Marsha A. Sovada

Differential effects of coyotes and red foxes on duck nest success

Low recruitment rates prevail among ducks in the Prairie Pothole Region of North America, primarily because of high nest depredation rates. The red fox (Vulpes vulpes) is a major predator of duck eggs, but fox abundance is depressed by coyotes (Canis latrans). We tested the hypothesis that nest success of upland-nesting ducks is higher in areas with coyotes than in areas with red foxes. We conducted the study during 1990-92 in uplands of 36 areas managed for nesting ducks in North Dakota and South Dakota. Overall nest succes averaged 32% (95% CI = 25-40) on 17 study areas where coyotes were the principal canid and 17% (CI = 11-25) on 13 study areas where red foxes were the principal canid (P = 0.01). Both canids were common on 6 other areas, where nest success averaged 25% (CI = 13-47). Habitat composition, predator communities with the exception of canids, and species composition of duck nests in coyote and red fox areas were similar overall. Upon examining only nests with ≥6 eggs on the last visit prior to hatch or depredation, we determined nests with evidence characteristic of fox predation accounted for 4% of depredated nests in coyote areas and 27% in fox areas (P = 0.001). An expanding coyote population is contributing to higher overall nest success. Management of coyotes may be an effective method for increasing duck nest success

Predator selection of prairie landscape features and its relation to duck nest success

Mammalian predation is a major cause of mortality for breeding waterfowl in the U.S. Northern Great Plains, and yet we know little about the selection of prairie habitats by predators or how this influences nest success in grassland nesting cover. We selected 2 41.4-km 2 study areas in both 1996 and 1997 in North Dakota, USA, with contrasting compositions of perennial grassland. A study area contained either 15-20% perennial grassland (Low Grassland Composition LGC]) or 45-55% perennial grassland (High Grassland Composition [HGC]). We used radiotelemetry to investigate the selection of 9 landscape cover types by red fox (Vulpes vulpes) and striped skunk (Mephitis mephitis), while simultaneously recording duck nest success within planted cover. The cover types included the edge and core areas of planted cover, wetland edges within planted cover or surrounded by cropland, pastureland, hayland, cropland, roads, and miscellaneous cover types. Striped skunks selected wetland edges surrounded by agriculture over all other cover types in LGC landscapes (P-values for all pairwise comparisons were <0.05). Striped skunks also selected wetland edges surrounded by agriculture over all other cover types in HGC landscapes (P < 0.05), except for wetland edges within planted cover (P = 0.12). Red foxes selected the edge and core areas of planted cover, as well as wetland edges within planted cover in LGC landscapes (i.e., they were attracted to the more isolated patches of planted cover). However, in HGC landscapes, red foxes did not select interior areas of planted cover (i.e., core areas of planted cover and wetland edges in planted cover) as frequently as edges of planted cover (P< 0.05). Red foxes selected core areas of planted cover more frequently in LGC than in HGC landscapes (P< 0.05) and selected pastureland more frequently in HGC than in LGC landscapes (P< 0.05). Furthermore, red foxes selected the isolated patches of planted cover more than pastureland in LGC landscapes (P< 0.05). Duck nest success was greater in HGC landscapes than in LGC landscapes for planted-cover core (P < 0.0001), planted-cover edge (P< 0.001) and planted cover-wetland edge (P< 0.001). Both the increased amount of planted-cover core area and the increased pastureland selection in HGC landscapes may have diluted predator foraging efficiency in the interior areas of planted cover and contributed to higher nest success in HGC landscapes. Our observations of predator cover-type selection not only support the restoration and management of large blocks of grassland but also indicate the influence of alternative cover types for mitigating nest predation in the Prairie Pothole Region.

Relationships of habitat patch size to predator community and survival of duck nests

We studied duck nest success and predator community composition in relation to size of discrete patches of nesting cover in the Prairie Pothole Region (PPR) of the United States in 1993-95 We focused on nests in uplands that were seeded to perennial grasses and forbs and enrolled in the Conservation Reserve Program (CRP) in Minnesota, North Dakota, and South Dakota. We estimated daily survival rates (DSRs) of upland duck nests and indices of activity for red foxes (Vulpes vulpes), coyotes (Canis latrans), American badgers (Taxidea taxus), striped skunks (Mephitis mephitis) and Franklins ground squirrels (Spermophilus franklinii) and related these variables to habitat patch size. The effect of patch size (small vs. large) on estimated annual mean DSR was dependent on date of nest initiation (early vs. late) and year. Examination of within-year comparisons for early and late nests suggested that DSR was generally greater in larger habitat patches Activity indices for the 5 mammalian nest predators were influenced differently by year, location, and patch size. Activity indices of the red fox were greatest in small patches. Coyote indices were the most inconsistent, demonstrating a year × location × patch size interaction. Activity indices of the striped skunk and American badger varied only among years. Franklins ground squirrel indices were affected by study area location, with higher indices in the southeast than the northwest. Red fox activity was weakly correlated with that of the striped skunk and coyote. Although a positive relationship between habitat patch size and nest success probably exists, we believe the experiment to fully test this hypothesis will continue to be elusive.

Causes and rates of mortality of swift foxes in western Kansas

Knowledge of mortality factors is important for developing strategies to conserve the swift fox (Vulpes velox), a species being considered for listing under the Endangered Species Act, but available information about swift fox mortality is inadequate. We used radiotelemetry techniques to examine the magnitude and causes of mortality of swift fox populations in 2 study areas in westem Kansas. One study area was predominantly cropland, the other rangeland, Mortality rates, calculated using Kaplan-Meier estimation techniques in a staggered entry design, were 0.55 ± 0.08 (x ± SE) for adult and 0.67 ± 0.08 for juvenile swift foxes. We did not detect differences between study areas in mortality rates for adults or juveniles. Predation by coyotes (Canis latrans) was the major cause of mortality for adult and juvenile swift foxes in both study areas, and vehicle collision was an important mortality factor for juveniles in the cropland study area. No mortality was attributed to starvation or disease.

Analysis of Predator Movement in Prairie Landscapes with Contrasting Grassland Composition

Abstract Mammalian predation influences waterfowl breeding success in the U.S. northern Great Plains, yet little is known about the influence of the landscape on the ability of predators to find waterfowl nests. We used radiotelemetry to record nightly movements of red foxes (Vulpes vulpes) and striped skunks (Mephitis mephitis) in two 41.4-km2 study areas in North Dakota. Study areas contained either 15–20% grassland (low grassland composition) or 45–55% grassland (high grassland composition). Grasslands included planted cover, pastureland, and hayland. We predicted that the type and composition of cover types in the landscape would influence both predator movement across the landscape (as measured by the fractal dimension and displacement ratio) as well as localized movement (as measured by the rate of movement and turning angle between locations) within patches of different cover types. Red fox movements were straighter (lower fractal dimensions and higher displacements) across landscapes with a low grassland composition, indicating directed movement between the more isolated patches of planted cover. Striped skunk movements did not differ between landscape types, illustrating their movement along wetland edges, which had similar compositions in both landscape types. The high variability in turning angles by red fox in planted cover and pastureland in both landscape types is consistent with restricted-area foraging. The high rate of movement by red foxes in planted cover and by striped skunks in wetland edges suggests that spatial memory may influence movement patterns. Understanding the behavior of predators in fragmented prairie landscape is essential for managing breeding habitat for grassland birds and for predicting the spatial and temporal dynamics of predators and their prey.

LANDSCAPE COMPOSITION, PATCH SIZE, AND DISTANCE TO EDGES: INTERACTIONS AFFECTING DUCK REPRODUCTIVE SUCCESS

Prairies and other North American grasslands, although highly fragmented, provide breeding habitat for a diverse array of species, including species of tremendous economic and ecological importance. Conservation and management of these species re- quires some understanding of how reproductive success is affected by edge effects, patch size, and characteristics of the landscape. We examined how differences in the percentage of grassland in the landscape influenced the relationships between the success of nests of upland-nesting ducks and (1) field size and (2) distance to nearest field and wetland edges. We collected data on study areas composed of 15-20% grassland and areas composed of 45-55% grassland in central North Dakota, USA during the 1996 and 1997 nesting seasons. Daily survival rates (DSRs) of duck nests were greater in study areas with 45-55% grassland than with 15-20% grassland. Within study areas, we detected a curvilinear relationship between DSR and field size: DSRs were highest in small and large fields and lowest in moderately sized fields. In study areas with 15-20% grassland, there was no relationship between probability of hatching and distance to nearest field edge, whereas in study areas with 45-55% grassland, there was a positive relationship between these two variables. Results of this study support the conclusion that both landscape composition and config- uration affect reproductive success of ground-nesting birds. We are prompted to question conservation strategies that favor clustering moderately sized patches of nesting habitat within agricultural landscapes because our results show that such patches would have low nest success, most likely caused by predation. Understanding the pattern of nest success, and the predator-prey mechanisms that produce the pattern, will enable design of patch configurations that are most conducive to meeting conservation goals.

Seasonal Food Habits of Swift Fox (Vulpes velox) in Cropland and Rangeland Landscapes in Western Kansas

Abstract Food habits of swift foxes (Vulpes velox) occupying two distinct landscapes (dominated by cropland versus rangeland) in western Kansas were determined by analysis of scats collected in 1993 and 1996. Frequencies of occurrence of prey items in scats were compared between cropland and rangeland areas by season. Overall, the most frequently occurring foods of swift foxes were mammals (92% of all scats) and arthropods (87%), followed by birds (24%), carrion (23%), plants (15%) and reptiles (4%). No differences were detected between landscapes for occurrence of mammals, arthropods or carrion in any season (P ≥ 0.100). Plants, specifically commercial sunflower seeds, were consumed more frequently in cropland than in rangeland in spring (P = 0.004) and fall (P = 0.001). Birds were more common in the swift fox diet in cropland than in rangeland during the fall (P = 0.008), whereas reptiles occurred more frequently in the diet in rangeland than in cropland during spring (P = 0.042). Variation in the diet of the swift fox between areas was most likely due to its opportunistic foraging behavior, resulting in a diet that closely links prey use with availability.

MARKOV CHAIN MONTE CARLO ESTIMATION OF SPECIES DISTRIBUTIONS: A CASE STUDY OF THE SWIFT FOX IN WESTERN KANSAS

Abstract Accurate maps of species distributions are essential tools for wildlife research and conservation. Unfortunately, biologists often are forced to rely on maps derived from observed occurrences recorded opportunistically during observation periods of variable length. Spurious inferences are likely to result because such maps are profoundly affected by the duration and intensity of observation and by methods used to delineate distributions, especially when detection is uncertain. We conducted a systematic survey of swift fox (Vulpes velox) distribution in western Kansas, USA, and used Markov chain Monte Carlo (MCMC) image restoration to rectify these problems. During 1997–1999, we searched 355 townships (ca. 93 km2) 1–3 times each for an average cost of

ASSESSING SMALL MAMMAL ABUNDANCE WITH TRACK-TUBE INDICES AND MARK-RECAPTURE POPULATION ESTIMATES

7,315 per year and achieved a detection rate (probability of detecting swift foxes, if present, during a single search) of θ̂ = 0.69 (95% Bayesian confidence interval [BCI] = [0.60, 0.77]). Our analysis produced an estimate of the underlying distribution, rather than a map of observed occurrences, that reflected the uncertainty associated with estimates of model parameters. To evaluate our results, we analyzed simulated data with similar properties. Results of our simulations suggest negligible bias and god good precision when probabilities of detection on ≥1 survey occasions (cumulative probabilities of detection) exceed 0.65. Although the use of MCMC image restoration has been limited by theoretical and computational complexities, alternatives do not possess the same advantages. Image models accommodate uncertain detection, do not require spatially independent data or a census of map units, and can be used to estimate species distributions directly from observations without relying on habitat covariates or parameters that must be estimated subjectively. These features facilitate economical surveys of large regions, the detection of temporal trends in distribution, and assessments of landscape-level relations between species and habitats. Requirements for the use of MCMC image restoration include study areas that can be partitioned into regular grids of mapping units, spatially contagious species distributions, reliable methods for identifying target species, and cumulative probabilities of detection ≥0.65.

Foods of American Badgers in West-central Minnesota and Southeastern North Dakota During the Duck Nesting Season

Abstract We compared track-tube sampling with mark–recapture livetrapping and evaluated a track-tube index, defined as the number of track tubes with identifiable small mammal tracks during a 4-night period, as a predictor of small mammal abundance estimates in North Dakota grasslands. Meadow voles (Microtus pennsylvanicus) were the most commonly recorded species by both methods, but were underrepresented in track-tube sampling, whereas 13-lined ground squirrels (Spermophilus tridecemlineatus) and Franklins ground squirrels (S. franklinii) were overrepresented in track-tube sampling. Estimates of average species richness were lower from track tubes than from livetrapping. Regression models revealed that the track-tube index was at best a moderately good predictor of small mammal population estimates because both the form (linear versus curvilinear) and slope of the relationship varied between years. In addition, 95% prediction intervals indicated low precision when predicting population estimates from new track-tube index observations. Track tubes required less time and expense than mark–recapture and eliminated handling of small mammals. Using track tubes along with mark–recapture in a double sampling for regression framework would have potential value when attempting to estimate abundance of small mammals over large areas.