracked from vehicles using null-peak systems which consisted of dual, 4-element Yagi antennas. Most radio-tracking (>90%) occurred during 1800-0900 hours, when coyotes were likely to be most active (Andelt, 1985). We calculated location estimates using the maximum likelihood estimation option in program Locate II (Pacer, Inc., Truro, Nova Scotia, Canada). Mean error for reference collars (known locations) was 84 m (95% of errors were <145 m).

Home range analysis.-We determined annual (January to December) home range sizes for coyotes using the minimum convex polygon (MCP) method (Mohr, 1947), as calculated by Animal Movement program (Hooge and Eichenlaub, 1997). We calculated home range sizes for resident coyotes with >39 locations and >7 mo of radio-tracking and transient coyotes with >29 locations and >6 mo of radio-tracking. Area observation curves generated by Animal Movement program showed that 23-29 locations were needed to effectively determine home ranges for individual coyotes. If coyotes lived 2 y, only the annual home range during the first year was used in analyses, unless a coyote shifted its home range to a new territory during the second year (n = 1). Mean home range sizes were compared between residents and transients using Hests (Zar, 1996) and deemed significant when P < 0.05.

Habitat use analysis.-Habitat use data were divided into two seasons defined as summer (May to October) and winter (November to April). The seasons were defined in this way to parallel major changes in climate and two major biological periods for coyotes (summer = birthing and young rearing; winter = dispersal and pair bonding). Habitat types were delineated using geographic information system (GIS) data obtained from quadrangle maps and were ground-truthed by visual inspection. Habitat selection (use vs. available) of residents and transients was determined at two spatial scales: within home ranges and within study area. At the home range scale, we determined habitat composition by overlaying individual home range polygons onto the CIS map of habitat types using ArcView (version 3.2, Environmental Systems Research Institute, Inc., Redlands, CA). Percentage of different habitat types within home-range polygons was considered available. We determined habitat use by plotting all locations for each coyote onto the GIS map. Expected number of locations for each habitat were calculated by multiplying total locations by percentage of habitat types within the home-range polygon. We compared actual habitat use to expected with chi-square goodness-of-fit tests (Neu et al., 1974; White and Garrott, 1990). Selection indices (observed/expected) were also calculated for illustrative purposes.

At the study area scale, we used locations from all radio-collared individuals to create (using MCP method) an availability polygon (Miller et αι., 1999) for each social class. Percentage of different habitat types within polygons was considered available. To determine seasonal use, we plotted all locations for each social class onto the GIS map. Expected number of locations for each habitat type was calculated by multiplying total locations by percentage of available habitat types. For each season, we then compared total locations within habitat types to expected using chi-square goodness-of-fit tests. We also compared overall use between residents and transients in each season and compared seasons within each social class, using chi-square goodness-of-fit tests.

Habitat selection of natal den sites was determined for resident coyotes by radio-tracking and walking up to coyotes during the day in the early pup-rearing period (late April-early May). Natal dens were distinguished from non-natal dens by presence of pups, food scraps, worn trails and bed sites (Althoff, 1980). Although coyotes often changed den sites, only one natal den per family group year was used in analyses, as subsequent dens usually were in the same field (hence, same habitat type). Because of these restrictions, number of dens used in analyses were relatively low (n = 10). However, our data met the minimum requirements outlined by Roscoe and Byars (1971) and Koehler and Larntz (1980), thus habitat types at natal dens were compared to expected (determined from availability polygons) using chisquare goodness-of-fit tests.

RESULTS

We obtained 819 locations for 7 resident and 5 transient coyotes during the study. Mean (± SE) annual home-range size differed significantly (P = 0.021) between residents (10.1 ± 1.1 km^sup 2^) and transients (84.5 ± 16.8 km^sup 2^; Fig. 2). At the home range scale, most residents selected for native prairie, whereas transients selected for CRP or farmland (Table 1 ).

At the study area scale, the availability polygon for residents (15,293 ha) included 29.8% native prairie, 39.7% farmland and 30.6% CRP fields. The availability polygon for transients (25,271 ha) included 32.9% native prairie, 48.3% farmland and 18.7% CRP fields. For residents, there was a strong selection for native prairie in both seasons (Table 2). For transients, there was strong selection for CRP fields in both seasons (Table 2).

Overall habitat use differed between seasons for residents (X^sub 2^^sup 2^= 17.90, P < 0.001), but not transients (X^sub 2^^sup 2^ = 1.88, P = 0.391). Residents used more native prairie and CRP, and less farmland, in winter than summer. Overall habitat use between residents and transients differed during summer (X^sub 2^^sup 2^ = 9.98, P = 0.007) and winter (X^sub 2^^sup 2^= 19.79, P < 0.001). Residents used more native prairie, less farmland and less CRP (summer only) than transients.

Ten natal dens of resident coyotes were located during the study. Habitat type at natal den sites differed (X^sub 2^^sup 2^ = 12.67, P = 0.002) from expected, as most (8 of 10) dens occurred in CRP fields.

DISCUSSION

Resident and transient coyotes selected for habitat types on our study site. During both seasons, residents selected for native prairie, whereas transients selected for CRP fields. Prairies were the primary habitat that coyotes occupied in North America prior to European settlement (Moore and Parker, 1992), thus it is not surprising that resident coyotes selected for this habitat type. Resident coyotes might have selected native prairie because it contained more prey species than the other habitat types, although we did not determine prey abundances on our study area. During our study, the most common food items in the coyote diet were small rodents in winter (45%) and insects in summer (31%; calculated from data in Lemons, 2001). Previous research in western Texas showed that, compared to native shortgrass prairie, CRP fields contained similar diversity and abundances of small mammals (>2y CRP fields; Hall and Willig, 1994), but significantly lesser diversity and abundances of arthropods (McIntyre and Thompson, 2003). Thus, native prairie likely contained more prey species for coyotes than the other two habitat types.

Transients likely selected CRP fields for several reasons. Firstly, CRP provided the best available cover for transients, as this was the only tall permanent vegetation on our study site. Because transients are solitary, they might need more cover than residents when resting, hunting or for other activities. Human persecution of coyotes on our study site was high (Kamler et al., 2003), therefore more cover provided greater protection from hunters. secondly, transients are subordinate to residents and, consequently, are often excluded from optimal resources by residents (Kamler and Gipson, 2000; Gese, 2001). Because residents selected for native prairie, transients selected for what was probably the next best available habitat, CRP fields. Comparisons of overall habitat use showed a similar trend, as residents used more native prairie and less farmland and CRP fields than transients. These findings were similar to that found in Kansas, where resident coyotes selected native grasslands and transients selected more suboptimal habitats such as woodland and farmland (Kamler and Gipson, 2000).

Seasonal changes in habitat use were exhibited by resident coyotes, but not transients. During winter, resident coyotes selected more CRP and native prairie, and less farmland, than in summer. Reasons for these seasonal changes in habitat use were not clear, but they could have been related to several factors, such as changes in vegetative cover and prey species. Farmland vegetation exhibited the most seasonal changes of all habitat types on our study site. For example, tall and green vegetation (e.g., corn) grew during summer, whereas during winter most fields were barren because corn stubble was burned or plowed after harvest. Because this habitat type became so barren in winter, resident coyotes preferred to use habitat with more cover. Additionally, prey abundance might have been severely reduced during winter when vegetation was minimal in farmland habitats. In other regions, coyotes also used agricultural fields less in winter than summer, due to the changes in vegetative cover (Andelt and Andelt, 1981; Person and Hirth, 1991).

Resident coyotes also showed habitat selection for natal den sites, as 80% were located in CRP fields. This indicated that, although resident coyotes preferred native prairie for hunting during nocturnal hours (when most telemetry locations were obtained), they preferred the habitat with the most cover for natal den sites. Previous research also showed that coyotes prefer sites with tall grasses or other substantial cover for dens (Rock, 1978; Althoff, 1980; Hallett et al., 1985). For example, in areas containing farmland and grassland, coyotes nearly always located their dens in grassland (Rock, 1978; Althoff, 1980; Hallett et al., 1985), where den entrances can often be concealed by grasses (Althoff, 1980). In our study site, CRP fields had much taller and thicker vegetation than native short-grass prairie, thus coyote dens could be better concealed than in native prairie. Additionally, native prairie on our study sites contained large numbers of cattle, and had more human activity (related to ranching), at least during daylight hours, than CRP fields. Coyotes generally avoid human activity (Gese et al., 1989), especially in areas where they are heavily persecuted (Kitchen et al., 2000).

Our study indicated CRP fields provided important cover for coyotes, probably because CRP was the only tall permanent vegetation on our study site. Although resident and transient coyotes used CRP for different reasons (e.g., dens for residents, foraging habitat for transients), this habitat appeared to be an important component of coyote ecology. It was unknown if CRP fields increased the carrying capacity of coyotes in our study area. However, the high use of CRP by both types of coyotes suggest this may have been possible, at least within the mosaic of farmland and native short-grass prairie.

Acknowledgments.-This project was funded by Texas Tech University, Texas Parks and Wildlife Department, U. S. Forest Service, U. S. U. A.-Wildlite Services program and section 6 Grant E-1-12 from the U. S. Fish and Wildlife Service's Endangered Species Program. Kansas Department of Wildlife and Parks and BWXT Pantex loaned us equipment. We thank C. C. Perchellet, B. Kamler and E. B. Fish for assistance with the project. We also thank F. Pronger and other landowners in Sherman and Dallam counties who allowed us to conduct research on their land. Our research protocol, no. 00979BX, was approved by the Texas Tech University Institutional Animal Care and Use Committee. This is Texas Tech University, College of Agricultural Sciences and Natural Resources technical publication T-9-1005.

[Reference]

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SUBMITTED 23 FEBRUARY 2004 ACCEPTED 10 MARCH 2004

[Author Affiliation]

JAN F. KAMLER1, WARREN B. BALLARD2 AND PATRICK R. LEMONS

Department of Range, Wildlife, and Fisheries Management, Box 42125, Texas Tech University, Lubbock 79409

RICKEY L. GILLILAND

USDA-Wildlife Services, Box 60277, West Texas A&M University, Canyon 79016

AND

KEVIN MOTE

Texas Parks and Wildlife Department, 301 Main Street, Brownwood 76801

[Author Affiliation]

1 Present address: Polish Academy of Sciences, Mammal Research Institute, 17-230 Bialowieza, Poland

2 Corresponding author e-mail: warren.ballard@ttu.edu

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