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The nonbreeding period is the longest and perhaps most hazardous time in the annual dynamics of migratory songbirds (Norris and Marra, 2007). Yet, we know little about the ecology of many species after they leave their breeding location (Webster et al., 2002). Linkages between individuals and their locations used on seasonal ranges can help us understand seasonal interactions influencing annual population dynamics (Norris, 2005; Monroy-Ojeda et al., 2013; Hostetler et al., 2015).

Shrubland birds in western North America face double stressors from habitat loss on breeding and wintering ranges (Knick et al., 2003; Ruth et al., 2008; Macias-Duerte et al., 2009). Brewer's (Spizella breweri) and sagebrush (Artemisiospiza nevadensis) sparrows are obligate to landscapes dominated by sagebrush (Artemisia species) on their breeding range (Martin and Carlson, 1998; Rotenberry et al., 1999). Black-throated sparrows (Amphispiza bilineata) breed in arid shrublands that include sagebrush and salt desert shrub communities (Johnson et al., 2002). All three species are middle-distance to short-distance migrants to winter ranges dominated by open shrublands in the southwestern united states and central Mexico.

We report recaptures of sparrows in subsequent years after banding on their wintering areas in the southwestern United States. Although these recaptures were incidental as part of a larger study on breeding and winter range connectivity in sagebrush-obligate songbirds (Knick et al., 2014), they provide insights into the tendency of individual birds to return to a wintering site from 1-4 y following initial capture.

We recaptured birds at 15 of 79 locations separated by >15 km (Fig. 1). We first surveyed an area of approximately 1-[km.sup.2] for flocks that were herded and flushed into strategically placed nets. We attempted to capture any bird within a 300-m flush zone surrounding a net wall consisting of 8-10 mist nets of 12-m length and 30-mm mesh size. We marked each bird with a numbered aluminum leg band registered by the U.S. Geological Survey, recorded body mass and standard measurements, and collected the primary P1 feather for isotope analysis prior to release at the location. Feathers are formed during the postbreeding molt for these species and contain an isotopic signature that reflects broad geographic and environmental gradients (Hobson, 2005). The level of deuterium in feathers ([[delta].sup.2][H.sub.f]) was the deviation ([delta]) in [sup.2]H/[sup.1]H relative to Vienna standard mean ocean water (Wassenaar and Hobson, 2003).

Initial captures in 2003-2005 were opportunistic because our primary objective was to identify characteristic habitats and primary locations used by sparrows over their winter range in the southwestern United States. We revisited 66 of the initial 79 capture locations generally at the same time each winter season (December-March) from 2005-2008. After 2008, some of our capture locations were sampled by banders not affiliated with the initial project.

Our study sites in southern California occurred in the overlap zone between sagebrush and Bell's sparrows (Artemisiospiza belli). After our field seasons, the progenitor sage sparrow (Amphispiza belli) was split into separate species as sagebrush sparrows and Bell's sparrows (Chesser et al., 2013). We did not conduct morphological or mtDNA analyses in a post hoc attempt to distinguish species but instead used S2Hf to predict breeding range and distinguish migratory sagebrush sparrows from resident Bell's sparrows (Fig. 1). We used a threshold of [[delta].sup.2][H.sub.f] > -[86%.sub.0] in the relationship derived for feathers of sagebrush sparrows with growing season precipitation (Knick et al., 2014).

Five Brewer's sparrows, seven sagebrush sparrows, and three black-throated sparrows were recaptured at their initial capture location in subsequent winters (Table 1). Most were recaptured the following winter. Of all birds banded, only 0.5% of Brewer's, 1.4% sagebrush (an additional five sagebrush sparrows that could have been resident Bell's sparrows were not included), and 1.2% black-throated sparrows were recaptured and all were at their initial capture location.

One Brewer's sparrow (Band #2340-51492) was recaptured for two successive winters. Two Brewer's sparrows (#2220-92919, #2220-92935) were first banded together in 2003 and again were together when recaptured 4 y later (2007). All three black-throated sparrows were recaptured the following winter. One sagebrush sparrow (#910-93019) was recaptured in the second and third subsequent winter. Sagebrush sparrow #1601-07138 was first captured in 2007 and recaptured 4 y later in 2011.

Isotopic signatures ([[delta].sup.2][H.sub.f]) indicated that two individuals in each of two groups of Brewer's sparrows (#2340-51492 and #2340-51449; #2220-92919 and #2220-92935) likely came from different breeding ranges (within-year difference in [[delta].sup.2][H.sub.f]: 9-29%; Table 1). Both individuals of the group, which included the group banded in 2003 and recaptured in 2007, were again captured together in following years.

Basic information about the winter ecology of Brewer's, sagebrush, and black-throated sparrows is lacking. These shrubland sparrows experience a different suite of anthropogenic and climate-related habitat changes on their winter range (Ruth et al., 2008; Macias-Duarte et al., 2009) that are in addition to changes altering their breeding landscape (Knick et al., 2003). The tendency to return to winter sites, coupled with information on environmental characteristics at those locations, can help us understand carry-over effects from winter stressors to breeding populations (Cresswell, 2014; Hostetler et al., 2015).

We were unable to statistically assess recapture rates and infer site fidelity because we did not standardize methods, effort, or locations. Birds that were not recaptured could have died, evaded recapture within the flush zone, used other winter locations, been in a different flock than that targeted, or could simply have been at the same location in subsequent years but outside of our flush zone. Nonetheless, a low but unknown proportion of individuals of these species return to winter sites in subsequent years. For two Brewer's and two sagebrush sparrows, use of a specific winter location extended at least 3 and 4 y following their initial capture.

Rate of return to a winter location does not equal site fidelity but indicates only that an individual visits a known location in what otherwise might be an itinerant strategy. Site fidelity might be weak in the southwestern United States for granivorous sparrows because seed production can be highly variable in space and time due to vagaries in precipitation (Pulliam and Brand, 1975; Swetnam and Betancourt, 1998). Winter area requirements and movement patterns for these species are unknown. In southeastern Arizona, only 1.4% of 72 Brewer's sparrows were recaptured the following winter (Gordon, 2000). Relative to higher recapture rates for more-sedentary Cassin's (Aimophila cassinii) and grasshopper (Ammodramus savannarum) sparrows, individual Brewer's sparrows likely were mobile and using areas outside our study sites. If the three species of sparrows in our study have adopted an itinerant wintering strategy, fidelity needs to be evaluated at larger spatial extents than our sites because birds likely have multiple locations in an area that they prospect each winter.

The U.S. Geological Survey (USGS), Forest and Rangeland Ecosystem Science Center and the U.S. Department of Defense supported our study. The Colorado Plateau Stable Isotope Laboratory at Northern Arizona University conducted isotope measurements and accuracy assessments. P. Blom recaptured sagebrush sparrow #1601-07138 and reported banding information. We appreciate funding from the U.S. Department of Defense Legacy Program and the USGS, Forest and Rangeland Ecosystem Science Center. C. Bock provided helpful comments on the manuscript. Any use of trade, product, or firm names is for descriptive purposes only and does not imply endorsement by the U.S. Government.

Literature Cited

Chesser, R. T., R. C. Banks, F. K. Barker, C. Cicero, J. L. Dunn, A. W. Kratter, I.J. Lovette, P. C. Rasmussen, J. V. RemsenJr., J. D. Rising, D. F. Stotz, and K. Winker. 2013. Fifty-fourth supplement to the American Ornithologists' Union Checklist of North American Birds. Auk 130:1-14.

Cresswell, W. 2014. Migratory connectivity of Palaearctic-African migratory birds and their responses to environmental change: the serial residency hypothesis. Ibis 156:493-510.

Gordon, C. 2000. Movement patterns of wintering grassland sparrows in Arizona. Auk 117:748-759.

Hobson, K. A. 2005. Stable isotopes and the determination of avian migratory connectivity and seasonal interactions. Auk 122:1037-1048.

Hostetler, J. A., T. S. Sillett, and P. P. Marra. 2015. Full-annualcycle population models for migratory birds. Auk 132:433-449.

Johnson, M. J., C. Van Riper, III, and K. M. Pearson. 2002. Black-throated sparrow (Amphispiza bilineata). The Birds of North America online (A. Poole, editor). Cornell Lab of ornithology, Ithaca, New York. Available at: http://bna.birds.cornell. edu/bna/species/637. doi:10.2173/bna.327. Accessed 1 December 2015.

Knick, S. T., D. S. Dobkin, J. T. Rotenberry, M. A. Schroeder, W. M. Vander Haegen, and C. van Riper, III. 2003. Teetering on the edge or too late? Conservation and research issues for avifauna of sagebrush habitats. Condor 105:611-634.

Knick, S. T., M. Leu, J. T. Rotenberry, S. E. Hanser, and K.A. Fesenmyer. 2014. Diffuse migratory connectivity in two species of shrubland birds: evidence from stable isotopes. oecologia 174:595-608.

Macias-Duarte, A., A. B. Montoya, C. E. MEndez-Gonzalez, J. R. Rodriguez-Salazar, W. G. Hunt, and P. G. Krannitz. 2009. Factors influencing habitat use by migratory grassland birds in the state of Chihuahua, Mexico. Auk 126:896-905.

Martin, J. W., and B. A. Carlson. 1998. Sage sparrow (Artemisiospiza belli), The Birds of North America Online (A. Poole, editor). Cornell Lab of Ornithology, Ithaca, New York. Available at: doi:10.2173/bna.326. Accessed 1 December 2015.

Monroy-Ojeda, A., M. Grosselet, G. Ruiz, and E. Del Valle. 2013. Winter site fidelity and winter residency of six migratory Neotropical species in Mexico. Wilson Journal of Ornithology 125:192-196.

Norris, D. R. 2005. Carry-over effects and habitat quality in migratory populations. Oikos 109:178-186.

Norris, D. R., and P. P. Marra. 2007. Seasonal interactions, habitat quality, and population dynamics in migratory birds. Condor 109:535-547.

Pulliam, H. R., and M. R. Brand. 1975. The production and utilization of seeds in plains grassland of southeastern Arizona. Ecology 56:1158-1166.

Rotenberry, J. T., M. A. Patten, and K. L. Preston. 1999. Brewer's sparrow (Spizella breweri). The Birds of North America Online (A. Poole, editor). Cornell Lab of Ornithology, Ithaca, New York. Available at: species/390. doi:10.2173/bna.390. Accessed 1 December 2015.

Ruth, J. M., T. Brush, and D.J. Krueper (editors). 2008. Birds of the US-Mexico borderlands: distribution, ecology, and conservation. Studies in Avian Biology 37. Cadmus Professional Communications, Ephrata, Pennsylvania.

Swetnam, T. W., and J. L. Betancourt. 1998. Mesoscale disturbance and ecological response to decadal climatic variability in the American Southwest. Journal of Climate 11:3128-3147.

Wassenaar, L. I., and K. A. Hobson. 2003. Comparative equilibration and online technique for determination of non-exchangeable hydrogen of keratins for use in animal migration studies. Isotopes in Environmental and Health Studies 39:211-217.

Webster, M. S., P. P. Marra, S. M. Haig, S. Bensch, and R. T. Holmes. 2002. Links between worlds: unraveling migratory connectivity. Trends in Ecology and Evolution 17:76-83.

Submitted 22 December 2015. Accepted 24 April 2017.

Associate Editor was Michael Scott Husak.

Steven T. Knick, * Matthias Leu, and Steven E. Hanser

U.S. Geological Survey, Forest and Rangeland Ecosystem Science Center, 970 Lusk Street, Boise, ID 83706 (STK, ML, SEH)

Present address of STK: 2140 White Pine Place, Boise, ID 83706

Present address of ML: Biology Department, College of William and Mary, Williamsburg, VA 23815

Present address of SEH: U.S. Geological Survey, 12201 Sunrise Valley Drive, MS 300, Reston, VA 20192

* Correspondent:

Caption: Fig. 1-Site locations where Brewer's sparrows (Spizella breweri), sagebrush sparrows (Artemisiospiza nevadensis), and black-throated sparrows (Amphispiza bilineata) were banded and recaptured between 2003 and 2011 from one to four subsequent winters following initial banding. We used levels of feather deuterium in sparrows captured in southern California to distinguish resident Bell's sparrows (Artemisiospiza belli) from migratory sagebrush sparrows overwintering from predicted breeding areas in shaded regions.
Table 1--Recaptured Brewer's, black-throated, and sagebrush sparrows
on wintering ranges in the southwestern United States between 2003
and 2011. (a)

                    U.S. Geological                  Capture
     Species        Survey band no.     Age            date

Brewer's sparrow    2340-51492        Juvenile   26 February 2005
                                      Adult      16 January 2006
                                      Adult      9 January 2007
                    2340-51499        Adult      27 February 2005
                                      Adult      16 January 2006
                    2350-10339        Adult      14 January 2006
                                      Adult      7 January 2007
                    2220-92919        Adult      19 February 2003
                                      Adult      17 February 2007
                    2220-92935        Adult      19 February 2003
                                      Adult      17 February 2007
Black-throated      1941-08961        Adult      13 February 2006
                                      Adult      8 December 2006
                    1941-08851        Adult      25 February 2006
                                      Adult      4 February 2007
                    1941-08865        Adult      1 March 2006
                                      Adult      20 December 2007
Sagebrush sparrow   1941-09100        Juvenile   11 January 2007
                                      Adult      9 January 2008
                    910-93019         Juvenile   18 December 2004
                                      Adult      12 January 2006
                                      Adult      10 January 2007
                    1601-07138        Adult      20 January 2007
                                      Adult      2 March 2011
                    1691-69130        Adult      22 January 2003
                                      Adult      10 December 2004
                    1831-85595        Adult      3 February 2006
                                      Adult      8 February 2007
                    1941-09163        Adult      27 January 2007
                                      Adult      25 January 2008
                    1941-09225        Juvenile   2 February 2007
                                      Adult      18 December 2007

                    U.S. Geological      No.
     Species        Survey band no.   banded (b)   Cocapture (c)

Brewer's sparrow    2340-51492            14
                                          13             A
                    2340-51499            6
                                          13             A
                    2350-10339            18
                    2220-92919                           B
                                          10             C
                    2220-92935                           B
                                          10             C
Black-throated      1941-08961            11
                    1941-08851            2
                    1941-08865            3
Sagebrush sparrow   1941-09100            5
                    910-93019             11
                    1601-07138            44

                    1691-69130            3
                    1831-85595            7
                    1941-09163            27
                    1941-09225            10

                    U.S. Geological
     Species        Survey band no.       Latitude

Brewer's sparrow    2340-51492        31[degrees]46'12"
                    2340-51499        31[degrees]46'12"
                    2350-10339        31[degrees]39'0"
                    2220-92919        33[degrees]39'0"
                    2220-92935        33[degrees]39'0"

Black-throated      1941-08961        33[degrees]6'0"
                    1941-08851        32[degrees]0'0"
                    1941-08865        32[degrees]28'12"

Sagebrush sparrow   1941-09100        34[degrees]52'12"
                    910-93019         33[degrees]22'48"
                    1601-07138        35[degrees]0'36"
                    1691-69130        35[degrees]40'0"
                    1831-85595        35[degrees]17'24"
                    1941-09163        35[degrees]51'0"
                    1941-09225        32[degrees]41'24"

                    U.S. Geological
     Species        Survey band no.        Longitude          State

Brewer's sparrow    2340-51492        --109[degrees]6'0"     Arizona
                                      --109[degrees]6'0"     Arizona
                                      --109[degrees]6'0"     Arizona
                    2340-51499        --109[degrees]6'0"     Arizona
                                      --109[degrees]6'0"     Arizona
                    2350-10339        --111[degrees]3'36"    Arizona
                                      --111[degrees]3'36"    Arizona
                    2220-92919        --107[degrees]7'48"    New Mexico
                                      --107[degrees]7'48"    New Mexico
                    2220-92935        --107[degrees]7'48"    New Mexico
                                      --107[degrees]7'48"    New Mexico

Black-throated      1941-08961        --114[degrees]18'0"    Arizona
                                      --114[degrees]18'0"    Arizona
                    1941-08851        --106[degrees]15'0"    New Mexico
                                      --106[degrees]15'0"    New Mexico
                    1941-08865        --106[degrees]31'12"   New Mexico
                                      -106[degrees]31'12"   New Mexico

Sagebrush sparrow   1941-09100        --114[degrees]31'48"   Arizona
                                      --114[degrees]31'48"   Arizona
                    910-93019         --112[degrees]46'48"   Arizona
                                      --112[degrees]46'48"   Arizona
                                      --112[degrees]46'48"   Arizona
                    1601-07138        --117[degrees]17'24"   California
                                      --117[degrees]17'24"   California
                    1691-69130        --117[degrees]40'0"    California
                                      --117[degrees]40'0"    California
                    1831-85595        --117[degrees]52'48"   California
                                      --117[degrees]52'48"   California
                    1941-09163        --117[degrees]40'0"    California
                                      --117[degrees]40'0"    California
                    1941-09225        -106[degrees]31'12"   New Mexico
                                      --106[degrees]31'12"   New Mexico

                    U.S. Geological   Mass    [[delta].sup.2]H
     Species        Survey band no.   (g)    ([per thousand]) (d)

Brewer's sparrow    2340-51492        10.5          -99.7
                                      10.0          -103.5
                                      10.6          -88.2
                    2340-51499        10.6          -90.5
                                      10.2          -74.2
                    2350-10339        10.9          -88.2
                                      10.5          -90.3
                    2220-92919        10.4          -111.6
                                      10.8          -105.8
                    2220-92935        11.0          -123.1
                                      12.3          -90.5

Black-throated      1941-08961        12.2

                    1941-08851        10.7
                    1941-08865        14.3
                                      15.2          -55.5

Sagebrush sparrow   1941-09100        16.2          -77.9
                                      16.6          -60.1
                    910-93019         14.8          -74.3
                                      14.7          -71.3
                                      16.0          -66.5
                    1601-07138                      -99.5
                    1691-69130        16.7          -101.7
                                      16.4          -85.9
                    1831-85595        16.7          -99.4
                                      15.9          -87.7
                    1941-09163        18.0          -107.3
                                      18.1          -89.7
                    1941-09225        19.5          -91.2
                                      20.8          -93.6

(a) We banded 1,047 Brewer's, 249 black-throated, and 509 sagebrush

(b) Number of individuals from the captured flock that received
U.S. Geological Survey bands.

(c) Individuals captured together in the same flock have the same

(d) Blank cells indicate that isotope data was not collected.
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Title Annotation:Notes
Author:Knick, Steven T.; Leu, Matthias; Hanser, Steven E.
Publication:Southwestern Naturalist
Geographic Code:1U800
Date:Jun 1, 2017

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