Burrowing Owl (Athene Cunicularia) Nesting Ecology In Western Kansas

Fort Hays State University FHSU Scholars Repository Master's Theses Graduate School Fall 2012 Burrowing Owl (Athene Cunicularia) Nesting Ecology In Western Kansas Carol D. Grover-Mariner Fort Hays State University, caroldmariner@gmail.com Follow this and additional works at: https://scholars.fhsu.edu/theses Part of the Biology Commons Recommended Citation Grover-Mariner, Carol D., "Burrowing Owl (Athene Cunicularia) Nesting Ecology In Western Kansas" (2012). Master's Theses. 116. https://scholars.fhsu.edu/theses/116 This Thesis is brought to you for free and open access by the Graduate School at FHSU Scholars Repository. It has been accepted for inclusion in Master's Theses by an authorized administrator of FHSU Scholars Repository.

BURROWING OWL (ATHENE CUNICULARIA) NESTING ECOLOGY IN WESTERN KANSAS being A Thesis Presented to the Graduate Faculty of the Fort Hays State University in Partial Fulfillment of the Requirements for the Degree of Master of Science by Carol Grover-Mariner B.S., Fort Hays State University Date Approved Major Professor Approved Chair, Graduate Council

This thesis for The Master of Science Degree By Carol D. Grover-Mariner has been approved Chair, Supervisory Committee Supervisory Committee Supervisory Committee Supervisory Committee Chair, Department of Biological Sciences i

PREFACE This thesis is written in the style of the Wilson Journal of Ornithology, to which a portion will be submitted for publication. ii

ABSTRACT The Burrowing Owl (Athene cunicularia) is a common occupant of Smoky Valley Ranch located in Logan County, Kansas, where the abundance of black-tailed prairie dog (Cynomys ludovicianus) burrows provides ideal habitat for the Burrowing Owl. The objectives of the study were to 1) estimate the densities of black-tailed prairie dog colonies through visual counts, 2) estimate nesting success and nest survival probabilities of the Burrowing Owl with the use of a burrow camera, and 3) estimate post-fledging dispersal and return rate of juvenile Burrowing Owl the following season. I located 96 potential nests from 24 March through 2 July 2007 and 64 potential nests from 28 May through 11 August 2008. Data collected at each nest burrow included a GPS location, clutch size, number of nestlings, number of juveniles fledged, bearing of nest burrow entrance, and 5 nearest habitable bearings of non-nest burrows. Juveniles also were trapped and banded. In 2007 mean clutch size equaled 5.16, mean number of nestlings equaled 2.43, mean number of fledged individuals equaled 2.61, apparent nest success was 33%, and Mayfield nest success was 67%. In 2008, mean clutch size equaled 4.05, mean number of nestlings equaled 2.79, mean number of fledged individuals equaled 2.56, apparent nest success was 39%, and Mayfield nest success was 57%. Densities of black-tailed prairie dog were then obtained on randomly selected locations within various colonies by using visual counts. Paired t-tests were used to compare Burrowing Owl nest success and survival probabilities to densities of black-tailed prairie dog and were found to be significant for Burrowing Owl nest success for 2007 (t = 12.966; df = 58; P < 0.001), 2008 (t = -8.247; df = 45; P < 0.001) and both years combined (t = -14.728; df = iii

104; P < 0.001) and Burrowing Owl nest survival probabilities for 2007 (t = 13.613; df = 58; P < 0.001), 2008 (t = -7.097; df = 45; P < 0.001) and both years combined (t = - 13.770; df = 104; P < 0.001). A paired t-test, comparing Burrowing Owl nest success and nest burrow entrance bearing, was found to be significant for 2007 (t = 11.67; df = 26; P < 0.001), 2008 (t = 8.795; df = 21; P < 0.001), and both years combined ( t = - 14.465; df = 49; P < 0.001). iv

ACKNOWLEDGMENTS First, I thank my grandmother, Melicent Grover, her enthusiasm for wildlife and the outdoors inspired me to pursue what I enjoy. Second, I thank my father, Karl Grover, for pointing me in the right direction and encouraging me to pursue a masters degree, my mother, Cathy Grover, for her never ending love and support. Third, I thank my major advisor, Dr. Elmer J. Finck, for his patience, guidance, and advice through all stages of my research. I thank my thesis committee, Dr. Rob Channell of Fort Hays State University (FHSU), Dr. Greg Farley (FHSU), and Charlie Lee of Kansas State University, for their contributions and suggestions on my research. I also thank faculty and fellow graduate students of FHSU Department of Biological Sciences for their support and help. I am especially thankful for the dedication and hard work of my field assistants and volunteers. I thank Jessica Bender, Jason Black, and Kim Goering for their assistance in the field and around the table entering endless data; Dave Bender, Ken Brunson, Ben Grover, Cathy Grover, Karl Grover, Jenny O Neill, Dr. Rob Penner, and Dustin Tacha for volunteering their time and hard work. My research was funded by a grant from The Nature Conservancy (Kansas Chapter), coordinated by Rob Manes. Additional funding was provided by the Department of Biological Sciences (FHSU) and by the Chickadee Checkoff Small Grants Kansas Nongame Wildlife Improvement Fund 2008, administered by the Kansas Department of Wildlife and Parks and coordinated by Ken Brunson. The Peeper Video v

Probe was purchased from Sandpiper Technologies, Inc., and I thank them for their help in determining which camera system would be best for my research. I thank Sheila Pfeifer for all she did in making sure that everything ran smoothly within the Department of Biological Sciences during my graduate research. I thank Randy Martin and Michelle Martin for help with schedules, housing arrangements, equipment, and field support. Finally, I thank my loving and supportive husband, Adrian Mariner. I would not have been able to achieve this without him. His unending support and motivation for me to finish got me here. Thank you and I love you so much! vi

TABLE OF CONTENTS Page GRADUATE COMMITTEE APPROVAL... i PREFACE... ii ABSTRACT... iii ACKNOWLEDGMENTS...v TABLE OF CONTENTS... vii LIST OF TABLES... viii LIST OF FIGURES... ix LIST OF APPENDICIES...x INTRODUCTION...1 METHODS...5 Site Description...5 Field Methods...7 Data Analyses...11 RESULTS...13 DISCUSSION...14 CONCLUSIONS...17 LITERATURE CITED...20 vii

LIST OF TABLES Table Page 1 Burrowing Owl Mayfield results for 2007 for the Smoky Valley Ranch, Logan County, Kansas...26 2 Burrowing Owl Mayfield results for 2008 for the Smoky Valley Ranch, Logan County, Kansas...27 3 Statistical analyses results of paired t-tests for the Smoky Valley Ranch, Logan County, Kansas...28 4 Statistical analyses results of Wilcoxon Signed Ranks test for the Smoky Valley Ranch, Logan County, Kansas...29 viii

LIST OF FIGURES Figure Page 1 Location of Smoky Valley Ranch, Logan County, Kansas (modified from Haug et al. 1993)...30 2 Aerial view of Smoky Valley Ranch, Logan County, Kanas with the Smoky Hill River bisecting it...31 3 Burrowing Owl breeding distribution (yellow), year-round range (green) and winter range (brown) in North America...32 4 Peeper TM Video Probe (Sandpiper Technologies, Inc.), the burrow camera, consisting of a handheld screen, battery pack and 4.5m cable with infrared camera at the distal end...33 5 A two door one-way push-door trap set up at a Burrowing Owl nest burrow...34 6 Tail-mount transmitter on a juvenile Burrowing Owl...35 7 Count site configuration for estimating black-tailed prairie dog densities...36 8 Gopher snake (Pituophis catenifer) in a Burrowing Owl nest burrow eating a nestling Burrowing Owl...37 9 American badger (Taxidea taxus) digging out a black-tailed prairie dog burrow.38 10 Burrowing Owl tracked down a burrow with tail-mount transmitter antennae shown in dirt...39 ix

LIST OF APPENDICIES Appendix Page A Black-tailed prairie dog data for 2007 and 2008 for the Smoky Valley Ranch, Logan County, Kansas...40 B 2007 Burrowing Owl data with Mayfield calculations for the Smoky Valley Ranch, Logan County, Kansas...50 C 2008 Burrowing Owl data with Mayfield calculations for the Smoky Valley Ranch, Logan County, Kansas...52 x

2 INTRODUCTION North American tallgrass, mixed, and short grass prairies are considered to be among the continent s most endangered ecosystems (Ricketts et al. 1999), and are possibly the highest priority relative to conserving biodiversity in the region (Samson and Knopf 1996). Only 52% of the historical extent of short grass prairie remains (Samson et al. 2004). The availability of land through the United States Homestead Act of 1862, the Canada Dominion Land Act of 1872, and the direct sale of land to private landowners, resulted in significant loss of native prairie in North America. Most loss has been primarily through conversion to agricultural land (Samson et al. 2004). These anthropogenic changes to the landscape have homogenized (Smith and Lomolino 2004) and fragmented the prairie ecosystem, leading to declines in biodiversity. Lands that remained prairie, now most commonly used for raising livestock (Bovidae), still provide some habitat for wildlife (Kie e al. 1996). Habitat fragmentation as well as habitat loss, like that of the short grass prairie community, has an impact on the stability of populations it supports (Turner et al. 2001). All populations have a probability of going extinct, but fragmentation and loss of required habitat increases extinction probability (Turner et al. 2001). Some species require larger areas of habitat for survival and if the habitat in which they live is reduced or fragmented, populations are reduced. Overall, loss of prairie has negative effects on native plants and animals (Samson and Knopf 1996). The prairie dog (Cynomys spp.), has undergone population declines as much as 98% throughout North America (Miller et al. 1994). One cause of the declines and

fragmentation of prairie dog colonies is because of human control (Hoogland 1996). The majority of Kansans think that prairie dogs compete with cattle for forage, or that cattle break their legs in prairie dog holes (Lee and Henderson 1989). Some reduction in prairie dog numbers has been through federal and state sponsored prairie dog control programs (Miller et al 1994). Due to fragmentation of the short grass prairie, as well as eradication or control methods, the remaining prairie dog colonies are smaller and more isolated (Miller et al. 1994). Prairie dogs affect the grassland ecosystem (Fahnestock and Detling 2002), and because of this are considered to be a keystone species. A keystone species activities and abundances influence the other species such that their absence significantly changes the structure and function of that community (Paine 1969). A keystone species in the grasslands is usually a species that significantly alters community structure and function, which in turn results in food and habitat structure for other species (Vinton and Collins 1996). Landscape fragmentation limits the ability of species (prairie dog) to disperse and colonize new areas. This also leaves the remaining colonies more susceptible to extinction (Miller et al. 1994) and results in loss of habitat they provide for other species. Prairie dog colonies provide habitat diversity by altering soil structure and chemical composition through mixing soils by burrowing, regulating vegetation species diversity through clipping and foraging of vegetation, and the addition of plant materials and excrement. Prairie dogs pose several management challenges to landowners and resource managers (Witmer and Fagerstone 2003). Landowners are challenged with keeping prairie dogs from extreme modification of rangeland without eradicating the prairie dog 3

4 populations that are present on their lands. In the western Great Plains, vertebrate biodiversity has declined, leading resource managers to spend increasing amounts of time and money to save species that depend on prairie dogs (Miller et al. 1994). Black-tailed prairie dog (Cynomys ludovicanus) presence is critical to the Burrowing Owl (Athene cunicularia). Burrowing Owl numbers are declining (Powell et al. 1994), therefore, it is considered a species of special concern through much of its range (Desmond et al. 2000). The Burrowing Owl inhabits dry, open, short grass plains (Haug et al. 1993) and tends to occupy areas with high densities of rodent burrows (Haug et al. 1993, Plumpton 1992), and higher densities of active burrows during the nesting season (Desmond et al. 2000). It is often associated with colonial sciurids in the Great Plains, most commonly black-tailed prairie dog (Butts 1973). Nesting within active prairie dog colonies might benefit the Burrowing Owl in several ways. Hoogland (1981) found that predator detection by prairie dogs increased as the number of prairie dogs present in the colony increased. Burrowing Owl might benefit from this through the dilution effect when nesting in an active prairie dog town and through the alarm calls by black-tailed prairie dog (Desmond et al. 2000). The dilution effect is a reduced probability of depredation due to the increased number of individuals serving as alternative prey sources. The black-tailed prairie dog also helps maintain habitat conditions suitable to Burrowing Owl through the clipping and grazing of vegetation as well as providing a number of burrows for use as satellite burrows (Desmond et al. 2000). Because the Burrowing Owl is associated closely with prairie dogs, when the prairie dog colonies are reduced in size, fragmented, or eradicated, Burrowing Owl habitat loss is

5 also likely to occur (McDonald et al. 2004). In a fragmented landscape, small, isolated habitat patches are less likely to be recolonized by Burrowing Owl (Todd et al. 2007). This unpredictability of habitat increases the probability of extinction of populations of prairie dog and the Burrowing Owl, as well as various other species relying on the microhabitats created by prairie dogs. This is a good example of how management for an ecosystem might have greater benefits than management for a single species. My goal was to provide data and results to The Nature Conservancy (TNC) that might be of significant value in determining management strategies for black-tailed prairie dog on Smoky Valley Ranch (SVR). The objectives of my study were to 1) estimate the densities of black-tailed prairie dog colonies through visual counts, 2) estimate nesting success and nest survival probabilities of the Burrowing Owl, and 3) estimate post-fledging dispersal and return rate of juvenile Burrowing Owl the following season. METHODS Site Description Research was conducted at SVR owned and managed by TNC and located in central Logan County, Kansas (Fig. 1). Smoky Valley Ranch is in the Central Short Grass Prairie and in TNC s Chalk Bluffs ecological site (Manes 2006). North American short grass prairie extends about 320 kilometers east from the Rocky Mountains where it meets the mixed grass prairie and extends south from central Alberta to central Texas (Samson and Knopf 1996). It is distinguished from other grassland ecoregions by low

6 rainfall, warm summer temperatures and relatively long growing seasons (Ricketts et al. 1999). Smoky Valley Ranch was 6,800 hectares, and was bisected by the Smoky Hill River into a northeastern portion and a smaller southwestern portion (Fig. 2). The Smoky Hill River has mature cottonwood trees (Populus sp.) along most of its riparian area (Manes 2006). The river retains water year-round in a few depressions and only has flowing water during periods of precipitation (Manes 2006). The land located to the northeast of the Smoky Hill River is primarily short grass prairie (Bouteloua sp. and Bouteloua dactyloides) with some steeply rolling hills and chalk bluffs. The land to the southwest of the river is primarily sandsage prairie (Artemisia filifolia) in the river valley, and re-seeded short grass prairie dominated by little bluestem (Schizachyrium scoparium) and side-oats grama (Bouteloua curtipendula) across the remainder (Manes 2006). Today the primary influences on the landscape of SVR include grazing by cattle (Bos taurus), American bison (Bison bison), and black-tailed prairie dog; absence of fire; localized groundwater and surface-water depletion (Manes 2006); climate patterns; sod breaking (Samson and Knopf 1996) by animals such as black-tailed prairie dog; and invasive species (Samson and Knopf 1996) such as musk thistle (Carduus nutans) and salt cedar (Tamarix spp.). As of September 2006, the northern two-thirds of SVR contained approximately 80% of the black-tailed prairie dog colonies by area that exist on the ranch (Manes 2006). The Nature Conservancy plans to maintain a black-tailed prairie dog population with at

least 810 occupied hectares on SVR (Manes 2006). Due to the close association between the black-tailed prairie dog and Burrowing Owl, TNC s goal of maintaining active black-tailed prairie dog colonies, and the goal to provide sufficient habitat for stable populations of Burrowing Owl, SVR was a good location for studying ecological relationships between black-tailed prairie dog and Burrowing Owl. Smoky Valley Ranch is within the breeding range of the Burrowing Owl (Fig. 3). The Burrowing Owl returns to SVR beginning in mid to late March. The Burrowing Owl tends to be monogamous for a season, however not permanently monogamous between seasons, especially in migratory populations (Haug et al. 1993). It is a semi-colonial species often associated with burrowing mammals primarily in dry, open, short grass, treeless plains (Haug et al. 1993). The Burrowing Owl entire nesting period is 75 days, i.e., eggs are laid at a rate of >1 per day, incubation period 30 days, and nestling period 43 days (Haug et al. 1993). Incubation is done by the female only while the male brings food items to the incubating female. Nest burrow entrances and cavity are lined with dried cow or horse (Equus caballus) manure. Burrowing Owls also will adorn the entrance with highly visible objects and prey items. Burrowing Owl nest survival probabilities were calculated for each nest and each nest was assigned success or failure based on whether or not young fledged from that nest. Field Methods Data were collected from 24 March 2007 through 28 July 2007 and 28 May 2008 through 11 August 2008. Burrowing Owl nest searches were conducted by driving along fence lines and roads through SVR throughout breeding season. Potential owl nest 7

8 burrows were located by observing flushed individuals around burrows and usually confirmed by the presence of shredded manure lining the burrow and scattered at the entrance, as well as the presence of prey items. The locations of potential nests were noted by using a Global Positioning System. Nests were checked once every seven days. Numbers of individuals flushed from the area when approaching a nest were recorded. Clutch and brood size was observed by using a burrow probe camera (Sandpiper Technologies ) (Fig. 4), which allowed me to see into the burrow without causing major disturbance to the burrow or its occupants. The burrow probe camera was illuminated with an infrared light and at the end of a 4.5 meter cable. The occupants of the burrow were visible on a handheld screen or headmount video display. Prey items in the burrow as well as presence of female on the nest also were recorded with the burrow probe camera. A compass bearing was recorded at the nest burrows entrance, as well as the five nearest habitable non-nest burrow entrances to determine if orientation of burrow entrance influenced nest success and/or survival probability. At the nest burrow and neighboring satellite burrows for each nest, pellets and prey item remains were collected, while non-collectible prey items, i.e., whole rodents, snakes, and toads, were recorded in notes and photographed for possible future diet analysis. Once the nestlings were seen at the entrance of the burrow, a double one-way push-door trap (modified from Botelho and Arrowood 1995 and Winchell 1999) was used to capture individuals. The trap was placed at the entrance of the burrow, covered with burlap to provide shade and maintain the illusion of a burrow (Fig. 5), and allowed

9 to sit until several individuals entered the trap; if possible the entire brood was captured. Traps were checked every 30-60 minutes. If cattle were near the set traps, they were monitored from a distance with binoculars to ensure cattle did not trample the traps or any occupants. After capture, individuals of the Burrowing Owl were banded with a size 4 U.S. Fish and Wildlife Service aluminum band (Pyle 1997). In 2007, ten offspring were radio tagged with a tail mount transmitter (Biotrack ) (Fig. 6) in an attempt to measure post-fledging dispersal and return rate the following season. Transmitters weighed 3.5 g (2.3% of a 150g adult) (Pyle 1997), which was less than the recommended 5% of the birds mass. A tail mount style was chosen so when juveniles molted their tail feathers the transmitter would be removed. Transmitters were attached by using super glue and anchor strings to secure the antenna to the tail feather rachis. The individuals of Burrowing Owl with the transmitters were then tracked by using a receiver (Wildlife Materials ) and Yagi antenna once every 3-5 days. When an individual was located, or a transmitter recovered a GPS coordinate was taken at the location. In 2008, mass, wing chord, and longest rectrix also were recorded for captured individuals. Nest fate was determined based on status of the nest at the previous check, condition of the burrow itself, or the observation of adults or fledglings in the area when observed from a distance of 100 m or more with binoculars. A nest was considered successful if at least one offspring survived to fledging, depredated if the presence of owl remains or evidence of digging by American badger (Taxidea taxus) or black-tailed prairie dog were observed, and unknown for all nests with an undeterminable fate.

10 The densities of the black-tailed prairie dog colonies were estimated mid-june through August, after the young of the year emerged from burrows and yearlings dispersed. The young of the year for the prairie dog emerge from the burrows at the end of May (Powell et al. 1994), and yearlings disperse about mid-june (Severson and Plumb 1998). I estimated the density of black-tailed prairie dogs by doing above ground counts of individuals within a marked area or count site (Fig. 7). The count sites were 200 x 200 m (4 ha) (Severson and Plumb 1998), corners were marked with 1m fiberglass poles, which had been spray painted a fluorescent orange/red. Poles were placed at each of the four corners of the count site, and in the middle of the sides opposite the corner of the observer location. The fiberglass poles aided in determining whether or not an individual black-tailed prairie dog was within in the count sites boundaries. To count the prairie dogs, I used binoculars (10 x 50 mm) and I was stationed at a corner atop a vehicle, ATV or in a chair from the highest point of elevation at the site. Visual counts for a single site were conducted for three consecutive days (Severson and Plumb 1998) at approximately the same time of day. I waited for 30 min. after arriving at the count site before beginning the counts to allow the black-tailed prairie dogs to become accustomed to my presence (Severson and Plumb 1998). The counts took 15 min. total and consisted of three 5 min. counts (C. D. Lee pers. comm.). Within the first 5 min. interval, several scans of the count site were conducted. The maximum number of black-tailed prairie dogs seen were recorded for each interval. This procedure was repeated at the 10 min. mark and 15 min. mark (C. D. Lee pers. comm.), resulting in 3 counts per site, per day. All visual counts were done between 0700 and 1000 (CST) (Severson and Plumb 1998)

11 or if unable to count in the morning period on one of the 3 consecutive days (due to weather) the evening period of 2 hours prior to sunset was used (Powell et al. 1994). However, if for some reason the evening period could not replace the interrupted morning period, the 3 day count was restarted for that count site. Above ground counts were not conducted during rain or if wind exceeded 32 km per hour (Severson and Plumb 1998). Weather readings were taken at the beginning of each count with a Kestrel pocket weather station to ensure wind speed was not greater than the 32 km/hr limit. Temperature, relative humidity, percent cloud cover, presence/absence of cattle, and visibility estimate also were recorded prior to each count. In 2008, count sites were conducted on the same locations as the previous season, and in addition, near Burrowing Owl nests located that season. Data Analyses The Mayfield method (Mayfield 1975) was used to estimate species nest success. Mayfield nest success is calculated as a percentage of nests failed per day. Mortality rates were calculated by dividing the number of losses by the total number of exposure days for any period, (i.e., incubation period, nestling period, and entire nesting period. Survival rates then were calculated by subtracting the mortality rate from one. Exposure days were calculated by backdating to the midpoint between the last active visit, and the first inactive observation (Mayfield 1975). A nest was considered successful if at least one nestling fledged. A Rayleigh s test (Zar 1999) was used to determine if burrow entrance bearings of nest burrows were uniformly distributed for 2007, 2008, and both years combined. A

12 Rayleigh s test also was used to determine if the bearing of paired random non-nest burrows were uniformly distributed for 2007, 2008, and both years combined. The paired random non-nest burrow was selected randomly by the computer from the 5 nearest non-nest burrows entrance bearings for each nest. Black-tailed prairie dog colonies were assigned to categories based on natural major breaks in density (very low (< 5 prairie dogs/4 ha), low (5-14 prairie dogs/4 ha), medium (15-20 prairie dogs/4 ha), high (21-49 prairie dogs/4 ha), and very high ( 50 prairie dogs/4 ha), for 2007, 2008, and both years combined. The estimated black-tailed prairie dog density associated with each owl nest was based on the closest black-tailed prairie dog count site to that nest or the count site the nest was located in. A paired t-test was used to compare Burrowing Owl survival probability to prairie dog density for 2007, 2008, and both years combined; and Burrowing Owl nest success to prairie dog density for 2007, 2008, and both years combined. All statistical analyses were conducted by using SPSS Statistics GradPack 17.0 with an significance level of 0.05. RESULTS From 24 March through 28 July 2007, 96 potential Burrowing Owl nests were located. Of these, 59 had sufficient data to allow Mayfield method (Mayfield 1975). Of these 33 were successful. From these data the calculated daily survival probability of a nest equaled 0.98 (daily mortality can be calculated by subtracting the daily survival probability from 1) and Mayfield nest success of the Burrowing Owl at Smoky Valley Ranch was 66%. The mean clutch size equaled 5.4 (± 2.23 standard deviation, n = 49); the mean number of nestlings equaled 3.4 (± 1.56 standard deviation, n = 93); and the

mean number of fledged juveniles equaled 2.6 (± 1.34 standard deviation, n = 33) (Table 1.). From 28 May through 11 August 2008, 64 potential Burrowing Owl nests were located. Of these, 46 had sufficient data to be used in the Mayfield method (Mayfield 1975), and of these 18 were successful. From these data the calculated daily survival probability of a nest equaled 0.98 (daily mortality can be calculated by subtracting the daily survival probability from 1) and Mayfield nest success of the Burrowing Owl at Smoky Valley Ranch is 57%. The mean clutch size equaled 4.1 (± 2.43 standard deviation, n = 44); the mean number of nestlings equaled 2.8 (± 1.44 standard deviation, n = 24); and the mean number of fledged juveniles equaled 2.6 (± 1.76 standard deviation, n = 18) (Table 1). The burrow entrance bearings of Burrowing Owl nest burrows were not uniformly distributed for 2007 (z 0.05,27 =1.60), 2008 (z 0.05,23 =1.93), and both years combined (z 0.05,50 =0.05). The burrow entrance bearings for paired non-nest burrows were not uniformly distributed for 2007 (z 0.05,27 =1.12) and both years combined (z 0.05,50 =0.61). However, for 2008 the burrow entrance bearings for paired non-nest burrows were uniformly distributed (z 0.05,23 =3.56). Black-tailed prairie dog density had a positive significant effect on Burrowing Owl nest survival probability for 2007 (t = 13.6; df = 58; P < 0.001), 2008 (t = -7.1; df = 45; P < 0.001), and both years combined (t = -13.8; df = 104; P < 0.001) (Table 3). Black-tailed prairie dog density also had a positive significant effect on Burrowing Owl 13

14 nest success for 2007 (t = 13.0; df = 58; P < 0.001), 2008 (t = -8.3; df = 45; P < 0.001), and both years combined (t = -15.0; df = 104; P < 0.001) (Table 3). DISCUSSION Burrowing Owl nest success at Smoky Valley Ranch was 66% (n=59) in 2007 and 57% (n=46) in 2008. This value was equal to or higher than the 57% and 50% nest success in Oregon in 1980 and 1981, respectively (Green and Anthony 1989). Their study area was in the shrub-steppe zone of northern Gilliam, Morrow, and Umatilla counties in north central Oregon. My value was lower than the 79% nest success found in Oklahoma (Butts 1971) in Beaver County, in 1970. I observed 3.4 chicks per pair (±1.56) in 2007 and 2.8 chicks per pair (±1.44) in 2008, which are comparable to studies in New Mexico (Arrowood et al. 2001), Florida (Mealey 1997), and Montana (Restani et al. 2001). In New Mexico (Arrowood et al. 2001), studies were conducted on New Mexico State University campus, Las Cruces, NM from 1993 to 2000, and the mean number of nestlings per pair ranged from 1.4 to 4.5 during these years. In Florida (Mealy 1997), studies were conducted in Dade and Broward counties, and observed 2.56 (±0.44) chicks per pair in 1988, 2.46 (±0.38) chicks per pair in 1989, and 2.8 (±0.32) chicks per pair in 1990. In Montana (Restani et al. 2001), studies were conducted in Custer and Prairie counties, and observed 2.6 (±0.40) chicks per pair. The observed number of 2.6 fledglings per pair (±1.34) in 2007 and 2.6 fledglings per pair (±1.76) in 2008 at SVR also was comparable to studies in Florida (Mealey 1997). In Florida (Mealy 1997), the observed number of fledglings per pair was 2.37 (±0.40) in 1988, 2.46 (±0.38) in 1989, and 2.73 (±0.34) in 1990.

I found that compass bearings for Burrowing Owl nest burrow entrances were not uniformly distributed, therefore nest burrow entrance bearing might influence nesting success. This is different than what Belthoff and King (2002) found in Idaho. I also found that burrow entrance bearings of paired random non-nest burrows were not uniformly distributed for 2007, similar to findings by Belthoff and King (2002), but were uniformly distributed for 2008. The burrow entrance bearing might be a result of preference by the prairie dog upon burrow excavation rather than a factor selected for by the Burrowing Owl for use as a nest burrow. However since it is not uniformly distributed for the most part, determining the optimum nest entrance bearing for nest success, could be beneficial when installing artificial nest burrows. The Burrowing Owls relies heavily on colonial, burrowing mammals (Lantz and Conway 2009), more specifically active black-tailed prairie dog colonies (Sidle et al. 2001, Desmond 1991, Butts 1973, and Butts and Lewis 1982). Burrowing Owl will use burrows in inactive colonies but use them only until the burrow begins to collapse due to lack of maintenance by the black-tailed prairie dog (Butts and Lewis 1982, pers. obs.). My research showed that nest success and survival probability was influenced significantly by the density of black-tailed prairie dogs within the colonies, which is similar to findings by Desmond (1991). This could be due to availability of food, less vegetative visual obstruction by clipping and grazing of prairie dog, stability of the burrows from black-tailed prairie dog maintenance, as well as the number of individuals to watch for predators and serve as alternative food sources to predators, i.e., dilution effect (Desmond et al. 2000). 15

16 The Burrowing Owl has several predators so the dilution effect probably aids in nest success (Desmond et al. 2000). Seven nests were lost to a known form of predation (three to American badger and four to black-tailed prairie dog). I did not know whether the digging by black-tailed prairie dogs was the cause of nest failure, or if the prairie dog only reclaimed the burrow following nest failure. In July 2008, a gopher snake (Pituophis catenifer) was observed eating a nestling Burrowing Owl (Fig. 8), which resulted in nest success of only one fledgling. At a different nest in July of 2008, an American badger was seen digging out a neighboring prairie dog burrow (Fig. 9). Both of these Burrowing Owl nests were successful, but they also were located within active black-tailed prairie dog colonies with a density category of very high, so the dilution effect could have aided in their success. The tail mount transmitters used during the 2007 field season to track post-fledging dispersal in 2007 and return rate in 2008, failed. The Burrowing Owls were apparently able to pull the transmitters off after several days. Most transmitters were recovered on the ground near the nest site. Only one transmitter, which was later lost, was tracked successfully and led to observing a fledgling with a lubber grasshopper (Brachystola magna) it had caught (Fig. 10). The tail mount transmitter style was chosen to reduce the effect upon the individuals. Since the purpose was to track post-fledging dispersal and return rate the following season, the transmitters only needed to last until individuals returned in March of the following year. Given the molt process of the Burrowing Owl, the tail mount transmitters could be shed in July-September of the following year (Pyle 1997). If I used

a backpack style transmitter then success would have been greater. However, this style is permanent unless the individual is recaptured. In the 2007 field season only one adult Burrowing Owl was captured in the push door trap so recapturing all juveniles upon their potential return in 2008 as adults would have been highly unlikely. CONCLUSIONS Further study of juvenile Burrowing Owl return rate at Smoky Valley Ranch is needed to better understand nesting site fidelity. The push-door trap is very effective for trapping juveniles when they first emerge, when transmitters and/or bands can be attached. However, the use of different types of traps for capturing adult Burrowing Owls is recommended. I suggest use of a noose carpet or Bal-chatri. Migratory populations of Burrowing Owl are known to return to the same site, in some cases the same burrows, the following nesting season especially if they were successful the prior season (MacCracken et al. 1985, Belthoff and King 1997). If they returned to a black-tailed prairie dog colony that underwent control methods and all the prairie dogs are gone, they might have a lower nest success due to lack of black-tailed prairie dog activity. This lower nest success might not be apparent the nesting season immediately following control methods because of the delay in burrow collapse from lack of prairie dog maintenance (Butts and Lewis 1982, Lantz et al. 2007). However, if the colony that was controlled is near an active colony, or the controlled colony was not entirely treated, prairie dogs might recolonize the area and provide an active colony for the Burrowing Owl to nest in. Burrowing Owl needs prairie dog activity for maintaining available burrows, low vegetation cover and to provide alternative prey sources to 17

18 predators (Desmond et al. 2000, Lantz et al. 2007). Timing of control is more complicated. Most control methods are done during the winter months at SVR, however, if control methods are applied during summer months, they could negatively affect Burrowing Owl nesting. If control methods require that burrows be filled in, like they do with the use of Phostoxin, I recommend educating the person doing the treatment to recognize an active Burrowing Owl nest so they do not fill it in. The use of Rozol can still negatively affect nesting Burrowing Owls if used during breeding season through secondary poisoning (Witmer and Fagerstone 2003). If the owl population is migratory, control methods could be done during late fall (end of October to November for SVR) or winter months after owls have migrated. Managing for an ecosystem like the one the black-tailed prairie dog provides, will be more effective than putting all efforts and money into managing for a single species. If landowners and resource managers can determine the best timing and level of control for prairie dogs on their properties, the microhabitats they create will flourish and in turn support the animals that rely on them, like the Burrowing Owl.

19 LITERATURE CITED Arrowood, P. C., C. A. Finley, and B. C. Thompson. 2001. Analyses of Burrowing Owl populations in New Mexico. Journal of Raptor Research 35:362-370. Belthoff, J. R., and R. A. King. 1997. Monitoring Between-Year Movements and Assessment of Artificial Burrow Features Useful in Conservation and Management of Burrowing Owls. A Cooperative Project between Boise State University and the Idaho Bureau of Land Management, Technical Bulletin 2000-03. Boise State University, Boise, Idaho, USA. Belthoff, J. R., and R. A. King. 2002. Nest Site characteristics of Burrowing Owls (Athene cunicularia) in the Snake River Birds of Prey National Conservation Area, Idaho, and applications to artificial burrow Installation. Western North American Naturalist 62:112-119. Botelho, E. S., and P.C. Arrowood. 1995. A novel, simple, safe and effective trap for Burrowing Owls and other fossorial animals. Journal of Field Ornithology 66:380-384. Butts, K. O. 1971. Ecology of Burrowing Owls in Western Oklahoma. A preliminary report. Proceedings of the Oklahoma Academy of Science 51:66-74. Butts, K. O. 1973. Life history and habitat requirements of Burrowing Owls in western Oklahoma. Thesis, Oklahoma State University, Stillwater, Oklahoma. Butts, K.O., and J.C. Lewis. 1982. The importance of prairie dog towns to Burrowing Owls in Oklahoma. Proceedings of the Oklahoma Academy of Science 62:46-52.

Desmond, M. J. 1991. Ecological aspects of burrowing owl nesting strategies. Thesis. University of Nebraska, Lincoln, USA. Desmond, M. J., J. A. Savidge, and K. M Eskridge. 2000. Correlations between Burrowing Owl and black-tailed prairie dog declines: A 7-year analysis. Journal of Wildlife Management 64: 1067-1075. Fahnestock, J.T., and J.K. Detling. 2002. Bison-prairie dog plant interactions in a North American mixed-grass prairie. Oecologia 132,86 95. Green, G. A., and R. G. Anthony. 1989. Nesting success and habitat relationships of Burrowing Owls in the Columbia Basin, Oregon. The Condor 91: 347-354. Haug, E. A., B. A. Millsap, and M. S. Martell. 1993. Burrowing Owl (Athene cunicularia). The Birds of North America. Number 61. Hoogland, J. L. 1981. The evolution of coloniality in white-tailed and black-tailed prairie dogs (Sciuridae: Cynomys leucurus and C. ludovicianus). Ecology 62:252-272. Hoogland, J. L. 1996. Cynomys ludovicianus. Mammalian Species. 535:1-10. Kie, J. G., V. C. Bleich, A. L. Medina, J. D. Yoakum, and J. W. Thomas. 1996. Managing rangelands for wildlife. Pages 663-688 in: Research and Management Techniques for Wildlife and Habitats. Bookhout, Theodore A. editor. Wildlife Society, Bethesda, Maryland. Lantz, S. J., C. J. Conway, and S. H. Anderson. 2007. Multiscale habitat selection by Burrowing Owls in black-tailed prairie dog colonies. The Journal of Wildlife Management 71:2664-2672. 20

Lantz, S. J., and C. J. Conway. 2009. Factors affecting daily nest survival of Burrowing Owls within black-tailed prairie dog colonies. The Journal of Wildlife Management 73:232-241. Lee, C. D., and F.R. Henderson. 1989. Kansas attitudes on prairie dog control. Paper presented at the ninth Great Plains Wildlife Damage Control Workshop, Fort Collins, Colorado, April 18-20. MacCracken, J. G., D. W. Uresk, and R. M. Hansen. 1985. Vegetation and soils of Burrowing Owl nest sites in Conata Basin, South Dakota. The Condor 87:152-154. Manes, R. (editor). 2006. A Management Plan for Smoky Valley Ranch: a Preserve of The Nature Conservancy of Kansas. The Nature Conservancy. Mayfield, H. F. 1975. Suggestions for calculating nest success. The Wilson Bulletin 87:456-466. McDonald, D., N. M. Korfanta, and S. J. Lantz. 2004. The Burrowing owl (Athene cunicularia): A technical conservation assessment. USDA Forest Service, Rocky Mountain Region, Species Conservation Project, Fort Collins, Colorado. Mealey, B. 1997. Reproductive ecology of the Burrowing Owls, Speotyto cunicularia floridana, in Dade and Broward Counties, Florida. Journal of Raptor Research 9:74-79. Miller, B., G. Ceballos, and R. Reading. 1994. The prairie dog and biotic diversity. Conservation Biology 8:677-681. 21

Paine, R. T. 1969. A note on trophic complexity and community stability. The American Naturalist 103:91-93. Plumpton, D. L. 1992. Aspects of nest site selection and habitat use by Burrowing Owls at the Rocky Mountain Arsenal, Colorado. M.S. Thesis. Texas Technical University, Lubbock, Texas, USA. Powell, K. L., R. J. Robel, K. E. Kemp, and M. D. Nellis. 1994. Aboveground counts of black-tailed prairie dogs: temporal nature and relationship to burrow entrance density. The Journal of Wildlife Management 58:361-366. Pyle, P. 1997. Pages 67-68 and 85-86 in Identification Guide to North American Birds: Part I - Columbidae to Ploceidae. Restani, M., L. R. Rau, and D. L. Flath. 2001. Nesting ecology of Burrowing Owls occupying black-tailed prairie dog towns in southeastern Montana. Journal of Raptor Research 35:296-303. Ricketts, T.H., E. Dinerstein, D.M. Olson, C.J. Loucks, W. Eichbaum, D. DellaSala, K. Kavanagh, P.Hedao, P.T. Hurley, K.M. Carney, R. Abell, and S. Walters. 1999. Terrestrial ecoregions of North America: a conservation assessment. Island Press. Washington, DC, USA. Samson, F. B., and F. L. Knopf. 1996. Prairie conservation. Preserving North America s most endangered ecosystem. Island Press, Washington D.C., and Covello, California, USA. Samson, F. B., F.L. Knopf, and W.R. Ostlie. 2004. Great Plains ecosystems: past, present, and future. Wildlife Society Bulletin 32:6-15. 22

23 Sawin, R., J. McCauley, R. Buchanan, and W. Lebsack. 1999. Smoky Valley Ranch Preserve; Geologic Reconnaissance. Kansas Geological Survey, Open-file Report 1999-36. Severson, K. E., and G. E. Plumb. 1998. Comparison of methods to estimate population densities of black-tailed prairie dogs. Wildlife Society Bulletin 26:859-866. Sidle, J. G., M. Ball, T. Byer, J. J. Chynoweth, G. Foli, R. Hodorff, G. Moravek, R. Peterson, and D. N. Svingen. 2001. Occurrence of Burrowing Owls in blacktailed prairie dog colonies on Great Plains National Grasslands. Journal of Raptor Research 35:316-321. Smith, G. A., and M. V. Lomolino. 2004. Black-tailed prairie dogs and the structure of avian communities on the short grass plains. Oecologia 138: 592-602. Thomsen, L. 1971. Behavior and dcology of Burrowing Owls on Oakland Municipal Airport. The Condor 73:177-192. Todd, L. D., R. G. Poulin, R. M. Bringham, E. M. Bayne, and T. I. Wellicome. 2007. Pre-migratory movements by juvenile Burrowing Owls in a patchy landscape. Avian Conservation and Ecology 2:4-15. Turner, M. G., R. H. Gardner, and R. V. O Neill. 2001. Organisms and landscape pattern. Pages 201-247 in: Landscape ecology in theory and practice: pattern and process. Springer Science + Business Media, Inc., New York, USA. Vinton, M. A., and S. L. Collins. 1996. Landscape heterogeneity gradients and habitat structure in native grasslands of the central Great Plains. Pages 3-19 in F.L.

24 Knopf and F.B. Samson, editors. Ecology and conservation of Great Plains vertebrates. Springer Verlag, New York, New York, USA. Winchell, C. S. 1999. An efficient technique to capture complete broods of burrowing owls. Wildlife Society Bulletin 27:193-196. Witmer, G. W., and K. A. Fagerstone. 2003. The use of toxicants in black-tailed prairie dog management: an overview. USDA National Wildlife Research. Fort Collins, Colorado, USA. Zar, J. H. 1999. Biostatistical Analysis. Prentice Hall, Upper Saddle River, New Jersey, USA.

Table 1. Burrowing Owl Mayfield results for 2007 for the Smoky Valley Ranch, Logan County, Kansas. 2007 Total potential nests found 96 Total used in Mayfield 59 Total successful nests 33 Mayfield nest success 67% Daily survival probability 0.98 Daily mortality probability 0.02 26 min. clutch size (eggs/pair) 2007 max. obs. number nestlings/pair max. obs. Fledged/pair Sample size (n) 57 44 33 mean 4.6 3.4 2.6 Standard deviation 2.22 1.56 1.34

27 Table 2. Burrowing Owl Mayfield results for 2008 for the Smoky Valley Ranch, Logan County, Kansas. 2008 Total potential nests found 64 Total used in Mayfield 43 Total successful nests 17 Mayfield nest success 57% Daily survival probability 0.98 Daily mortality probability 0.02 min. clutch size 2008 max. obs. number nestlings max. obs. fledged Sample size (n) 44 24 18 mean 4.1 2.8 2.6 Standard deviation 2.43 1.44 1.76

Table 3. Statistical analyses of paired t-tests for the Smoky Valley Ranch, Logan County, Kansas. t df P Burrowing Owl survival probability relative to prairie dog density 2007 13.6 58 < 0.001 2008-7.1 45 < 0.001 both years combined -13.7 104 < 0.001 Burrowing Owl nest success relative to prairie dog density 2007 13.0 58 < 0.001 2008-8.2 45 < 0.001 both years combined -14.7 104 < 0.001 28

Table 4. Statistical analyses of Wilcoxon Signed Ranks test for the Smoky Valley Ranch, Logan County, Kansas. Burrowing Owl nest burrow entrance bearing relative to paired random non-nest burrow entrance bearing z df P 2007-0.5 25 0.620 2008-1.3 21 0.200 both years combined 0.7 49 0.475 29

Figure 1. Location of Smoky Valley Ranch, Logan County, Kansas, (38.854375; -100.976928). The yellow represents Burrowing Owl breeding range, green represents year-round range and brown represents winter range. 30

Figure 2. Aerial view of Smoky Valley Ranch bisected by the Smoky Hill River. 31

Figure 3. Burrowing Owl breeding distribution (yellow), year-round range (green) and winter range (brown) (modified from Haug et al. 1993). 32

Figure 4. Peeper TM Video Probe (Sandpiper Technologies, Inc.), the burrow camera, consisting of a handheld screen, battery pack and 4.5m cable with infrared camera at the distal end. 33

Figure 5. A two door one-way, push-door trap set up at a Burrowing Owl nest burrow. 34

Figure 6. Tail-mount transmitter on a juvenile Burrowing Owl. 35

Figure 7. Count site configuration for estimating black-tailed prairie dog densities. 36

Figure 8. Gopher snake (Pituophis catenifer) in a Burrowing Owl nest burrow eating a nestling Burrowing Owl. 37

Figure 9. American badger (Taxidea taxus) digging out a black-tailed prairie dog burrow. 38

Figure 10. Burrowing Owl with a tail-mount transmitter antenna showing in the dirt, within a burrow. 39

Appendix A. Black-tailed prairie dog density data for 2007 and 2008 for the Smoky Valley Ranch, Logan County, Kansas. Density categories: VL-very low, L-low, M- medium, H-high, VH-very high. Maximum Year-Colony ID LATITUDE LONGITUDE Density Density Category 07-10b 38.90027-101.00992 0 VL 38.90041-101.01105 38.89937-101.01005 38.89951-101.01122 07-11b 38.88000-100.99285 64 VH 38.87918-100.99300 38.87932-100.99417 38.88019-100.99389 07-11c 38.87836-100.99060 64 VH 38.87752-100.99072 38.87849-100.99169 38.87764-100.99185 07-12a 38.86961-101.00061 13 L 38.86967-101.00177 38.86873-101.00173 38.86874-101.00062 07-12b 38.86871-101.00186 10 L 38.86974-101.00290 38.86881-101.00306 38.86967-101.00179 07-14a 38.86243-100.97236 3 VL 38.86339-100.97205 38.86351-100.97335 38.86253-100.97360 07-14b 38.86304-100.97270 4 VL 38.86348-100.97172 38.86226-100.97212 40

41 38.86269-100.97108 07-14c 38.84790-100.96932 9 L 38.84780-100.97041 38.84703-100.96912 38.84696-100.97030 07-15 38.85766-100.94631 7 L 38.85790-100.94743 38.85674-100.94642 38.85703-100.94759 07-17a 38.84023-100.95852 18 M 38.84079-100.95956 38.83935-100.95939 38.84002-100.96033 07-17b 38.85746-100.97768 12 L 38.85749-100.97880 38.85830-100.97755 38.85839-100.97868 07-18 38.82397-100.96978 5 L 38.82412-100.97088 38.82306-100.96984 38.82320-100.97109 07-19a 38.81607-100.99603 6 L 38.81521-100.99600 38.81524-100.99728 38.81616-100.99719 07-19b 38.81610-100.99887 15 M 38.81608-100.99765 38.81520-100.99776 38.81525-100.99130 07-19c 38.81611-100.99889 22 H

42 38.81604-101.00016 38.81510-100.99899 38.81515-101.00042 07-1 38.93032-100.95473 0 VL 38.93040-100.95597 38.92942-100.95481 38.92947-100.95597 07-20 38.87650-100.96420 12 L 38.87661-100.96564 38.87556-100.96458 38.87569-100.96595 07-23 38.90678-100.97958 2 VL 38.90690-100.98071 38.90593-100.97972 38.90599-100.98087 07-28 38.91742-100.95454 2 VL 38.91743-100.95585 38.91650-100.95464 38.91653-100.95580 07-2 38.90981-100.96337 4 VL 38.90981-100.96457 38.90887-100.96346 38.90887-100.96460 07-30 38.87410-100.98415 13 L 38.87381-100.98298 38.87293-100.98324 38.87310-100.98443 07-3a 38.90141-100.93857 0 VL 38.90147-100.93965 38.90051-100.93863 38.90061-100.93967

43 07-3b 38.90137-100.93737 0 VL 38.90037-100.93638 38.90128-100.93628 38.90014-100.93759 07-41a 38.84428-100.92718 11 L 38.84339-100.92837 38.84432-100.92836 38.84344-100.92723 07-41b 38.84531-100.92707 7 L 38.84522-100.92824 38.84428-100.92719 38.84438-100.92838 07-5-b 38.89413-100.92683 0 VL 38.89412-100.92808 38.89312-100.92688 38.89328-100.92835 07-6a 38.89237-100.95951 15 M 38.89132-100.95975 38.89163-100.96088 38.89250-100.96071 07-6b 38.88384-100.96271 15 M 38.88372-100.96153 38.88464-100.96136 38.88471-100.96255 07-7 38.88524-100.98262 21 H 38.88536-100.98370 38.88441-100.98262 38.88431-100.98380 07-9 38.89433-100.99736 16 M 38.89452-100.99856