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International Wader Studses 9:6-12

Shorebirds of the western of : overview and importanceto continental populations

Lewis W. Oring & J. Michael Reed

Oring, L. W., and J.M. Reed. 1996. Shorebirdsof the WesternGreat Basin of North America: Overview and importanceto continentalpopulations. International Wader Studies 9: 6-12. Large-scalestudies of North Americanshorebirds have, for the mostpart, emphasizedcoastal .Nevertheless, interior regions also are critical to shorebirdpopulations. Our aim herewas to reviewthe status of shorebirdsin oneimportant inland , the western Great Basin of North America. We identifiedspecies on whichmanagement efforts should be concentrated,identified importantwetlands, and estimatedthe globalimportance of westernGreat Basin shorebird populations.Nine speciesof shorebirdsregularly breed in thewestern Great Basin and 24 species occuras migrants. The western Great Basin is an importantbreeding area for SnowyPlovers (Charadrius alexandrinus), Long-billed (Numenius americanus), American Avocets (Recurvirostraamericana), Black-necked Stilts (Himantopus mexicanus), Wilson's Phalaropes (Phalaropustricolor), Killdeer ( C. vociferus),and Willets ( Catoptrophorus semipalmatus). Among migratingbirds, the western Great Basin supports large numbers, and significant proportions of the world'spopulations of Wilson'sand Red-neckedphalaropes (P. lobatus), Long-billed Dowitchers, WesternSandpipers (Calidris mauri) and American Avocets. Finally, we suggestpriorities for conservation efforts for shorebirds in the western Great Basin.

LewisW. Oring,, Evolution, and Conservation Biology Program, and Department of Environmental andResource Sciences, University of , 1000 Valley Rd., Reno, NV 89512USA J.Michael Reed, Biological Resources Research Center and Department of Environmental and Resource Sciences,University of Nevada, 1000 Valley Rd., Reno, NV 89512USA •Currentaddress: Department of Biology, Tufts University, Medford, MA 02155USA

Introduction , [Shuford et al. 1994], over 60,000 Wilson's Phalaropes[Phalaropus tricolor] at , [Jeh11988]; and 100,000Long-billed Shorebirdsare amongthe world's mostmobile Dowitchers[Limnodromus scolopaceus] in Lahontan organisms.In many cases,individuals spend parts Valley,Nevada [Neel & Henr)• thisvolume]). of their livesin differentnations and ecotypes. Perhapseven more important, however, is that the Hence,shorebird conservation is a complex vastmajority of individualsof certainspecies rely on international issue. In order to conserve these inlandwetlands, particularly during migration importantaspects of our fauna, it is essentialto (Senner& Howe 1984;Morrison & Myers 1989). understandthe dynamicsof shorebirdmovements acrossthe landscape.The mobilityof shorebirds If shorebirdmanagement and conservationin North yieldsan addedelement of complexitywhen it Americaare to be efficient,a thorough comesto determiningpopulation size, i.e, changes in understandingof thebiogeographic regions of the numbersmay resulteither from populationchanges interiorof the continentis needed.In thispaper we or from shiftsin spaceuse. Althoughestimates have addressshorebirds in oneof theseimportant regions, beenmade of numbersof someshorebird species in the westernGreat Basin. Our goalsare to review the regionsof the westernhemisphere (e.g., western statusof shorebirdsof thisregion, to identifythose USA, Page& Gill 1994;, Morrison & specieson whichmanagement efforts should Ross1989a, b), for most speciesthere are no accurate concentrate,and to estimatethe globalimportance of estimatesof globalpopulation sizes. westernGreat Basin shorebird populations. Large-scaleshorebird studies in North Americahave, The westernGreat Basin region (Figure 1), bordered for the mostpart, emphasizedcoastal regions (e.g., to the westby the SierraNevada and Cascade Morrison & Ross 1989a, b; Helmers 1992). This is ranges,is amongthe driestregions in North due in part to the largenumber of individualsof America. The mountains to the west form a someshorebird species that congregatein coastal shadow,and throughoutthe basin evaporation stagingor overwinteringsites. Nevertheless,interior exceedsprecipitation, sometimes by an order of regionsalso are critical to North Americanshorebird magnitude.As a consequence,water is in short populations(Eldridge 1992; Skagen & Knopf 1993). supplyand biodiversityis concentrated In some cases, numbers of shorebirds at interior sites disproportionatelyin (Brussard et al. in are similarto thoseat majorstaging areas on the press).Ironically, and perhaps because of the extreme coast(e.g., 200,000+ recurvirostrids at Great Salt premiumon surfacewater, wetlands of the region Oring & Reed:Great Basinshorebirds overview

L, Summ• L. & L.Abert

Nat'lo ! IlifeRefuge Alkalai

=yramidL. H•jmboldtWildlife Mgmt. Area Salt Lake • CarsonSink •'tillw3terWildlife IVIgrnt. Area "Carson L

WalkerL.

Owens

An'zona

Figure 1. Major wetlandsof the westernGreat Basin and the GreatSalt Lake of the easternGreat Basin. are of greatimportance to populationsof a number interyearvariability in existence(e.g., Engilis & Reid, of waterbird species. thisvolume). Types 2, 3, and 4 areespecially importantto shorebirds. Wetlandsof the westernGreat Basinregion include sixgeneral types: (1) ribbon-likeriparian areas, (2) Although important to North Americanwaterbirds, largedeep lakes, such as Pyramid and WalkerLakes, wetlandsin the westernGreat Basin are increasingly Nevada and Mono Lake, California, (3) shallow threatened.Formerly extensiveand diverseareas salinelakes of varyingsizes, including large lakes suchas Stillwater/CarsonLake (Nevada), and Mono suchas , Oregonand GooseLake, Lake (California),are recognizedas Hemispheric /California,and numeroussmall saline (former)or International(latter) sites by the Western playasand pans,(4) complexnatural and managed HemisphereShorebird Reserve Network (1993). Still, wetlands(marshes), e.g., Warner Wetlands, Oregon, theseareas are of greatconcern due to increased (5) upland bogsand wet meadows,and (6) artificial salinityand chemicalloading caused by urban and reservoirs.These wetlands are characterizedby agriculturaluses (Jehl 1988; Rubega & Robinson,this springrecharge from snowmeltand enormous volume). Otherwetlands of comparableimportance, International Wader Studies 9:6-12

but not as well understood, also are threatened. For 1994). In fact,birds disappeared from many of the example,while LakeAbert is undergoing sitesin which they had beenpresent: breeders at 20/ consideration as an Area of Critical Environmental 53 Californiasites, 6/29 in Oregon,and 2/6 in Concernby the Bureauof Land Management(Kerma .Counts in northwesternNevada have etal. 1990),there are proposalsto mineminerals, paralleledthis decline. Herman et al. (1988) harvestbrine shrimp(Artemia monica), as well as a estimatedthat therewere probably2000 breeders in requestfor licensefrom the FederalEnergy southeasternOregon and westernNevada in 1980, RegulatoryCommission license to alterwater and Bradleyet al. (1991)reported numbers had managementof the lake (Devaurs1990; Keister 1992; droppedfrom 878 in 1980to 139in 1991. The 1991 M. Stern,pers. comm.). countmissed one historically important breeding site, but even if the 1991 estimate were doubled, it showed a severe decline.

Shorebird Populationsof the Althoughthere are only moderatenumbers of Long- Western Great Basin and their billed Curlews in the western Great Basin, because of Relative Importance to Continental their range-widedecline (Page & Gill 1994)this area Populations mightbe importantto continentalpopulations. Locallydense populations of breedershave been recorded:4-5,000 pairs of AmericanAvocets and Nine speciesof shorebirdsregularly breed in the 1,000pairs of Black-neckedstilts (Table 2). westernGreat Basinand 24 speciesoccur as migrants (Table1). Sevenof the 24 occurin small to very small The westernGreat Basinis alsoimportant as numbers.At leastsix additionalspecies have been breedingand migratinggrounds for American reportedone or moretimes as accidentals (Ryser Avocetsand Black-neckedStilts. Thesespecies occur 1985;Alcorn 1988;LWO and N. Warnockper. obs.). commonlyand widely throughoutwetlands of the With severalnotable exceptions, the majorityof region,and in someyears large numbers of young which comefrom Mono Lake, California(e.g., Jehl are produced(Neel & Henry,this volume). Similarl}4 1986,1988; Herman et al. 1988;Rubega & Inouye Killdeerand Willetsbreed in largenumbers across 1994;Page et al. 1995;Shuford et al. 1995),there are the region,sometimes achieving high densities.For few publishedstudies of shorebirdsin the western example,up to 80 pairsof Killdeersand 50 pairsof Great Basin. In addition to studies at Mono Lake, Willetsbred around100 ha of pondsin oneyear. Hainline (1974)surveyed shorebirds in this areain Theselocally dense populations imply thatnumbers the early 1970s,and variousgovernment agencies in the westernGreat Basin as a wholecould be quite have conductedsurveys (e.g., Neel & Henry,this high. volume). Migration Breeding Birds The westernGreat Basinis probablyeven more The westernGreat Basinis an importantbreeding importantto shorebirdsduring migration (Table 3). areafor Long-billedCurlews, American Avocets, Thisarea is extremelyimportant to Wilson'sand Black-neckedStilts, Wilson's Phalaropes, Killdeer, Red-neckedphalaropes, Long-billed Dowitchers, and Willets,and westernSnowy Plovers (Charadrius AmericanAvocets. Some recent local peak counts alexandrinusnivosus) (Table 2). Surveysof Snowy supportthis assertion. Counts of bothWilson's and Ploversin Washington,Oregon, California, and Red-neckedphalaropes have been as high as50,000- Nevada showed a 20% decline in breeder numbers 100,000(July) at the LahontonValley, Nevada (Neel from the late 1970sto the late 1980s(Page & Gill & Henry,this volume), Lake Abert, Oregon, and

Table1. Habitatsused by shorebirdsin thewestern Great Basin.

Species HabitatNeeds

Breeding: SnowyPlover Charadrius alexandrinus alkalai flats with freshwater inflows Killdeer C. vociferus broad,open areas near water, especially with dikesor roads Black-neckedStilt Himantopus mexicanus vegetatedshallows or islandsin compexwetlands American Avocet Recurvirostra americana openislands, wetlands with predatorprotection, freshwater inflows for chicks Willet Catoptrophorussemipalmatus complexwetlands with nearbygrasslands or GreatBasin Long-billedCurlew Numeniusamericanus short-grassprairie, grazed pastures, flooded meadows, irrigated pastures;some tall vegetationfor chicksto hide Wilson'sPhalarope Phalaropus tricolor floodedmeadows, irrigated pastures, grasslands near complexwetlands

Migrating and Post-breeding: WesternSandpiper Calidris mauri mudflats with soft substrate LeastSandpiper C. minutilla marshedges, some vegetation Long-billed Dowitcher mudflatsadjacent to marshes Red-neckedPhalarope Phalaropus lobatus openwater, especially in salinelakes Wilson'sPhalarope openwater, especially in salinelakes American Avocet mudflats with soft substrate, shallow water Orlng & Reed:Great Basinshorebirds overview

Table 2. Importantcontributions of breedingshorebirds in thewestern Great Basin Region to North AmericanPopulationsL

Species WetlandType EstimatedStatus in the EstimatedImportance Western Great Basin

American Avocet shallowsaline lakes and playas, World populationunknown; Major contributor to complexvegetated wetlands 4,000-5,000pairs breeding at a speciespersistence. singlesite (Neel & Henry, this volume). Black-necked Stilt complexvegetated wetlands World populationunknown; Major contributorto 1,000pairs breeding at a single speciespersistence. site (Neel & Henry,this volume). SnowyPlover shallowsaline lakes and playas In 1980, 671 individuals were Major contributorto reportedin the Lahontan persistenceof Valley,accotinting for 69% westernsubspecies. of thisspecies counted in westernNevada (Page et al. 1991);over 5% of western U.S. population(Page & Gill 1994). Long-billedCurlew complexvegetated wetlands Commonbreeder, surveys Might be important lack comprehensiveness. contributor to speciespersistence.

•Thewestem Great Bash• region hosts five additionalspecies of breedh•gshorebirds: Killdeer, Common Snipe (Gallinago gallinago), Spotted Sandpiper(Actiris rnacularia), Willet, and Wilson'sPhalarope. However, we do not considerthe regionto be of criticalimportance to the persistenceof thesepopualtions due to theirgeneral abundance and the widespreadnature of theirbreedh•g distributions.

Mono Lake, California (Jeh11981,1986; LWO pers. SummerLake: This 10,100 ha shallow, alkaline lake obs.). On 29 April 1987,100,000 Long-billed supports10,000-100,000 migrating shorebirds (Fall) Dowitchers were seen at Carson Lake and Stillwater (Pageand Gill 1994). National Wildlife Refuge(Neel & Henry, this LakeAbert: Jehl (1988) reports an estimated150,000 volume). This is 20% of thosereported in western migrantWilson's Phalaropes in 1981in this 14,800ha North America(Page & Gill 1994). Lake Abert has shallow,alkaline lake. W. Devaurs (U.S. Bureauof had over 28,000avocets present on a singleday Land Management,pers. comm.) recorded peak, (unpubl.data), and 64,030were seenin the Lahontan single-dayshorebird counts of 89,288,with 34,819 Valleyin 1987.This is over25% of thebirds estimated American Avocets(October, 1993), 19,313 Least and to be in westernNorth America (Page& Gill 1994). Westernsandpipers combined (August, 1994), and Other migrantsfor which the westernGreat Basinis 27,600Wilson's and Red-neckedphalaropes alsoimportant include Western and Leastsandpipers combined(August, 1994). (especiallyin the spring). Therewas a combined WarnerWetlands: There are no publisheddata on countof 58,949Least and Westernsandpipers on the shorebirduse of this 13,000ha complex,but sameday (speciesnot distinguished)in the Lahonton thereis little doubtthat the areais importantto Valley (Neel et al. 1990). Basedon the ratio of birds westernGreat Basinshorebirds, especially Black- identifiedto species,approximately 50,200 were neckedStilts, Willets, and Long-billedCurlews. WesternSandpipers. In August 1986,the Nevada Divisionof Wildlife reported10,000 'peeps' that were Nevada mostlyLeast Sandpipers and 3,300leasts in October 1987 (Neel, unpubl.data), and LWO saw 8,000Least LahontanValley: The LahontanValley wetlands Sandpipersnear ,California (unpubl. includeStillwater National Wildlife Refuge,Alkali data, spring1994). Springcounts of over 23,000 Lake,Carson Lake State Game Management Area, LeastSandpipers have been reported for Summer and HumboldtNational Wildlife Refuge. In 1988, Lake, Oregon(Paulson 1993). thissite was declaredof Hemisphericimportance in the WesternHemisphere Shorebird Reserve Network (Neel et al. 1989;Neel & Henry thisvolume). This Wetlands of Critical Concern for designationresults from the siteharboring over Shorebirds in the Western Great Basin 250,000shorebirds during migration. Althoughthe LahontanValley hasnot had this many shorebirds Many wetlandsin the westernGeat Basinprovide recentlybecause of water diversionsand drought,it still actsas a stagingarea for many shorebirds,and importanthabitat for largenumbers of breedingor might returnto its formerimportance with increased migratingshorebirds. Below we summarizethe water allocation(for detailssee Neel & Henry, this limited informationavailable from published accountsand a few unpublishedaccounts. In the volume). During wet years,as much as 80% of all Long-billedDowitchers might usethis area during following list, reportedlake sizesrepresent those in migration. During April 1987,100,000 were seenat yearsof averageprecipitation (Grayson 1993). CarsonLake and StillwaterNational Wildlife Refuge (Neel et al. 1990). During someyears, the Lahontan Oregon Valley hassignificant numbers of other species,such MalheurLake: This complexlake/marsh is 18,400ha, as phalaropes,stilts, and avocets(Rawlings et al. and 1000-10,000migrating shorebirds have been 1987;Neel & Henry,this volume). For example,Neel recordedannually (Fall) (Pageand Gill 1994). & Henry (thisvolume) reported67,000 Wilson's International Wader Studles 9:6-12

Table3. Importantcontributions of transientshorebirds in the westernGreat Basin Region to North AmericanPopulationsL

Species WetlandType EstimatedStatus in the EstimatedImportance 2 Western Great Basin

Long-billed shallow, saline lakes Countsof 100,000locally 3 Veryimportant to speciespersistence Dowitcher and playas,complex vegetatedwetlands American Avocet shallow, saline lakes Countsto 64,000locally 3 Very importantto speciespersistence and playas,complex vegetatedwetlands Red-necked shallow, saline lakes Countsto 75,000locally 3 Criticallyimportant to speciespersistence Phalarope and playas,complex vegetatedwetlands Wilson'sPhalarope shallow, saline lakes Countsto 150,000locally; Likelyof importancefor species and playas,complex approximately10% of persistence vegetatedwetlands continentalpopulation 4 WesternSandpiper shallow, saline lakes Approximately60,000 Might be of importancefor species and playas,complex locally• persistence vegetatedwetlands LeastSandpiper shallow, saline lakes 23,150counted locally s Probablyimportant for persistence and playas,complex of westernNorth Americanpopulations vegetatedwetlands

•At least24 speciesrecorded as at leastoccasional transients in additionto 9 speciesof breedingshorebirds. Only thosespecies in which transientpopulations are of contintentalimportance are listed. Sixor moreadditional species could be consideredaccidental. Zrhereare no goodhemispheric estimates of numbersof thesespecies, and theseestimates are presented as hypotheses. 3Neel& Henry, thisvolume 'Jehl (1988) SPaulson(1993)

Phalaropesat StillwaterNational Wildlife Refuge in the westernGreat Basin is exceedinglydifficult (July,1987). becausespecies track water availabilityacross broad areas.Therefore, to properlymonitor species' California numbersrequires a coordinatedeffort that coversa largespatial scale (Jeh11988; Reed et al., thisvolume) MonoLake: Red-necked Phalaropes use Mono Lake becausefrom yearto yearindividuals might move duringmigration, with single-day counts as high as from onesite to anotherdepending on water levels. 21,600(Winkler et al. 1977). Jehl(1988) reported peak countsof migratingWilson's Phalaropes exceeding Thereare no federally(U.S.A.) listed endangered or 60,000(July, 1981), and Winkler (1977)reported threatened shorebirds in the western Great Basin, but 93,000(July, 1976). Only GreatSalt Lake (Utah) has the Long-billedCurlew, Mountain Plover (Charadrius moreWilson's Phalaropes during migration(Jehl montanus),and westernSnowy Plover are listedas 1988). During the 1930s,this deep,saline lake speciesof concern.All shorebirdspecies for which averagedover 22,000ha. Althoughthe lakehas thewestern Great Basin is importantto - declinedseverely, recent water allocationswill result wide populations,depend also on other in increased water levels. biogeographicareas; and each of thesespecies ranges acrosstwo or more countries(Table 4). Thus,we HoneyLake Valley: This shallow,alkaline lake has a cannotrely on managementefforts in theGreat Basin capacityfor covering23,000 ha. Page& Gill (1994) aloneto protectthese birds. One importantaspect of estimatedthat during migration,the lakeaverages future researchis movements of shorebirdswithin, 10,000-100,000shorebirds (Spring). Recently into and out of the Great Basin. Conservation efforts developedwetlands in the HoneyLake Valley shouldbe madein accordwith the following increasethe importanceof this area to shorebirds. priorities: GooseLake: On the borderof Oregonand California, 1. Identify and maintainwetlands of critical Page& Gill (1994)estimated that duringmigration concern for Great Basin shorebirds. this39,400 ha lakesupports 1,000-10,000 shorebirds; 2. Concentrate enhancement efforts on areas with this estimate is conservative. low cost,high qualitywater. Avoid relianceon deep ,irrigation return flow water, or water AlkaliLakes: Page & Gill (1994)estimated that during with high concentrationsof toxicelements or migration,these lakes have 1,000-10,000shorebirds; compounds. thisestimate is extremelyconservative for yearsin 3. Concentrate enhancement efforts in areas which lake levelsare up. wherethese efforts are compatible with existingland use practices. 4. Assure that natural freshwater inflows to Conclusions hypersalinewetlands are protected. 5. Supportinterstate and internationalefforts to Becausetime and moneyavailable for conservation establishwetland preserves and networksin accord are limited,conservation and managementefforts with items 2-4. shouldbe extendedto thosespecies that will benefit 6. Enhanceexisting Great Basin wetland the most. Monitoringand assessementof shorebirds preservesin accordwith items 2-4.

10 Oring & Reed:Great Basinshorebirds overview

Table 4. Shorebirdspecies in the westernGreat Basin that are of importanceto continentalpopulations, and the countriesin whichthey breed, migrate, and overwinter.

Species BreedingArea WinteringArea

Breeding: SnowyPlover w andcentral U.S. w coasts of U.S. and , , Cuba, , w and n coasts of South America American Avocet interior w North America, ne Gulf w and e coastsof Mexico, Gulf of Mexico, of Mexico, central e coast of U.S. Cuba, West Indies Black-necked Stilt sw, s central, and se U.S. coastal Mexico, , most of South America Long-billedCurlew interior w North America w coast U.S. and Mexico, w and n Gulf of Mexico Wilson'sPhalarope w and central North America s andw SouthAmerica, primarily centralArgentina

Migrant: WesternSandpiper nw coast , e coast Chukotski e and w coasts of U.S. Mexico, and Peninsula () Central America, n coast South America LeastSandpiper n North America s North Americathrough n South America Red-neckedPhalarope holarctic w central U.S., Pacific of South America coast, central and s , Arabian , Long-billedDowitcher ne coast Siberia, n and w Alaska, w coast North America, s U.S., Mexico, n n Central America

Herman, S. G., Bulger,J. B., & Buchanan,J. B. 1988. The Acknowledgments SnowyPlover in southeasternOregon and . J. Field Ornithol.59: 13-21. We thankJ. Jehl and N. Warnockfor commentingon Jehl,J. R., Jr. 1986. Biologyof Red-neckedPhalaropes this paper. LWO's researchwas supportedby the (Phalaropuslobatus) at thewestern edge of the Great National Science Foundation, the Nevada Basinin fall migration. GreatBasin Nat. 46: 185-197. AgriculturalStation, and the Forestand Rangeland Jehl,J. R., Jr. 1988. Biologyof the EaredGrebe and Wilson's EcosystemScience Center of the NationalBiological Phalaropein the nonbreedingseason: a studyof Service.JMR's research was supportedby NSF adaptationsto salinelakes. Stud.Avian Biol. 12: 1-74. Grant No. DEB-9322733 and the U.S. Forest Service. Keister,G. E, Jr. 1992.The ecology of LakeAbert: Analysis This is Contribution Number 50 of the Nevada of furtherdevelopment. Oregon Dept. of Fishand Wildlife Tech.Rep. 92-5-02. BiodiversityInitiative. Kenna,J., Devaurs, W., Troutman,D., King,G., Street,W., Cannon, B., Simontacchi, & Mondrell, V. 1990. Warner Wetlands Area of Critical Environmental Concern References (ACEC)management plan. U.S. Dept.Interior, Bureau of Land Management,Report BLM-OR-PT-90-28-1792. Alcorn,J. R 1988.The Birds of Nevada. Fairview West Morrison,R. I. G., & Myers,J. 1989.Shorebird flyways in Publishing,Fallon, Nevada. the . In: H. Goyd and J.-Y.Pirot (eds.), Bradley,P. V., Neel, L. A., & Turner,R.J. 1991.Population Flywaysand reserve networks for waterbirds, pp. 85-96. surveys,species distribution, and key habitats of selected International Waterfowl and Wetlands Research Bureau, nongamespecies. Nevada Division of Wildlife,Project SpecialPubl. No. 9. W-53-R-17,Study I, Jobs1-6. Morrison,R. I. G., & Ross,R. K. 1989a. Atlasofnearctic Brussard,P. F., Charlet,D. A., & Dobkin,D. S. In press.The shorebirdson thecoast of South America. Vol. 1. Canadian GreatBasin and MojaveDesert Region. In: P.Opler Wildlife ServiceSpecial Publications, Ottawa. (ed.), Statusand trends of biological resources in theUnited Morrison,R. I. G., & Ross,R. K. 1989b.Atlas ofnearctic States.National Biological Service, Washington, D.C. shorebirdson thecoast of SouthAmerica. Vol. 2. Canadian Devaurs,W.R. 1990. WarnerWetlands habitat management Wildlife ServiceSpecial Publications, Ottawa. plan.U.S. Dept.Interior, Bureau of LandManagement, Neel, L. A., Bradley,P. B., & Haley,R. D. 1989. Population ReportBLM-OR-PT-90-34-1792. surveys,species distribution, and key habitats of selected Eldridge,J. 1992. Management ofhabitat for breedingand nongamespecies. Nevada Division of Wildlife,Project migratingshorebirds in themidwest. Fish and Wildlife W-53-R-15,Study I, Jobs1-6. Leaflet13.2.12. Washington, D.C.: U.S. Departmentof Neel, L. A., Bradley,P. B., & Haley,R. D. 1990. Population the Interior. surveys,species distribution, and key habitats of selected Grayson,D. K. 1993.The desert's past: A naturalprehistory of nongamespecies. Nevada Divisionof Wildlife, Project theGreat Basin. Smithsonian Inst. Press, Washington, W-53-R-16,Study I, Jobs1-6. D.C. Page,G. W., & Gill, Jr.,R. E. 1994. Shorebirdsin western Hainline,J. L. 1974. Distribution,migration, and breeding of North America: Late 1800s to late 1900s. Stud. Avian shorebirdsin western Nevada. Unpubl. M. S. Thesis, Biol. 15: 147-160. Universityof Nevada,Reno. Page,G. W., Stern,M. A., & Paton,P. W. C. 1995. Helmers,D. L. 1992.Shorebird management manual. Western Differencesin winteringareas of SnowyPlovers from HemisphereShorebird Reserve Network, Manomet, inlandbreeding sites in westernNorth America. Condor Massachusetts. 97: 258-262.

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Paulson,D. 1993. Shorebirdsof thePacific Northwest. Universityof WashingtonPress, Seattle. Rawlings,M. S.,Haley, R. D., & Neel,L. A. 1987.Population surveys,species distribution, and key habitats of selected nongamespecies. Nevada Division of Wildlife,Project W-53-R-13,Study I, Jobs1-6. Rubega,M. A., & Inouye,C. 1994. Preyswitching in red- neckedphalaropes Phalaropus lobatus: Feeding limitations,the functionalresponse and water managementat MonoLake, California, U.S.A. Biol. Conserv. 70: 205-210. Ryser,E A. 1985.Birds of the Great Basin: A naturalhistory. Universityof NevadaPress, Reno. Senner,S. E., & Howe, M. A. 1984. Conservation of Nearctic shorebirds.In: J.Burger and B. Olla (eds.),Behavior of marineanimals, Vol. 6, Shorebirds:Migration and foraging behavior,pp. 379-421.Plenum Press, New York. Shuford,W. D., Page,G. W., & Hickey,C.M. 1995. Distributionand abundanceof SnowyPlovers winteringin the interiorof Californiaand adjacent states. Western Birds 26: 82-98. Shuford,W. D., Roy,V. L., Page,G. W., & Paul,D. S. 1994. A comprehensivesurvey ofshorebirds in wetlands at Great SaltLake, Utah, 10-11 August 1994. Unpublishedreport of PointReyes Bird Observatory, Stinson Beach, California. Skagen,S. K., & Knopf,E L. 1993.Toward conservation of midcontinentalshorebird migrations. Conserv. Biol. 7: 533-541. WesternHemisphere Shorebird Reserve Network. 1993. WesternHemisphere Shorebird Reserve Network Site Profiles.WA Publication No. 4. Wetlandsfor the , Manomet, Massachusetts. Winkler,D. W., Weigen,C.P., Engstrom, E B.,& Burch,S. E. 1977.Ornithology. In: D. W. Winkler(ed.), An ecological studyof Mono Lake, California, pp. 88-113.Institute of EcologyPublication No. 12. Universityof California, Davis.

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