Terry L. Shaffer

A UNIFIED APPROACH TO ANALYZING NEST SUCCESS

Abstract Logistic regression has become increasingly popular for modeling nest success in terms of nest-specific explanatory variables. However, logistic regression models for nest fate are inappropriate when applied to data from nests found at various ages, for the same reason that the apparent estimator of nest success is biased (i.e. older clutches are more likely to be successful than younger clutches). A generalized linear model is presented and illustrated that gives ornithologists access to a flexible, suitable alternative to logistic regression that is appropriate when exposure periods vary, as they usually do. Unlike the Mayfield method (1961, 1975) and the logistic regression method of Aebischer (1999), the logistic-exposure model requires no assumptions about when nest losses occur. Nest survival models involving continuous and categorical explanatory variables, multiway classifications, and time-specific (e.g. nest age) and random effects are easily implemented with the logistic-exposure model. Application of the model to a sample of Yellow-breasted Chat (Icteria virens) nests shows that logistic-exposure estimates for individual levels of categorical explanatory variables agree closely with estimates obtained with Johnson (1979) constant-survival estimator. Use of the logistic-exposure method to model time-specific effects of nest age and date on survival of Blue-winged Teal (Anas discors) and Mallard (A. platyrhynchos) nests gives results comparable to those reported by Klett and Johnson (1982). However, the logistic-exposure approach is less subjective and much easier to implement than Klett and Johnsons method. In addition, logistic-exposure survival rate estimates are constrained to the (0,1) interval, whereas Klett and Johnson estimates are not. When applied to a sample of Mountain Plover (Charadrius montanus) nests, the logistic-exposure method gives results either identical to, or similar to, those obtained with the nest survival model in program MARK (White and Burnham 1999). I illustrate how the combination of generalized linear models and information-theoretic techniques for model selection, along with commonly available statistical software, provides ornithologists with a powerful, easily used approach to analyzing nest success.

Duck Nest Success in the Prairie Pothole Region

We estimated nest success of mallard (Anas platyrhynchos), gadwall (A. strepera), blue-winged teal (A. discors), northern shoveler (A. clypeata), and northern pintail (A. acuta) for 5 regions in North Dakota, South Dakota, and Minnesota, for 1-3 periods between 1966 and 1984, and for 8 habitat classes. We obtained composite estimates of nest success for regions and periods by weighting each habitat proportional to the number of nest initiations. The distribution of nest initiations was derived from estimates of breeding populations, preferences of species for nesting habitats, and availability of habitats. Nest success rates ranged from 15,000 records of nests found in the United States portion of the Prairie Pothole Region (Fig. 1). Supplemental information on breeding population levels, nesting habitat availability, and habitat use by nesting 432 DUCK NEST SUCCESS * Klett et al.

TIME-SPECIFIC VARIATION IN PASSERINE NEST SURVIVAL: NEW INSIGHTS INTO OLD QUESTIONS

Abstract Understanding nest survival is critical to bird conservation and to studies of avian life history. Nest survival likely varies with nest age and date, but until recently researchers had only limited tools to efficiently address those sources of variability. Beginning with Mayfield (1961), many researchers have averaged survival rates within time-specific categories (e.g. egg and nestling stages; early and late nesting dates). However, Mayfield’s estimator assumes constant survival within categories, and violations of that assumption can lead to biased estimates. We used the logistic-exposure method to examine nest survival as a function of nest age and date in Clay-colored Sparrows (Spizella pallida) and Vesper Sparrows (Pooecetes gramineus) breeding in north-central North Dakota. Daily survival rates increased during egg laying, decreased during incubation to a low shortly after hatch, and then increased during brood rearing in both species. Variation in survival with nest age suggests that traditional categorical averaging using Mayfield’s or similar methods would have been inappropriate for this study; similar variation may bias results of other studies. Nest survival also varied with date. For both species, survival was high during the peak of nest initiations in late May and early June and declined throughout the remainder of the nesting season. On the basis of our results, we encourage researchers to consider models of nest survival that involve continuous time-specific explanatory variables (e.g. nest age or date). We also encourage researchers to document nest age as precisely as possible (e.g. by candling eggs) to facilitate age-specific analyses. Models of nest survival that incorporate time-specific information may provide insights that are unavailable from averaged data. Determining time-specific patterns in nest survival may improve our understanding of predator-prey interactions, evolution of avian life histories, and aspects of population dynamics that are critical to bird conservation.

Impact of the Conservation Reserve Program on duck recruitment in the U.S. Prairie Pothole Region

The U.S. Department of Agriculture (USDA)s Conservation Reserve Program (CRP) resulted in the conversion of about 1.9 million ha of cropland to perennial grass cover in the Prairie Pothole Region of North Dakota, South Dakota, and northeastern Montana by 1992. Many wildlife managers believed this cover would provide benefits to wildlife, including upland nesting ducks. During 1992-1995, we evaluated success of 5 duck species nesting in CRP fields and nearby Waterfowl Production Areas (WPA) throughout the region. We examined relationships between daily survival rates (DSR) of duck nests in CRP cover and landscape-level habitat and population parameters. We computed DSR of duck nests in other major cover types in our study area from data collected during 1980-1984 (pre-CRP) and 1990-1994 (CRP) periods. We then applied recruitment models to estimate duck production in our study area during peak CRP years (1992-1997) and compared these results with those that simulated the scenario in which cropland was in place of CRP cover (i.e., the CRP had not occurred). DSR were higher in all habitats combined during the CRP period compared to the pre-CRP period. Regressions of DSR in CRP cover on the percent of each study plot in perennial cover and geographic location were significant (P < 0.01) for 4 of 5 duck (Anas spp.) species. Estimated nest success and recruitment rates for the 5 species combined during 1992-1997 were 46% and 30% higher, respectively, with CRP cover on the landscape compared to a scenario where we simulated cropland in place of CRP. Our model estimated an additional 12.4 million recruits from our study area to the fall flight as a consequence of the CRP during 1992-1997. Our results document benefits to 5 duck species in the northern plains associated with a farm program that provided financial incentives to landowners for planting undisturbed grass cover as an alternative to annual crops.

The Farm Bill and Duck Production in the Prairie Pothole Region: Increasing the Benefits

Abstract The Food Security Act of 1985 contained provisions that affected wildlife conservation nationwide. Two provisions that most benefited waterfowl populations in the Prairie Pothole Region (PPR) were the Conservation Reserve Program (CRP) and “Swampbuster” (wetland conservation). Permanent cover established under the CRP provides attractive nesting habitat for upland-nesting ducks that is more secure than other major habitats. Swampbuster has prevented drainage of wetlands vital to breeding duck pairs. In 2007 many CRP contracts will expire. Deliberations will begin in late 2006 regarding the next Farm Bill. The United States Department of Agriculture needs sound biological information and scientific analyses to help establish wildlife priorities in the Farm Bill. We used data from breeding duck population and wetland habitat surveys to develop models for 5 species of upland-nesting ducks and applied these models to >2.6 million wetlands in a digital database for the PPR in North and South Dakota, USA. We used geographic information systems techniques to identify locations in the PPR where CRP cover would be accessible to the greatest number of nesting hens. We then summarized distribution of current CRP contracts relative to distribution of upland-breeding ducks. We also used our models to predict change in the breeding duck population (landscape carrying capacity) that might occur if certain wetlands were exempt from the Swampbuster provision. Our analyses showed that 75% of CRP contracts as of July 2005 were in areas accessible to high or medium numbers of breeding ducks and 25% were in areas of low populations. We suggest a method to prioritize CRP extensions and reenrollment of current contracts or target new contracts to maintain or increase duck production. Additionally, our models suggested that if the Swampbuster provision were removed from future Farm Bills and protected wetland were drained, this area of the PPR could experience a 37% decline in the waterfowl populations we studied.

FACTORS AFFECTING SONGBIRD NEST SURVIVAL IN RIPARIAN FORESTS IN A MIDWESTERN AGRICULTURAL LANDSCAPE

Abstract We investigated factors affecting nest success of songbirds in riparian forest and buffers in northeastern Missouri. We used an information-theoretic approach to determine support for hypotheses concerning effects of nest-site, habitat-patch, edge, and temporal factors on nest success of songbirds in three narrow (55–95 m) and three wide (400–530 m) riparian forests with adjacent grassland-shrub buffer strips and in three narrow and three wide riparian forests without adjacent grassland-shrub buffer strips. We predicted that temporal effects would have the most support and that habitat-patch and edge effects would have little support, because nest predation would be great across all sites in the highly fragmented, predominantly agricultural landscape. Interval nest success was 0.404, 0.227, 0.070, and 0.186, respectively, for Gray Catbird (Dumetella carolinensis), Northern Cardinal (Cardinalis cardinalis), Indigo Bunting (Passerina cyanea), and forest interior species pooled (Acadian Flycatcher [Empidonax virescens], Wood Thrush [Hylocichla mustelina], Ovenbird [Seiurus aurocapillus], and Kentucky Warbler [Oporornis formosus]). The effect of nest stage on nest success had the most support; daily nest success for Gray Catbird and Indigo Bunting were lowest in the laying stage. We found strong support for greater nest success of Gray Catbird in riparian forests with adjacent buffer strips than in riparian forests without adjacent buffer strips. Patch width also occurred in the most-supported model for Gray Catbird, but with very limited support. The null model received the most support for Northern Cardinal. Riparian forests provided breeding habitat for area-sensitive forest species and grassland-shrub nesting species. Buffer strips provided additional breeding habitat for grassland-shrub nesting species. Interval nest success for Indigo Bunting and area-sensitive forest species pooled, however, fell well below the level that is likely necessary to balance juvenile and adult mortality, which suggests that when riparian forests are located within agricultural landscapes, the potential even for wide riparian forests with adjacent buffer strips to provide high-quality breeding habitat is severely diminished for some species.

Nest Survival of Clay-Colored and Vesper Sparrows in Relation to Woodland Edge in Mixed-Grass Prairies

Abstract The quantity and quality of northern mixed-grass prairie continues to decline because of conversion to agriculture, invasion of woody and exotic plants, and disruption of important ecological processes that shape grasslands. Declines in grassland bird populations in North Dakota, USA, have coincided with these largely anthropogenic alterations to prairie habitat. In grasslands of north-central and northwestern North Dakota, woody plants have increased due primarily to fire suppression, extirpation of bison (Bos bison), and widescale planting of tree shelter belts. In northern grasslands, effects of woody vegetation on survival of grassland birds are poorly understood, and conclusions are based mainly on studies conducted outside the region. We examined nest survival of clay-colored sparrows (Spizella pallida) and vesper sparrows (Pooecetes gramineus) relative to the distance nests were located from aspen (Populus tremuloides) woodland edges and relative to other habitat features near the nest. Clay-colored and vesper sparrow nest survival was higher for nests located near woodland edges, nests with greater cover of Kentucky bluegrass (Poa pratensis), and nests more concealed by vegetation. Vesper sparrow nest survival increased as the percent cover of tall shrubs near the nest increased. Based on video-camera data, the 13-lined ground squirrel (Spermophilus tridecemlineatus) was the most common predator of sparrow eggs and young. Thirteen-lined ground squirrels were more common far from woodland edges than near, and this pattern may, in part, explain clay-colored and vesper sparrow nest survival in relation to woodland edges. In contrast to our results, studies conducted in other grassland systems generally report lower nest survival for grassland birds nesting near trees and shrubs. This disparity in results demonstrates the need to identify specific nest predators and their distributions with respect to important habitat features because these data can be important in explaining—and perhaps predicting—patterns of nest predation.

FLORISTIC QUALITY ASSESSMENT OF ONE NATURAL AND THREE RESTORED WETLAND COMPLEXES IN NORTH DAKOTA, USA

AbstractFloristic quality assessment is potentially an important tool for conservation efforts in the northern Great Plains of North America, but it has received little rigorous evaluation. Floristic quality assessments rely on coefficients assigned to each plant species of a region’s flora based on the conservatism of each species relative to others in the region. These “coefficients of conservatism” (C values) are assigned by a panel of experts familiar with a region’s flora. The floristic quality assessment method has faced some criticism due to the subjective nature of these assignments. To evaluate the effect of this subjectivity on floristic quality assessments, we performed separate evaluations of the native plant communities in a natural wetland complex and three restored wetland complexes. In our first assessment, we used C values assigned “subjectively” by the Northern Great Plains Floristic Quality Assessment Panel. We then performed an independent assessment using the observed distributions of species among a group of wetlands that ranged from highly disturbed to largely undisturbed (data-generated C values). Using the panel-assigned C values, mean C values (

Effects of Prescribed Fire on Vegetation and Passerine Birds in Northern Mixed-Grass Prairie

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