J. Field Ornithol. 77(4):372–383, 2006 DOI: 10.1111/j.1557-9263.2006.00067.x Community composition and migration chronology of shorebirds using the saline lakes of the Southern Great Plains, USA Adrian E. Andrei,1,4 Loren M. Smith,1 David A. Haukos,2 and James G. Surles3 1 2 Department of Range, Wildlife, and Fisheries Management, Texas Tech University, Lubbock, Texas 79409, USA U.S. Fish and Wildlife Service, Department of Range, Wildlife, and Fisheries Management, Texas Tech University, Lubbock, Texas 79409, USA 3 Department of Mathematics and Statistics, Texas Tech University, Lubbock, Texas 79409, USA Received 25 May 2006; accepted 10 September 2006 ABSTRACT. Shorebirds migrating through the Southern Great Plains (SGP), USA, use freshwater playas and saline lakes as stopovers. The importance of playas is well documented, but the role of saline lakes is not clearly understood. During 2002 and 2003, we conducted surveys to determine the extent to which the saline lakes serve as stopovers. Twenty-eight species were recorded, and total seasonal abundance ranged from 6779 to 29,924 birds. Potential shorebird abundance for extant saline lakes was estimated at 37,000–71,000 shorebirds annually. American Avocets (Recurvirostra americana), Western Sandpipers (Calidris mauri), Baird’s Sandpipers (C. bairdi), Least Sandpipers (C. minutilla), Snowy Plovers (Charadrius alexandrinus), Killdeer (Charadrius vociferus), and Wilson’s Phalaropes (Phalaropus tricolor) were the most abundant species. Community composition of shorebirds differed between saline lakes and regional freshwater playas. Peak spring abundance was generally in April, whereas summer/fall migration was more protracted and shorebird abundance peaked during 6–8 weeks in August and September. Migration chronologies differed among morphologically similar species, and among representative species from different guilds. Such patterns of temporal separation permit partitioning of resources by shorebirds migrating through the SGP. The saline lakes of the SGP should be regarded as stopover sites of regional and international value. To ensure that saline lakes function as stopovers and to help maintain those unique communities that inhabit them, conservation of saline lakes should focus on preserving spring flows and conserving water. SINOPSIS. Composición de la comunidad y cronologı́a migratoria de playeros que utilizan los lagos salobres de las grandes planicies del sur de los EUA Los playeros que migran a través de las grandes planicies del sur de los Estados Unidos, utilizan como lugares de parada playas de agua fresca y lagos salobres. La importancia de las playas ha sido muy bien documentada, pero no se entiende bien el rol que puedan tener los lagos salobres. Durante el 2002 y el 2003, realizamos trabajos para determinar hasta que punto los lagos salobres servı́an como lugares importantes de parada. En estos, enontramos la presencia de 28 especies y una abundancia que varı́a entre 6779 y 29,924 aves. El uso potencial de estos lagos salinos es de 37,000 a 71,000 individuos. Las especies más abundantes en este tipo de habitat resultaron ser Recurvirostra americana, Caladris mauri, C . bairdi, C . minutilla, Charadrius alexandrinus y Phalaropus tricolor. La composición de playeros resultó ser diferente en las playas de agua fresca y en los lagos salobres. El pico en la abundancia durante la primavera, se llevó a cabo generalmente en Abril, mientras que el pico de los migrantes de verano/otoño ocurrió por un periodo de 6–8 semanas de agosto a septiembre. La cronologı́a migratoria tiene sus diferencias entre especies de morfologı́a similar, al igual que en especies de otros grupos. Este patrón de separación temporera, permite la repartición de recursos entre las aves que migran a través de las grandes planicies. Los lagos salobres en dicha área deben ser considerados como lugares de paradas de gran importancia a nivel nacional e internacional. Para asegurar que dichos lagos salobres mantengan su función como lugares de paradas, la conservación de los mismos debe enfocar el preservar el flujo de agua a estos durante la primavera. Key words: migration chronology, playas, saline lakes, shorebirds, Southern Great Plains Many shorebirds migrate through interior North America between Arctic nesting grounds and wintering areas in Central and South America (Morrison 1984, Skagen and Knopf 4 Corresponding author. Email: andreia@lincolnu. edu C 2006 1993, Davis and Smith 1998a). These migrants need to replenish and accumulate energy reserves (Morrison 1984, Senner and Howe 1984, Myers et al. 1987) at stopover sites along the migration route (Skagen and Knopf 1993). Millions of shorebirds use the 25,000 playas of the Southern Great Plains (SGP) (Davis and Smith 1998b) as important “stepping stones” (Skagen and Knopf C 2006 Association of Field Ornithologists The Author(s). Journal compilation 372 Vol. 77, No. 4 Shorebird Use of Saline Lakes 1993) necessary to complete their migration. In addition to freshwater playas (Smith 2003), there are approximately 45 saline lakes (Reeves and Reeves 1996) in the SGP. The breeding biology of shorebirds on saline lakes (Conway et al. 2005) has been documented, but the extent to which migrant shorebirds use saline lakes as stopovers is unknown. Unlike playas that are freshwater-recharge wetlands, the saline lakes of the SGP are large, sparsely vegetated, saline (>4 ppt) discharge wetlands (Brüne 1981, Andrei 2005). Historically, numerous springs fed by the Ogallala aquifer discharged in the saline lakes (Brüne 1981), and these lakes and associated springs once served as perennial aquatic habitats for wildlife. By the 1950s, spring flows had been severely reduced due to declining aquifer levels, and several lakes completely dried (Brüne 1981). Since then, water levels of the Ogallala aquifer have continued to decline (Triplet 1998, Sophocleous 2000). The freshwater playas of the SGP are filled by rainfall runoff and serve as recharge points for the Ogallala aquifer (Osterkamp and Wood 1987, Smith 2003). Playas are generally smaller than saline lakes (Guthery and Bryant 1982, Reeves and Reeves 1996). Shorebird communities using saline lakes during migration may differ from those using playas (Davis and Smith 1998a) because of ecological differences between these two types of habitats. Data concerning composition, abundance, and migration chronology of shorebirds using the saline lakes will aid the United States Shorebird Conservation Plan (Brown et al. 2001) by identifying potentially important stopovers in the Playa Lakes region. Information about shorebird migration chronologies may be important for developing conservation strategies for saline lakes. Further, a comparison of migration chronologies of shorebirds using saline lakes may help explain how shorebirds use the unpredictable resources (Haukos and Smith 1992) of the spatially dynamic wetlands (Skagen and Knopf 1993) in the Great Plains. A temporal separation of migrant shorebirds, noted for their dietary flexibility (Skagen and Oman 1996), may allow partitioning of foraging resources by morphologically similar shorebirds (Davis and Smith 1998a). Our objectives were to (1) document the relative abundance and community composition of shorebirds using saline lakes of the SGP during spring and summer/fall migra- 373 tions, (2) examine differences between shorebird communities using freshwater playas and those using saline lakes, (3) examine migration chronologies of shorebirds in saline lakes and test for differences in migration chronologies among species within guilds as well as among species from different guilds, and (4) propose conservation recommendations for shorebirds using saline lakes of the SGP. METHODS Study area. Our study was conducted in the SGP of Texas and New Mexico, USA. (Fig. 1). Formed by wind erosion and dissolution of salts (Reeves and Reeves 1996) and ranging in size from approximately 4 ha (Frost Lake, 32◦ 49 34.77 N and 102◦ 00 42.77 W) to over 6000 ha (Cedar Lake, 32◦ 49 06.64 N and 102◦ 16 21.89 W), the saline lakes are generally larger than playas (x¯ = 6.3 ha) of the SGP (Smith 2003). Vegetation cover (≤1%; Andrei 2005) in the shallow (0–100 cm) saline lakes consists of plants such as Saltgrass (Distichlis spp.), Pickleweed (Salicornia spp.), Bulrush (Scirpus spp.), and Saltcedar (Tamarix spp.). Originally covered by short- to mid-grass prairies, the SGP are among the most intensely cultivated regions in the world (Bolen et al. 1989). Preceding and during our study, precipitation recorded in Lubbock, Texas, was below the yearly average of 48 cm (32.9 cm in 2001, 47.6 cm in 2002, and 20.9 cm in 2003; National Oceanic and Atmospheric Administration 2004). Shorebird surveys. Saline lakes were located and identified following Reeves and Reeves (1996; Fig. 1). We asked for permission to access lakes from landowners and surveyed shorebirds on all lakes where permission was granted. Because obtaining permission was random, we surveyed a random sample of saline lakes in the SGP. Shorebird surveys were conducted weekly during spring (10 March–15 June 2002 and 2 March–7 June 2003) and summer/fall (7 July– 9 November 2002 and 7 July–8 November 2003). In 2002, 21 lakes were surveyed during spring and 25 during summer/fall. In 2003 27 lakes were surveyed during both spring and summer/fall. During surveys we either counted all birds from one location at the edge of the lake or by walking along the edge of the lake when all shorebirds were not visible from one location. We used 10 × 50 binoculars and a 20–50 × 80 374 A. E. Andrei et al. J. Field Ornithol. Fall 2006 Fig. 1. Study area and locations of the saline lakes of the Southern Great Plains (after Reeves and Reeves (1996)). There are two or more lakes at some locations, including Yellow House, Cunavea Basin, Twin Lakes, Double Lakes, Four Lakes, and Muleshoe National Wildlife Refuge. spotting scope. Large flocks were counted by dividing them into groups of 5, 10, or 20 individuals and using landmarks to divide flocks into groups that could be counted accurately. We performed repeated counts on large flocks (several hundred individuals) and averaged the results (Colwell and Cooper 1993). Because lakes were relatively open and unvegetated, we Vol. 77, No. 4 Shorebird Use of Saline Lakes assumed all shorebirds at each lake were visible and counted. To minimize bias associated with conducting surveys on the same lake at the same time on consecutive weeks, we assigned weekly survey times randomly for each lake to one of three diurnal periods: early day (sunrise–11:00), midday (11:00–15:00), and late day (15:00– sunset; Davis and Smith 1998a). Data analysis. We summed survey data across lakes and within each season of each year to determine abundance (number of shorebirds observed per season) and species composition (percent of total number of observed birds per season for each species). We estimated annual shorebird use days for all saline lakes throughout the SGP by: (1) assuming a turnover of shorebirds of 7 d (Skagen and Knopf 1994b), (2) estimating annual shorebird use days for the surveyed lakes, and (3) extrapolating relative to the total number of saline lakes in the SGP. For each season (spring or summer/fall), we averaged the total number of birds of the more common species (>5 individuals) observed during a migration season for the 2 years of study (2002 and 2003). Similarly, we averaged the seasonal abundances of each shorebird species observed in playas of the SGP for the two study years for each season and each species, as reported by Davis and Smith (1998a) for 1993 and 1994. For each season (spring and summer/fall), we used averaged seasonal abundances of the more common species to compare community composition of shorebirds of saline lakes and playas with 2 tests for independence (Zar 1996). Pearson’s 2 was the test statistic. We also examined possible differences between average seasonal abundance and species composition (percent of total number of birds per season) of shorebirds in saline lakes and playas. We used the statistical analysis system (SAS Institute 2000) for all analyses. To test for differences in migration chronologies of shorebirds, we categorized species into foraging guilds according to Skagen and Oman (1996), and used 2 tests for independence (Sokal and Rohlf 1981) to compare small prober/gleaners (Least, Semipalmated, Baird’s, and Western sandpipers; scientific names of all shorebirds mentioned in the text are provided in the Appendix) and representative species from different foraging guilds, including a small prober/gleaner (Least Sandpiper), a medium gleaner/prober 375 (Lesser Yellowlegs), a gleaner/sweeper (Wilson’s Phalarope), a gleaner/prober (American Avocet), a large gleaner/prober (Long-billed Curlew), and two medium gleaner/probers (Long-billed Dowitcher and Stilt Sandpiper). We used the numbers of birds of each species observed during each week of migration to compare migration chronologies for each season of each year (spring and summer/fall of 2002 and 2003) with 2 . Pearson’s 2 was the test statistic (Zar 1996). RESULTS In 2002 16 lakes were used by shorebirds during the spring and 20 were used during the summer and fall. In 2003 17 lakes were used by shorebirds during the spring and 10 during the summer and fall. We observed 28 species of shorebirds during our study (spring: 26 species in 2002 and 24 in 2003; summer/fall: 26 species in 2002 and 28 in 2003; see Appendix). The most frequently observed species during spring migration were Snowy Plover, Killdeer, American Avocet, Western Sandpiper, Baird’s Sandpiper, and Wilson’s Phalarope (Table 1). During summer/fall migration, the most common species were American Avocet, Western Sandpiper, Baird’s Sandpiper, and Wilson’s Phalarope (Table 1). Shorebirds were observed using 23 of the 45 saline lakes of the SGP during 2002 and 2003, years with below average precipitation. Assuming a turnover of shorebirds of 7 d (Skagen and Knopf 1994b), conservative annual estimates of shorebird use days for surveyed saline lakes are 36,703 for 2002 and 18,628 for 2003. An extrapolation to all saline lakes of the region (×2), assuming similar habitat conditions in unsurveyed lakes, suggests a range of 37,256 to 73,406 shorebird use days annually. The length of stay at the saline lakes likely varies among species and between seasons and years depending on available habitat conditions. Skagen and Knopf (1994b) reported average lengths of stay between 3.4 and 9.7 d for Semipalmated and White-rumped sandpipers in Kansas. Shorebird communities using playas and saline lakes differed during both spring ( 2 16 = 2769.3, P < 0.001) and summer/fall ( 2 16 = 19,797.4, P < 0.001) migration. In spring J. Field Ornithol. Fall 2006 A. E. Andrei et al. 376 Table 1. Relative abundance and species composition (% of total number of observed birds per season) of migrant shorebirds using saline lakes of the Southern Great Plains, Texas and New Mexico, during spring (2 March–15 June) and summer/fall (7 July–9 November) of 2002 and 2003. Spring 2002 a Summer/Fall 2003 2002 2003 b Species N % N % N % N Snowy Plover 878 12.95 504 6.73 693 2.32 914 Killdeer 761 11.23 411 5.49 531 1.77 581 Black-necked Stilt 195 2.88 48 0.64 100 0.33 71 American Avocet 1673 24.68 809 10.80 1101 3.68 1775 Spotted Sandpiper 36 0.53 13 0.17 28 0.09 19 Greater Yellowlegs 21 0.31 44 0.59 56 0.19 109 Lesser Yellowlegs 42 0.62 4 0.05 39 0.13 68 Long-billed Curlew 46 0.68 12 0.16 28 0.09 175 Semipalmated Sandpiper 301 4.44 155 2.07 280 0.94 443 Western Sandpiper 942 13.90 487 6.50 2593 8.67 1713 Least Sandpiper 574 8.47 565 7.54 1078 3.60 1057 White-rumped Sandpiper 119 1.76 30 0.40 40 0.13 67 Baird’s Sandpiper 377 5.56 1053 14.06 5788 19.34 1957 Pectoral Sandpiper 35 0.52 45 0.60 59 0.20 12 Stilt Sandpiper 24 0.35 96 1.28 242 0.81 32 Long-billed Dowitcher 44 0.65 45 0.60 380 1.27 32 Wilson’s Phalarope 652 9.62 3138 41.90 16,746 55.96 2058 Other speciesc 59 0.86 31 0.42 142 0.48 55 Total 6779 7490 29,924 11,138 a See Appendix for scientific names and a list of all species observed at the saline lakes. b N = number of birds of each species observed per season. c See Appendix for a complete list of shorebirds observed in the saline lakes of the Southern Great during spring and fall migration, 2002–2003. % 8.21 5.22 0.64 15.94 0.17 0.98 0.61 1.57 3.98 15.38 9.49 0.60 17.57 0.11 0.29 0.29 18.48 0.51 Plains Table 2. Mean abundance (x¯) and species composition (% of the seasonal mean) of the common (>5 individuals observed per season) shorebirds observed per migration season during spring surveys at playas (1993 and 1994; Davis and Smith 1998a) and saline lakes (2002 and 2003) of the Southern Great Plains, Texas and New Mexico. Playas Species x¯ % x¯ a Snowy Plover 132 1.67 691 Killdeer 726.5 9.19 586 Black-necked Stilt 181.5 2.30 121.5 American Avocet 1893 23.95 1241 Spotted Sandpiper 92.5 1.17 24.5 Greater Yellowlegs 61.5 0.78 32.5 Lesser Yellowlegs 77.5 0.98 23 Long-billed Curlew 16 0.20 29 Semipalmated Sandpiper 84.5 1.07 228 Western Sandpiper 436 5.52 714.5 Least Sandpiper 227.5 2.88 569.5 White-rumped Sandpiper 5 0.06 74.5 Baird’s Sandpiper 182 2.30 715 Pectoral Sandpiper 103.5 1.31 40 Stilt Sandpiper 368.5 4.66 60 Long-billed Dowitcher 1401.5 17.73 44.5 Wilson’s Phalarope 1914.5 24.22 1895 Total 7903.5 7089.5 a See Appendix for scientific names and a list of all species observed at saline lakes. Saline lakes % 9.75 8.27 1.71 17.50 0.35 0.46 0.32 0.41 3.22 10.08 8.03 1.05 10.09 0.56 0.85 0.63 26.73 Vol. 77, No. 4 Shorebird Use of Saline Lakes 377 Table 3. Mean abundance (x¯) and species composition (% of the seasonal mean) of the common (>5 individuals observed per season) shorebirds observed per migration season during summer/fall surveys at playas (1993 and 1994; Davis and Smith 1998a) and saline lakes (2002 and 2003) of the Southern Great Plains, Texas and New Mexico. Playas Saline lakes x¯ % x¯ Species Snowy Plover 91 0.58 803.5 Killdeer 771.5 4.93 556 Black-necked Stilt 323.5 2.07 85.5 American Avocet 2560 16.35 1438 Spotted Sandpiper 46.5 0.30 23.5 Greater Yellowlegs 301 1.92 82.5 Lesser Yellowlegs 1352 8.63 53.5 Long-billed Curlew 2618 16.72 101.5 Semipalmated Sandpiper 111 0.71 361.5 Western Sandpiper 560.5 3.58 2153 Least Sandpiper 895 5.72 1067.5 White-rumped Sandpiper 5 0.03 53.5 Baird’s Sandpiper 464 2.96 3872.5 Pectoral Sandpiper 98.5 0.63 35.5 Stilt Sandpiper 1329 8.49 137 Long-billed Dowitcher 3493.5 22.31 206 Wilson’s Phalarope 640.5 4.09 9402 Total 15, 660.5 20, 432.5 a See Appendix for scientific names and a list of all species observed at saline lakes. a Wilson’s Phalaropes and American Avocets were dominant (>10%) both at playas and saline lakes, Long-billed Dowitchers were dominant only at playas, and Baird’s Sandpipers and Western Sandpipers were dominant only at saline lakes. Least Sandpipers (8.03% of the seasonal total at saline lakes and 2.88% at playas) and Snowy Plovers (9.75% of the seasonal total at saline lakes and 1.67% at playas) were also more dominant in saline lakes (Table 2). During summer/fall migrations (Table 3), Long-billed Curlews, Long-billed Dowitchers, and American Avocets were dominant (>10%) at playas, whereas Wilson’s Phalaropes, Western Sandpipers, and Baird’s Sandpipers were dominant at saline lakes. Snowy Plovers were found more frequently at saline lakes (3.93%) than playas (0.58%), whereas, Lesser Yellowlegs (8.63% vs. 0.26%) and Stilt Sandpipers (8.49% vs. 0.67%) were observed more frequently at playas (Table 3). Overall, migration (all shorebirds combined) was longer in the summer/fall than the spring (Fig. 2). Peak numbers were generally observed during 4–6 weeks in the spring and 6–8 weeks in the summer/fall. % 3.93 2.72 0.42 7.04 0.12 0.40 0.26 0.50 1.77 10.54 5.22 0.26 18.95 0.17 0.67 1.01 46.01 Spring migration chronologies of small prober/gleaners (Baird’s, Least, Semipalmated, and Western sandpipers) differed (2002: 2 27 = 1896.4, P < 0.001; 2003: 2 36 = 607.2, P < 0.001) (Fig. 3). Least Sandpipers and Semipalmated Sandpipers were the earliest migrants, followed by Baird’s Sandpipers in early April. Western Sandpipers were late migrants, reaching peak abundance during mid-April. Spring migration chronologies among the four representative species of shorebirds (small prober/gleaner—Least Sandpiper, medium gleaner/prober—Lesser Yellowlegs, gleaner/ sweeper—Wilson’s Phalarope, and gleaner/ prober—American Avocet) differed in 2002 ( 2 33 = 1396.9, P < 0.001) and 2003 ( 2 36 = 1244.1, P < 0.001; Fig. 3). American Avocets and Lesser Yellowlegs reached peak abundance during early April, whereas Wilson’s Phalaropes reached peak abundance during late April. Migration chronologies of Long-billed Curlews, Long-billed Dowitchers, and Stilt Sandpipers differed in spring 2002 ( 2 14 = 114.6, P < 0.001) and 2003 ( 2 24 = 67.3, P < 0.001). Long-billed Curlews were early migrants and departed by mid-April, whereas 40 40 2002 2002 Percent J. Field Ornithol. Fall 2006 A. E. Andrei et al. 378 30 30 20 20 10 10 0 0 1 15 1 15 1 15 1 1 15 1 15 1 15 1 15 1 15 1 15 1 15 1 40 40 2003 2003 30 30 20 20 10 10 0 0 1 15 March 1 15 April 1 15 1 1 M ay 15 July 1 Aug Sept Oct Date Fig. 2. Migration chronologies (percent of total number of observed birds for each season) of shorebirds (all species combined) using the saline lakes of the Southern Great Plains, spring and summer/fall 2002–2003. Stilt Sandpipers migrated during the second half of April and in May. Long-billed Dowitchers had a protracted migration lasting from mid-March to mid-May (Fig. 3). Summer/fall migration chronologies of Baird’s Sandpipers, Least Sandpipers, Semipalmated Sandpipers, and Western Sandpipers differed in 2002 ( 2 27 = 1896.4, P < 0.001) and 2003 ( 2 36 = 607.2, P < 0.001) (Fig. 4). Baird’s, Least, and Western sandpipers exhibited similar migration chronologies (July–September), whereas most semipalmated Sandpipers migrated from mid-August to mid-September. Summer/fall chronologies differed among American Avocets, Least Sandpipers, Lesser Yellowlegs, and Wilson’s Phalaropes in 2002 ( 2 51 = 3673.8, P < 0.001) and 2003 ( 2 48 = 2700.1, P < 0.001) (Fig. 4). In 2002 all four species migrated between mid-July and midSeptember and exhibited some overlap. In 2003 most Lesser Yellowlegs arrived in mid-September and departed by the end of September. Migration chronologies of Long-billed Curlews, Long-billed Dowitchers, and Stilt Sandpipers differed during summer/fall in 2002 ( 2 14 = 114.6, P < 0.001) and 2003 ( 2 24 = 67.3, P < 0.001; Fig. 4). Stilt Sandpipers were early migrants, reaching peak abundance during late July, followed by Long-billed Dowitchers and Long-billed Curlews during August. DISCUSSION Several wetland complexes are important stopover sites for shorebirds in interior North America. These include Horsehead and Sibley Lakes in North Dakota, Cheyenne Bottoms in Kansas, Great Salt Plains National Wildlife Refuge (NWR) in Oklahoma (Senner and Howe 1984), the prairie potholes (Deleon and Smith 1999), and playas of the SGP (Davis and Smith 1998a). At least 37,000 shorebirds use the saline lakes in the SGP annually. Because these lakes are scattered over a vast landscape, they should Shorebird Use of Saline Lakes Vol. 77, No. 4 American Avocet Wilson's Phalarope Lesser Yellowlegs 60 60 Percent 50 2002 40 30 30 20 20 10 10 0 15 1 15 1 15 Long billed Curlew Percent 1 1 15 1 15 Long-billed Dowitcher 1 15 1 Stilt Sandpiper 60 2002 50 50 40 40 30 30 20 20 10 10 2003 0 0 1 15 1 15 1 Baird's Sandpiper Percent 2003 0 1 60 Least Sandpiper 50 40 60 379 15 1 1 15 1 Semipalmated Sandpiper 15 1 15 1 Western Sandpiper 60 2003 2002 50 50 40 40 30 30 20 20 10 10 0 0 1 15 March 1 15 April 1 15 May 1 1 15 March 1 15 April 1 15 1 May Date Fig. 3. Migration chronologies (percent of total number of observed birds for each species) of 10 common shorebird species using the saline lakes of the Southern Great Plains, spring (March–June) 2002–2003. be considered as Western Hemisphere Shorebird Reserve Network (WHSRN) landscape sites of regional importance (Skagen 2005). Moreover, communities of birds using saline lakes differ from those using the 25,000 freshwater playas that are used by millions of shorebirds when they are wet (Davis and Smith 1998a). Thus, the mosaic of isolated wetlands in the entire SGP should be considered as a WHSRN landscape site of hemispheric importance (Skagen 2005). American Avocet Wilson's Phalarope Lesser Yellowlegs 60 Percent 50 50 40 40 30 30 20 20 10 10 15 1 15 1 15 1 Long billed Curlew 15 1 1 1 15 1 15 1 15 1 Stilt Sandpiper 60 2002 Percent 15 Long-billed Dowitcher 60 50 50 40 40 30 30 20 20 10 10 2003 0 0 1 15 1 15 1 Baird's Sandpiper 50 2003 0 1 60 Least Sandpiper 60 2002 0 Percent J. Field Ornithol. Fall 2006 A. E. Andrei et al. 380 15 1 15 1 1 15 1 15 Semipalmated Sandpiper 1 15 1 15 1 Western Sandpiper 60 2002 50 40 40 30 30 20 20 10 10 0 2003 0 1 15 July 1 15 Aug 1 15 1 Sept 15 Oct 1 1 15 July 1 15 Aug 1 15 Sept 1 15 1 Oct Date Fig. 4. Migration chronologies (percent of total number of observed birds for each species) of 10 common shorebird species using the saline lakes of the Southern Great Plains, summer/fall (July–November) 2002– 2003. Species composition and abundance of migrant shorebirds at saline lakes in our study varied between seasons and years. Seasonal and yearly variation in the composition of migrant shorebirds have also been reported at other wet- lands in the Great Plains (Skagen and Knopf 1994a, Davis and Smith 1998a). Variation in community composition and abundance of migrant shorebirds is likely a function of regional availability of habitats and food along migration Vol. 77, No. 4 Shorebird Use of Saline Lakes routes, nesting success and survival on the breeding grounds, and different migration routes during spring and fall (Morrison 1984, O’Reilly and Wingfield 1995). Comparison of our results with those of Davis and Smith (1998a) suggests differences among species in the use of playas and saline lakes of the SGP, with several species, including Snowy Plover (a highly imperiled species, USSCP 2004), Western Sandpiper, Wilson’s Phalarope, and Baird’s, Least, and Semipalmated sandpipers, observed more frequently at the saline lakes than at playas. Sandpipers (Calidris spp.), small-bodied shorebirds with many potential predators (Barnard 1980, Whitfield 1985), may prefer saline lakes because their sparsely vegetated mudflats offer good visibility for predator detection and avoidance (Metcalfe 1984). Differences among species of shorebirds using saline lakes and playas may also be due to changes in shorebird populations. Playas were surveyed in 1993 and 1994 (Davis and Smith 1998a) and the saline lakes were surveyed in 2002 and 2003 (our study). Some species, such as Wilson’s Phalaropes, may have expanded their range (Colwell and Jehl 1994), and populations of highly imperiled species, such as the Longbilled Curlew (USSCP 2004), have declined. Our results revealed differences in migration chronologies among species within guilds and among species in different guilds. Specifically, small prober/gleaners (Least, Semipalmated, Baird’s, and Western sandpipers), representative species from different foraging guilds (Least Sandpipers, Lesser Yellowlegs, Wilson’s Phalaropes, and American Avocets), and large and medium gleaner/probers (Long-billed Curlews, Long-billed Dowitchers, and Stilt Sandpipers) exhibited temporal segregation at the saline lakes during spring and summer/fall migration. As a result fewer individuals use food resources at the saline lakes at any given time and reduce the likelihood that resources will become limiting, particularly in the spring when invertebrate prey are less abundant (Davis and Smith 1998a, Andrei 2005). When food resources are depleted (Schneider and Harrington 1981) or potentially limiting (Székely and Bamberger 1992), shorebirds may not be able to replenish energy reserves and experience reduced survival and recruitment (Duffy et al. 1981, Schneider and Harrington 1981). 381 Unlike playas of the SGP (Smith 2003), most saline lakes are created and maintained by springs fed by the Ogallala aquifer that is heavily used for irrigation of crops and domestic uses (Triplet 1998, Sophocleous 2000). Further, the Ogallala aquifer is primarily recharged through playas of the SGP (Osterkamp and Wood 1987, Wood and Osterkamp 1987, Wood 2000). Therefore, conservation of playas, important stopover habitats for migrant shorebirds (Davis and Smith 1998a), may also help conserve saline lakes and the springs discharging into them. Preserving saline lakes of SGP as stopovers for shorebirds would likely require conservation of water on local and regional scales and purchase of easements for irrigation rights. Maintaining and restoring grasslands in areas surrounding playas and the saline lakes would reduce use of aquifer water for irrigation near saline lakes, likely reduce sedimentation in the recharge areas of playas, and restore and improve spring flows. ACKNOWLEDGMENTS We thank W. Johnson and S. McMurry for comments on the manuscript. L. M. Smith was supported by the Caesar Kleberg Foundation for Wildlife Conservation. This is manuscript T-9-1113 of the College of Agricultural Sciences and Natural Resources, Texas Tech University. 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Playa-lake basins in the southern High Plains of Texas and New Mexico-Part II, a hydrologic model and mass-balance arguments for their development. Geological Society of America Bulletin 99: 224–230. ZAR, J. H. 1996. Biostatistical analysis. Prentice Hall, Inc., Englewood Cliffs, NJ. Vol. 77, No. 4 Shorebird Use of Saline Lakes 383 Appendix. Shorebirds observed (“x”) in saline lakes of the Southern Great Plains of Texas and New Mexico during spring and summer/fall migrations, 2002–2003. Common name Black-bellied Plover American Golden-Plover Snowy Plover Semipalmated Plover Killdeer Black-necked Stilt American Avocet Spotted Sandpiper Solitary Sandpiper Greater Yellowlegs Willet Lesser Yellowlegs Long-billed Curlew Hudsonian Godwit Marbled Godwit Sanderling Semipalmated Sandpiper Western Sandpiper Least Sandpiper White-rumped Sandpiper Baird’s Sandpiper Pectoral Sandpiper Dunlin Stilt Sandpiper Long-billed Dowitcher Wilson’s Snipe Wilson’s Phalarope Red-necked Phalarope Scientific name Pluvialis squatarola Pluvialis dominica Charadrius alexandrinus Charadrius semipalmatus Charadrius vociferus Himantopus mexicanus Recurvirostra americana Actitis macularius Tringa solitaria Tringa melanoleuca Tringa semipalmata Tringa flavipes Numenius americanus Limosa haemastica Limosa fedoa Calidris alba Calidris pusilla Calidris mauri Calidris minutilla Calidris fuscicollis Calidris bairdii Calidris melanotos Calidris alpina Calidris himantopus Limnodromus scolopaceus Gallinago delicata Phalaropus tricolor Phalaropus lobatus Spring × × × × × × × × × × × × × × × × × × × × × × × × × × Summer/Fall × × × × × × × × × × × × × × × × × × × × × × × × × × × ×