Response of Grassland Birds in Sand Shinnery Oak Communities Restored Using Tebuthiuron and Grazing in Eastern New Mexico Lindsay A. Smythe1,2,3 and David A. Haukos1 Abstract Sand shinnery oak (Quercus havardii) communities are a unique component of grassland bird habitat in eastern New Mexico and have been impacted by human activities for decades. These communities are frequently managed with livestock grazing and herbicide application for shrub control, strategies that potentially can be used to restore the historical shrub–grass composition of this plant community. During spring migration and the breeding seasons of 2004 and 2005, we compared density and community structure of grassland bird species among four combinations of tebuthiuron application and grazing treatments that were being evaluated for restoration of shinnery oak communities. We performed biweekly point transects on sixteen 65-ha study plots in these communities. Density of all avian species combined did not differ between grazed and ungrazed plots. Tebuthiuron-treated plots had a 40% higher average density for combined species than untreated plots. There was a 41% higher average density of all species during spring 2005 than 2004, but density was similar during the breeding season of both years. These trends were predominantly influenced by densities of migratory Cassin’s Sparrow (Aimophila cassinii), which were greater in tebuthiuron-treated plots in both years. Densities of resident Meadowlarks (Sturnella spp.) exhibited little response to tebuthiuron or grazing treatments. Avian species richness, evenness, and diversity were only minimally affected by the tebuthiuron and grazing treatments. This study occurred over a period of highly variable precipitation, so future assessments, spanning longer wet–dry cycles and maturing plant communities, may be necessary to completely determine avian response to these restoration efforts. Introduction The short- and mixed-grass prairie ecosystem of eastern New Mexico provides habitat for many species of grassland birds, including numerous migratory species of concern (Rich et al. 2004; NMPIF 2007). Data from the North American Breeding Bird Survey demonstrate consistent declines across the breeding range of most grassland bird species between 1966 and 2005 (Peterjohn & Sauer 1999; Sauer et al. 2005). Furthermore, populations of grassland birds showed the most widespread and greatest decline of all surveyed avian groups (Herkert 1995). Habitat destruction, fragmentation, and degradation appear to be the underlying influences in these population trends, although these effects may manifest differently among species (Peterjohn & Sauer 1999). Grassland birds tend to select habitats based on vegetation structure (Wiens 1973), and thus modification of grassland habitat may affect the abundance and distribution of bird populations (Martin 1995). Sand shinnery oak (Quercus havardii) communities are a unique vegetation type interspersed in the more traditionally defined grasslands of eastern New Mexico, exemplified by shrubs such as Sand sage (Artemisia filifolia) and Sand shinnery oak, as well as several grasses including Sand bluestem (Andropogon hallii), Big bluestem (An. gerardii), Purple three-awn (Aristida purpurea), Hairy grama (Bouteloua hirsuta), Fall witchgrass (Leptoloma cognatum), Little bluestem (Schizachyrium scoparium), and Sand dropseed (Sporobolus cryptandrus) (Martin 1990; NMPIF 2007; plant names follow Hitchcock 1971). These communities were historically codominated by shrubs and grasses (Martin 1990; NMPIF 2007). There is no evidence that shinnery oak invades overgrazed rangeland, but this species is an effective water gatherer and, when given an advantage, may nearly eliminate associated plants due to the effects of shading and moisture competition (Peterson & Boyd 1998). Thus, unmanaged grazing can change the community composition, resulting in decreased grass production and greater frequency of shinnery oak (Peterson & Boyd 1998). In many areas, shinnery oak has become dominant or nearly a monoculture, and there is interest in restoring the communities to a more historic grass/shrub balance. Tools available for restoring shinnery communities to their historical vegetation composition and structure 1 Department of Natural Resources Management, Texas Tech University, Box 42125, Lubbock, TX 79409, U.S.A. 2 Address correspondence to L. A. Smythe, email lindsay_smythe@fws.gov 3 Present address: Kofa National Wildlife Refuge, 9300 E. 28th street, Yuma, AZ 85365, U.S.A. Ó 2008 Society for Ecological Restoration International doi: 10.1111/j.1526-100X.2008.00443.x Restoration Ecology Key words: density, DISTANCE, grassland birds, grazing, herbicide, shinnery oak. 1 Response of Grassland Birds in Sand Shinnery Oak Communities include shinnery oak control and managed livestock grazing. In New Mexico, the predominant method of shinnery oak control is herbicide application (Peterson & Boyd 1998). Between 1981 and 1993, the U.S. Bureau of Land Management treated 40,469 ha of shinnery oak in New Mexico with the herbicide tebuthiuron (N-[5-(1,1-dimethylethyl)-1, 3, 4-thiadiazol-2-yl]-N, N9-dimethylurea) (USDI BLM 1997). Although grassland bird communities evolved in conjunction with seasonal ungulate grazing, the effects of cattle grazing systems on bird communities and their habitats are neither uniform nor easily defined (Wiens & Dyer 1975). According to Peterson and Boyd (1998), about 20 songbird species nest in Sand shinnery oak communities and approximately 80 additional species historically used this habitat, but none are endemic to the community. NMPIF (2007) has identified eight bird priority species of concern using Sand shinnery oak communities. These communities have as much value for their support of prairie breeding species during migration and winter as they do for birds that breed in shinnery oak communities (Rich et al. 2004). Thus, use of grazing and herbicides to restore shinnery habitats may impact migratory bird populations throughout the southwestern United States. Also, many previous studies assumed that shinnery oak communities were pristine, when historically these communities supported a greater grass component (Peterson & Boyd 1998). Our objective was to compare density, richness, evenness, and diversity of breeding grassland bird species among four combinations of herbicide and grazing treatments to determine if any of these treatments negatively impacted grassland bird community parameters. Methods Study Area Our study site was in Roosevelt County, eastern New Mexico, U.S.A., centered at lat 33.66°N, long 2103.13°W, NAD27. The study area consisted of 16 plots of approximately 65 ha each (one plot was 80 ha). Tebuthiuron (0.75 kg/ha) was applied aerially to 532 ha of private land in 2000, which was adjacent to 518 ha of the state-owned North Bluit Prairie Chicken Area that was not treated and represented the extant shinnery oak/grassland community. A year after the tebuthiuron application, a 6.5fold increase in herbaceous plant production and a 29-fold increase in grass seed production was recorded on the treated areas (Dixon, unpublished data). The control area had not been grazed for at least 7 years before the study began; the tebuthiuron-treated areas had not been grazed for at least 5 years before the study began, 2 years pretebuthiuron treatment, and 3 years posttebuthiuron treatment. In 2003, the treated and untreated areas were crossfenced into eight plots each, and managed grazing was applied randomly to half of the tebuthiuron-treated and 2 untreated areas. The grazing treatment was a short-duration system in which plots were grazed once during the dormant season (January and February) and once during the growing season (July). Stocking rate was calculated each season based on measured forage production and designed to take 25% of the available herbaceous material per season. From 2003 to 2005, stocking rate ranged from 584 to 2,224 animal-days on the tebuthiuron-treated plots and from 147 to 556 animal-days on the untreated plots (Smythe 2006). Higher stocking rates occurred during the dormant season. Study plots consisted of two treatments arranged in four combinations: tebuthiuron with grazing; tebuthiuron without grazing; no tebuthiuron with grazing; and a control of no tebuthiuron or grazing (Smythe 2006). Point Transects In each of the 16 treatment plots (four replications of four treatment combinations), we randomly placed four 4-ha subplots. Point transects for avian sampling were set up in a grid of four points, 100 m apart, per subplot (Ralph et al. 1993; Buckland et al. 2001). To estimate avian density in each treatment combination, we surveyed at least two of the four points in each subplot every month (three per subplot were surveyed in February to ensure adequate detections for distance sampling). Points in the same grid were not sampled on the same day to maintain independence. We conducted point transects from February through July of 2004 and 2005 to record spring migrant species and breeding birds. From February through April, we conducted point transects from sunrise until four were completed (approximately 0600–1000); from May through July, we conducted transects from sunrise until two were completed (approximately 0445–0800) because the birds became inactive more quickly as the weather grew warmer. This sampling design differed slightly from the recommended survey period (May–July) and time frame (0500– 0900) for stable detectability of breeding birds (Ralph et al. 1993), but distance sampling provides a reliable estimate of density in the face of variability in detection due to factors such as cue production (e.g., singing) and environmental differences (Buckland et al. 2001). We recorded all avian species seen or heard within 5 minutes at the point. To reliably estimate absolute density, we recorded distance to each bird seen in the point transect with a Leica laser rangefinder (Ransom & Pinchak 2003). Birds that were heard but not seen were not included in density calculations. Density was reported as individuals per hectare for all species except Grasshopper Sparrows (Ammodramus savannarum) and Cassin’s Sparrows (Aimophila cassinii), which were reported as singing males per hectare because generally only singing males were detected during surveys. We grouped Eastern Meadowlarks (Sturnella magna) and Western Meadowlarks (S. neglecta) for analysis due to difficulty in identifying nonsinging individuals by sight in the field. Restoration Ecology Response of Grassland Birds in Sand Shinnery Oak Communities Statistical Analyses Bird density was estimated with program DISTANCE (Buckland et al. 2001; Thomas et al. 2003). The detection function model with the lowest AICc value (Akaike’s information criterion adjusted for small sample sizes) was selected as the best fit (Anderson & Burnham 2002). Density of each species was estimated for each month. The tebuthiuron-treated and untreated areas were analyzed as combined completely randomized designs to allow for inference beyond the study site (Cochran & Cox 1957). Densities were compared among treatments, months, and years using analysis of variance in a mixed linear model after the data were tested for normality and homogeneous variances (Cochran & Cox 1957). Tebuthiuron treatment, grazing occurrence, month, and year were analyzed as fixed effects, with year as the first repeated measure and month as the second repeated measure. We separated means using the least significant difference test with pairwise comparisons of least squares means if the F test on marginal means was significant (p < 0.05). Regression analysis of density by month was conducted using contrast coefficients within the model. We performed statistical analyses using SAS 9.1 (SAS Institute Inc. 2003). Means are reported ± SE. Species richness was the number of species observed during a survey, excluding incidentals such as birds on fencelines or flying over the survey area. We used relative abundance of bird species to calculate diversity with the P Shannon–Weiner index (H9 ¼ pi ln pi ; Magurran 1988: 35) and then calculated evenness as the ratio of observed diversity to maximum diversity (E ¼ H9/Hmax; Magurran 1988:37). F[1,12] ¼ 11.95, p ¼ 0.005). Average total density was 41% greater in 2005 than in 2004 (0.21 ± 0.01 vs. 0.15 ± 0.01, respectively), but because of an interaction between month and year (F[5,132] ¼ 3.19, p ¼ 0.01), the increase in density from 2004 to 2005 was not consistent across months. Density was greater in 2005 from March through May and then converged to 2004 levels for June to July (Fig. 1). Regression analysis of bird density by month revealed a quadratic relationship in both years for both tebuthiuron-treated (2004: F[1,60] ¼ 10.0, p ¼ 0.003; 2005: F[1,60] ¼ 47.42, p < 0.0001) and untreated plots (2004: F[1,60] ¼ 16.60, p ¼ 0.0001; 2005: F[1,60] ¼ 30.43, p < 0.0001; Fig. 1). On all plots, in both 2004 and 2005, the highest density of birds (0.24 ± 0.03 and 0.37 ± 0.04, respectively) occurred in April (Fig. 1). Density by Species Cassin’s Sparrow and Meadowlark were the only bird species for which we were able to estimate species-level densities in all survey months during both years. The study area is located in the boundary region between wintering and Results This study occurred during a period of unusually variable precipitation. Average yearly precipitation in the region is 31.5 cm (USDC NCDC 2005); however, 2003 represented the end of a 15-year period of drought at 26.2 cm of precipitation for the year in the study area. In 2004, cumulative precipitation was 85.9 cm, the second highest ever recorded in this region. Precipitation in 2005 was again below average at 27.5 cm (Dixon, unpublished data). A summary of vegetation response to tebuthiuron and grazing treatments (Dixon, unpublished data) can be found in Smythe and Haukos (2008) and Smythe (2006). In this paper, ‘‘migration’’ is defined as late February through the end of March, and ‘‘breeding season’’ as April through August. Total Bird Density Density (individuals per hectare) of all species (n ¼ 28) (Appendix) was not affected by the grazing treatment (F[1,12] ¼ 0.20, p ¼ 0.66). Average total density was 40% higher in tebuthiuron-treated plots than in untreated plots (X treated ¼ 0.21 ± 0.01, X untreated ¼ 0.15 ± 0.01; Restoration Ecology Figure 1. Density of all avian species from February to July in tebuthiuron-treated and untreated Sand shinnery oak communities in Roosevelt County, New Mexico, in 2004 and 2005. 3 Response of Grassland Birds in Sand Shinnery Oak Communities breeding areas for Cassin’s Sparrow, and although the occasional individual was observed during the winter, large numbers were not present until the breeding season. Singing Cassin’s Sparrows were first observed in the study area in April in 2004; however, they were first recorded a month earlier, during March, in 2005. Average density (singing males per hectare) of Cassin’s Sparrows on tebuthiurontreated plots (X ¼ 0.11 ± 0.009) was twice that on untreated plots (X ¼ 0.05 ± 0.007; F[1,12] ¼ 19.31, p ¼ 0.0009; Fig. 2). Average density was not affected by grazing (F[1,12] ¼ 2.42, p ¼ 0.15). Average density of Cassin’s Sparrows was greater in 2005 (X ¼ 0.10 ± 0.01) than in 2004 (X ¼ 0.06 ± 0.01), but because of an interaction between year and month (F[4,96] ¼ 2.99, p ¼ 0.02), the differences were observed only in March, April, and May (Fig. 2). Regression analysis across months revealed that in 2004, Cassin’s Sparrows increased linearly in treated plots (F[1,36] ¼ 16.55, p ¼ 0.0002) and remained constant in untreated plots (F[1,36] ¼ 1.24, p ¼ 0.27). In 2005, the relationship was quadratic in both the tebuthiuron-treated (F[1,48] ¼ 9.48, p ¼ 0.003) and untreated plots (F[1,48] ¼ 11.79, p ¼ 0.001) (Fig. 2). Meadowlark density (individuals per hectare) was not affected by tebuthiuron treatment (F[1,12] ¼ 0.04, p ¼ 0.85) or grazing (F[1,12] ¼ 0.88, p ¼ 0.37). On average, there was a greater density of Meadowlarks in 2004 (X ¼ 0.06 ± 0.009) than 2005 (X ¼ 0.03 ± 0.004; Fig. 3); however, because of an interaction between year and month (F[5,132] ¼ 5.51, p ¼ 0.0001), the difference was not consistent over all months. Meadowlark density was similar in February and July of both years but greater in 2004 for all other months (Fig. 3). There was a linear relationship between density and month in both 2004 and 2005 (F[1,60] ¼ 18.32, p < 0.0001 and F[1,60] ¼ 22.60, p < 0.0001, respectively; Fig. 3); this was predominantly influenced by the increase in density from February to March (Fig. 3). For a few species (Chihuahuan Raven [Corvus cryptoleucus], Grasshopper Sparrow [Ammodramus savannarum], Loggerhead Shrike [Lanius ludovicianus], Mourning Dove [Zenaida macroura], Vesper Sparrow [Pooecetes gramineus], Western Kingbird [Tyrannus verticalis], and WhiteCrowned Sparrow [Zonotrichia leucophrys]), we were able to calculate density for only part of the sampling period. These species were either migrants, not present in all months, or residents present in low numbers (Smythe 2006). Species Richness, Diversity, and Evenness There was no overall tebuthiuron treatment (F[1,12] ¼ 4.45, p ¼ 0.06) or grazing (F[1,12] ¼ 0.04, p ¼ 0.85) effect on species richness. On average, species richness was greater in 2005 than 2004 (X 2004 ¼ 6.46 ± 0.40, X 2005 ¼ 8.14 ± 0.48), but because of an interaction between month and year (F[5,120] ¼ 9.33, p < 0.0001), the difference between years was present only during late winter and spring migration (February, March, and April) (Fig. 4). Species diversity exhibited an interaction among tebuthiuron treatment, grazing treatment, and month (F[5,120] ¼ 2.33, p ¼ 0.04) and between year and month (F[5,120] ¼ 5.92, p < 0.0001). Thus, although diversity varied from month to Figure 2. Density of Cassin’s Sparrows (Aimophila cassinii) from March to July in tebuthiuron-treated and untreated Sand shinnery oak communities in Roosevelt County, New Mexico, in 2004 and 2005. 4 Figure 3. Density of Meadowlarks (Sturnella spp.) from February to July in Sand shinnery oak communities in Roosevelt County, New Mexico, in 2004 and 2005. Restoration Ecology Response of Grassland Birds in Sand Shinnery Oak Communities Figure 4. Species richness, diversity, and evenness of avian species from February to July in Sand shinnery oak communities in Roosevelt County, New Mexico, in 2004 and 2005. Uppercase letters indicate differences (p < 0.05) between years. month, the variation was not consistent across grazing and tebuthiuron treatments. Diversity was lower on ungrazed, untreated plots than other treatment combinations in February and March of both years. On average, diversity was greater in 2004 than 2005 (X 2004 ¼ 1.43 ± 0.07, X 2005 ¼ 1.29 ± 0.09), but the difference between years was observed only during February, March, and May (Fig. 4). Evenness varied similarly to species richness in that there was an interaction between month and year (F[5,132] ¼ 15.70, p < 0.0001) and there were no effects from tebuthiuron (F[1,12] ¼ 0.38, p ¼ 0.55) or grazing (F[1,12] ¼ 0.16, p ¼ 0.70) treatments. On average, evenness was closer to Restoration Ecology 1 (where 1 represents a situation in which all species are equally abundant) in 2004 than 2005 (X2004 ¼ 0.78 ± 0.03, X 2005 ¼ 0.64 ± 0.04), but the difference was present only during February, March, and May (Fig. 4). The decline in evenness in February and March of 2005 was caused by large flocks of Chestnut-collared Longspurs (Calcarius ornatus), and the decline in May was caused by large flocks of Lark Buntings (Calamospiza melanocorys). The influence of these flocks on species diversity is apparent: the dominance of these species caused the corresponding reduction in species diversity in the same months of 2005. 5 Response of Grassland Birds in Sand Shinnery Oak Communities Discussion This study was part of a larger, long-term experiment in which the goal was to restore the Sand shinnery oak community to a more historical grass/shrub balance and develop a grazing regime that would maintain that balance. Our results reflect the initial response of grassland birds to these restoration efforts, 4–5 years posttebuthiuron application. Our results are limited to the most common species in this community type and complicated by variable rainfall. Under the climate conditions of this study, application of tebuthiuron at 0.75 kg/ha to restore Sand shinnery oak communities in Roosevelt County, New Mexico, resulted in increased density of grassland birds. Increased density on tebuthiuron-treated plots was present in both wet and dry years. This differs from Martin (1990), who found no difference in relative abundance of all species between tebuthiuron-treated (using 0.5 kg/ha) and untreated shinnery oak communities in southeastern New Mexico. In our study, migratory species appear to respond differently to tebuthiuron treatment than resident species; the influx of migratory Cassin’s Sparrows had the greatest influence on the trends observed in density of all species. The grazing regime had very little impact on avian density. Month and year were the other major factors affecting grassland bird density; this temporal variation was likely in response to habitat conditions that resulted from precipitation patterns. The quadratic relationship observed in the density of all species stems largely from the patterns of migration and breeding in the study area. In addition to these temporal patterns, abnormally abundant rainfall appears to produce similar conditions to tebuthiuron treatment (denser grass within the shrub component), which migratory species appear to key into more than resident species. In this manner, the abundant rainfall of 2004 may have masked treatment effects that might have appeared had rainfall remained at or below average. A migratory species in this study, Cassin’s Sparrow, exhibited the greatest response to the treatments. Although the rare individual was observed overwintering on the study site, the conspicuous arrival of migratory males was evident in April 2004 and March 2005, which is when the density of all species on the treated and untreated areas diverged. This indicates that density of Cassin’s Sparrow was the greatest influence on overall density trends on the study site. The linear density relationship in 2004 versus the quadratic relationship in 2005 suggests a habitat saturation effect: in 2004, untreated areas were quickly saturated and remained at a constant density, whereas density in the tebuthiuron-treated areas expanded linearly. After the abundant rainfall in 2004, vegetation in the study area was denser in the beginning of 2005 (Smythe 2006; Dixon, unpublished data). Untreated plots supported more birds in 2005 than 2004, resulting in a quadratic relationship that mirrored that of the tebuthiuron-treated plots. The peak density of Cas- 6 sin’s Sparrow in tebuthiuron-treated plots was similar in 2004 (0.31 singing males/ha) and 2005 (0.29 singing males/ ha), but occurred a month earlier in 2005, again suggesting habitat saturation at a certain density. The maximum density of Cassin’s Sparrow in this study was comparable to the density (0.27 singing males/ha) recorded by Schnase (1984) in Mesquite mixed-grass prairie, but approximately 50% of the average density (0.57 singing males/ha) previously reported by Maurer (1985) in grasslands. Meadowlarks, the resident species for which we had consistent data, exhibited little response to either treatment. The neutral response of Meadowlarks to tebuthiuron treatment was consistent with Martin’s (1990) results for Meadowlarks in treated and untreated shinnery oak areas. Gruver and Guthery (1986) also found Meadowlark densities to be equal on treated and untreated areas of Mesquite (Prosopis spp.) 13 years after Mesquite control, and Castrale (1982) recorded no population response to sagebrush control by Western Meadowlarks. The change in density from February to March in both years likely represents an influx of migrating Meadowlarks into the study area for the breeding season, but more information on Meadowlark movements is needed to support this assertion. Year and month also influenced density of all species because manifestations of migratory patterns or improved habitat conditions from increased precipitation. The increase in density from 2004 to 2005 stems from increases in migratory species, predominantly Grasshopper Sparrow (which were not present during migration in 2004; Smythe 2006) and also from the earlier arrival of Cassin’s Sparrow. The increase in density during the breeding season might be related to the above-average rainfall in 2004. This type of response by Cassin’s Sparrow to precipitation was hypothesized by Hubbard (1977). Similarly, Igl and Johnson (1995, 1999) reported the change in status of Le Conte’s Sparrow (Ammodramus leconteii) from an uncommon breeding species to the most abundant over a period of 6 years that coincided with the amelioration of drought conditions and associated changes in habitat in the northern Great Plains. Ruth (2000) summarized anecdotal evidence suggesting similar distribution changes in response to precipitation by Cassin’s Sparrows. After the peak migration period, density of all species was similar between years, indicating that the arrival of migratory breeding sparrows had the greatest influence on overall density trends. Grasslands generally have low densities of birds, but densities on this study site were considerably lower than those reported in several other grassland studies (e.g., Giezentanner 1970; Wiens 1973; Cody 1985; Igl & Ballard 1999; DeJong 2001). Current low densities in Sand shinnery communities might indicate an ecological sink or reduced habitat carrying capacity, although the normal productivity for these communities is not well documented. Avian species richness, evenness, and diversity were only minimally affected by the tebuthiuron and grazing treatments. This partially agrees with Lueders et al. (2006), who found that neither species richness nor bird densities were Restoration Ecology Response of Grassland Birds in Sand Shinnery Oak Communities higher in structurally more diverse habitat in North Dakota. It appears that tebuthiuron treatment at this application rate and resulting vegetation response affect mainly the numbers of birds present, not which species or in what proportions they occur. Tebuthiuron-treated plots supported a greater density of the same community of birds as untreated plots (Smythe 2006), although additional longterm monitoring, spanning wet–dry cycles and maturing vegetation communities, will be necessary to determine the full community structure response to treatments. The moderate grazing regime (Holechek et al. 2001) implemented in this study appears to have little effect on grassland bird populations in this region, likely because the mosaic of grazed and ungrazed plots offered structural diversity. It is also important to emphasize that grazing was deferred on the study plots for 3 years after the tebuthiuron treatment, longer that the 1- to 2-year deferrals recommended by most current management guidelines (Peterson & Boyd 1998). Wiens and Dyer (1975) suggested that because grassland birds are so ecologically plastic, avifaunal shifts occur only after some threshold of habitat change has been passed, and therefore, many species of grassland birds would be unresponsive to moderate levels of grazing, such as occurred in this study. These results indicate that short-duration grazing regimes, based on the correct stocking rate and knowledge of available forage, are not detrimental to grassland bird populations. Bourliere and Hadley (1970) and Wiens (1973) suggested that migratory grassland birds live and reproduce off of the excesses produced by grasslands. Large seasonal or annual fluctuations in weather and production may produce surplus organic and nutrient matter that can be exploited by migrant species. Cody (1985) suggested that in response to climatic and production variations, grassland birds maintain a generalized morphology and behavior and practice resource tracking. This phenotypic plasticity in behavior and diet allows grassland birds to select geographic areas from year to year based on available resources. Igl and Johnson (1999) described how erratic population fluctuations are characteristic for most grassland passerines because they are highly dependent on specific vegetation features that vary dramatically in response to local moisture conditions. Given that seed and herbaceous production was considerably higher on treated plots than on untreated plots (Dixon, unpublished data), this resource tracking may explain the differences in density patterns between treated and untreated plots. This also might explain why migrant sparrows responded differently to shinnery oak control than did the year-round resident Meadowlarks. Meadowlarks are exclusively insectivorous (Wiens 1973) and Cassin’s Sparrow largely so during the breeding season, but Cassin’s Sparrow also consume seeds and plant matter (Ruth 2000). Our results would likely be more pronounced in average or below-average precipitation years. In this study, Sand shinnery oak plots treated with tebuthiuron to restore a more historical grass/shrubland combined with a moderate grazing system supported a greater Restoration Ecology density of Cassin’s Sparrow (a migratory species) than untreated plots and equal densities of Meadowlarks (a resident species) as untreated plots. This suggests that restoring Sand shinnery oak communities to a more historical grass/ shrub balance can create vegetation heterogeneity that benefits migratory grassland birds and does not appear to harm residents. A carefully managed grazing regime also does not appear to negatively impact grassland birds; however, grazing must be managed to maintain restoration efforts, and continued monitoring is needed to assess the long-term effects of restoration. There are approximately 1,068,370 hectares of shinnery oak in New Mexico (Garrison & McDaniel 1982) and about 2 million hectares of Sand shinnery communities across the southern Great Plains (Peterson & Boyd 1998). In addition, these Sand shinnery communities are important in winter to breeding birds of other prairie communities (Rich et al. 2004). Although our results may not apply directly to most of the priority species in these communities, they may provide general guidance on how species may respond to these management practices. The exception to this was Cassin’s Sparrow, a species of continental importance (Rich et al. 2004), which may benefit greatly from the management described in this study. If drought conditions continue to be the norm in this region, tebuthiuron treatment may help provide the optimal grass/shrub mix for an increased density of grassland birds that would otherwise be dependent on normal or above-normal precipitation. Finally, vegetation variables influencing grassland passerine density can differ from those affecting nesting success (Winter et al. 2005), and thus avian density responses to these treatments do not imply similar effects on nesting success. Nest productivity and reproductive success are other important factors to consider when evaluating habitat quality and the impacts of management practices in Sand shinnery oak communities. Implications for Practice Sand shinnery oak plots restored to a grass/shrub mix with low doses of tebuthiuron (0.73 kg/ha) supported a greater density of spring migrants and breeding birds than untreated plots. d Tebuthiuron treatment may create a grass/shrub mix normally restricted to years of above-average precipitation. This could increase densities of some migratory grassland bird species such as Cassin’s Sparrow and does not appear to harm resident species such as Meadowlarks. Further research, spanning longer wet–dry cycles and maturing plant communities, will be necessary to completely determine avian response to these restoration efforts. d A carefully managed, moderate grazing regime also does not appear to negatively impact grassland bird density; however, grazing must be managed to maintain restoration efforts, and continued monitoring is needed to determine the long-term effects of restoration. d 7 Response of Grassland Birds in Sand Shinnery Oak Communities Acknowledgments We thank D. B. Wester for assistance with statistical analyses and C. L. Dixon of Wildlife Plus Consulting for sharing vegetation and rainfall data. We also thank A. Andrei and J. Hull for their conscientious data collection and hard work in the field. 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Common Name Scientific Name American Kestrel Ash-throated flycatcher Blue Grosbeak Brewer’s Sparrow Bullock’s Oriole Burrowing Owl Cassin’s Sparrow Chestnut-collared Longspur Chihuahuan Raven Chipping Sparrow Eastern Meadowlark Field Sparrow Grasshopper Sparrow Lark Bunting Lark Sparrow Loggerhead Shrike Mourning Dove Northern Bobwhite Northern Mockingbird Prairie Falcon Savanna Sparrow Sage Thrasher Scissor-tailed Flycatcher Swainson’s Hawk Western Kingbird Western Meadowlark White-crowned Sparrow Vesper Sparrow Falco sparverius Myiarchus cinerascens Guiraca caerulea Spizella breweri Icterus bullockii Athene cunicularia Aimophila cassinii Calcarius ornatus Corvus cryptoleucus Spizella passerina Sturnella magna Spizella pusilla Ammodramus savannarum Calamospiza melanocorys Chondestes grammacus Lanius ludovicianus Zenaida macroura Colinus virginianus Mimus polyglottos F. mexicanus Passerculus sandwichensis Oreoscoptes montanus Tyrannus forficatus Buteo swainsoni T. verticalis Sturnella neglecta Zonotrichia leucophrys Pooecetes gramineus Species Nests in Sand Shinnery Oak Communities Species-Specific Density Estimated in This Study Species of Continental Importance in Southwest Regiona Priority Species in New Mexicob Yes Yes Yes No Yes Yes Yes No Yes No Yes No No No Yes Yes Yes Yes Yes No No No Yes Yes Yes Yes No No No No No No No No Yes No Yesd No Yesc No Yesd No No Yesd Yesd No No No No No No No Yesd Yesc Yesd Yesd No No No Yes No No Yes No No No No No No No No No No No No No No No No Yes No No No No No No No No No No No No No No No No No No No Yes No No No No No No No Yes No No No Yes a As defined by the Partners in Flight North American Landbird Conservation Plan (Rich et al. 2004). As defined by the New Mexico Partners in Flight Bird Conservation Plan (NMPIF 2007). c Species grouped for density estimate. d Results presented in Smythe (2006). b Restoration Ecology 9