The Ecology and Behavior of the Long-Billed Curlew in Southeastern Washington Author(S): Julia N

Allen Press

The Ecology and Behavior of the Long-Billed Curlew in Southeastern Washington Author(s): Julia N. Allen Source: Wildlife Monographs, No. 73, The Ecology and Behavior of the Long-Billed Curlew in Southeastern Washington (Oct., 1980), pp. 3-67 Published by: Wiley on behalf of the Wildlife Society Stable URL: http://www.jstor.org/stable/3830659 . Accessed: 27/08/2013 17:50

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This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions ' r t FRO,SITISPIECE. Adultmale long-billedcurlew. (Photo by MaryM. Tremaine)

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions THE ECOLOGY AND BEHAVIOR OF THE LONG-BILLEDCURLEW IN SOUTHEASTERNWASHINGTON Julia N. Allen

Battelle,Pacific Northwest Laboratories, Ecological Sciences Department,Richland, Washington 99352

CONTENTS

INTRODUCTION 6 Ground-Calling 36 Objective 6 Scraplng 36 Status of theLong-billed Curlew 6 Tossing ------36 ACKNOWLEDGMENTS 7 Courtship-Run 37 TAXONOMYOF THE LONG-BILLED CURLEW 7 Shaking 37 DESCRIPTION OF STUDY AREA 7 Mounting 37 GeneralDescriptior 7 Copulation 38 PhysicalFeatures 8 PostcopulatoryBehavior 38 Vegetation ---- 8 MIGRATIONAND BREEDING CHRONOLOGY 38 Weather 9 ARRIVALAND TERRITORIALITY 39 SpecificStudy Sites 9 DAILY ROUTINE ------42 PROCEDURES AND METHODS 12 SOLICITATIONAND INITIAL PAIR ScientificNames 12 FORMATION 43 Summaryof Records 12 COURTSHIP, MATINGAND PRELIMINARY StudyAreas 14 NESTING 44 Habit(ltAnalysis 14 NESTING 45 WeatherData 14 Nest Site Selection 45 Observationson CurlewBehavior 14 Nest Constructionand Materials 46 Nest Study ------15 Scrape Size ------46 Capturingand Banding 15 Nest Site Characteristics 46 Censusing 15 Nest Density 48 Weightsand Measurements 15 EGG LAYING 49 PredationStudy 16 LayingBehavior 49 Sex and Age Determination 17 ClutchInitiation and LayingIntertals 49 Duta Analysis 17 ClutchSize and Egg Description 50 PLUMAGE,MORPHOLOGY, AND SEXUAL Egg Losses and RenestingAttempts 50 MATURITY 17 INCUBATION 51 Plumage ---- 17 Behavior 52 Weightsand Measurements 19 Responsesto Disturbances 54 Age at FirstBreeding 21 NestingSuccess 56 DISTRIBUTION 21 HATCHING 56 BreedingRange 22 Behavior 57 WinterRange 22 BROOD PERIOD 58 BREEI)ING POPULATIONNUMBERS 23 Nest Departure 58 MAINTENANCEBEHAVIOR 24 Brooding 58 ComfortMovements 24 Behavior 58 Resting 27 Territoriality 61 Feeding ------28 Responsesto Disturbances 61 LOCOMOTION 30 Accidents,Injuries, and Mortality 62 AGONISTIC DISPLAYS AND POSTURES 31 STAGINGAND DEPARTURE 63 Mild Interactions 31 LITERATURE CITED 66 ViolentInteractions 34 APPENDIX INTERSPECIFIC INTERACTIONS 34 List of Common and ScientificNames of SEXUAL DISPLAYS AND POSTURES 35 PlantsMentioned in the Text 67 Bounding-SKKFlight 35

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INTRODUCTION ordsof long-billed curlews in those areas. Itis hopedthis information willbe useful The long-billedcurlew Numenius toagencies or individuals responsible for americanus is thelargest of the shore- thepreservation ofthe long-billed cur- birdsand yet is oneof North Americas lewand its habitat. lesserknown bird species. Its life historyS ecologyand behavior have until recently,received little attention. The paucity Statusof the Long-billed Curlew ofknowledge about the long-billed cur- Atone time, the long-billed curlew oc- lewrelates to its isolated and heretofore curred in large numbers over most of the remoteranges and to its status as a non- prairieregions of the United States and gamespecies; nongame wildlife have southernCanada. However, by the early onlyrecently begun to receivemanage- 1900s the species had already declined mentconsideration from governmental markedly. That reduction was a resultof agencies.The breeding range is restrict-(1) overzealoussport hunters, who took ed to scatteredlocations in thewestern advantage of the cllrlew's natmre of re- UnitedStates and parts of eastern Wash- turningto cireleover wounded com- ingtonstill provide suitable nesting hab- rades(2) markethunting (curlew wings itat. brought74¢ apiece in 1900,Stone 1901)> Thereis littlepublished information and (3) the"reclamation" ofvast areas of on the distributionand ecologyof the theirformer breeding range for farming long-billedcurlew, and until recently al- andranching (Wolf 1931 Sugden1933). mostall ofits known biology was based Today,curlews are restricted toscattered on fairlygeneral works written around populationsinthe West. They are rarely theturn of the century (e.g.> Bailey 1904 encounteredeast of the Mississippi Riv- Bent1907; Cameron 1907; Dawson 1909; er,whereas formerly they were a com- Dice 1918; Ferry1910; Grinnel]and montransient along the Atlantic Coast. Hunt1929; Oberholser 1918; Shufeldt The numbers observed on marshes flats 1913;Silloway 1902, 1909 Wickershamand fieldstoday represent a eorlcentra- 1902).More sophisticated long-billed tion in a restrictedrange rather than an curlewstudies are becomingmore nu- inereasein the number of birds (Sudgers merous(e.g., Campbell 1972> Forsythe 1933). Changing conditions in landuse 1967,1970, 1971, 1972, 1973; Graul 1971, southof the United States where some McCallumet al. 1977;Sadler and Maher curlewswinter have undoubtedlyalso 1976;Timken 1969)> but as wasnoted by contributedto theirreduced numbers. Palmer(1967) and Pettingill (1968)> there Iong-billedcurlews nest from southern is stillscant information on their breed- * 1 1 BritishColumbia, Alberta Saskatche- ng oenavlor. wanXand Manitobasouthward to Utah NewMexico, and Texas. They no longer Objective nestin Michigan,Minnesota, Wisconsin, Illinois Iowa, easternNebraska and The objectiveof this study was to ex- Kansas(American Ornithologists Union aminein depththe nesting ecology and 1957,Oberholser 1918). They winter behaviorof the long-billed curlew on a fromCalifornia? western Nevada Texasy breedingarea relatively free of disrup- and Louisiana southward toBaja Califor- tivehuman activity. Two summersof nia,Oaxaca7 Mexico, and Guatemalaw and fieldworkwere devoted to that endn and aIso fromSouth Carolina to Florida in 1976a postbreeding season survey of (AmericanOrnithologists Union 1957). themajor National Wildlife Refuges in Othermembers ofthe genus NuinenX4s Washington,southern Idaho, Utah Ne- in NorthAmerica are the whimbrel Nu- vadaSCalifornia, and Oregonenlarged menius phaeopus, the eskimo curlew N. thescope by providing unpublished rec- borealisand the bristle-thighed curlew

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N. tahitiensis.On rareoccasions, Euro- NORCUSLaboratory Graduate Partici- peancurlews N. arquatamay be sightedpant Program. Battelle, Pacific Northwest on theAtlantic coast, but they are dis- Laboratories,Ecological Sciences De- placedmigrants and notnative to the partmentprovided the use oftheir facil- UnitedStates. It is interestingtocompare ities and equipment on theArid Lands therelative abundance of the whimbrel, Ecology Reserve, Hanford National En- eskimoand long-billedcurlews today vironmentalResearch Park. withtheir status 100 years ago. At that I thankthe personnel at Battellewho time,the whiInbrel was the rarest of the providedtheir assistance and coopera- 3; today,it is themost common. The es- tion.I alsothank the U.S. Fish and Wild- kimocurlew is virtuallyextinct; the last lifeService and those National Wildlife confirmedsighting having been in 1972 Refugepersonnel who provided infor- (Iversen1976). Possibly, the long-billed mation from their Quarterly Reports. curlewcan hold its own, but only if mea- I am indebtedto MaryTremaine for surestaken to preserve it are soon forth- sharing her knowledge oflong-billed cur- coming. lewswith me, and forsuggesting tech- Althoughthe curlewis a protectedniques for making field observations and speciesthroughout North America, the photographsofthe birds. The pen and processof habitat destruction continues ink illustrationswere drawn by Dick at an everincreasing rate causing their r1tzner. numbersto decline. In bothsoutheastern Washingtonand northeastern Oregon, an TAXONOMYOF THE estimated30,000 acres per year of curlew LONG-BILLED CURLEW habitatare lost to cultivation (J.E. Kurtz 1976,Refuge Manager, Umatilla National The long-billedcurlew (family Scolo- WildlifeRefuge Umatilla, Oregon 97882, pacidae,order Charadriiformes) is closely pers.comm.). In view of that,public relatedto the snipe sandpipers, and yel- landssuch as NationalWildlife Refuges lowlegs. In theearly 1900s, considerable andthe Department ofEnergy Hanford debate was going on concerning the sta- Sitewhere curlews occur are bound to tusof the 2 racesof the long-billed cur- becomearid islands of refuge in a sea of lew andover correct names to be used irrigatedfarmland. For the foreseeable(Bishop 1910, Grinnell 1921, Oberholser filtureSlack of habitat protection threat- 1918, Ridgway 19197. Evidence preens thesurvival of the long-billed cur- sentedpoints toward a smallernorthern lew.Apparently, in response to this sit- raceversus a largersouthern race, but uation,the U.S. Departmentof the thereis notablesize overlapbetween Interior( 1973) has categorized the long- them.The AmericanOrnithologists' billedcurlew as 4extinction Numenius americanus americanus and hutsufficient information is not available N. a. parrus. Thebird I studiedin south- todetermine its status. More information eastern Washington belongs to the small- is needed. errace N. a. parv1ls.

AcKN( )WLEDGMENTS DESCSPTION OF STUDY AREA This researchwas supportedby the Description U.S . DepartmentofEnergy Division of General Biologicaland Environmental Research, TheU.S. Department ofEnergy's Han- underContract DE-AC06-76RLO 1830, fordSite comprises1,476.3 km2 or andby the Northwest College and Uni- 147,715.5ha ofshrub-steppe vegetation versityAssociation for Science under the atthe southern end of the lower Colum-

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largebasaltic rock outcroppings that rise to a heightof 338.94 and 224.94 m, re- spectively,between the central plateau andthe northern reaches of the river. The northernshoreline ofthe Columbia River is markedby the steep-walled"White Bluffs"rising 70-100 m abovethe bank forabout 11 km;steep banks are also foundalong the eastern border of the river abovethe city of Richland. Two sets ofdunes cross the Hanford Site. About 3,000ha ofactive sand dunes form a belt 6.4 kmwide immediately south of Han- FIG. 1. WashingtonState showing Columbia River fordtownsite to a pointopposite Ringold Basinand thelocation of the U.S. Departmentof onthe eastern bank of the river. The sec- EnergyHanford Site. ondgroup of dunes focuses on thearea betweenHorn Rapids on the Yakima Riv- erand the 300 Area, with the dune belt bia Basinin southeasternWashington dying out to the northwest and southeast. (Fig. 1). Formerlyin privateland hold- Thedunes trend northeastward. ings,the Atomic Energy Commission ac- TheColumbia River is the largest body quiredthe land in 1943as a nationalse- ofwater in theregion and flows within curityarea that was closed to agriculture, the Hanford Site for 83 km.Twenty is- grazing,and unofficialtravel. For the landsthat range in sizefrom 2.7 to 135.0 mostpart, all ofthe original residential ha lie within that stretch ofthe river. The andcommercial buildings have been re- riveris approximately800 m acrossand movedand only remnant trees and con- flowis manipulatedat a seriesof hydro- creteweir boxes stand as testimonytothe electricdams upstream. Other bodies of irrigationthat once serviced the area. To- waterinclude 4 smallponds on ornear day,there are 12 controlledor limitedthe centrally located plateau, Rattlesnake accessareas on theHanford Site identi- Spring at thebase of Rattlesnake Moun- fiedby area numbers (e.g., 300 Area, 200- tain,and severalflowing wells in the E Area,100-H Area, etc.). Each of those northeasterncorner of the site. areasis totallyenclosed by a highanti- personnelfence. Vegetation

P1>ysicalFeatures Commonand scientificnames of all plantsmentioned are listedin theAp- The moststriking topographic feature pendix. ofthe Hanford Site is RattlesnakeMoun- Threemajor vegetational types (Fig. 2) tainwhich bounds the site on the south- occur on theHanford Site west of the west.For 5 km,the crest of the mountain Columbia River (Cline et al. 1975):(1) is 1,100m high. The northern slope drops the sagebrush-bitterbrush/Sandberg's steeplyto 650 m and then eases down to bluegrass-cheatgrasstypecovers the low Cold CreekValley at 150 m. Northof elevationsin theeastern and southeast- ColdCreek Valley, the land surface again ernparts of the siteand occupiesthe risesto a plateauat about 225 m near the sandiestsoil short of dunes, (2) thesage- centerof the Hanford Site. From there, brush/bluebunch wheatgrass-Sandberg's theterrain slopes downward toward the bluegrasstype is confinedto theRattle- ColumbiaRiver through a series of snakeHills at elevationsabove 274 m, benchesor terraces.The GableMoun- and (3) thevegetation that occurs as a tain-GableButte complex consists of 2 broadzone between those 2 typesis the

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SAGERRUSHlBLUEBUNCH9TBffiSS- sagebrush/cheatgrass-Sandberg'sblue- SANDBERGSBLUEGRASS grasstype. A generalpaucity of herba- SAGEBRUSH-KIERERU9ISANDBER4 S BLUEGMSS-CHSTGMSS ceouscover tends to favorinvasion by 3 SAGE8RUSHJCHZTG@SS-SANDBERS S Russianthistle. ,r X Althoughmuch of the Hanford Site was r undevelopedatthe time of its inception, severalfarming communities were dis- placed.The fieldswere left fallow and weresoon invaded by cheatgrass. Nota- blyenough, those abandoned fields, sur- roundedlike islands by stands of native vegetationand isolatedover time from humanintrusion, provide the suitable UMBIA nestinghabitat for the long-billed curlew on the HanfordSite. Furthermore, the presenceof a loose substratein those fieldsseems to be an importantfactor in 0 5 determiningwhich areas are utilized by MIES curlews. FIG. 2. Boundariesof the 3 majorvegetational typeson theHanford Site, southeastern Washing- Weather ton. The climateof the Hanfordarea is greatlyinfluenced by the Cascade Moun- ordin theHanford area. The spring and tainRange to thewest. Annual precipi- summer of 1976were also windierand tationat theHanford Meteorology Sta- cloudierthan normal. Even so, the spring tion(at 223 m elevationnear the center growthof plants was forthe most part of the site)averages 16.5 cm but has unaffected,and theweather in general rangedfrom about 7 to30 cmduring the didnot appear to be detrimentalto suc- past30 years.On the average, 60 percent cessful curlew nesting. ofthe precipitation falls between Octo- Theweather during 1977 was bizarre, ber and February.Precipitation de- withmany months being substantially creasesafter the 3 wettestmonths of No- warmeror coolerthan normal and with vember,December, and January,but precipitationduring usually dry months. increasesagain to a secondarymaximum Coupled with the shortage of precipita- in June.June is also themonth of the tionin 1976,drought conditions persist- highestaverage wind speed, and only ed untilAugust when sudden rains final- oncein 7 yearswill it haveeven 1 day lyinitiated the "spring growing season" witha peakgust under 21 kph.Decem- formany plants. Consequently, when the ber,on theother hand, is themonth of curlewsarrived in Marchmany plants lowestaverage wind speed and has an hadnot germinated orbloomed, includ- averageof 10 suchdays. The climateof ingthe cheatgrass. The only nesting cov- theHanford region can thus be describeder in thefields was the dead vegetation as havingwindy springs, hot and dry fromthe previous year. summers,and moderatelycold winters. Julyis the hottestand driestmonth, SpecificStudy Sites whereasJanuary is the wettest and cold- estmonth. Althoughthe entire Hanford Site west Temperaturesduring 1976 averaged ofthe Columbia River was surveyed for 0.6 C coolerthan usual. Total precipita- curlews, only 3 studysites were selected tionfor the year was 46 percentof nor- forconcentrated behavioral observations mal,making 1976 the driest year on rec- (Fig.3). (Becauseof its relative inacces-

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* "r r--

r t

r-X WAHLUKE StOPE

rJ 100-H/100-[

300AREA STU DY S lTE

FIG. 3. The HanfordSite in southeasternWashington with asterisks (*) toshow location of concentrated studyareas. sibility,the Hanford Site north of the Co- (1) The300 Area study site was the pri- lumbiaRiver was notincluded in this marystudy area. Located west of the 300 study.However, groups of birds from the Areaand approximately 0.8 km from the WahlukeSlope area north of the Colum- Columbia River, that site encompassed biaRiver are included under the topic of approximately10.36 kmS of sagebrush- STAGINGAND DEPARTURE.) bitterbrush/Sandberg'sbluegrass-cheat-

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FIG. 4. Habitatat the 300 Area study site. Upper, South Field; Lower, Mosaic of shrub and field areas.

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions 12 WILDLIFE MONOGRAPHS grassvegetation and former homesteads drops off abruptly near the 100-H Area, witha fewremnant trees. Sandy, shrub butit descendsmore gradually towards coveredridges and small dunes are pres- the 100-DArea where a largecobble entand separate the relatively flat nest- beachof riverwash extends out along the ing fields(Fig. 4). The topographyis water. gentlyundulating and broken and lacks (3) The 100-Fstudy site of approximate- well-definedsurface drainage channels. ly 5.18km2 was southeastof the 100-F In thespring, the ridges and dunes dis- Areaalong the Columbia River. Though playa burstof flowers: long-leaf phlox, notvisited as *equentlyas thefirst 2 prickly-pearcactus, pale eveningprim- areas,it was usedmost consistently for rose,balsamroot, desert parsley, lupine, observationsduring the broodperiod. yarrow,fleabane, antelope bitterbrush, There is 1 largemain field of very dense broadiaea,wallflower, and tarweedfid- cheatgrass,and a testwell accessroad dleneck.Later in thesummer, Russian runsnorth and south through the site di- thistlegerminates in thefields and be- vidingthe field. Along the northern edge comesmore prominent. is an oldburn now covered with black- The fieldsthemselves are cheatgrassened big sagebrush stumps, Russian this- mixedwith Sandberg's bluegrass with tle,and grasses.Sagebrush/cheatgrass- occasionalpatches of Jim Hill mustard.Sandberg's bluegrass vegetation sur- Variationsof this include 2 fieldswith roundsseveral smaller cheatgrass fields. bluebunchwheatgrass sparsely distrib- Sand dropseed is veryprominent in one uted,and a fieldwith gray rabbitbrush field and asparagus continues to come up and big sagebrushsparsely distributed. each year in another.Jiln Hill mustard The mainvehicle entrance to the site is occursover much of the site, and several viaa testwell access road. treesgrow along the river bluff. The to- (2) The 100-H/100-Dstudy site was be- pographyisgently undulating with sandy tweenthe 100-Hand 100-DAreas di- ridgesexcept for the fields leveled when rectlyadjoining the ColumbiaRiver. originallyfarmed. Thatlocation is markedlydifferent from the300 Areastudy site in physicalap- pearance(Fig. 5). It is larger(approxi- PROCEDURES AND METHODS mately15.48 km2), more open, and lacks ScientificNames thelushness of vegetation atthe 300 Area site.The major vegetational type is sage- Commonand scientific names of birds brush/cheatgrass-Sandberg'sbluegrass. follow the AmericanOrnithologists' It is comprisedalmost exclusively of UnionChecklist ofNorth American Birds abandonedfields and orchards with only (1957),names of mammalsfollow Hall scatteredareas of sparselydistributed and Kelson (1959), and names of amphib- shrubs(big sagebrush,spiny hopsage, ians and reptiles follow Stebbins (1966). grayrabbitbrush, and buckwheat). Most Plantidentification and namesfollow of the fieldsare cheatgrass-Sandberg'sHitchcock and Cronquist (1973). bluegrass;Jim Hill mustard is extensive in someareas and Munro globe-mallow Summaryof Records is common,blooming profusely in the spring.There are 2 grovesof trees along In additionto the data collected in the theriver, several old fruitorchards, and field,I compiled all the available records individualtrees planted at old homesites. for the long-billed curlew on themajor The entiresite is criss-crossedwith old nationalwildlife refuges in Washington, roadways,most of them originally aban- southernIdaho, Utah, Nevada, Califor- donedby theAtomic Energy Commis- nia, and Oregon. Census information was sion.The surface is relativelyflat and de- takenfrom the refuge quarterly reports clinestoward the river.The shorelinefor 1960 and 1970 to June of 1976. I also

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FIG. 5. Habitatat the 100-H/100-Dstudy site. Upper, adjacent to ColumbiaRiver showing White Bluffs; Lower,vicinity of Study-plotRoad territoryshowing 100-D Area.

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions 14 WILDLIFE MONOGRAPHS drewupon communications with Han- andtotal litter canopy coverage were also fordSite personnel, U.S. Fishand Wild- estimated.As classificationnumbers are lifeService personnel, and reports of pri- oftenmodified by individualinvesti- vateindividuals. gatorsusing that method, they are given here:1 = 0-5 percent,2 = 5-25percent, StudyAreas 3 = 25-50 percent,4 = 50-75 percent, 5= 75-95percent, and 6= 95-100per- The 300 Areastudy site was selected cent.Frequency of occurrence was cal- becauseit supports the highest density of culatedas percentageofthe total number nestinglong-billed curlews on theHan- ofexamined plots in whicha particular fordSite. It alsocontains the major hab- taxonwas present. All field data on hab- itattypes used by long-billed curlews in itatanalysis were obtained between 6 othersoutheastern Washington breeding and 13 June 1977. areas. Fieldworkstarted 15 March1976 and WeatherData coveredthe breeding seasons of 1976 and 1977.I visitedthe 300 Areastudy site Summarizationsof the climatological dailyfrom early March until mid-July data from the fIanford Meteorology Sta- whenall thecurlews were gone. I spent tionat 223 m elevation near the center of timeat the secondary study sites on a ro- theHanford Site have been used. tatingbasis, the time of day,duration, andfrequency ofthose visits depending Observationson Curlew Behavior uponthe phase of thebreeding cycle. Otherareas of the Hanford Site were cen- Observations,aided by 7 x 50 power susedweekly or biweekly. binocularsand a 15-60variable power Descriptivedetails of the Hanford Site spottingscope, were made in 2 ways.The weredeveloped from my field notes, U.S. first,and most productive prior to hatch- GeologicalSurvey maps, and Battelle,ing, was from 6 stationaryblinds placed PacificNorthwest Laboratories' docu- in strategiclocations that overlooked 8 ments(Cline et al. 1975,Stone et al. territorynest fields on the 300 Area study 1972). site. The blindswere constructed of brownburlap and measured 1.2 x 1.2x Habitat Analysis 1.5 m witha windowon eachside that couldbe pinnedopen or closed. At the Replicated50-m transects were laid 300Area study site, my truck was always outin each of the major curlew utilization parked in the same location, and set path- areas.The transect lines were chosen so waysto the blinds were used. That rou- as to characterizethe nesting fields and tinewas intendedto reduceimpact on thefeeding habitats of each area. Canopy thevegetation and to habituate the birds coverageprovided by herbaceoustaxa tomy presence as muchas possible. was measuredby reading50 2 x 5-dm The secondmeans of observation was plotframes systematically spaced at l-m froma pickup truck parked along a road- intervalson each line transect. From 100 wayand was used during the entire 1976 to300 plots were read in eacharea, de- breedingseason. During 1977, it was pendingon the variabilityand utiliza-used on all curlewareas other than the tion.Canopy coverage for each taxon was 300Area study site and for all broodpe- estimatedvisually using a methoddevel- riodobservations. The majority ofthe be- opedby Daubenmire(1959). A modifi-havioral observations were made in the cationof the methodwas to separate open fieldswhere the birdscould be speciesinto litter and live categories and seenreadily, and descriptions ofdisplays totreat the 2 categoriesas separate "taxa" arebased on what was considered typical foranalysis. Total live canopy coverage for the species.

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NestStudy werecaught by handlater. I withheld markingthe birds in anyway that might Afterthe long-billed curlews had start- adversely affect their natural camouflage ed incubatingat the 300 Area study site andmake them more susceptable to pre- in 1976,the fields were searched system- dation. Notes requesting band returns aticallyfor nests. However, sitting and weresent to North American Bird Band- watchingthe birds proved less destruc-er, National Audubon Society, The Mur- tiveto the grass cover and far more pro- relet,The Auk, The Wilson Bulletin, The ductivein locatingnests. A nestlocation Wildlife Society, and The Condorfor was firstapproximated using the sur- publication.Flyers were also sentto roundingfeatures during nest relief, then Mexicoand to all thenational wildlife approachedand markedwith a small refugesin Washington,Oregon7 Idaho, whiteflag placed about 5 maway. California,Arizona, Nevada, Utah, New Theeggs in each nest were weighed in Mexico,and Texasrequesting informa- a plasticbag usinga 300-gspring scale tionon band sightings. andindividually marked with a colored doton the tip of the small end. The eggs weresubsequently weighed weekly until Censusing hatching.By the identifying dots, I was Censusingof long-billedcurlews on able to studythe fate of each egg,the theHanford Site was donethroughout hatchingsequence within the nest, and thebreeding season by makingweekly theduration of hatching. After each ex- roadcounts that consisted of stopping amination,the area around the nest and everyquarter mile (0.4 km),walking a alongthe approach route was sprinkledshort way off the road, imitating a curlew withparadichlorobenzene granules to call,and recording the number and sex maskmy scent that might otherwise lead of the curlewsthat responded. Birds coyotesCanis latransor other mamma- were also counted at loafingand staging lian predatorsto the nest site. Afterareas on a dailybasis toward the end of hatching,the broken shells were collect- the 1977 season. Also in 1977 1 Colum- ed *om each nest.Eggs that failed to bia Rivercensus was madeby boatin hatchwere also collected. whichthe islandsand shorelinesbe- Duringthe 1977 breeding season, ac- tweenVernita Bridge and Richland were tivenests were not approached and the searched. nestfields were not traversed. Walking throughthe fields left scars for the re- mainderof the season and tended to dis- Weightsand Measurements ruptbehavioral patterns. At the end of Each chickcaptured for banding was thebreeding season scrapes were mea- alsoweighed and measured. Small chicks sured,photographed, and described. wereweighed in a plasticbag with a 300- g hand-heldspring scale. Larger juve- Ca7oturingand Barzding nileswere weighed on a 1,610-gplatform balancewith their wings secured with StandardU.S. Fishand WildlifeSer- Velerostraps. Measurements were taken vicebands (size 5) wereplaced above the on all hirdswith vernier calipers or a footon all capturedlong-billed curlew millimeterstraightedge. The measure- chicks.A numbered,colored, plastic leg mentsare described here as somedo not band(National Band and Tag Co., New- followconventional methods. ( 1) Length port,Kentucky) placed on theopposite of tarsus (tarsometatarsus)as measured legwas used to identifyeach individual diagonally from the point of thetibia/ bird,year, and place of origin.Most metatarsusjoint behind to thepoint of chickswere weighed,measured, and themetatarsus/middle toejoint (Fig. 6a). bandedas soonas theyhatched. Others (2) Length of exposed culmen fromthe

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions : \ >with placed'\>taken.X gA thewith stretched F Natural tarsusin(6)feathers the Length wingstoHistory, Pugetthe was from of tip measuredTacoma, closedSoundtheof farthestthe Washington. aswingMuseumtoe the primaryexclu- (wing dis- of

16 WILDLIFE MONOGRAPHS

8| pointwhere the feathers of the forehead + t endstraight tothe tip of the culmen (Fig. :5 3 8 6b).(3) Lengthof bill from gape as mea- v suredby the chord, straight from the tip 9> \ , ofthe maxilla to the corner of the mouth (Fig. 6c). (4) Lengthof middletoe as : ;; measuredon the dorsal surface with the , b footbent back from the point of the joint siveof the claw (Fig. 6d). (5) Lengthof , ,,Gij =';,? S .S halluxas measuredalong the ventral sur- Gm C -<> facebetween the point of juncture with e thetarsus and thetip exclusiveof the claw(Fig. 6e). On olderjuveniles, wing chordand tail measurements were also

A %9\ chord)as measuredfrom the farthest an- g _ teriorpoint to the tip of the longest pri- W1 1 /X - mary,without attempting to straighten /9 the curveof the feather(Fig. 6fl.(7) d e Lengthof tail as measuredfrom the in- sertionpoint of the 2 middlerectrices to _ = ,yj_ thetip of the longest tail feather when thetail is closed(Fig. 6g). Three chicks againmeasured and weighed f Fouradult long-billed curlews were collected,weighed, and also measured as above.In addition,the extent of wings \ 1 tance betweenthe tips of the out- { / t - tipon one side to the farthest primary tip < __ onthe other by laying the bird flat on its 9 backand grasping each wing at the carpal jointto spread the wings out along a ruler k as faras possiblewithout injuring the 8 birdor flatteningthe wing quills (Fig. X X ; 6h). Studyskins were preparedand

u h PredationStudy Wl L Time-lapsecameras (Fitzner 1977) 4 n wereset up in several blinds overlooking nestsat the 300 Area study site to record FIG. 6. Measurementsof long-billedcurlews a, predation.The cameras were mounted in Lengthof tarsus, b, Lengthof exposed culmen, c Lengthof bill from gape; d, Lengthof middle toe. ra1nproofboxes l1ned with foam, then e, Lengthof hallux;f, Length of closedwing; g, mountedon woodenstakes inside the Lengthoftail; h, Extentofwingswith feathers. blinds.The cameraswere set to take1 frameevery 60 sec.When the predator at

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions THE LONG-BILLEDCURLEW IN SOUTHEASTERNWASHINGTONIIen 17 a nestsite was notseen or filmed,the remainingevidence was examinedfor possibleidentification ofthe predator.

Sex and Age Determination I judgedsex ofadult long-billed cur- lewsby configuration and length of the billin proportiontothe head. In profile, thebill of the female is morethan 3 times thelength of the head; in themale, the billis 3 timesor less than the length of thehead (Fig. 7). In addition,the bill of a malehas a moreperfectly symmetrical curvewhereas the female bill is rather flaton top with a morepronounced curve towardsthe tip. That method of distin- guishingsex was devised by M. M. Tre- maine( 1976,Ornithologist, P.O. Box 31152,Omaha, Nebraska 68131, pers. comm.)for curlews in Nebraskaand is accurateand easy to use aftera littleex- perience.Also, the femaleof a pairis oftenup to one-thirdlarger than the male,but is a difficultcriterion touse un- lessboth members ofthe pair are present forcomparison. Juveniles (birds of the year)can be separatedfrom adults by theirnoticeably shorter bill length.

FIG. 7. Comparisonof bill lengthof male and fe- Data Analysis malelong-billed curlews. Female in foreground. ProgramCOVER (Sauer and Owzarski 1979)written specifically tohandle hab- itatanalysis data obtained with Dauben- (Bailey1904, Bent1962, Brooks1920, mire's(1959) method was usedto com- Dawson1909, Grinnell and Hunt 1929, bine the habitatdata. That programGriscom 1939, Palmer1967, Ridgway providesthe following: percentage cov- 1919,Suckley 1860, Wilsonand Bona- erageby species,relative coverage and parte1831). Palmer(1967) andRidgway speciesdiversity, and frequencyof oc- (1919)made a distinctionbetween nup- currence.Weights and measurementstial and winter plumages, and both noted werecompared with t-tests. thatin the winterplumage the under partsare darker or deeper pinkish in col- or. Bent(1962) reportedthe juvenile PLUMAGE,MORPHOLOGY, AND plumageto be somewhatmore tawny SEXUAL MATURITY thanthe winter adult plumage, especially below,and the streaks on theneck and Plumage thebreast to be fewerand narrow. Suck- Adults ley (1860) thoughtthe overallrufous shadeof the adult plumage to be more The plumageof adult long-billed cur- distinctin the young. On the basis of the lewsis welldescribed by many authors 4 adultstudy skins in nuptialplumage I

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions WILDLIFE MONOGRAPHS 18 TABLE 1.-WEIGHTS (G) AND MEASUREMENTS(MM) OF LONG-BILLED CURLEWS AT HATCHING,HANFORD SITE, SOUTHEASTERNWASHINGTON, 1976

Culmen Chick Hatching no. date Weight Tarsus Fromgape Exposed Middletoe 1 11 May 52 47.1 27.0 20.3 33.8 2 11 May 51 45.3 27.0 20.2 34.4 3 11 May 50 43.3 25.3 20.7 33.8 4 11 May 56 44.3 26.0 19.6 33.5 5 11 May 51 45.2 25.2 18.5 33.5 6 12 May 50 43.6 25.3 20.6 35.8 7 12 May 44 45.1 23.9 21.5 37.7 8 13 May 56 45.2 26.5 21.3 35.4 9 13 May 52 46.5 27.8 20.3 35.1 10 13 May 53 42.8 24.8 19.0 34.8 11 14 May 47 44.2 24.5 19.5 38.0 12 14 May 50 42.5 24.7 20.2 35.9 13 14 May 49 44.7 26.2 21.5 36.9 14 25 May 48 46.7 25.8 21.1 36.0 15 25 May 50 43.8 25.2 22.0 35.7 16 25 May 50 45.5 26.5 22.3 35.5 17 29 May 45 39.7 23.2 18.5 30.6 18 29 May 47 42.5 26.5 20.8 35.0 19 29 May 46 40.4 28.5 22.1 34.2 20 29 May 47 41.6 25.6 21.9 32.4 23 3Jun 53 48.5 28.5 24.5 37.0 24 3 Jun 46 46.5 30.0 25.0 37.1 Mean + SE 49.7 + 0.69 44.3 + 0.47 26.1 + 0.34 21.0 + 0.35 35.1 + 0.38 preparedand 4 juvenilesexamined dur- thanthe male, and that as a rulethe male ingthe postnatal molt, I found the follow- shows more white on thewing coverts. ing: I cannotpick out either characteristic in (1) thestreaks on theneck were fewer the 4 studyskins I prepared(1 female andless pronounced (narrow and small- and3 male),and the sexes appear indis- er) in thejuvenile plumage than in the tinguishableonthe basis of plumage. In nuptialplumage (per Bent) addition,my field observations did not (2) exceptfor a fewlateral feathers, the revealany sexual dimorphism in plum- breastof thejuveniles was completelyage. Individual birds could be identified unstriped,unlike the adults that are con- by featheraberrations and/or variations spicuouslystriped, and in thefeather patterns, but could not be (3) thejuvenile plumage was moreru- usedas a keyto the sex of the bird. fousor with a deeperpinkish-cinnamon castthan the nuptial plumage (per Suck- Chicks ley).If the adultwinter plumage is a The downyyoung were described deeperpinkish or rufous than the adult brieflyby Allen and Kyllingstad (1949), nuptialplumage, then the juvenile plum- Bent(1962), Palmer (1967), and Ridgway age maybe moretawny when compared (1919) as basicallybuffy colored with ir- atthat time (per Bent). regulardark brown markings. According Grinnelland Hunt (1929) reported that toBent (1962), the buffy color varies from thefemale long-billed curlew has a much "warmbuff' on thebreast and flanks to pinkerflush over the bodyand wings "creambuff' on theface, upper parts,

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TABLE 2. WEIGHTS (G) AND MEASUREMENTS(MM) a largernumber of chickswere exam- OF LONG-BILLED CURLEWS AT HATCHINGIN UTAH ined,one probablywould find a grada- (FORSYTHE 1973) AND COLORADO (GRAUL 1971) tionof colors from one extreme tothe oth- Chick Hatching Ex osed er. no. date Weight Tarsus cu5)men Utah Weightsand Measurements 1 24 May 1966 56.6 35 22 2 24 May 1966 55.6 31 21 The weightand measurementsof22 3 24 May 1966 57.6 41 21 long-billedcurlews at hatching are given Colorado inTable 1. Table2 givesthose measure- 1 3 Jun 1969 66.0 46.0 22.0 mentsat hatchingpublished by 2 other 2 3 Jun 1969 62.0 47.0 22.0 authors,but neither of them described 3 3 Jun 1969 62.0 47.5 22.0 theirmeasurement procedures. I believe weightand exposedculmen measurementsare comparablebetween us, but andbelly, and to "creamcolor" on the notthe tarsus length because the wide throatwith the crown even paler. variationsuggests different measuring Afterbanding several chicks at the 300 techniques.A Bonferronit-test (Miller Areastudy site, I noticedwhat at first ap- 1966)indicates that the weights from the pearedto be 2 colorphases in the newly 3 studiesare significantly different from hatchedchicks. Of 6 broods,3 chicks one another(cx= 0.05); the long-billed wereentirely "cream colored" on the curlewchicks from Washington were sig- breastand bellyand 10 were"warm nificantlylighter at hatchingthan those buff'to varyingdegrees on thebreast fromUtah or Colorado.On the other andbelly. The 3 "creamcolored" chicks hand,an analysisof varianceon the werefrom separate nests. I thoughtthis lengthof the exposed culmen does not mightindicate a difference insex, but all showsignificant variation among the 3 12 chicksfrom 5 broodsbanded at the groups(a = 0.05). 100-H/lOO-Dstudy site were cream col- Table3 givesthe weight and measure- ored,and the probability is remote that mentsof juveniles and adults. Forsythe all thechicks would be thesame sex. If (1973)reported the growth in lengthof

TABLE 3. WEIGHTS (G) AND MEASUREMENTS(MM) OF JUVENILEAND ADULT LONG-BILLED CURLEWS, HANFORD SITE, SOUTHEASTERNWASHINGTON, 1976-1977

Culmen Bird Hatching Wing From Middle Wing no. date Sex Agel Weight chord Tarsus gape Exposed Tail toe Hallux span 21 3Jun 1976 Unk Juv 164 72.5 45.2 38.5 - 43.0 10 22 3Jun 1976 Unk Juv 187 73.0 47.2 39.7 - 43.4 10 25 7 Jun1976 Unk Juv 129 63.5 43.7 38.7 - 42.0 10 52 19 May 1976 Unk 8 days 55 49.6 30.7 22.8 - 37.7 11 213 lOJun1976 Unk Juv 250 81.9 53.4 48.4 44.4 10 132 14Jun1976 Unk 31 days 383 182 89.2 62.8 56.0 47.1 12 26 9 Jun1977 9 Ad 533.9 269 88.4 129 131 108 45.8 10 944 27 9 Jun1977 d Ad 494.4 255 88.2 124 120 99 44.8 11 887 28 9Jun 1977 d Ad 502.2 269 87.9 116 113 109 43.4 11 909 29 15Jun1977 d Ad 541.4 272 86.8 120 120 108 46.2 10 918 30 13 Jun1977 Unk Juv 308.3 163 89.4 59 54 62 43.0 11 l Ad = adult,Juv = juvenilechick of *lnknown age. 2 Recapturedafter banded as chick,see Table 1. 3 Recapturedafter banded as juvenile.

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions ALLENOBERHOLSERRIDGWAYGRALLEN-N.A.RIDGWAY119191-N.A.OBERHOLSERGRINNELL{1921)-N.OBERHOLSER(19B}-N.A.OBERHOLSERRRIDGWAYALLEN-N.A.INNELL I INNELLDGEWAY- 119211N.119191- (A. PARVUS 11910(19B111919)1919}-1190-N.A.- 1191QIPARVUS{1921) N. (1918)119181-11918) N.A.AMER N. 119181-- -N.A...... AMERICANUSN. -ICANUS ...... AMERICANUS A. - A.N.A. AMERICANUS- PARVUSPARVUS - N.A...... -PARVUSN. N.A. N.A. N.A.PARVUSN.A. N.A...... AMERICANUS...... AMERICANUS ...... AMERICANUS ,,,AMERICANUS ...... PARVUSPARVUS --PARVUSPARWS} .. W 100+- 100+ ------10D+100+, , 1>1F ...... 1F 1>100+1F1> .. ,,, -200+200+20Df200+ 2>2F2F

WILDLIFE MONOGRAPHS 20

WINGe 17SPEC} GR I NNELL( 19211 - N. AMER ICANUS ...... 200+ 248 2|1 273 OBERHOLSER ( 190 - N.A. AMERICANUS ...... , 2F {3 SPEC} 268 274 281

8 . {10 SPEC) 26&2 27-3 288

(7SPEC} 253|.52§?61

RIDGWAY ( 1919) - N. A. PARVUS...... 200+ {10+ SPEC) 253;5 285 287

ALLEN-N.A.PARVUS ...... 200+ (3SPEC) 255 265.3 272

WING99

18SPEC} 2 28} 5 291

{3 SPECt 26a5 28|6 298 308 RI DGWAY 119191 - N. A. AMERICANUS ...... 20> {11 SPEC 268 291.3

(3SPEC} 252 266 275 . I . /ll+SPEC} 251 26|5 274.6

{1SPEC} 269 a

EXPOSEDCULMAdY 12e2 13Lg 123 I 139 14Z8 155

137 145|.3 1552

106 §1 145

105;4 12&1 1448

113 117j712o

EXPOSEDCULMEN 99

- l - 203 9 159;6 17 166 196 222

s l - - l 219.2 z 163 184 118 1407 16,2

118.1 1 7 1T7

131 b

TAILdY

GRINNELL(1921) - N. AIVIERICANUS...... {7 SPEC} 95.2 1 §*610Bu5

OBERHOLSER(19181- N.A. AMERICANUS...... 13SPECI 109 181 1?8

RlDGWhY11919) - N.A. AMERICANUS...... (10 SPEC} 105.5 114.2 12&3

OBERHOLSER(1918} - N.A. PARVUS...... (J SPEC) 105 11|2 llt

RIDGWAY{1919} - N.A. PARVUS ...... (10+ SPEC) 96 1 | 6 123;7

{3 SPEC) 99 108.3 109

TAILvQ GRINNELL(1921) - N.AMERICANUS...... ( Z S PEC) 9z?6 105.4108.6

OBERHOLSER(19181- N.A. AMERICSUS ...... {3SPEC} 121 126 136

RIDGWAY41919) - N.A. AMERICANUS...... {11 SPEC} 104 1184 135.6

OBERHOLSER{lql8} - N.A. PARWS ...... {3 SPEC} 104;5 1g1 1}6

RIDGWAY119191- N.A. PARVUS...... {ll+SPEC} 104 11z2 116

ALLEN-N.A.PARVUS ...... <1 SPEC} 108

c FIG. 8. Presentationof measurements(mm) of adultlong-billed curlews. a, Wingchord; b, Exposed culmen;c, Tail.Values are maximum, minimum, and mean.

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TABLE 4. COMPARISONOF AVERAGE(MEAN + SE) WEIGHTS (G) AND MEASUREMENTS(MM) OF ADULT MALE AND FEMALE LONG-BILLED CURLEWS FROM WASHINGTON(THIS STUDY) AND CALIFORNIA (GRIN- NELL 1921). FIGURES IN PARENTHESESINDICATE THE NUMBERSOF SPECIMENS

Weight Culmen Wingchord Tail Males California(7) 693.0 + 21.85 132.91+ 3.59 261.0 + 2.94 103.62+ 1.72 Washington(3) 512.67 + 14.54 117.67+ 2.33 265.33 + 5.24 105.33+ 3.18 Females California(8) 815.64 + 36.34 175.81+ 6.11 282.63 + 2.67 105.84+ 1.09 Washington(1) 533.9 131 269 108 ' Only7 specimens. theexposed culmen, tarsus, and middle Age at First Breeding toeof chicks to be linearbut not at equal rates.I donot have enough data points to I observedsmall flocks of birdsthat determinelinearity. wouldpass through the nesting areas af- In the sameformat as presentedby ternesting was well underway.Those Grinnell( 1921), Fig. 8 comparesthe smallgroups would frequent the areas adultmeasurements found by me, Grin- adjacentto the territories forup to a week nell(1921), Oberholser (1918), and Ridg- ata timewhere they would be seenwalk- way(1919). The latter 2 authors classified ing and feedingleisurely as a group. theirspecimens into subspecies on the Theyshowed no migratory restlesssness. basisof size, but as Grinnell(1921) point- Occasionally,such a groupwould alight ed out,there is a significantdegree of inone of the nest fields to feed, but were overlapbetween the 2, andmost of the rarelydriven out by the territory holders. measurementsare formigrants. The In additionto this group behavior which specimensfrom Washington tend to align is distinctivefrom that of migrants, the moreclosely with the N. a. parsus spec- groupmernbers all appearedto havea imensof Ridgway and Oberholserthan shorter,male length bill. Taverner (1934) withtheir N. a. americanus specimens.stated that the bills of curlews continue Thisis particularlyapparent for exposed togrow for some time after apparent ma- culmen,and the shortness of the bill in turity.Possibly, the birdsI observed parsus as comparedwith americanus is were1-year-old subadults that would not thecharacteristic emphasized most. reachsexual maturity until the following Comparingthe measurements ofthe 3 yearand whose bills were, therefore, not malesfrom Washington (collected on yetfully developed. theirbreeding grounds) with the 7 males (collectedas migrants)measured in Cal- iforniaby Grinnell (1921), those for body weightand length of exposed culmen dif- DISTRIBUTION feredsignificantly (t-test, a = 0.05).Wing The presentdistribution ofthe long- chordand tailmeasurements, however, billed curlew has notbeen adequately showedno significantdifference at the described;although common in partsof 0.05level. Table 4 summarizeshis data itsrange, its distribution and abundance and mine.In summary,those weights throughoutits rangeare poorlyunder- and measurementsconfirm the place- stood.Included in thissection is infor- mentof Washington bred long-billed cur- mationfrom national wildlife refuge rec- lewsinto the smaller, northern subspe- ords,that is tenuousat bestbut does cies,parrus. providedistributional data on long-billed

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those,nesting populations are thought notto occurat Imperialor Havasu,the birdsbeing uncommon migrants. Al- thoughcurlews are year-round residents in theSacramento Valley (where Sacra- mento,Colusa, and Delevan are located), theyare not known to nest (Manolis and Tangren1975). Long-billed curlews are alsopresent throughout the year in vary- ingnumbers in the Imperial Valley (Sal- tonSea), but again are not known to nest (McCaskie1970). Washington Withinthe state of Washington, most of thecurlew population is restricted tothe ColumbiaRiver Basin. Fig. 10 shows the areaswithin and adjacentto the state whereknown breeding populations exist. Additionalpopulations undoubtedly ex- istelsewhere in thestate as well.The distributionofcurlews on theHanford Siteis shownin Fig.11.

WanterRange Long-billedcurlews do notwinter in Washington,Oregon, Idaho, Utah or Ne- FIG. 9. Locationsof national wildlife refuges from vada.As previouslynoted, curlews are whichrecords were obtained. year-roundresidents in theSacramento andImperial valleys of California. Spring migrantsapparently move northward up curlewsin the areas whereit was col- thewestern coast of Mexico and concen- lected. tratein the Gulf of California. The Salton Sea,just north of the head of the Gulf of CaliforniaXis the last large body of water BreedingRange thosebirds find before they encounter WesternUnited States the desertareas of easternCalifornia. Springand fallconcentrations of long- Nationalwildlife refuge records I have billedcurlews at theSalton Sea suggest summarizedshow curlewsduring the thatthey are wintering in Mexico or far- breedingseason on ornear the following ther south. refuges (Fig. 9): Columbia, Saddle Stenzelet al. (1976)also identifieda Mountain,McNary, Toppenish, Cold winteringpopulation ofabout 40 birds on Springs,Umatilla, Deer Flat (Sand Hol- BolinasLagoon northwest ofSan Fran- low areabetweerl Ontario and Caldwell7 cisco,California, on the PacificCoast. Idaho),Minidoka, Malheur, Clear Lake, Forsythe(1970) observed wintering cur- Upper Klamath Lower Klamath,Tule lewsin Texasat AransasNational Wild- Lake, Modoc,Sacramento, Colusa, De- life Refuge)Aransas County, and at levan, Salton Sea, Imperial,Havasu, Kingsvilleand PadreIsland National Stillwater,Ruby Lake, and Bear River. Of SeashoreMonument, Kleberg County.

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| ' A

- 49o

47o -

1 l o

------118°

FIG. 10. Areasin Washingtonwhere curlews were recorded in 1976-1977.

Bystrak(1974) mappedthe winteringlogistics of surveying a range character- areas of bird species summarizedfrom izedby widely scattered pockets of suit- AudubonChristmas Bird Counts nation- ablehabitat within forbidding terrain re- wide(a 1971-1972average in thecase of motefrom centers of human habitation. thelong-billed curlew). In additionto the Moreover,the pairs disperse widely dur- Sacramentoand Imperialvalleys in Cal- ingthe breeding season and their char- ifornia,he showedcurlews in the San acteristicshyness makes them difficult to JoaquinValley, the San FranciscoBay locate.None of those factors has encour- area,and along much of the coast. In Tex- agedan efforttomake a completecount. as, the birdsconcentrate inland around McCallumet al. (1977)reported 73 Midland(Midland County), around San sightingsinvolving 226 individualcur- Angelo(Tom Green County), and in the lewsin 13 countiesfor 2 breedingsea- areabetween those cities. The Texasand sonsin Colorado, but whether that figure LouisianaGulf Coast regions are shown is thebreeding population ofthe state or tobe verypopular. The onlyother sight- thesum of birds counted in 2 yearsis not ingsshown are in Florida:Monroe Coun- clear. ty,the Tampa Bay area,the MerrittIs- The HanfordSite and adjoining Wah- landarea, and aroundJacksonville. lukeSlope together support a long-billed curlewpopulation of approximately 300 NUMBERS birdsduring the breeding season. The BREEDING POPULATION HanfordSite west of the Columbia River The factthat no totalcensus of long- hasa residentpopulation ofapproximate- billed curlewsexists reflects formidable ly 100birds. Of those, approximately 60

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions 24 WILDLIFE MONOGR PHS arepaired, 20 areunpaired but territorial wing Stretch is performed slowly and de- malesand 20 are unattached individuals. liberately and lastsfor several seconds. It doesnot appear to have any social sig- MAINTENANCEBEHAVIOR nificanceand was mostcommonly ob- servedfollowing resting. ComfortMovements Wingand Leg Stretch.-Another form ofstretch is theWing and Leg Stretch Comfortmovements are those mainte- (Fig. 12c). In thatmovement a leg is nanceactivities that aid in the care of the stretchedbackward, extended, and held bodyand are not concerned with feeding, as thewing on the same side is partially locomotionor resting.Some of those extendedand stretcheddownward and movementsdiffer in formfrom group to backwardadjacent to it. The stretch also groupand have been used in classifica-does notappear to haveany social sig- tion.For example,Simmons (1957), by nificance.Only one birdat a timewas usingthe method of head scratching as a observedto give the Wingand Leg character,placed the recurvirostrids near Stretch,and whenit was observedthe thecharadriids rather than near the scol- birdswere walkingand feedingand oEracids.Many of the comfort movements merely paused briefly to stretch. exhibitedby long-billed curlews are al- Scratching.In thesuborder Charad- mostidentical in formto thoseof other rii,2 methodsof head scratchingare species)but have notpreviously been found.The scolopacidsscratch directly described. (i.e.,the foot is broughtdirectly to the Two-wingFlap. TheTwo-wing Flap head)and thecharadriids scratch indi- is a movementwhereby the head and rectly(i e., thebird lowers a wingand neckare stretched upwards and the angle bringsthe correspondingleg overthe ofthe back is inereasedslightly. Simul- shoulder)(Simmons 1961). Long-billed taneously,the wings are extended verti- curlews scratch the head directlywith callyinto the air over the head to their thefoot (Fig. 12d)and werenever ob- fullextent and flutteredbriefly. The servedto use the indirectmethod. wingsare then held extended in a pause Scratchingappeared to alleviate some ir- (Fig.12a). The bird rises on its toes, and ritatingstimulus and as sucha stimulus commonlyan individualwill lift off the was likelyto be presentat anytime, groundseveral centimeters while per- anotheractivity usually is interruptedfor formingthe fluttering phase of the move- scratchingpurposes. The foot is 1lsedfor ment.The Two-wing Flap was observed scratching all parts of the head and those regularlyinflocks of birds at loafing sites areasof the breast and neck the bird can whereit appearedto be preparatoryfor reach. Otherareas of the body are flight.It alsoappeared to signal impend- "scratched' with the bill, the tip of which ingflight to neighboringbirds and thus is directedto the location of the stimu]us was an aid in coordinatingdeparture. and the mandibles move back and forth TheTwo-wing Flap was also observed in rapidlyat the site. Some individuals birdson theirterritories, and in those tendedto scratchmore than others, cases it presumablyfunctioned in possiblya reflection ofthe presence, ab- stretchingcramped muscles. sence,or abundance of ectoparasiteKs the Two-wingStretch.-The Two-wingindividual harbored. Stretch(Fig. 12b) is a movemerltwhere- Bathing.-A bathirlgbird (Fig. 12e) bythe body is broughtto a nearhorizon- lowers itself into knee-deep water by talposition with the head and neck ex- bendingthe legs until the breast is at tended.Simultaneously, the wingsare waterlevel. The bird then wets itself by extendedvertically into the air over the repeatedlyrocking forward rapidly and backand to theirfull extent. The bird submergingthe head in a dippingmo- standssquarely on bothfeet. The Two- tion.Simultaneously the wings are quiv-

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r s

r- ._L_1 rl rX L_t rJ

r-> WAHtUKEStOPE rX

= FIG. 11. Locationsof areas on theHanford Site, southeastern Washington, diagonally lined, utilized by long-billedcurlews, 1976-1977. eredslightly. Once wet, the bird briefly preening, the bird will extend and flap its beatsthe waterwith its foldedwings. wingsseveral times. Bathing frequently Thisis donein a head-upposition. After- was observedto be performedsimulta- ward,the bird stands, shakes off the ex- neouslyby 2 ormore birds at thesame cesswater, moves to shallow water or to location.As bathing was a relativelyun- shore,and preens.Frequently during commonactivity, the sight of 1 bathing

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daythe birds stop and preen one partic- ulararea briefly before proceeding with anotheractivity. Preening was most com- monlyobserved interspersed with feed- ingor preceded by feeding and followed by resting.In flocksof curlewsloafing alongthe river, many birds are often observedpreening simultaneously, dem- onstratingthe social nature of the activity.In that situation, birds were sometimes observedpreening while standing on one leg,and occasionally a bird would lose itsbalance resulting in a "hop-and-flap" sequenceas thebird would attempt to recoverequilibrium without putting the otherfoot down. A hop-and-flapconsist- ed ofa birdhopping a shortdistance to theside on one leg while simultaneously extendingand flapping its wings several times. FeatherShaking. FeatherShaking is a generalizedruffling of the feathers broughtabout by shakingof thebody FIG. 12. Typicalpostures for comfort movements whilestanding on both feet with the head oflong-billed curlews. a, Two-wingFlap; b, Two- andneck extended out in front (Fig. 12f). wingStretch; c, Wingand Leg Stretch;d, Direct Asa comfort headscratch; e, Bathing;f, movement,Feather Shaking FeatherShaking. wasclosely associated with preening and resting,and both of those activities were commonlypreceded and followed by it. birdlikely Duringcold or rainy weather, the birds stimulatedothers to bathe. wouldstop and shake Solitarybathers, however, were also ob- frequently,pre- served. sumablyto enhanceheat retention. FeatherShaking was also observed in the Preening. Preeningconsists of ma- morningafter nipulating thebirds stood up from and arrangingthe feathers roostingforthe night and after bathing as withthe bill. Oil is obtainedfrom the previouslynoted. uropygialgland with the bill and is ma- Other nipulated Comfort Movements. Bill intothe feathers. While preen- shaking(not to be confusedwith Shaking ing,birds frequently will rub the top of behavior thehead discussedunder SEXUAL DIS- overthe back and sides.This PLAYSAND POSTURES)consists ofa violent helpsto spreadthe oil overthe feathers shakeof and thehead, usually only once. probablyserves to preenthe top of Birdswere observed to stop momentari- thehead which cannot be reachedwith ly,shake the thebill. head,and often scratch the Preeningbirds usually stand on headbefore proceeding. Therefore, this bothfeet with their feathers ruffled up. movementmay Apair will be in responseto some standwithin a meterof each irritatingstimulus. Additionally, a male other,facing the same direction,and wouldsometimes preen.During shakehis bill following windyand/or hot weather, aprolonged and unsuccessful mating at- thebirds seek the shelter of a largeshrub tempt. for protectionand shade. Complete Footshaking consists of rapidly shak- preeningofthe body lasts a minimumof inga foot and was 5min, but observedin 2 typesof manytimes throughout the situations.First, the movement was per-

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THE LONG-BIT.T.F*DCURLEW IN SOUTHEASTERNWASHINGTON-Allen 27 formedas a legwas being raised into the one-legresting position and was presum- ablyto removedebris. Second, shaking ofthe trailing foot was seen occasionally whilea birdwas walkingand feeding, andappeared to rid the foot of debris or entanglementinthe vegetation. Featherarranging movements, such as theshaking of the tail or lowering of the tail,are oftenobserved especially after bathingor when a birdis preening.Shak- ,.a"> ingof the tail was also sometimesper- formedby one or both members of a pair followinga courtship/matingsequence andby participants at the conclusion of an agonisticinteraction. In addition,a A birdwould often shake its tail while V walkingand feeding. < A b Whileresting or walking and feeding, birdsoccasionally were observed to raise theirfeathers in a waythat produced the samevisual effect as FeatherShaking. \

However,the plumage would often be fli8e',,,,m',,, heldin theraised position for a minute or so beforethe bird would resume its FIG. 13. Restingpostures of long-billed curlews. usualprofile. a, Bill-back;b, Hunch-down;c, Sitting.

Resting On theirterritories, they will seek wind Thebasic characteristic ofresting is a protectionand/or shade behind any avail- lackof or a minimizingofactivity that ableshrubs, even though such a position probablyresults in minimal expenditures appears to block their view in at least one ofenergy. The mostcommon behavior direction. thatcan be includedin restingis sleep- Restinglong-billed curlews assume ing,but unfortunatelyit is notalways severaldistinct postures. In termsof en- possibleto tell if a birdis, in fact, sleep- ergyexpenditure, the different postures ing.Consequently, noattempt was made probablyare roughlyequivalent and I to differentiatesleeping from resting. I wasunable to associate the different pos- consideredas resting(1) all birdsin one tureswith particular situations ortime in- of3 restingpostures and (2) birdsthat tervalsexcept that during inclement werestationary and preening at very low weatheronly the hunch-downposture intensities.Birds preening at lowinten- was observed. sitieswere considered as restingbecause In thebill-back position, the bill is preeningfrequently immediately pre- placedunder the scapulars; the legs may cedesand follows resting, and the tran- be in eitherof 2 positions.In one,the sitionfrom one to the other is difficultto birdstands on both legs, whereas in the identify.Also, when birds were mildly other the bird stands on 1 leg withthe disturbedwhile resting they frequently other tucked under the breast (Fig. 13a). preenedbefore resuming their rest. In theone leg method, the leg is brought Curlewsrest facing into the wind so up intoposition sometimes before and thatin any given area, all resting individ- sometimes after the bill is placedon the ualsare found facing the same direction. back. When the head is turnedaround,

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions 28 WILDLIFE MONOGRAPHS thewing on theopposite side is liftedbirdX usually the male, will fly over to the slightlytoallow the bill to slip under the otherand land nearby,then continue scapulars,but there was no correlationfeeding. Once incubation has begun, the betweenthe directionthe head was unattendingindividual feeds alone when turtledand the leg that was raised. When onthe territory. restingin that position is terminated,the Analysisof the different habitat areas headis turnedforward before the leg is usedfor feeding has revealed that at the lowered. 300Area study site the partially exposed In thehunch-down posture (Fig. 13b), duneareas and ridge areas had the great- thebird stands on bothfeet facing for- est variety of plant species. Those areas wardwith the head and neckretracted were also the most preferred for feeding. intothe shoulders. The feathers are often In anattempt tobe consistentwith oth- fluffedup, particularly during inclement er authors,I have assigned the feeding weather.Often too, the tail is droppedmovements ofcurlews to 2 categories:(1) belowthe level of the primaries. pecks,in whichonly the bill tip touches Long-billedcurlews often employ the substrate, and (2) probes,in which anothermethod of resting in whichthey thebill is partiallyor fully inserted into siton theground with both legs tucked the substrate. undertheir breasts. While sitting (Fig. Pecksform a largeproportion ofthe 13c),the bird usually faces forward with feedingmovements, but theirsignifi- theneck semiretracted. Less frequently,cance is notalways easy to assess. They thebill is turnedback and placed under areused to obtain prey on thesubstrate thescapulars. Curlews are likely to rest surface;however, most pecks do notre- sittingon hotdays, and migrantsmost sultin anyobvious capture. Curlews commonlyrest in that position. "walking-and-feeding"within a territory are in constantmotion as theyzigzag acrossan areawith bill pointeddiago- Feeding nallydownward, pecking here and there) and headsbobbing. Occasionallyy they Likeother waders, long-billed curlews may be seento walk with head held high are opportunists,feeding on whateverthen to run rapidly, sometimes for several foodsare available. Consquently, one ex- meters,ending with a peck(or a probe) pectsconsiderable differences in the forprey that presumably has been seen. dietsof birds at differentlocalities or at Huntingby sight has also been reported differenttimes of the year. by Burton(1974), Stenzel et al. (1976), Ontheir breeding grounds, long-billed and Hibbert-Ware and Ruttledge (1944). curlewsappear to feedentirely by day. Probescommence directly in frontor Theyfeed within and throughouttheir diagonallyto oneside. As thebill is in- territoriesas well as elsewhereon the sertedin a burrowor hole during a probe HanfordSite and in theirrigated fields a greatdeal ofmovement involving the nearby.Prior to incubation,a pair will headand neck and the legs and feet may feedtogether with one following the oth- be seenas thebird maneuvers to locate er orboth wandering in separatedirec- the prey. Vigorous side-to-side, forward- tionsand eventually rejoining. Common- and-back, and up-and-down movements ly I sawthem 50 mor more apart when ofthe head are used in those endeavors, feedingin the open fields. Usually, visual andoftentimes a bird will appear to stand contactis maintained)but vocal commu- on its head with tail tipped skyward inits nicationalso plays an importantrole es- efforttocapture a prey item. Turns of up peciallyin the shrub areas; a soft"Curlee to 180°with the bill inserted full length Curlee"or "Wheet Wheet" call is used. intothe ground are common.Subterra- Whentoo great a distanceis reached,one neanprey items are gripped in thebill

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions THE LONG-BILLEDCURLEW IN SOUTHEASTERNWASHINGTON Allen 29 tip and extractedbefore being swal- violentmovements with the bill near the lowed.Curlews walking and feeding in horizontal. a fieldprobe into existing holes or crev- icesas theycome upon them and at the PreySpecies basesof grass clumps. They give partic- ularattention tothick patches of vegeta- Long-billedcurlews were observed to tion,often probing in themrepeatedly. eat largeblack beetles and also other In sandysoil theywalk along pecking smaller insects that could not be identi- andprobing in the loose substrate, some- fiedfrom a distance.Superficial exami- timesviolently stabbing into holes and nationof exeretasamples revealed an othertimes calmly exploring depressions abundance of adult tenebrionid and cara- thatmay indicate a submergedbeetle or bid beetleexoskeletons. Other insect a rodentcache. Most searches are con- partswere also present but could not be centratedaround the bases of small forbs so easilyidentified. Subterranean insect and shrubs,and each bird makes many larvaeprobably constitute an important exploratoryprobes for each successfulfraction of the dietjudging from the catch. amountof time spent probing in loose Stenzelet al. (1976) described a "pause substrate. probe"used by long-billedcurlews to Timken(1969) and Sadler and Maher obtainprey in submerged areas. On 2 oc- (1976)reported long-billed curlew pre- casions,I observeda similar"pause dationon altricial nestlings of lark bunt- probe"technique used to captureprey ing Calamos piza melanocorys and fromburrows in the fields.The bird hornedlark Eremo phila alpestris, re- walkedalong, then stopped suddenly spectively. Horned larks are verycom- andposed with the tip of its bill at the monon the Hanford Site, but I neverob- openingof a burrowor withits bill in- serveda curlewpreying on a nest. sertedinto the burrow. It frozein that positionfor several moments, presum- RegurgitatedPellets ablyuntil detecting some movement below,then made a quickthrust into the Manywading birds regurgitate pellets burrow.Most attempts were fruitless and thatcontain the undigested hard parts of otherswere aborted as presumablyno theirprey. These can be usefulin study- preywas detected.After each one, the ingthe diet,and severalauthors have birdmoved on to another burrow and re- doneso withcurlews (Goss-Custard and peatedthe sequence. Jones1976, Hibbert-Ware and Ruttledge Anothertechnique for obtaining prey 1944,Stenzel et al. 1976).According to is to climbup intothe bases of shrubs Hibbert-Ware andRuttledge, the number andpoke among the branches with the andsize of the pellets are cyclic and vary bill.Where Munro globe-mallow occurs, with the time of year and with changes thebirds pay particular attention toit. An in diet.Their field observations showed individualmay circle a plantseveral thatwhen food changed from insects and timespoking among the branches, pre- grainsto thatof earthworms and other sumablygleaning insects from the fo- softfoods utilized during the winter and liage. springmonths, the pellets diminished in Aftercapture, the passage of small prey numberand size to a markeddegree. itemsup thebill is effectedby rapid, Thissuggests that pellets are rare or ab- smallopening and closing movements of sentduring the breeding season. the mandibles,accompanied by back- On theHanford Site, long-billed cur- wardjerks of the head as it is graduallylews werenot observed to regurgitate raised.Larger items are conveyed to the pellets,and no suchpellets were found mouthand swallowed by fewer and more in roostingor loafing areas even though

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FIG. 14. Normalflight posture of the long-billed '<,(t- / 1''--Xa\41¢, i i<41XX d curlew. FIG. 15. Posturesof long-billedcurlews used in aggressiveinteractions. a, Upright; b, Crouch-Run; c, Wing-Raising;d,Appeasement, appeasor on left, theirdiet is highlyinsectivorous through- opponenton right. outthe breeding season.

LOCOMOTION head beforetouching down. This puts theminto a stalland allows them to drop Innormal flight, the neck is heldslight- gracefully tothe ground. Or they may run ly foldedin a mannersomewhat analo- a meteror so with the wings raised before gousto that of herons, but not as extremecoming to a halt.In bothforms, the tail (Fig.14). Hamilton (1975), in comparingfans out as the birdtouches down. If the Americanavocet Recurvirostra alightinginshallow water, curlews hover americanaand the black-necked stilt Hi- withtheir legs dangling and lower them- mantopushimantopus, reported distinc- selves gingerly. Other types of special- tiveneck positions for each species. The izedcurlew flights are discussed later. long-billedcurlew holds its neckin a I neverobserved long-billed curlews mannersimilar to the stiltbut witha to swim,and wadingwas notcommon morepronounced fold. exceptwhen bathing. Most of the loco- Whentaking off, curlews customarily motion occurs on theground where the use theirlegs to obtainthe necessarybirds walk or run. Running was normally spring.The legsare broughtup to the observedduring agonistic interactions. flightposition soon after take-off and are Theexact postures used during walking, loweredjust before landing. When land- running,or wadingare determinedby ing,the birds swing upwards with the theother activities being performed si- wingsmomentarily fluttering high over- multaneously.

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AGONISTIC DISPLAYS intricatepatterns ofdives, swoops, turns, AND POSTURES andtwists are performedas 2 birdsfly arounda territory.Flying sequences are I havecategorized agonistic intraspe- commonly initiated following other ago- cificinteractions as mild (in whichno nisticinteractions, and the birdsmay physicalcontact occurs between individ- make numerous short flights from place uals)and violent (in which physical con- toplace before the sequence is terminat- tactbetween individuals does occur). Al- ed. In general,aerial pursuit continues thoughagonistic interactions arecommon until the individual being chased is driv- duringthe prenesting season, they been away. comeinfrequent once nesting has begun Hovering. Hoveringisa formof aerial and do noterupt again until the brood behaviorused by an aggressive male and season. Throughoutthis paper such mayfunction in locatingan opponent termsas aggression,aggressive, and ag- hidingon theground. The maleflutters gressorrefer to offensive behavior. overheadon rapidlybeating wings in a TheUpright posture used during such wayresembling the hovering ofan Amer- interactionsis characterized bythe birds icankestrel Falco sparveriusor a belted standingimmobile with the angle of their kingfisherMegaceryle alcyon. The hov- backsabove the horizontal and with their eringis ofshort duration (20 sec orless) necksextended and theirheads raised andconcludes with the aggressor landing high(Fig. 15a). It is oftenheld for several nearthe opponent. minutesbefore other postures are as- Crouch-Run. A displayoften used sumed.Frequently, when 2 birdsare in- duringmild interactions is the Crouch- teracting,both will assume an UprightRun. In thatdisplay, the head is lowered, posture,or as one bird approaches theback is heldat an anglebelow the anotherthe other will assume an Upright horizontal, the body feathers are raised, posture. thetail is fanned,and the bill is directed Duringintraspecific interactions, the outin front (Fig. 15b). The bird crouches orientationofthe bodies of the interact- down with the wings slightly raised from ingbirds is veryimportant tothem. If the the body.This displayis directedat birdsface each other, it indicates aggres- another bird and is accompaniedby sionor a tendencytoattack. Facing away movementtoward the other bird, either (orlooking away) indicates appeasement. by walking or (more frequently) byrun- Interactingbirds frequently stand paral- ning.It is usedby an aggressive bird and lel to eachother with their heads either is a runningthreat that serves to lessen facingthe same or oppositedirections. the distance between the aggressor and The bodies' parallelrelationship be- itsopponent. The Crouch-Runwas ob- tweencombatants probably is advanta-served to be used whenthe opponent geoussince such a positionwould enable waswithin about 10 mof the aggressor. eitherindividual to strike with its wings The displayshows an attacktendency orfeet. Attack or retreat are equally pos- andusually was continuouswith an at- siblefrom the bodies' parallel position. tempt to peck (or stab) the opponent, followedby a returnto an Uprightposture. Mild Interactions A stationaryform of the Crouch-Run is oftenperformed when the opponent is at Mildinteractions occur most common- close range, e.g., when a territorialbird lywhile a birdis stationaryorwhile it is is joinedby an outsider.In suchcases, runningorwalking on land. But they also theaggressor assumes the Crouch-Run occurin flight.Frequently, one bird will posture,aiming its bill at itsopponent, pursueanother that is alsoin flight.Ae- but withoutthe accompanyingmove- rialpursuit is generallyand for the most menttoward the opponent. partone bird chasing another, but often Theopponent's response to the Crouch-

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32 WILDLIFE MONOGRAPHS

Runvaries. Departure of the intruderof another bird. If the nonaggressor fails usuallyresults from a stationaryCrouch- to defendits position and fliesor runs Runor from a walkingapproach. A run- away,Supplanting has been effected and ningapproach concludes with (1) flightthe relinquishedposition will be as- ofthe intruder (2) flightof the intruder sumed by thesupplanter. If, however, andsubsequent pursuit by the aggressor, the nonaggressor does notrespond and (3) theintruder running or fluttering out remainsin theimmediate area, ground of immediatestriking range in which pursuitby the aggressive bird or violent casethe threat would be repeatedX(4)the interactionmay result. intruderassuming an appeasementpos- Wi7zg-Raising.In Wing-Raising(Fig. tllrethereby delaying or aborting a peck 15c),the bird essentially maintains an bythe aggressor or (5) violent interaction TJpright posture but withthe wings ofsome kind. raisedover the back.It is a standing u prtgnt-nun.-1le uprlgnt-nun 1S a treat cisp ay performec. Dy an aggres- runningthreat commonly used by an ag- sivebird and directed at another bird in gressivebird to pursue an opponent or to closerange. Wing-Raising is commonly lessenthe distance between it and its op- associatedwith Supplanting and the Up- ponentIt replacesthe Crouch-Runat right-Runand also functions as a means distancesof more than about 10 m.The of establishingindividual distance be- Upright-Runis essentially an Uprighttween flock members atloafing sites* Ab- postllreput in motion and may terrninate breviated Wing-Raising, in which the in anyof the other mild or violentdis- wingsare onlypartially raised, is more playsdescribed hereinX depending upon cornmonthan the full display and often thesituation and the response of the op- precedesthe full display. The intruders ponent. responseto Wing-Raisingusually is to Concealmerzt. Concealmentis a dis- departor to move further away. playin whichthe aggressor approaches Feather-Raising.Feather-Raising is a itsopponent then suddenly flops down generalizedraising of the body feathers in thegrass and disappearsfrom view. thatresults in a visualeffect similar to The birdassllmes a positionwith the thatof Feather Shaking. The pose is held headand neck low, the bill resting on the forseveral moments ata time,and the tail groundand the body flattened. The op- is oftenfanned simultaneously. Feather- ponentoften appears unable to locate the Raisingis usuallyterminated with Feath- concealedindividual and will walk back er Shakingafter which the usual profile andforth across the area as ifsearching is resumed.Feather-Raising was ob- forhim. When the opponent approaches served in a fewinstances among males. theconcealed bird theconcealed indi- Underthe circllmstancesinvolved 2 vidualsuddenly springs up at it in a maleswere competing for a femaleand Crouch-Runposture then instantly drops eachtime all 3 birdscame together the Dacq< into the grass out of sight. The op- maleswould perform Feather-Raising. ponentstands and watches as theaggres- TerritorialBoundary Display.-Terri- sor repeatsthe displaythe aggressortorial Boundary Displays involve2 eachtime moving closer to the opponent neighboring territory holders near the Eventuallyeither the concealingbird territorialline. Both birds assume an Up- will springup and proceedwith the rightposture and make a seriesof short Crouch-Runorthe opponent will depart. dashes directly towards or parallelto Howeverthe sequenceis oftenpro- eachother. Each dash, or charge, is fol- longedand sometimes results in violentlowed by an abrupt halt and violent grass interaction. pullingor displacementpecking in Supplunting Supplantingis anotherwhich chunks of duffand litterare formof mild interaction in which an ag- snatchedup in thebillS "chewed>> into gressivebird flies or runs to the position little pieces thendropped. Sometimes

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions THE LONG-BILLEDCURLEW IN SOUTHEASTERNWASHINGTON Allen 33 ratherlarge pieces of sagebrush or dead displayends when the paths of both in- floweringstalks of Jim Hill mustardare dividualshave diverged to a pointwhere pickedup in the bill and carried for short neithershows further aggression. distances.Each timea birdhalts and Appeasement. Appeasementbya bird strikesthe ground with its bill, its tail si- maybe indicatedby a generalcrouched multaneouslyfans out. In addition,the posturein whichthe bill is pointeddi- tipsof the wings are often dropped below agonallydownward, the neck is retract- thelevel of thetail. Some birds carry ed, andthe bird turns at an angleside- theirwings in that manner throughout an waysand away from its opponent (Fig. entireencounter while others drop them 15d).In responseto a runningthreat, an onlywhen making a charge.Still others appeaser will often remain stationary in neverseem to dropthem. The positionthat posture, but then "duck" down sud- varieswith the individual and with the denlywhen the aggressor approaches as intensityofthe encounter. ifin a reflex-likeresponse to an expected Neitherbird crosses the territorialblow. Other times an Appeasement-Run boundaryline nor comes within 1 m of willresult in whichthe same basic pos- itsopponent in mostcases. During such tureis maintained,but the birdruns interactions,the birdsalmost always away.Invariably, the aggressor pursues walkwith their heads turned at an angle ineither the Crouch-Run orthe Upright- towardsone another;looking away by Runposture, and suchchases are often one individualtriggers a charge on the protractedas those being pursued usu- partof its opponent. When one bird ini- allyare unwilling to leave. An Appease- tiatesa charge,the other stands Upright ment-Run was never observed to termi- andwatches. When a chargeis directlynate in violentinteraction, probably towardsan individual within close range, becauseof the nature of the display, but thatindividual takes a stancefacing the in all cases,intruders were eventually aggressor.The sudden halt at theend of drivenaway. suchdirect charges is sometimesaccom- Appeasementis also indicatedby sit- paniedby a swingingto the side instead tingor Iyingdown. That behavior was ofdisplacement pecking, so that the birds observedin females as a responseto male windup standingperpendicular to each aggression.The femalewould sit down other.Occasionally, both birds run at one withthe body flattened ina posturesim- anothersimultaneously and eitherhalt ilarto thatassumed for incubation. Her andsnatch up debris,or turn to the bod- headwould either be low,with the tip of ies'parallel position when they reach the the bill restingon the ground,or boundaryline. huncheddown but still in theupright Occasionally,a Territorial Boundary position. The male's response was either Displayerupts into violent interaction. to standand watch or to walkback and Whenit does, it usually is froma bodies' forthin frontof her. The female usually parallelposition in which one bird sidles remainedsitting for less thana minute, upto the other, raises its wings, and flips and immediatelyupon rising remained up to attackthe opponent with its feet. crouched.Feather Shaking usually was Often,however the opponentmoves performedbythe female before proceed- awaysideways rather than fight. ingwith another activity. Asa displayprogresses, the opponents TheBill-Down Display probably serves movealong the boundary line. Eventu- an appeasementfunction, presumably ally,their paths begin to diverge and they minimizingthe threatening potential of begindisplacement feeding. One indi- thebill. The body is heldwith the long vidualmay turn and surgeback to the axisparallel to theground. The neckis linein an Upright-Run after the other has retractedand the head is inline with the movedsome distance away, but then will body.The bill chattersand is directed againturn and also wanderaway. The downward,perpendicular tothe ground.

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions WILDLIFEMONOGRAPHS 34 interac- neitherparticipant in a violent showsany indications ofap- tioninitially demon- peasement;rather, both birds stratea definite tendency to attack. positionfor fighting Themost common attack isfacing front to front.The birds withtheir bills, claw- eachother stabbing withtheir ingwith their feet, and striking oftenflapping 0.5 m or upraisedwings, in a moreoff the ground as theyclash Violentfighting may also noisyruckus. thefeet beside to side. In thatposition, weaponsthough the arethe primary mayalso wingsare upraised and the bills Sidewaysencounters last beengaged. either onlya few seconds as one bird bothbirds turn to the movesaway or the front-to-frontposition to continue fight. maneuveris jumping, r Anotherattack onthe back of the op- of the long- withwings raised, Interspecificinteractions violentkicks with FIG.16. b, ArcDisplay; ponentand delivering billedcurlew. a, The Alert-Attitude; peckingwith the bill. c, Enticement-Run. thefeet while by Oftentimesa back attack is preceded the opponentwith runningtowards attacks wingsraised over the back. Such position,some birds remain still always,result in a recip- Inthat movementsusually, but not and/or butothers make little jerking rocalback attack by the opponent andback with the bill, while still forward move- front-to-frontfighting. com- othersmake very slow rhythmic Peckingor attempting topeck is andback with the bill. I aggressivebirds fol- mentsforward display monlyperformed by mademany observations ofthe a runningthreat. Avoidance or but only 1 between lowing ofthe op- involvingpairs, a fe- departureis the usual response nonpairedbirds. In thelatter case, a runningthreat toward an ponent. maleceased demonstrated intrudingmale when he INTERACTIONS Sinceall observations ofBill- INTERSPECIFIC thedisplay. birdsclose to- to DownDisplays involved Escapeis themost usual reaction in pairs,perhaps the display thebreeding season gether tendenciesin disturbancesoutside On servesto reduce aggressive re- oraway from the nesting territories. matethat permits the 2 birdsto duringthe breeding sea- the theterritories aredif- mainclose to each other. son,reactions toman or predators time,the Alert-Attitude ferent.At either bird ViolentInteractiorws (Fig. 16a),in whichthe alarmed heightwith elon- actual stretchesup toits full Violentinteractions involve is the firstresponse. The opponent.The gatedneck, Simmons physicalcontact with an Alert-Attitudewas named by andthe feet are all used itas a preparatory wings,the bill, But al- (1955)who described as weaponsby fightingbirds. movementindicating "suspi- interactionsare sometimesescape is oftenaccom- thoughsuch observedciousness." That attitude prolonged,an injurywas never witha strong"Wheet Wheet" call aresimilar panied in thearea toresult. Violent interactions but whichalerts other curlews inmany respects to mild interactions,

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions THE LONG-BILLEDCURLEW IN SOUTHEASTERNWASHINGTON-Allen 35 whothen also assume the Alert-Attitude. Cortus corax. It is uttered,most com- TheAlert-Attitude is held for a timeand, monlytwice, when the bird is approach- ifnecessary, escape by flight or a threating. The "karee"sound begins harshly ordistraction display follows. Otherwise) but then becomes more of a whistle;it is previousactivity is resumedas soonas utteredas thebird reaches the peak of its thedisturbance has passed. climband is followedby the"Curlee" Flyingthreats toward avian predators call as thebird heads away. The Arc Dis- areoften very elaborate. One or a fewof playis a maximumintensity attack re- thebreeding birds fly to the level of the sponseand appears to serve to intimidate predatorand fly at it withraucous cries a predatorand also to attract conspecifics of('WheetWheet'2 attempting tostrike it to the site. Similardisplays are per- onthe back with their feet. The predator formed by otherlarge shorebirds (e.g., mustusually twist and stoop to dodge the Americanavocet, Hamilton 1975; oyster- attackswhich continue until the predator catcher Hematopus ostralegusnblack- hasdeparted. tailedgodwit Limosa limosa, European The Chirpingcall signifiesan intensecurlew, and whimbrel,Simmons 1955). responseto a c.isturbance. I believe it to The Enticement-Runis used to lure a be thesame vocalization as 4'cackling"ground predator away from the nest or andthe long, rattling call, "que-he-he-he- young. The performing bird lands about he-he,"mentioned by other authors (e.g., 10 m in frontof the intruder, then runs Bent1962). The Chirping call is alsobe- alongthrough the grass with head turned lievedto be thesame as the"Ki-keck" sideways enticing the intruder tofollow. call describedby Forsythe(1970). That Ifthe intruder gets too close to the bird. callis an alarmcall given in responseto thebird flies further ahead, lands, and groundpredators and often attracts con- runsthrough the grass again. During that * n speclucs. display,the body is horizontaland the Injuryfeigning to distracta predatorhead is heldclose to the body (Fig. 16c). involvesmovement away from a nest site withthe toes just touching the ground, thewings flapping against the grass, the SEXUALDISPLAYS AND POSTURES neckoutstretched, and the head turned Bounding-SKKFlight sidewayswatching the intruder behind. A behaviorknown as Demonstrationin In theBounding-SKK Flight displawr, wadersis essentiallyan aerialand vocal thehird rapidly flutters upward, rising formof reaction akin to the"mobbing" almost perpendicularly, then sets its behaviorof othergroups (Simmons wingsin a downwardcurved arch resem- 1955).Demonstrating long-billed cur- blingan umbrella(Fig; 17a). In thatpo- lews circlean intruder,making short sition,it slowlyglides back down, pin- flightsfrom place to place giving repeat- ions motionless onthe breeze, sometimes ed '4WheetWheet,," "Curlee Curlee,X' coming to within 0.3 m of the ground be- andChirping calls. forerising again. The Soft"Kerr Kerr" In theArc Display, so namedby For- call consistsof a high-pitchedand very sythe(1970) and also notedby LaFave melodious<'kerr" note that tapers off at (1954)and Silloway (1902), the bird (usu- theend and is repeatedin a series.Each allythe male) flies rapidly straight atthe timethe call is repeated,the bird opens intruder(Fig. 16b). When within several itsbill only slightly and the sound is like metersof the intruderthe bird angles thatof wind blowing through a pipe. The straightupward, then cireles back to re- Soft"Kerr Kerr" call beginsas thebird peatthe performance. The "GuaahKa- reachesthe peak of the flight and contin- ree"call is givenduring this display. The ues untilit reachesthe bottom. Some- "guaah"is a harshguttural sound which times,a birdperforming bounding flight hasqualities similar to the caw of a raven givesthe <'Hee-who" call intermingled

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36 WILDLIFE MONOGR PHS

pitchedand very melodious (yet mourn- fulsounding) call that has several variations.The basic note is a longdrawnout "whee"that fades in, rises in the middle, andthen fades out. It is utteredwith the billonly slightly opened and is repeated in a series.The followingare variations on the basic "Whee" call: "whee," cs wnee-a-ee-a-ee, ,, Cs w neer, ,, Cs wnee-a-ee- a-eer,,, ss ee-a-ee-a-ee,,, ane 1 Cs eee.,, , U.s ne "Hee-who"call is sometimesintermixed withthe "Whee" call. Ground-Calling is performedby a maleto attracta female forbreeding. Occasionally, it is performedby a memberof a well-estab- lishedpair, but its function in suchin- stancesis unclear.

tcraptng

FIG. 17. Sexual displaysand posturesof the long- Scrapingis commonamong shorebird billed curlew.a, Bounding-SKKFlight display; b, Scraping;c, Courtship-Run;d, Shakingbehavior, speciesthat nest on the ground. In Scrap- male behind female;e, Mountingprior to copula- ing(Fig. 17b), the performing bird drops tion;f, Copulation. downonto its breast, wings slightly away fromthe body, with tail and wingtips pointedupwards. The bill is directedfor- withthe Soft"Kerr Kerr." The "Hee- wardand diagonallydownward as the who"call is anothermelodious call and feetare rapidlykicked backward alter- consistsof 2 drawnout notes, the second nately,pushing dirt up and outof the lowerpitched than the first. The Bound- shallowdepression (or "scrape") that is ing-SKKFlight display is a meansof so- formed.The birdmay rise and change licitationused by a maleto advertise his positionin thescrape frequently, each unpairedstatus. changeusually involving a rotationof Althoughno othercurlews have been about90 or 180°. Betweenscraping reportedto havethis exact display with movements,the bird will commonly in- itsaccompanying call, some species do spectthe scrape with its bill andpoke have aerialdisplays with similar ele- aroundin the bottom. Scraping produces ments.For example, the European cur- nestbowls and manyof them become lewhas a similarrising and falling flight scattered throughout a territory. Some of witha high-pitchedand flute-likecall thosesites are used repeatedlyduring thatis utteredon the descent (Bent 1962, courtship. Watson1972). The whimbrel gives a call describedas a verylong-drawn "kewiu" ,"1 osstng. thatis utteredwith scarcely opened bill and heardthroughout the day (Bent Tossingis a nest-buildingmovement 1962).Possibly, the call is similarto the thatappears to be essentiallythe same as long-billedcurlew's Soft "Kerr Kerr" the SidewaysThrowing Display de- call. scribedfor other shorebirds and gulls (Graul1973, unpublished dissertation, Ground-Calling Universityof Minnesota,Minneapolis, Minnesota).During Tossing, the bird In Ground-Callingthebird assumes an eitherstands with its back to the scrape appeasementposture and utters a high- andthrows nest material along one side

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions THE LONG-BILLED CURLEW IN SOUTHEASTERNWASHINGTON-Allen 37 ofthe body or alternately on eachside, malemoves closer to thefemale strad- or it standssideways to thescrape and dlingher tail, and climbing movements throwsnest material off to one side. The aremade with the feet. His bill at that nestmaterial is picked up in the bill then pointruffles the neck feathers of the fe- tossedinto the scrape in repeated, quick malealternately on each side,and she movements.The act mayalso be per- hasassumed a more horizontal body pos- formedwhile the bird is standingin the ture,though with head and neck still up- scrape. rightand looking around. The Shakingcall is givenby the male Coz4rtship-Run throughoutthe Shaking display. It is difficultto describe but sounds like an eve- TheCourtship-Run (Fig. 17c) is a pre- ningchorus of Pacific treefrogs Hyla re- copulatoryrunning approach posture gilla arounda pond. It also has a squeaky performedby themale. The neckis re- characteristic.When a birdis givingthe tractedunlike in the Upright-Run andthe call,the mandibles are not visibly sepa- backis notat an anglebelow the hori- ratedbut rathervibrate against one zontalas inthe Crouch-Run. The head is another.Forsythe ( 1970) briefly men- heldupright with the bill parallel to the tioneda "Sou"call he thoughtmight be groundand thebody feathers in their associatedwith copulation attempts. His usualposition. As themale approaches description ofthe behavior of the birds thefemale, the wings may be raisedpar- at the timethe call was heardcorre- tiallywith the primaries "flagging" in spondsclosely with the Shaking behavior preparationfor Shaking (see below) I observed,but his phonetic"sou sou whichfollows immediately. sou"rendition does not.

Shaking Mounting Shakingis a precopulatorydisplay per- Atthe height of his Shaking frenzy, the formedby the male. DuringShaking male mountsthe femaleby fluttering (Fig. 17d),the male assumes a positiononto her back. At that point, he flexeshis behindthe female with the wings raised legsuntil he is restingon histarsi with outto theside andthe primaries "flag- eachfoot cupped over a shoulderof the ging."The tail of the male is cockedup- female(Fig. 17e).While on thefemale, ward,the neck is outstretched,and the the malesmakes very rapid "marking angleof the back approaches the horizon- time" movements with his tarsi and feet tal.The footwork ofthe male is elaboratesuch that his body vibrates or oscillates as he movesback and forth behind the fromside to side. Simultaneously,he femaleon either side of her tail, violently holds his wingsabove his back and shakinghis head and bill out in front. As movesthem in a waythat helps to main- he shakes,the bill of the male ruffles the tainhis balance. The male's tail is cocked shoulderfeathers ofthe female. His head upwardsand the head and neck are up- bobs up and downsuch that his bill rightwith the bill pointeddiagonally tracesa U pattern,moving first down one downward. sideof her shoulders and then down the The femalestands with both feet other.The femalestands erect, looking squarely underneath. Her head and neck around.Occasionally, the femalewill arealso upright, but her bill is parallelto movea fewsteps ahead, but for the most theground and herback is horizontal. partshe stands still. Fromthe front, her trunk appears some- Asshaking progresses, the movements what dorsoventrally flattened, giving her ofthe male become more frenzied. The an ellipticalsilhouette. As the foot shuf- wingsare raised gradually until they are flingof the male continues, he gradually bentin a positionabove the back where pushesthe female's wings apart with his theyare fluttered ina jerkyfashion. The tarsisuch that her wing tips drop below

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PostcopulatoryBehavior ARRIVAL Thereare no postcopulatorydisplays PRENESTINGPHASE as such.The female Whistles as soonas themale dismounts, shakes her plumage, NESTINGPHASE and beginswalking and feeding,occa- sionallypausing and Feather Shaking a CHICK PHASE secondtime. The male dismounts, stands FLOCKINGPHASE briefly,preens, and thenalso begins walkingand feeding or, less commonly, l l l l l l l l l l l l l l l fliesto another part of the field. 142128 7142128 7142128 7 142128 7142128 MARCH APRIL MAY JUNE JULY MIGRATIONAND BREEDING FIG. 18. Breedingchronology of the long-billed CHRONOLOGY curlewon the Hanford Site, southeastern Washing- ton,1976-1977. The springmigration is a general northwardmovement, but actual routes havenot been identified. MY useof color thelevel of her tail and one tarsus slips bandswas intended to provide informa- slightlyunderneath each wing.Mount tionon migratorypathways, but no re- timeprior to copulation is about60 sec. turnshave been reported. Themale continues tocall after mount- Smallflocks and singlebirds sporad- ingand a rapidlyrepeated "wee" note is icallypass through on their way to other discernible.I believe the vocalizations areas after the long-billed curlews have heardduring Shaking to be thesame as startedbreeding on the Hanford Site, and thoseduring mounting, the only differ- as theseason progresses, migrants and encebeing a certainamount of distortion residents can easilybe confused.The causedby the action of the head during transientsrest or feedin or nearareas Shaking. wherethe residentsare established. Whenflying through a nesting area, mi- Copulation grantscommonly give a characteristic "WhortWhort" call that is repeatedcon- Themale's tail passes around the side tinuallyby 1 birdat 2-sec intervals. The ofthe female's just prior to dismounting. call appears to havethe effect of adver- His taildrops under one of her wings as tisingtheir migratory status and therefore shetips slightly forward (Fig. 17f). Cop- dispellsany agonistic displays by the ter- ulationis brief(about 2 sec); themale ritorialresidents . Migrants not using the fallsbackwards as his tail is pressed "WhortWhort" call give the "Wheet down,fanned, and his wingsare flut- Wheet"call as theycross the territories. tered.Simultaneously, thefemale "falls (Localpairs or individuals often fly over forward"a few steps moving out from onenesting area on their way to and from underthe male. theirown. Those birds fly along estab- If a copulatoryattempt fails, the male lishedroutes that do notcross directly regainshis position on the female's back overthe nest fields.) withfeet shuffling and tries to copulate Whenmigrants land in a territorydur- again5 to 10 sec later.Rarely, 3 or4 at- ingthe day, they alight in the open field temptsare madebefore coition occurs. first.They walk and feed very rapidly and Oftentimes,however, once an unsuccess- often cock their heads skyward. Their be- fulattempt has been made by the male, haviorconveys a certainrestlessness or the femalewill no longerstand and nervousnessthat is particularlyapparent movesout from under him before he can in thosethat are paired.Members of a continue. pairmove in unisonremaining within 1

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions THE LONG-BILLEDCURLEW IN SOUTHEASTERNWASHINGTON Allen 39 m of each otherand, occasionally,1 maybe seenresting or loafing in them for birdgives the "Whort Whort" call as they severalhours at a time.The birds usually feed.Migrants usually linger in an area just sit,but occasionally1 will stand onlybriefly, commonly less than an hour and preenor wandernearby feeding. duringthe day, but sometimes they stay Now and again,1 will cock its head to roostmost of the night.Those that andlook skyward. spendthe night Whistle back and forth In 1976,no curlewswere seen on the forabout a halfhour before departure at HanfordSite after 1 July.A laterecord dawn.Fig. 18 summarizesthe breeding was 1 birdin Richland,Washington, on chronologyofthe long-billed curlew on 24 October1976. Three curlews (2 at theHanford Site. LeadbetterPoint and 1 atOcean Shores) Arrivaldates were 17 March1976 and werealso reportedby birdwatchersfor 21 March1977 forthe Hanford area. At theWashington coast in the fall of 1976, the UmatillaNational Wildlife Refuge butlong-billed curlews are considered approximately40 km southward, arrival rare migrants west of the Cascade Moun- dateswere 21 March1968, 14 March tains.In 1977,a largeconcentration of 1969,24 March1970,13 March1971,20 long-billedcurlews left the Hanford Site March1975, 15 March1976, and 21 from15 to 23 June,and the last bird was March1977 foran averagedate of 20 seenon 22 July. Late records for the year March.For the years 1956-1971, Little- wereof birds feeding in alfalfa fields east field(1973) foundthe earliest and latest ofRed Mountain adjacent to the Hanford arrivaldates at MalheurNational Wild- Sitethe week of 20 August, and 2 curlews lifeRefuge in southeastern Oregon to be onthe McNary National Wildlife Refuge 22 March1970 and 17 April1967, re- on24 August. spectively.The averagearrival date at Malheuris 28 Marchfor that period. Mal- ARRIVALAND TERRITORIALITY heuris approximately335 kmsouth of Hanfordand one wouldexpect curlews Firstobservations were of single birds toarrive there first. Possibly, the 2 pop- (males),but within a week numerous in- ulationsof birds follow different migra- dividuals had arrived and adult females toryroutes. continuedto tricklein throughthe first The firsteggs were laid on about8 weekof April. In Utah,Fcrsythe (1970) Aprilin 1976and on about2 Aprilin reporteda period after arrival when small 1977.A fewadults may have still been flockswould frequent plowed fields and incubatingin June,but most eggs pastures.That period would last for sev- hatchedby the end of May. Chicks began eral weeksbefore agonistic behavior appearingon 11 Mayduring both years wouldincrease, leading to flock breakup andmost fledged by late June. andthe dispersal of pairs to theirterri- The fallmigration of long-billedcur- toriesto breed.On the HanfordSite, lewscommences before the onset of the however,flocks of adult birds were not driestand hottestmonths. Those pairs observedto remainintact after arrival, whoseeggs have been destroyedhead butrather the birds were first seen dis- southwardearly. I observed1 pairto persedon their territories. Only on 1 oc- stay8 dayson theterritory after losing casionwas agonisticbehavior observed theirnest, and M. M. Tremaine( 1976 amongmembers of a groupthat resulted pers.comm.) found departure within 10 ingroup breakup. daysfor lost nests. The overall movement Forsythe(1970) and Wolf (1931) both ofthe birds south is, therefore, more ex- reportedthat the birds seem to be paired tendedthan the move north, and, in fact, onarrival, and Wolf (1931) also believed theearly migrants also appear to be more themto be matedon arrival as he never leisurelyin theirbehavior. Small flocks observed mating. Territories at Hanford or pairsfrequent cheatgrass fields and werefirst occupied either by pairs or, and

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FIG. 19. 300Area study site showing locations of the central territory nest fields of long-billed curlews HanfordSite, southeastern Washington, 1976-1977.

equallyas commonly,byunpaired males. about2 ha in area,with the smallest In addition,courtship behavior, call ad- about0.4 ha. Total territory size was only vertising,and territorialbehavior were about6-8 ha. Nestingterritories were observedas soonas thebirds arrived. largestat the 100-H/100-Dstudy site Fig.19 shows the locations of the cen- wherethe featuresare moreuniform. tralterritory nest fields on the300 Area There,and at theother nesting areas as studysite. Territories were identified well, "nest fields" as suchdo notexist andnamed according totheir relative po- becausethe habitat is so open.Instead, sitionor to a prominentfeature within certain "nest areas" within the territories them.The map also shows the location of can be defined.In thoserelatively flat areasand structuresmentioned in the and openhabitats, the territories were text. about20 ha inarea. Initialterritory size varied with topog- Thesame territories were used in 1976 raphyand habitaton theHanford Site. and1977, and I believethem to be his- Theywere smallest at the 300 Area study torical in naturewith previously defined sitewhere topography and habitat are the boundaries.As such, I havetaken the lib- mostvaried, and the substrate isalso very ertyof referringto themas territories sandy.There, each territoryconsisted even though the boundaries are notal- roughlyof a nestfield with an adjoiningways defended against conspecifics. An shrubarea. Most of those nest fields were increasein thenumber of birds in 1977

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions THE LONG-BILLEDCURLEW IN SOUTHEASTERNWASHINGTON-Allen 41 over1976 did not result in a reductionof Wing-Raising,and Concealmentwere territorysize butrather in an increasedused to driveaway trespassers, and vio- utilizationof marginal areas, and some lentinteractions would erupt occasion- birdsseemed to not nest at all. Conceiv- ally. ably,territory size hadalready been re- Variationsoccurred in theinitial reac- ducedto a minimum.In 1976, all theter- tionof territorial birds to trespassers and ritorialmales at the300 Area study site dependedin parton the stageof the acquireda matefor the breeding season. breedingcycle and on whetherthe ter- Butin 1977,a shortageof females left al- ritorialindividual was present when the mosthalf of them unpaired. In addition,trespasser first arrived. Paired males manymales holding marginal territories were more tolerant of outsiders in their in 1977were unpaired for the season. territoriesforlonger periods of time after Apparently,atleast some birds may re- matinghad occurred. Ifa territorialmale turnto the same territories insubsequent or pair returned tothe territory tofind an years.This is basedon identifiable plum- intruderpresent, the male usually would age characteristicsand on behavioralfly directly tothe site of the intruder and idiosyneraciesofthe birds seen from one proceedwith agonistic displays. If a ter- yearto thenext. Head markingsin par- ritorialmale or pair was already present ticularwere recorded for members of whenan interloperarrived, an Upright eachpair, and in at least3 cases,birds posturewas assumed by all individuals withthe same markings returned the fol- andwas held for several moments before lowingyear. However, I do notknow anotherposture (e.g, Crouch-Run)was howcommon such head patterns are in assumed,or previous activity (e.g., feed- thegeneral population. ingby the female) was resumed. Agonisticencounters between early ar- Defenseof territorialboundaries for rivalswere common.The fewbirds themost part involved Territorial Bound- wouldrange over large overlapping areas aryDisplays between neighbors, though coveringparts of severalterritories violent encounters were not uncommon. thougheach bird or pair could be asso- TerritorialBoundary Displays were con- ciated with1 territoryin particular.sistently observed to occuronly along Boundarieswere not necessarily defend- those stretches ofthe boundary line that ed, especiallynot by unpairedmales separatedone nestfield from another, againstneighboring paired females, until and in those"zones," the line corre- morebirds had arrived. Once all theter- spondedto a physicalmarker of some ritorieswere occupied,paired birds kind(e g.,a deadfurrow, a road, a coyote wouldoften tolerate the association of path,a fenceline7 a ditch).Elsewhere, a otherbirds in areasoutside the actual certainamount of flexibilityexisted in nestfield, but unpaired males were more the territoriallines. Those neighbors defensiveof theirentire territory and whoserlest fields did notadjoin were wouldgive aerial pursuit to any male in- neverobserved to engagein Territorial trudersnot accompanied by a female. BoundaryDisplays with one another. Territorialdefense usually was per- A TerritorialBoundary Display was formedby the male. Supplanting was the initiatedwhen a neighborprovoked an mostcommonly effectual means of evict- encounterby entering the display zone ingtrespassers from territories but the and walkingalong the line. The other Crouch-Runfollowed by aerialpursuit neighbor then flew or ranover to the wasalso observed frequently in that en- zoneand took up a positionopposite his deavor.Following a flying chase, the ter- opponent,oftentimes sidling towards ritorialbird would return tothe field with himand sometimeswith the farwing a loud Whistleand sometimesbriefly raised overhead. Encounters most com- performBounding-SKK Flight before monlyinvolved just the males, but prior alighting.Additionally, theUpright-Run, tothe onset of incubation both members

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions 42 WILDLIFE MONOGRAPHS ofa pairwere often present though usu- lymove in and out of their nest fields all allyonly one memberof each pair was daylong. directlyinvolved in theinteraction. En- Courtshipdisplays are mostfrequent countersusually were either male-to- during the earlymorning hours from maleor, more rarely, female-to-female. dawn to about midmorning, butcommon- Only1 male-to-femaleinteraction was lycontinue through the morning and into observed.When 2 pairedmales were in- earlyafternoon during the initial period. teracting,the females either stood and Courtshipactivities by one pair appear to watchedor more commonly) continued to stimulatecourtship activities in neigh- walkand feed. When 2 femaleswere in- boringpairs (social facilitation) as fre- teracting,however, the males assumed quently3 or 4 pairswill be courtingwith- an appeasementposture and performed in sightof each other. Solicitation for a displacementfeeding while slowly but femaleby unpaired males also begins in continuallywalking around about a l-m2 the early morning,but continues area.They displayed that behavior until throughoutthe day. All reproductive ac- the TerritorialBoundary Display be- tivityis, however,stifled during incle- tweenthe females ended. With the pass- mentweather as thebirds tend to stay in ingof time on theterritories, the inten- the shrub areas where there is morepro- sityof interactionsbetween neighbors tection. graduallylessened to thepoint where Afterabout the first 2 weeks,but still fulldisplays were rarely observed. En- priorto incubation,daily activities be- teringthe zone by a neighborstill pro- comemore routine. Territorial pairs usu- vokedan encounter,but the birds often allyare in theirnest fields only during remainedfarther apart and engaged only the earlymorning, late afternoon,eve- indisplacement feeding. ning,and night.During the day,they feedquietly in the shrubareas within DAILY ROUTINE theirterritories, occasionally coming out intothe open for maintenance activities Each daybegins approximately one- orto pursue an intruder, orthey are gone halfhour before dawn with repeated vol- fromtheir territories entirely, having leysof Whistling as individual birds call movedto otherareas to feed.Courtship outone after another from their roosting displays in pairsare performedalmost sites. "Curlee Curlee" and ''Wheet entirelyduring the firstfew hours of Wheet"calls may also be heardbut Whis- light.Paired birds have developed pat- tlingis the mostcommon. Periods of ternsfor feeding, courtship, and mating quietare interspersed with such rounds thatthey follow routinely. The unpaired ofWhistles, but the vocalizations contin- males continue to maintain a high profile ue untilthe birds begin their other daily as before,though perhaps with a little activities. lessvigilance, and tend to be muchless Duringabout the first 2 weeksfollow- routine in theiractivities. As theseason ingarrival, the birds are very active on progressesand a mateis stillnot ob- theirterritories. There is rarelya peace- tained,solicitation becomes more in- fulmoment during the day as callsring tensewith each passing female. incessantlyacross the fields,birds fly Allbirds usually leave their territories backand forth across their territories, so- oncea dayfor several hours (less for un- licitationand courtship begin with a fren-paired males) during the heat of the day zy,and agonisticencounters occur fre- tofeed in irrigated fields or in other areas quently.Unpaired males are particularly of the Hanford Site, and/or to loaf on is- vocal,and themajority of their time is landsin theColumbia River. The daily spentmaintaining a visible profile. departureofa pairfrom their territory is Pairedand unpaired birds alike frequent- often preceded by "Wheet,""Curlee,"

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions THE LONG-BILLEDCURLEW IN SOUTHEASTERNWASHINGTON- Allen 43 and "CurleeWheet" calls for up to 10 mostcommonly flying in a patharound minwhile the birds walk and feed. Ac- theperimeter ofhis territory orin a fig- tualdeparture is initiated by the female, ure-8 pattern over the nest field, but at whotakes off quietly or calling <'Wheet times no particularpattern can be dis- Wheet."The maleassumes an Alert-At-cerned. If a femaleis in histerritory, a titude,then follows calling <'Curlee Cur- male usuallywill orienthis display lee' andcatches up withthe female in aroundher. Bounding-SKK Flights may flight.Commonly when a territoryis un- lastup to one-halfhour, but sequences attended,neighboring birds will trespass usually are shorter.Frequently though, flagrantly. thebirds will remain on the ground only Curlewsreturn to theirterritories in a minuteor 2 betweensequences. When lateafternoon and frequent the fields un- a malecompletes a seriesof flights he tilthey roost for the night. Occasionally, Whistles as he lands courtshipbehavior was observed during Bounding-SKKFlights become partic- the afternoon/eveningperiod, but was ularlyintense whenever a female passes notcommon. througha nesting area. At such times, all Atdusk, the birds become very quiet theunpaired males rise up andperform ontheir territories as they walk and feed thedisplay over their respective territo- throughthe fields, and eventually none ries.If the female is unpaired,she usu- are heard.As twilightnears, they stop allyalights in the nest field of one of the and assumean Alert-Attitudeoften lis- solicitingmales but often does not stay teningand watching for several minutes. there. Instead, she wanders from territory Thenjust at dark they fly quietly to their to territorywalking and feeding.Once roostingarea (up to 200 m away) and dis- sheleaves a male'sterritory, he becomes appearfrom sight. The samepart of the less intensein his solicitation,landing fieldis usedfor roosting each night and frequentlyand spendingmore time on membersof a pairroost 10-50 m apart. theground. For reasons not apparent to Whereasterritorial birds roost in their me,a femalewould eventually remain in fields,migrants stopping over often use oneterritory and tolerate the association theareas of transitional vegetation bor- ofthat male for up to a day,then move to deringthe fields. a neighboringterritory and male.She By searchingroosting areas covered mightswitch back and forthmore than withunusually thick stands of cheatgrass, once before a pairbond was formed and I was able to findoval-shaped depres- she remainedwith one ofthe males. A sionsin thegrass with curlew exereta at malepersists with Bounding-SKK Flights oneend. The narrowness ofthe ovals in- throughoutthefirst day of his association dicatedthat the birds do notflatten out witha female)and some were observed onthe ground when roosting as theydo tocontinue the displays for up to 3 days. whenconcealing. I had no way of ascer- Witheach succeeding day, however, the tainingthe head position during roosting. displays became shorter and fewerand usuallywere performed only when tran- SOLICITATIONAND INITIAL sientspassed through the area or in as- PAIR FORMATION sociationwith other pair formation behavior. Solicitationor call advertisingby un- As a formof call advertising, Ground- pairedmales is seen in the formof Callingis performedby an unpaired Bounding-SKKFlights and Ground-Call- male within sight of a femalewho is ing.Those displays serve to attract a fe- eitherassociated with another male or maleand are important ininitial pair for- who has recentlyarrived but is in a mationalong with the Scrape Ceremony. neighboring territory. The soliciting A maleexhibits Bounding-SKK Flight malewalks slowly through his field in an

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dencies,though the male maybe observedto flinch in response to them. Whilestanding in thescrape, the female maychange her directionmany times,and eachtime the male will re- positionhimself accordingly tomaintain the originalorientation. Also, while standingin thescrape, the female may inspectit withthe tip of her bill, make Tossingmovements, orperform Scraping herself.Sometimes, the female Whistles FIG. 20. Scrape ceremonyof the long-billedcurlew showingBill-Down Display and orientationof duringthe courseof thoseactivities. the male. However,she remains in the cavity only brieflybefore stepping out, though she usuallystands for a timeclose by before appeasementposture giving the "Whee" walkingaway. Once the female has va- call. Ground-Callingusually is inter-cated the scrape, the male slowly raises spersedwith bouts of Bounding-SKKhis bill and then his head as iftesting the Flight;it is also an integralpart of the femalefor aggressiveness before assum- ScrapeCeremony. inganother posture. The ScrapeCeremony is performed ScrapeCeremonies may be repeated manytimes during pair formation and in- bythe male numerous times throughout volvesritualized Scraping. Initially, the thefirst day of association with a female ScrapeCeremony is preceded by Bound- such that it seems all ofher time is spent ing-SKKFlight, after which the male walkingback and forth across the field in takesup a positionacross the field from response to his calls.Some males stop an unpairedfemale in his territoryand onlybriefly during the day to rest or feed performsScraping behavior while ordrive out an intruder. Ground-Calling.Between Scraping Atthat stage following the ceremonies, movementshe standsin thescrape and the male usuallyperforms Bounding- calls repeatedly.The femalewalks di- SKKFlight and the female begins walk- rectlytoward the male, often moving rap- ing and feeding.Gradually, this leads idly.When she is withinabout 10 m, her intothe next phase in whichprecopula- pace slowsand the maleassumes the torydisplays follow each ScrapeCere- Bill-DownDisplay posture and contin-mony, and the pair bond becomes well ues Ground-Callingsoftly. The femaleestablished. Birds that arrive already thenapproaches the scrape and steps into pairedbegin their courtship activities at it as themale moves out sideways still thenext phase. maintainingthe Bill-Down posture. The femalecommonly Whistles as sheenters COURTSHIP, MATING,AND thescrape. The male takes up a position PRELIMINARYNESTING withhis body oriented perpendicular to, and almostalways behind the female Throughoutthis paper terms such as (Fig.20). Onceout of the scrape cavity, << courtsnlp. ,, ana1 << courtlng. ,, reter^ to themale usually ceases to call. Duringevents leading to copulation. After initial initialScrape Ceremonies, the femalepair formation, theScrape Ceremony be- oftendisplays a certainamount of ag- comesa precopulatorydisplay. Scraping gressivenesstowards the maleand di- behavioreventually becomes disassociat- rectspecking "intention movements" at ed withmating and appearsto be the him.The Bill-Down Display of the male mechanismfor designating the most fa- appearsto dissipate those aggressive ten- vorablepotential nest locations.

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions THE LONG-BILLEDCURLEW IN SOUTHEASTERNWASHINGTON Allen 45 The basic ScrapeCeremony is de- closelyassociated with the courtship/ scribedin thepreceding section in its matingsequence is involved. contextwith pair formation. The Scrape The finalphase occurs when the fe- Ceremonyis thesame when performed male will tolerate the entire sequence of as a precopulatorydisplay with 1 ad- behaviorfrom precopulatory displays dition:the maledemonstrates Tossing throughcoition. Once thatphase is behaviorwhile calling the female over to reached,the Scrape Ceremony becomes thenest. Tossing behavior is widespreadisolated from courtship displays which amongshorebirds but its role in court-are then preceded by the Courtship-Run. shipis uncertain. (TheScrape Ceremony will be discussed Followingthe Scrape Ceremony when furtherunder Nest Site Selection.) How- themale raises his bill and head, he also ever,a malemay still be unsuccessfulat slowlyraises his wingsout to theside, his firstmating attempt each day,and "flagging"the primaries. In thatposition therefore repeated courtship displays he "cautiously"moves around behind stilloccur. I did notobserve more than thefemale and begins Shaking. 1 copulationin a pairon anygiven day, Initially,the female is unreceptiveto thoughsome males would attempt 2 or theadvances of the male. She doesnot more. tolerateShaking behavior, and either turnsin a Crouch-Runposture and pecks atthe male or runs away. The male often NESTING pursues,chasing along behind, or cir- Nest Site Selection clingone way and then another in an ef- fortto take up a positionbehind her tail Scrapingactivities result in several withoutbeing struck or grabbed.Once scrapes,usually in one general area of the thefemale is receptiveto Shaking, Shak- nestfield with 2 or3 occasionallybeing inglasts for about 3045 sec beforethe foundwithin 8 m of one another. Most of maleattempts tomount. More time is re- thosescrapes are used repeatedly during quired,however, before the female be- ScrapeCeremonies and one of them be- comestolerant of Mounting behavior. comesthe nest, but the means by which Eventually,the female permits Shak- theactual site is selectedis notclear. ingand Mounting, but runs out from un- Once Scrapingbehavior has become derthe male before copulation occurs. distinctfrom courtship behavior, each Shethen turns in a Crouch-Runposture Scrape Ceremony is followedby the andthreatens the male. The male usually maleTossing while the female wanders does notpersist with courtship at the away.Once the nestsite is selected, time,but tries again later. At that stage, ScrapeCeremonies cease butTossing someScrape Ceremonies are followed by continuesas themethod for lining the Tossingby the male (instead of further nest cup. courtship). The Tossing often begins Commonly,during the process of nest whilethe female is stillstanding in the siteselection, a birdwas seenTossing scrapeand continues until after she has andmissing the scrape altogether as a re- walkedaway. In addition,mating at- sultof either misorientation or standing temptsare sometimes made without the toofar away from the site. Presumably in precedingScrape Ceremony. At such thelatter case, the materialtossed to- timesthe Courtship-Run is displayed by wardsthe scrape might be pickedup lat- themale followed by Shaking. The point erand tossed into it. Even with repeated at whichthe ScrapeCeremony (with use andTossing by the male, however, Tossingbehavior) changes from a pre- the scrapeswere neversubstantially copulatorybehavior to a nestsite selec- builtand usually were devoid of nesting tionbehavior is vague.A gradualprocess materialwhen they were abandoned.

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SubsequentScraping behavior undoubt- Building movements discussed by Graul edlyremoved material previously placed (unpublisheddissertation) forthe moun- inthe bowl. tainplover Charadrius montanus. Incu- Duringnest site selection, the female bating individuals also reach out and pick oftenperformed Scraping and Tossing upmaterial, then drop it directly infront duringthe Scrape Ceremony, and I sus- ofthe breast in a forwardand back move- pectthat her behavior ultimately deter- ment. As a resultof these nest building minedwhich site would be used.During activities,the brim of the nest becomes thatperiod, females were observed to slightlyelevated above the surroundings. startscrapes and performScraping be- Eventually,the area surroundingsome haviorwithout the accompaniment ofthe nestsbecomes completely denuded of male.A malewas never observed to re- vegetationand litter. spondto Scrapinginitiated by a female, butrather continued with his present ac- ScrapeSize tivity.On the other hand, neither did the femalecall to the male or otherwise ap- Fifty-ninescrapes were measured for pearto be tryingto attract his attention. depth and diameter (major and perpen- dicularaxes). Depth in millimeters(58 Nest Constructionand Materials scrapes)was x = 46.04+ 10.67SE with a minimumof23 mmand a maximumof Thenest is initiallyformed as a scrape 66 mm.Diameter in millimeters( 118 duringa ScrapeCeremony. The bowl is measurements)was x = 201.16+ 37.04 madedeeper with subsequent Scraping SE withthe smallestmeasuring 130 x andwith repeated bill poking. Improve- 130 mm and the largest 275 x 245 mm. mentin theform of added nest material Those measurements aresimilar to those byTossing is madethroughout thelaying reported hy Graul (1971) for 1 nest(di- period. ameter190 mm and depth 45 mm)and The nestcup is linedwith whatever by Silloway (1902) for 1 nest(major axis materialis availablein the immediate vi- about220 mm, minor axis about 153 mm, cinity.Cheatgrass leaves and culms are anddepth about 51 mm).The variation the mostcommon and are used along in scrapediameter probably is a reflec- withrablit pellets,small stemsand tionof the size of the bird that made the twigs,seeds, Canada goose Branta can- scrape.Depth probably varies with the adensis exereta,and miscellaneous litter. extensiveness to which the scrape was Occasionally,a small stone was found in frequented. a nestalong with the eggs. Considerable differenceswere noted in theamount of NestSite Characteristics caretaken to constructthe nest. Some nestscontained sparse amounts of insu- Evidenceis accumulatingthat birds latingmaterial and bare soil couldbe basetheir choice of habitat on features of seenin the nest bottom. Other nests had the environmentsuch as slope of the thickbottoms and wallsthat afforded ground and structureof thevegetation moreprotection tothe clutch. Generally rather than on theplant species present too,nests were more substantial where (Wiens1969). nestingmaterial was locallyabundant thanwhere nesting material was scarce. VegetationType Incubatingbirds often picked up near- bymaterial and placed it in the perimeter Allnest fields were of a generalcheat- ofthe nest using the same nest building grass community of1 of2 types.The first movementsas in Tossing, except that the typewas a cheatgrass/Sandberg'sblue- birdis in a sittingposition. That activity grass association and the second type was is essentiallythe same as theSide-ways cheatgrass with no accompanyingblue-

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TABLE 5. TOTAL COVERAGE(%) AND FREQUENCYOF OCCURRENCE(%) OF BROMUSAND POA IN 2 TYPES OF NEST FIELDS, HANFORD SITE, SOUTHEASTERNWASHINGTON. ALL VALUES ARE MEANS+ SE

BromuslPoa association Bromus

Frequency of Frequency of Total coverage occurrence Total coverage occurrence Bromus tectorum 6.7 + 2.2 72.7 + 15.7 13.6 + 1.6 93.5 + 5.8 old Bromus tectorum 65.4 + 10.9 100 + 0 92.2 + 2.8 99.5 + 0.5 Poa sandbergii 16.6 + 4.2 81 + 7.9 Old Poa sandbergii 4.6 + 2 56.3 + 18.5 grass.Table 5 summarizestotal percent- munity were evaluatedby Cole and age coverageand percentage frequency Sharpe (1976). Theyfound that both of occurrenceof cheatgrassand Sand- speciesdiversity and total density were berg'sbluegrass for those types. Various increasedfor their grazed study site in forbsand/or shrubs in veryminor quan- partbecause long-billed curlews were titieswere associated with both types. absentfrom their ungrazed study site. Othercheatgrass communities (e.g., those Workdone by M. M. Tremainein Ne- thatinclude a preponderanceofJim Hill braska(1976 pers. comm.) supports those mustard)and othergrass communities observations that long-billed curlews (e.g.,bluebunch wheatgrass stands) were maybe restrictedto such grazing areas. notutilized by curlews for nesting. Wolf(1931) confirmed a preference for Of21 nestfields (or areas) identified, short grass. 6 (29'ho)were cheatgrass and 15 (71No) In summary,long-billed curlews prob- werecheatgrass/Sandberg's bluegrass. ablyselect sites because of the shortness Onthe Hanford Site, all areas with cheat- ofthe vegetation and also the spacing of grass/Sandbergsbluegrass fields in them thegrass clumps. Because they rely on wereutilized by curlews, whereas many camouflagefor protection ofthe eggs and areasof just cheatgrass were never used. themselvesduring incubation, the short In thenest fields, the average height of grasspresumably allows for better visi- cheatgrasswas less than 100 mm. Cheat- bilityof approaching danger and the ir- grassculms are robust, leafy, and spread- regularpattern of the grass clumps coming(as opposedto erect) and the panicle plementstheir cryptic coloration. is dense and drooping.It coversthe groundmuch more effectively than Sand- Slopeof the Ground berg'sbluegrass (see Table 5). Sand- berg'sbluegrass has veryslender erect Long-billedcurlews usually select rel- culmsthat rise *om a tuftof very fine ativelyflat areas for nest sites, but some shortbasal foliage. It averaged about 200 nestswere found in locationswhere the mmin heightin thenest fields, notice- groundis uneven;e.g., on top of a dead ablytaller than the cheatgrass, but due to furrow(3 nests)or on a roadsidefurrow itsfineness probably did notpresent a (1 nest).Wolf (1931) stated that the ideal visualbarrier to thebirds. In addition,location is ona slightrise of ground, and wherecheatgrass and Sandberg'sblue- Graul(1971) found the nest he observed grasswere associated, the grass clumps at theedge of a largevalley adjacent to weremore widely spaced with mosses a gentlysloping hill. I found3 nestson andlichens in between,and the cheat- a slightrise of ground,but most nests grassdid not reach the density found in weretoward the southernedge of the areasof pure stands (see Table5). fieldregardless ofslope. In fieldswith an Theeffects ofgrazing on breeding bird east-westmajor axis, the nest was located densityand diversityin a prairiecom- in eitherthe east quarteror the west

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TABLE 6. A COMPARISON OF THE DISTANCE (CM) 30 cmof an object,and 16 nests(2757o) TO THE NEAREST CONSPICUOUS OBJECT BETWEEN wereagainst an object. DIFFERENT SCRAPES OF THE SAME BIRD. O INDI- CATES THAT THE SCRAPE WAS AGAINST A CONSPIC- Individualbirds were not consistent in UOUS OBJECT, AND NA INDICATES NO OBJECTS IN selectingscrape sites in relationto con- THE VICINITY OF THE SCRAPE spicuousobjects (Table 6). However,I was unableto makecomparisons be- ScrapeNo. Bnod 1 2 3 4 5 6 tweenthe nest sites chosen by the same birdin differentyears, and individuals 1 O NA NA NA NA NA maybe moreconsistent in theirfinal 2 O O O 5.5 300 NA choice.Of 59 scrapes,14 wereused as 3 NA NA NA NA NA NA nestsand 9 ofthe 14 werenear a con- 4 O 41 NA NA NA NA though,con- S NA NA NA NA NA NA spicuousobject. In general, 6 O O 28.5 - - spicuousobjects are in shortsupply in 7 136 120 71 246 thecheatgrass fields on the Hanford Site. Surfacerocks and sage stumps were re- movedwhen the land was farmed,and quarter.On the100-H/100-D study site, thereare no grazing livestock to provide theoverall slope of the ground is down- manurepiles. Those objects present are wardto the Columbia River to the north, widely scattered and some nest fields ap- andso a southerlynest site gave the bird pearto be withoutany at all. Therefore, a sweepingview down the Seld. But on thetendency to select a nestsite close to the300 Area study site, the direction of a conspicuousobject may be limitedby slopein each field is variableand the ad- theavailability ofsuch objects. vantagesof a southerlylocation are not readilyapparent. Presenceof Other Curlews Thepresence of long-billed curlews in ProximitytoConspicuous Objects anarea undoubtedly serves to attract con- Silloway(1902) found several nests be- specificsto the siteand increasesthe sidepiles of dried cow manure, and Sug- probabilityofobtaining a mate, in addi- den (1933)included a photographof a tionto thesurvival value afforded in a long-billedcurlew nest adjacent to a pile groupsituation. Territorial behavior ofhorse manure. Cameron (1907) noted spacesthe birds over a givenarea, but thatcurlews look so like"buffalo chips" eventhough definite boundary lines are as tobe easilymistaken for them at a little established,nests in neighboringfields distance.In areasheavily grazed, and werenever adjacent. The nearest neigh- thereforecovered with only scant vege- bornesting distance was 250 m at the 300 tation,the selection of a sitenext to a Areastudy site. However, in particularly manurepile wouldseem advantageous favorable habitat, there is a tendencyfor bymaking the incubating bird less con- pairsto nest within sight of one another. SpiCUOUS . Nestsin other areas were 500 m or more On the HanfordSite, distances from apart which concurs with Forsythe (1970) scrapesto the nearest conspicuous object whofound nests about 462 m apart. weremeasured. The mostcommon ob- jectswere old big sagebrush limbs, rocks, Nest Density baredirt mounds, and dead furrows. Oth- er itemswere a steelcable, a horsema- Sadlerand Maher (1976) estimated ap- nurepile, a rusty5-gallon can, an old proximately1 pair of breeding curlews tumbleweed,and a largebunchgrass. per 6 to7 km2of suitable habitat in Sas- Twenty-twonests (37%) were within 1 m katchewan.Nesting density is much ofan object,18 nests(31Wo) were within higher on theHanford Site. Fifteen ter-

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THE LONG-BILLED CURLEW IN SOUTHEASTERNWASHINGTON-Allen 49 ritorieswere identified at the300 Area thenest bowl and stands over the eggs studysite that encompasses only 10.36 sitsdown, or alternately stands and sits. km2.The 5.18-km2100-F study site sup- Whenstanding over the eggs, the female portsat least 3 territories,and the 100-H/ usually assumes a bodyposture essen- 100-Dsite (15.48 km2) supports at least tiallythe same as thatdescribed for Ap- 10. peasement.On 2 occasions,a female was Differencesin nest density can be at- observedto be callingsoftly while stand- tributedto differencesin habitat and to ingin thatposition. The Bill-Down Dis- territoriality.Each territorymust have playis performedby thefemale when suitableareas for nesting and chick rear- andif the male approaches her while she ingthat include adequate food, cover, is standingin thenest bowl, and termi- andshelter. The 300 Area study site sup- natesas soonas he movesaway. Males portsthe highest nesting density of cur- werenever observed to stand over a nest lewsbecause the habitat there is prime. forany length of time, but females would Suchis notthe case at theother study oftendo so forup to30 min.Either bird sites,and consequently the nesting den- mightsit on the eggs for 1-2 hours during sitiesare lower. The prairies of Saskatch- a visit. ewan wherecurlews occur are much losslngoe nav1or 1S C. boselyassoc1atec . morevast than the area at Hanford,and, withnest attentiveness and occurs along therefore,one expects to find differences with repeated visual and tactile inspec- inbreeding density. tionof the bowl and itscontents while thebird is inor around the nest. Tossing is alsoperformed just before a birdsteps EGG LAYING outof the nest and again before it moves LayingBehavior awayfrom the site. Courtshipcontinues during egg laying, The behaviorof 3 femaleswas ob- butmating attempts were observed to be servedduring egg laying and was similar successful only on those mornings when in eachcase: the female approaches the anegg was not laid. The laying of the last nestto withina meter,stands alert mo- eggin a clutchwas observed in 1 pair, mentarilythen steps into the bowl. She andin thatcase thelast copulation was firstinspects the nest looking and poking seenthe day before. aroundwith her bill, and sometimes toss- es or pulls additionalmaterial inside. Oncethat is completed,the female set- Clt4tchInitiation and LayingIntervals tlesinto the bowl and sits quietly. After theegg is laid,the female rises from the Datesfor the laying of first eggs were nestand begins Tossing while still stand- calculatedby backdating athatching al- ingin the bowl. Eventually, she steps out lowing28 daysfor incubation (Graul ofthe nest but continues tossing behind 1971)and 6 daysfor laying. Most clutches orto theside for 5-10 min. Time spent wereinitiated within 2a5 weeks of when onthe nest in the3 casesobserved was thefirst clutch was started.In 1977,1 13,25 and87 min.The final egg in the clutchwas initiatedunusually late, 23 clutchwas laid during the 87-min obser- May. vation,and the onset of incubation may Eggswere laid in theearly morning haveinfluenced the extended time spent hours(within 2 hoursof dawn) on alter- onthe nest. natedays. I wasable to record the inter- Oncelaying has started, both birds be- valbetween the laying of 2 eggs(the first comevery attentive and return tothe nest andsecond) in only 1 case,and it was 47 numeroustimes throughout theday. Dur- hoursand 25 min.Graul (1971) reported ingsuch visits, the bird usually steps into a case of3 eggsbeing laid over 4 days

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thatto be true,though most of the eggs hadeither a lightbeige or a palebluish- greenground color with chocolate col- oredmarkings. The dark markings can be bold,large blotches or fine, small specks, ormixtures ofthe 2 (Fig.22). However, unlikewhat Bent (1962) reported, there was a tendencyfor the markings to be heavierand morenumerous at thebig endof the egg. I haveno evidencethat individualfemales tend to lay eggs with a distinctivepattern, and single clutches containedeggs thathad bothground colors. Two eggsweighed soon after laying weighed73 and 71 g. Table 7 summarizesthe egg weights from 4 clutches andgroups them by week.

Egg Losses and RenestingAttempts Coyotesand black-billed magpies Pica pica werethe major predators and were particularlyadept at locatingnests, es- peciallyif I hadpreviously approached the nests.Ravens and Pacificgopher snakesPituophis melanoleucus catenifer wereprobably next in importance;bad- gersTaxidea taxis mayalso havebeen FIG. 21. Nestsof long-billedcurlews with com- effectual. pleteclutches. Upper, Proximity to a conspicuous Ifthe nest is locatedby a groundpred- object;Lower, Usual arrangement ofeggs. ator,the birds' initial response is Dem- onstration,followed by displacement feedingsome distance away while callinga rapid"Whee-whee-wheet." No at- anda minimumlaying period of 5 days. temptis madeto drive the predator away Forsythe(1970) observed egg layingat fromthe nest. Flying threats are directed 2-dayintervals. towardavian predators that enter the territory,and a curlewwill not hesitate to Clutch Size and Egg Description drivea black-billedmagpie or a raven awayfrom the nest should it be located. Allnests in whichincubation was ini- Ifa black-billedmagpie pecks one of the tiatedcontained 4 eggs (Fig. 21). That is eggsin the nest, the curlews abandon the thenumber reported as theusual clutch entireclutch (4 observations),butas a ra- size forlong-billed curlews (e.g., Bent venmay fly away from the nest site with 1962,Sugden 1933, Wolf 1931). anegg (2 observations),possibly the cur- Theeggs are well described in the lit- lewscontinue to attend any that are left eraturefrom the early 1900s when oology behind.Pacific gopher snakes are com- was popular(e.g., Bent 1962, Shufeldt mon on the Hanford Site, and in the sand- 1913).Those descriptions indicate a con- hillsregion of Nebraska, bullsnakes Pi- siderablevariation in shape,prevailing tuophis melanoleucus sayi are a major groundcolor, and spotting. I also found predatorand raidnests for the eggs of

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions THE LONG-BII.T.F.nCURLEW IN SOUTHEASTERNWASHINGTON Allen 51 birdslike the American avocet and the long-billedcurlew (Tremaine 1975). Consequently,though I havenever ac- tuallyobserved such, I suspectPacific gophersnakes are predators on thecur- lewnests at Hanford. Their impact, however,is difEcultto ascertain. Ifeggshells remain in thenest after it hasbeen destroyed, as usually is thecase followingpredation by a black-billed magpie,the female curlew will often re- movethem in the same manner as shells are removedfollowing hatching (see Hatching).An aberrant form of shell re- movalis also oftendisplayed in which the femalepicks up and crushesthe shellsbetween her mandibles, dropping thefragments inand around the nest. In 1977,5 neststhat had been located duringthe egg layingperiod were de- stroyedprior to theonset of incubation. Threeof those losses were attributed to coyotesand 2 to black-billedmagpies. Only2 nestswere found during that pe- riodin 1976and both were destroyed by black-billedmagpies. Long-billedcurlews do notappear to renestonce their first attempt has been thwarted.However, I did observe1 fe- malelay 1 eggin a secondscrape followingthe destruction ofthe nest by a black- billedmagpie. The nestcontained only 1 eggat thetime of destruction, which was presumablythe first one laid.The egglaid in the scrape was also pecked by a black-billedmagpie and no further lay- ingattempts were observed.

INCUBATION The incubationperiod for the long- billedcurlew has been reported once by Graul(1971) as 27 daysand 14 hours (+9 hours)between the laying and hatching ofthe last egg. As the start of incubation wasunknown for all thesuccessful nests I observed,I was unable to verify those data. Observationsat 2 nestsindicate that the male initiatesincubation on the morningfollowing the laying of the last egg.Both male and femalelong-billed FIG. 22. Examplesof eggs of the long-billed curlew showingvariation in shapeand markings.

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions 52 WILDLIFE MONOGRAPHS he mayremain low on the nest until re- lievedby the female, popping his head upin response to her calls. In hotweather,the attending bird sits with head held highand pants. Duringthe day, the incubating adult addsnest material to the rim of the bowl in an idlemanner, though such activity is sometimesmore pronounced at nest FIG. 23. Normalincubation posture of the long- relieftime. The incubating bird also fre- billedcurlew. quentlyrepositions itself to face a differentdirection on the nest. Sometimes the bird accomplishesthat by nestling curlewsshare in incubation duties, but it aroundwhile remaining seated on the takesa dayor 2 forthem to establisha nest.More commonly, the birdrises, routineof nest relief where the female pokesin thebowl with its bill (presum- sitsduring the day and the male sits at ablyto rearrange orrotate the eggs), and night.Prior to thattime, they may ex- thenturns to facea differentdirection. changeplaces several times during the Occasionally,the breastfeathers are day,and the male may actually incubate preenedat that time. Sitting down again, morethan the female. On 1 occasiondur- however,usually is protractedas the bird ingthat early adjustment period, a female repeatedlyattempts to settleover the wasobserved to supplantthe male from eggs. The incubatingbird usually is the nestby performingWing-Raising. quiet, though Whistling from the nest is Alsoduring those first few days, the birds commonat nest-relieftime (see below). tendto fly offthe nest at the slightest dis- Whistlesare also occasionally exchanged turbance.Once the incubation phase is betweenneighbors, and a sittingbird underwayinan area, the fields are much willWhistle at a trespassingcurlew if its quieterand only the unpaired males and mateis notpresent to defendthe terri- transientindividuals participate in ex- tory. changingWhistles at dawn. The birds re- To risefrom the eggs, the incubating mainrelatively quiet unless disturbed. bird tips forward onto its breast bringing itslegs up behind,then pushes itself up Behavior andback with the tip of the bill until the feetare regained. When settling onto the The normaldaytime incubation pos- eggs,the bird first lowers itself onto the tureis an alert, head up position (Fig. 23). breastwith the wings slightly away from Duringinclement weather, however, the thebody. The rear is thenlowered as the incubatingbird modifies its posture and birdsimultaneously squirms into posi- sitswith its head low, often below the tionand adjusts the wings individually. levelof the grass. Birds using that low Duringthe day when the female is in- posturefrequently raise their heads up, cubating,the male may or maynot be lookaround, then sink back down. In the presenton the territory.Some males evening,just before dusk, the male ducks leavefor only 1-2 hours, spending most his head presumablyin thatsame low ofthe day feeding within the territory, position.I was unableto observethe whileothers are goneuntil nest relief birdsat nightbut assume the male re- timeand leave the female to fend for her- mainedlow on the nest throughout.self. When the female is relievedin the Shortlyafter dawn, the male may slowly afternoonorevening, she usually leaves raisehis head and assumethe normal immediatelyforthe river or another feed- headup positionused during the day, or ingarea and does not return until dusk.

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TABLE 7. EGG WEIGHTS (G) OF LONG-BILLED CURLEWSAT VARIOUSSTAGES IN INCUBATION,1HANFORD SITE, SOUTHEASTERNWASHINGTON

Eggs Firstweek Second2week Thirdweek Pipping Clutch A 1 68 64,65 63 2 66 62,64 63 3 66 64,63 64 4 68 64,64 64 Clutch B 1 74,71 67 66 2 70,68 64 64 3 70,68 63 63 4 73,71 67 67 Clutch C 1 63 59 58 2 65 61 60 3 65 61 61 4 64 62 59 Clutch D 1 67 61 2 60 58 3 62 58 4 65 62 Mean + SE 67.0 + 0.58 66.6 + 0.82 63.3 + 0.61 61.4 + 0.89 ' Weightsare groupedby weekof incubation calculated by backdating. 2 ClutchesA and B wereweighed twice during the second week.

Shedeparts again in themorning before tween 2 bigsagebrushes and rested there returningto incubate for the rest of the for20 minbefore departing. day. Typicallyat changeovertime the incomingbird flies into the nest field and NestRelief Whistlesas it alights.The incubating birdanswers the call with a Whistlefrom Nestrelief occurs about the same time thenest. The incomingbird then walks eachday for any given pair unless the orflies to within about 15 mof the nest. birdsare disturbed. For example, nest re- The incubatingindividual flies directly liefby one pair was delayed 2 hoursone offthe nest as itsmate approaches, and rnorningbecause a Swainson'shawk Bu- landselsewhere in thefield. Departure teo swainsoni wasperched in a treenear fromthe nest is generallyin thesame theirnest. The female curlew wandered direction each time, and on 1 occasion upand down the field displacement feed- a malewas observed to rotate 180° on the ingand watching until the hawk left. The nestbefore flying off. The incoming bird morningexchange period was either be- approachesthe nestalong a particular tween0500 and 0630 or between0900 routeand settles over the eggs. The eggs and 1015. Eveningexchange periods usuallyare left exposed for less then 60 werenot as clearlydefined but were ob- sec.However, a disturbance atthat time servedbetween 1700 and 1930, although will cause the relieving bird to delay its onone very hot day a malewas observed approach, and in hotweather that could torelieve the female at 1343.The female killthe embryos. The bird that has been immediatelyflew to a shadyspot be- relievedpreens its breastfeathers and

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions MONOGRAPHS WILDLIFE nest,the 54 mof the within10-15 dis- proaches birdceases with those nonincubatingdisplacement feeds playsandstands or callinga soft,rapid "Whee-whee-wheet" or"CurleeCurlee." normal onthe nest maintains Thebird whenits mate or incubationpostureeven It as- birdsare Chirping. neighboringposture, Iying mo- sumesa concealing possible tightto the ground as tionlessas theground (Fig. withthebill resting on within afterthe intruder comes 24),only up nearbycalling sightorits mate flies intruder Curlee."As long as the "Curlee to thenest, the in- ispotential a threat and birdmaintains that posture long-billedcurlew concealing cubating partnercontinues call- FIG.24.Incubating nonincubating from on nest. the furthercalls are heard ing.When no orit Whistles nonincubatingpartner the sittingbird slowly rais- a Two-wing an"allclear," the Ifthe non- its wingswith headand looks around. an stretches Flapbefore begin- es its is notpresent when Stretchora Two-wing 1 occasion,a incubatingmate on thenest anotheractivity. On appears,the bird ning flewvery rapidly low intruder forhalf an houror more malepreened, then area,swoop- mayremain low slowly groundaround a small intruderhas gone before tothe whiletipping from side afterthe observedsuch de- ingupand down I couldnot risingup.I commonly to a blind toside as ifchasing butterflies. extra- laysfollowing my approach thoughtthat the male's somebirds habituated resistthe a resultof pure de- thougheventually poptheir heads ordinarybehavior was even tomy routine and would freedfrom the nest minutesafter my disappear- lightat being onlystretching upwithin thoughhe was probably anceinto the blind. do not muscles. femalesusually cramped Incubating unlessthe intrud- flushwhen approached males, Disturbances withinabout 2 m.The Responsesto ercomes vacatethe nest be- however,sometimes tolocate Intruders intruderis close enough Responsesto Human forean (e.g.,Graul 1971, Sug- de- it.Several authors their humanintruders are Wolf1931) have touted Responsesto otherdistur- den1933, an incubatingbird separatelyfrom abilityto approach itoff the scribed did notobserve other iton the back or lift bancesbecause I nestsites, andstroke theend of the in- predatorsapproach particularlynear do ground responsesare sim- nest, I wasnever able to thoughI presume the cubationperiod. wasbecause andbelieve the reason ilar. fieldwhen the that thenests by search- Ifa humanenters a nest non- Ioriginally located incubating birdis nearby,the fieldsand flushing the nonincubating an Alert-Atti-ing the was flushedfrom birdassumes birds.Once a curlew seemed incubating givesthe Chirping andi a sensitivitythreshold tudeand usually Ifthe in- thenest, birdsubsequently WheetWheet" call. tobe brokenand the or"Wheet immediately,Dem- thefirst sign of approaching truderdoes not leave displace- flushedat mayattract a pred- interspersedwith Suchbehavior have onstration Alert-Attitudedanger. thatmight otherwise mentfeeding and the atorto the site sometimesthe Enticement- unnoticed. ensues,and theintruder ap- gone Runis perforrned.As

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Whenflushed from the nest at close buta paircan usuallykeep them from range,the incubatingbird bursts forth their eggs if not otherwise distracted. An witha loudsquawk, then feigns injury for incubatingcurlew does not conceal in re- 10-20m beforeperforming the Entice- sponseto a black-billedmagpie, but the ment-Runand calling"Wheet Wheet.' nonincubatingbird will pursue one on The birdeventually takes flight and, theground with the Crouch-Run, or in joinedby its mate, Demonstration inter- flight. Though gulls (Larus spp.) routine- spersedwith the Enticement-Runand lyfly over the 300 Area study site, they displacementfeeding is performed.are ignoredby thecurlews and them- Neighboringbirds that are not incubating selves show no interestin thenesting gathernearby and add their voices to the shorebirdsunless a disturbanceata nest ruckus. sitecreates a ruckusthat then attracts Oncethe intruder has departed, the at- them. tendingbird often takes up to an hour to One or 2 coyotescommonly cross settleback on the nest. During that time, throughthe nesting fields daily, some- thebird preens or feeds and may return times passing within a fewmeters of a to thenest several times to inspectthe nest.The incubating bird conceals as the contents.The birdmay settle over the nonincubatingbird flies to a vantage eggs,then fly off again a fewminutes lat- pointChirping. The nonincubatingbird eronly to wander back and resettle. That thendisplacement preens or displace- periodis crucialfor the eggs during hot mentfeeds while calling '

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions 56 WILDLIFE MONOGRAPHS ofa nearbytrain invariably elicited that eggshellsremain in the nest they are re- response. moved.The birdsat thatpoint are hy- persensitiveto anydisturbance and di- Nesting Success rect intensiveflying attacks toward potentialpredators. Demonstration and Nestingsuccess was good in 1976(no displacementfeeding are performed. losseswere observed) and poor in 1977. Withinseveral hours, the birdsbegin Allrlests I hadlocated prior to hatching walking and feeding or rest in thefield weredestroyed by predators,including near the nest. In time,the male begins all but2 at the300 Area study site. The Ground-Callingand thenScraping and successof the predators in 1977is attrib-Shaking, all intermixed.Incomplete utedto insufficient nesting cover that re- ScrapeCeremonies and Tossing behav- sultedfroln the dry winter of 1976. Ad- ior are aIso performed;the Bill-Down ditionally,coyotes were thought to be Display,however, was neverobserved. . we . - , morea zuncant ln . // t zan 1n Lfi /O DUt The femaleis notreceptive to renesved thatis subjective. courtingby the male but she doesper- Time-lapsefilm from 1 nestshows a formScraping at several different sites of coyoteapproaching and eating the eggs. herown. That behavior lasts for up to2 The coyotewas originallypassing by hours,interspersed with brief periods of about5 maway but then spotted the con- maintenanceactivity and standing cealingcurlew and turned to investigate. around. Eight hours after one nestwas In a yearof normal grass growth, a coyote destroyed,the pair finally left the nest woulcIbe unableto spot a concealingcur- fieldand flew toward the river. lewat 5 m. Duringthe incubation period, coyotes HATCHING appearto be themajor predator, but other potentialpredators include badgers, Pa- Starringofan eggcommonly appeared cificgopher snakes, and black-billed3 to4 daysprior to hatching, and a hole magpies.At 1 nest,trampling by feral usuallydeveloped 1 to2 daysbefore the horsesappeared to be thecause of de- chickemerged. "Clicking" noises from struction,but such damage may be offsetinside the egg were heard in pipped eggs bythe benefits gained from grazing dur- up to53 hoursprior to hatching. ingyears of overabundant plant growth. Forsythe( 1967) described the hatching No lossesby adverse weather were not- ofthe long-billed curlew. He observed ed. thatthe pip hole was startedapproxi- matelyone-third the distance from the BehaviorFollowing Nest Loss largeend of the egg, then was progres- sivelyenlarged until the chick pushed CCurlewsabandon the nestonce the outthe large end and emerged by split- eggsare damaged or destroyed.Once a tingthe remaining shell into 3 roughly nestis abandonedby a pair,they gradu- triangular pieces. I did notobserve the allybecome less and lessresponsive to completehatching process, but did find predatorsand disturbances. The territory the pip hole to be startedas hedescribed is stilldefended against conspecifics for (Fig.25). I also collectedseveral shells a weekto 10days following the destruc- as soonas thechicks were free. In each, tionof the nest but the birds spend more abouthalf of the large end and one side and moretime each dayaway from it. weremissing whereas the restof the Finally,they depart and presumably mi- shellwas intact, indicating a slightly dif- gratesouth. ferentmethod from that Forsythe found. Immediatelyafter the predator leaves Presumably,the eggs were rotated thenest, the curlews approach and if any throughoutincubation as a resultof pok-

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions THE LONG-BIT.I.F.DCURLEW IN SOUTHEASTERNWASHINGTON Allen 57 hatchingtime and both members of the pairremain present once the eggsare starred.Once hatching had actually begun,the female appeared to assumeall incubationduties as she was invariably on the nestwhen checks were made, evenat dawnwhen previously the male was in attendance.During hatching, however,itwas not unusual to find both adultsoff the nest for short periods of timefor reasons I was unableto ascertain.

Responsesto Intruders Hatchingof the young seems to elicit a protectiveresponse in the adults that is notseen previous to thattime. Arc Dis- playsare directed at human intruders as are Demonstrationand Enticement-Run displays,and, therefore, presumably at otherground predators as well. Avian predatorsthat fly over or perch within the territoryare confrontedwith the same flyingthreats, and marshhawks are at- tackedat that stage for the first time. Con- FIG. 25. Hatchingof long-billedcurlew chicks. specificsattracted to the sitejoin the Upper,Pipping; Lower, Downy chick. ruckusthough do not perform the Entice- ment-Runorthe Arc Display. If the nest ingand shufflingbythe adult bird, but has notbeen disturbed on previousoc- duringhatching they maintained a rela- casions,the femaleremains concealed tivelystable position, usually with the overthe eggs during the male's displays pippedarea upward. This would seem unlessdiscovered; then she too Demon- advantageous,making it easierfor the stratesand performsthe Enticement- chicksto switch to air breathing. Run.As soon as theintruder retreats, she Hatchingusually was synchronousor the male returns to thenest immedi- withina nest,all the chicksbeing ately. hatchedwithin 5 hours,and seemedto occurat anyhour of theday or night. EggshellRemoval Hatchingwas also synchronous within an area,unlike what Forsythe (1970) ob- Adultsremove eggshells soon after servedin Utah.In 1976,hatching on the hatchingby grasping them in the bill and 300 Areastudy site was 11-14May (5 flyingseveral hundred meters before nests);at the 100-H/100-D Area it was 25 alightingand dropping them. Each shell Mayto 2 June(4 nests). is droppedin a differentspot and not al- waysin the same direction from the nest. Behavior Graul(1971) observed the male (short- billedindividual) perform that function. Long-billedcurlews, like many other At2 nestswhere I observedthe behavior, birds,become especially attentive at thefemales removed the shells.

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is favorable,the entire brood is likelyto leavethe nest that afternoon. Ifthe last chickhatches later in theafternoon or evening,the brood usually stays at the nestuntil the followingday, leaving aboutmidmorning when the temperature hasrisen suffliciently forthe chicks not to requirebrooding. In rainyor cool weather,the brood may stay in the nest for up to24 hours(1 observation).Ifthe site is disturbedduring hatching, the chicks are movedaway from the nest prematurely, FIG. 26. Broodingposture of the long-billed cur- oftenbefore the youngest are fully mo- lew. bile. At the timeof departure,the male standswatch some distance away in the ChickBehavior directionthey intend to move as thefe- Chicksgenerally dry out completelymale gives a lowcall while standing near within3 hoursof hatching but even be- thenest. At the 300 Area study site, the forethey are dry they begin to toddle out chickswere moved directly to the south ofthe nest alone, resting in the grass and ridgearea (Fig. 19),and the male took returningtobe broodedfor short periods. up a positionon top of thefirst ridge Withina fewhours, coordination ofthe overlookingthe field and the rest of the youngimproves substantially, and those ridgearea. Normally, the chicks return thathatch first often wander too far from and are brooded at the nest site at night thenest before the others are ready and forseveral days following the hatching, becomelost. At 2 sites,I wasunable to andthe site is defendedduring that time findthe first hatched chick within a 10-m (M. M. Tremaine,1976 pers. comm.). radiusof the nest. Newly hatched chicks do notfollow or respond well to the calls Brooding ofthe adults, and once mobile may run fromthe nest when there is a disturbance Broodingis accomplishedby the same insteadof concealing. Upon approaching posture used in incubationor by the a nestsoon after hatching, I would usu- adultstanding with the wings dropped allyfind the chicks resting in or just out- downwith the young standing beneath sidethe bowl. When handled, the chicks (Fig. 26). Young are brooded regularly at wouldoften cry in distress,giving the differenttimes during their first few days "Squee" call whichwould throw the by bothsexes, particularly during the adultsinto a frenzyof displays at close coolearly morning hours. I do notknow quarters.One chick called "Peep Peep" theage of the young when brooding ter- and"Hee-who" while resting in the nest minated,but I neverobserved chicks priorto being banded. During the first 24 beingbrooded after they were about 2 hours,the chicksslowly peck at the weeksold. groundand at various objects and appear tostart feeding by the second day. Behavior

BROOD PERIOD Shading Nest Departure The chicksspend most of their time walkingand feedingeven duringthe Ifthe last chick hatches in the morning heat of the day, though when small they orvery early afternoon and the weather frequently rest for short periods in the

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions THE LONG-BILLED CURLEW IN SOUTHEASTERNWASHINGTON Allen 59 shadeof a shrubor a largeforb. Some- timesthe chickscrawl up intosuch plants,presumably gaining the addition- al advantageof the cooling effect of the wind.I neverobserved adults shading theirchicks, but on the Hanford Site the chickrearing areas all includeplant specieslarge enough to provide shade so adult shadingis not necessary.The adultsalso seek shade during the heat of theday, but one bird always maintains a lookout. BroodMovements Long-billedcurlew family units do not keeptogether the way some other shore- birdsdo, e.g., killdeer Charadrius voci- ferus.The chicks wander independently andoften become widely separated. The olderthey get, the more independent the chicksbecome and the farther apart they wander.One juvenile of about 3 weeks ofage was foundapproximately 563 m FIG. 27. Male long-billedcurlew perching in fromits parents and a sibling.On another shrub. occasion,the 2 chicksin that same brood wereseparated by approximately 805m. cificarea. Detailed observations were not Oneparent was with each chick and nei- madeon broodsin thesouth ridge area thercould see theother. The adults can becausethe topography and vegetation determinethe general direction oftravel inhibitedsightings. At the 100-H/100-D bythe male flying 50-100 m aheadand and 100-Fstudy sites, the broodsfol- standing,while the chicks and the female loweda dailyroutine of moving toward wandertowards him walking and feed- theriver from their nest area in the early ing.If thechicks are wanderingin the morning,feeding in thefields along the preferreddirection on theirown, the bluffsduring the day, and then heading adultsmay merely follow along behind backtoward their nest area in thelate them.If the adultsremain in 1 local- afternoonand evening.The chicksand izedarea, the chicks will center their ac- adultswould then roost for the night in tivitiesthere but may range 100 m out ornear their nest area. andback. Theadults maintain a close watch, usu- Perching ally50-100 m apart with the chicks some- wherein between.Daily brood move- Long-billedcurlews commonly alight mentswere of 2 patterns,presumably a in big sagebrush,antelope bitterbrush, functionof the habitat available. At the trees,or on topof dried tumbleweeds, 300 Areastudy site, the chickswere dirtmounds, rocks, stumps, fence posts, movedfrom the fieldsinto the south utilitypoles, rails, etc. (Fig. 27). Silloway ridgearea, where at first they were found (1909) discovereda male curlewthat in differentlocations each day. Then useda solitaryfence post as a vantage sometimewithin the first 2 weeks, chick pointfor guarding his home. Though that rearingareas wereselected and each behaviormay be seenat any time-during broodcould be founddaily in 1 spe- thebreeding season, it is particularlyad-

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions 60 WILDLIFEMONOGRAPHS *omplace to vantageousduring the brooding period maleresponded by flying their placearound the field as thechicks ran whenthe adultsare protecting in an at- flightlessyoung. At the 300 Areastudy firstone way and then another rearedin shrub temptto keepup, but the male always sitewhere the chicks are itgot so dark vegetationthat averages about 1.2 m landedtoo far away. Finally, possiblyspot in- Icould no longer see thechicks and pre- high,the adults cannot thegrass. trudersat a distancewhile standing on sumablythey sat down in theground. Hence, at least 1 adultperch- esin the top of a suitablytall shrub. Oc- Feeding casionally,a bird was observed to assume whileperched Thechicks walk and feed in a manner abill-back resting posture thatthey ina shrub,but that may have been a dis- similarto the adultsexcept chickrearing areas morefrequently run after insects and placementactivity. In Theypay par- whereshrubs are notpredominant, the probingwas not observed. adultsperch on whateveris available. ticularattention to Munroglobe-mallow River plantswhen they are available,poking Thoseareas along the Columbia at the havescattered trees and I haveseen cur- amongthe foliageand tugging lewsland in themas highas 8 moff the leaves. groundwhen disturbed by an intruder. Fromsuch vantage points, the parent cur- Flapping-Jump lewskeep close watch over the fields un- Thedowny young were frequently ob- tilall dangerhas passed. I have Thebirds touch down gracefully when servedto performa movement andthe wings are termedthe Flapping-Jump,in which alightingabove ground simulta- flutteredoverhead until balance is ob- theywould make short dashes, a ratherlengthy neously flapping their wings and jump- tained.Sometimes it is to fly. processif thebranches of a particularing up anddown as ifattempting weightof the bird Thebehavior occurred when the chicks bushare flimsy and the suchas a istoo great for them. Some birds are often werecrossing an openarea forseveral road,and each chick in turn would do the persistentand keep trying before minutes,breaking numerous branches Flapping-Jump fora fewseconds beforeselecting another perching site. goingon.

Roosting Fledging downywhen they The femalesusually abandon their Thechicks are still are2 to3 weeks areno longer brooded at night. They con- broodswhen the chicks aftersundown old, thoughit is notunusual for 1 to tinuewalking and feeding Afterthe females untilthe femalecalls themwith the remainuntil fledging. roostingarea. depart,only the males care for the chicks. "Wheet"call towardthe foundin which Soonafter sundown, the male flies from Twoyoung broods were ableto observe onlythe female was caring for the chicks, thefield but I wasnever hadpresum- wherehe went.At twilight, each chick andin those cases the male grasswhile the fe- ablybeen killed. disappearsinto the hatching,the ju- malestands alert. At dark, she flies low About31 daysafter also disappears.I venilesand attendingadult(s) change acrossthe field and beginmoving made 1 observationof chickbehavior their daily routine and theiraban- crosscountry away from their nesting on thefirst night following ofchicks donmentby the female. The chicks were area.The feather development 15min after sun- ofknown ages captured prior to that wan- alonewith the male and be within10 downbegan to followafter him. The deringperiod appeared to

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions THE LONG-BILLEDCURLEW IN SOUTHEASTERNWASHINGTON Allen 61 to 14 daysof completion. On thatbasis, hesitateto leave,resulting in prolonged long-billedcurlews fledge at 4145 days. Appeasement-Runbehavior. If a group Watson( 1972)reported that Eurasian fliesin and landsnear the chicks, the curlewsfledge in 3542 days. maleor femalethreatens or pecks each Oncethe birds start moving they may birdone at a timeand often repeatedly travelover 2 kma day,and tracking them beforethe entiregroup departs. The throughthe shrubby areas to makeob- chicksappear not to be botheredby the servationswas notfeasible since the adultinteractions and ignore the intrud- adultsat that stage no longer fly up at the ers. firstsign of disturbance.As a result,I haveno data for that period and was not able to watchthe process of"learning" Responsesto Disturbances tofly. However, on 21 July,I watched 1 Atthe first sign of approaching danger, juvenileflying from the Columbia River adultswith a newlyhatched brood give afterloafing for several hours on an is- a mixedseries of Chirping, "Curlee Cur- land.A strongbreeze was blowingand lee,"and "Wheet Wheet" calls. Demon- thebird had considerable difficulty mak- strationis then performed bythe female ingheadway, but eventually itwas able whilethe male remains with the chicks, toland on top of the bluffs above the riv- thoughflying to a vantagepoint. If the er.After resting for 7 min,the bird again intrudercontinues to approach, then the tookoff and with great difficulty disap- malealso Demonstrates,and sometimes pearedover the horizon headed north- the Enticement-Run is performed. The east.An adult bird could have easily ne- ArcDisplay is performedifthe intruder gotiatedthe climb over the bluffs in even is veryclose to theyoung; e.g., when a a strongwind. humanis searchingfor the chicks. Whenan avianpredator flies over the Territoriality chickrearing area, the adults pursue it intensely.Occasionally, a raptorwill Pairswith young spend a considerableperch near the chicks or will be perched amountof time evicting other long-billed in an areawhere the adults are trying to curlewsfrom the vicinity of the chicks. take the chicks. In suchsituations, the The areadefended by the parents, how- adultcurlews repeatedly perform the Arc ever,is a minimumdistance from the Display.On 2 occasions,I observed par- chicksthemselves rather than the entire entcurlews flying ata perchedred-tailed "territory"as was the case prior to hatch- hawk for over 35 min.Perching raptors, ing.Therefore, as the chicksmove so however,rarely appear to be annoyedby doesthe area defended. The interactions the aerial displays of long-billed curlews, withconspecifics usually are mildbut andultimately the curlews resort to chick violentencounters are also common. removalas a solution. The interlopersbecome increasingly The chicksusually do not conceal problematicthrough June. Their num- promptlyin response to the adult alarm bersincrease as do thesize of the groups calls,but they do concealif both adults as thosebirds without chicks are flock-leave the immediate area. Their cryptic ing,and migrants from other areas start coloration serves them well as they to arrive.In thelate afternoon and eve- crouchflat and motionless on the ground ning,they move into the fields to feed (Fig.28), making them difficult tolocate. androost for the night. They appear to be The adultscall themup witha soft attractedby thepresence of other cur- "Wheet" call when the dangerhas lewsand usually try to land with the par- passed. entbirds and theiryoung. Commonly, As thechicks get older, they wait to suchintruders are verypersistent and concealuntil they actually see the intrud-

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olderones run until they are out of sight ofthe intruder. The adults fly to a vantage pointand keep watch until the intruder hasdeparted.

Accidents,Injuries, and Mortality Twofledglings were involved in colli- sionswith man-made structures. One was founddead, presumably having flown intoa utilitywire. The otherwas found stunnedafter an apparentcollision with a car;it FIG. 28. Long-billedcurlew chick recoveredand was released. concealing. Threechicks were captured that were sufferingfrom some kind of injury. The firstwas about a weekold and had a frac- erapproaching. Also, those older more turedtibiotarsus. experiencedchicks Thechick was found in tendto hide when an emaciatedcondition and died within possiblewith at leastthe head under a 2 hours.The shrubor forb, this often secondwas 2-3 weeksold involvinga run andhad 2 cheatgrassflorets lodged in 1 toa suitablepoint. As they run, they turn eye.The eye was theirheads to watch completelyclosed, but the intruderand afterI removedthe florets the bird was holdtheir wings out to the side. I located able to a chickthat had partiallyopen it. The eye ap- crawledinto a small pearednormal when the same chick was depression,covered with grass and bro- recaptured1 kenJim Hill mustard weeklater. The thirdcase stems.Another was involveda 4-5 -week-old chick. It had an located13 cm offthe ground in a big unidentifiedgrass sagebrush.Corresponding floretembedded in the roughlywith feathershaft of 1 primary,and that pri- thechange in chick behavior, long-billed mary had not curlewadults gradually emergedas fullyas theoth- stopperforming ers. I removedthe floret before releasing threatdisplays and give only alarm calls thebird. inresponse to disturbances. Mortalityis highest in chicksunder 1 Theolder the chicks get, the more they weekof age and runbefore concealing, and seemsto level off after oftenI could that.A veryunusual situation is to see a approachwithin 10 m before losing sight pairof adults with ofthem. Chicks within morethan 2 youngin 1-2 weeksof theircare and mostadults have none. fledgingwould conceal, then just as I was Thechicks are reachingfor them they lostmost frequently at wouldjump up nightwhen observations are made with andtake off running. Once in hand,the difficulty. chickwould give a call similarto the Many adult"Guaah" ofthe causes of chick mortality call. areuncertain, but predation is Whena chickis firstcaptured by a hu- no doubt man,the partlyresponsible. R. E. Fitzner(1977, adultsDemonstrate and per- ResearchScientist, formthe Arc Display. After Battelle,Pacific thechick has NorthwestLaboratories, Richland, Wash- beenin hand for several minutes, itusu- ington99352, pers. allyceases to give its alarm comm.)found the re- noteand the mainsof 1 downychick and 1 adultat a adultsdisplacement feed or assume the Swainson'shawk Alert-Attitudenearby. As nest,feathers from 3 ju- soonas the venileor adult birds in ferruginous hawk chickis released the adults begin again castings,and withtheir Demonstrations. feathersfrom 1 juvenileor The chick adultbird in a greathorned owl casting. runsaway: younger ones usually conceal Isaw a inthe nearest shrub if one is black-billedmagpie carry off 1 available; chickthat was less then 24 hoursold. I

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions THE LONG-BILLED CURLEW IN SOUTHEASTERNWASHINGTON-Allen 63 suspectcoyotes are ofprimary concern, tips of islands in theColumbia River. A thoughI neveractually saw one takea spiton the south side of Island 3 between chick. 100-Hand 100-D Areas serves as thepri- Somechicks may perish early by wan- maryloafing and staging area for curlews deringtoo far from the nest too soon or fromthe Hanford Site and the Wahluke by notbeing able to keepup withthe Slope. adultsand other brood members. A dis- Birdsusually arrive at Island3 in the turbanceat hatchingtime resulting in morningbetween 1000 and noon. They chickremoval most certainly increases arrive singly or in groups, some of which chicklosses in thisway. One chickless includeup to 100 birdsby mid-June. than2 daysold was found dead in a nest Time on the islandis spentresting, field,presumably ofexposure. Exposure preening, and occasionally bathing. The duringthe heat of the day or at night may majorityof the birds remain at thesite causemortality. untillate afternoon or evening,usually Adverseweather conditions are un- leavingbetween 1600 and 1800;all are doubtedlystressful for the chicks. The goneby 2000. nightsare still cool when the chicks are Asthe daily departure time grows near, downyand June is wetand windy. I have "WheetWheet Wheet" calls and bouts of watchedchicks walking and feeding for Two-wingFlap movementsripple hoursat a timein a highwind while it throughthe flock.The birdsbecome rainedcontinuously. Cold, wet, windy more restless, shifting about calling nightsmust certainly take their toll. The "WheetWheet Wheet," and groups be- windalso poses a problemfor the adults ginto leave forthe fields where they whena predatorapproaches because roostfor the night. theycannot fly at it effectively. Eachgroup leaves in a flurryof Two- Manybroods are stillon theground wing Flap behavior.A fewbirds lift off afterthe flocking phase is wellunderway. first, calling "Wheet Wheet Wheet," and Thosechicks that hatch late in Maywill arejoined immediately byothers. Their stillbe downyafter the bulk of the adults flightis erratic as theyare buffeted bythe has departed,and thatabsence of con- wind,but eventually they rise up and fly specificswhich would otherwise provide away in strong,even flight though in no additionaleyes and ears to guard against particular formation. Often shortly after predatorsmay also contributeto chick takingoff, individuals were observed to mortality. dive straightdown, twisting back and In 1976,no estimateof fledging suc- forth,then to swoop up just before hitting cesswas made.In 1977,9 juvenilesre- thewater as ifthey were playing in the mainedalong the riverafter the main wind. congregationoflong-billed curlews from The curlewsapparently can identify theHanford Site and the Wahluke Slope theother birds that roost in their area be- had departed.I was unableto estimatecause each grouplifts off together and ageor ascertain the sex of the birds at the headsin a particulardirection. When sev- stagingarea because the staging island is eralgroups leave simultaneouslythere toodistant from the mainland. doesn'tseem to be anyconfusion among groupmembers as towhich way to go or STAGINGAND DEPARTURE withwhom. However, when a groupini- tiatesits departure, other birds may also Stagingand departurewere not ob- flyup tojoin them but then return to the servedin 1976.In 1977,the first small flock. flockswere observed in the fields in early As theflocking phase progresses, the June. birdsarrive earlier and leave later each Flocksspend the heat of the day rest- day;the flocks also become larger. Apeak ingin fieldsor loafingon theupstream population of approximately250 birds

This content downloaded from 128.193.8.24 on Tue, 27 Aug 2013 17:50:28 PM All use subject to JSTOR Terms and Conditions 64 WILDLIFE MONOGRAPHS waspresent on 17June, following which in one particulardirection they began theirnumbers decreased to merely a few millingaround over a nearbyfield, call- stragglersby theend ofthe month. In ing"Wheet Wheet Wheet" continuously July,only a smallgroup of juveniles was as they spiraled upwards.I watched present. those groupsthrough binoculars until Duringthat period of flock reduction, they went up so highthat I couldno long- groupsof up to 30 birdswere observed er see them.Presumably those birds mi- to leaveIsland 3 at theusual daily de- gratedfrom the Hanford Site and south- parturetime, but instead of heading out easternWashington.

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A CURLEW S FAREWELL I triedbut I can tryno more I criedbut I can cryno more I failedto bringa youngchickys cry Intothis world. Timenow bids me sayfarewell The sunis settingand I mustgo ButI willcome again next year . . . and try UntilI die.

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inthe egg young of the long- CITED 1971.Clicking LITERATURE billedCurlew. Wilson Bull. 83 441442. . 1972. Long-billedcurlew with supernu- The meraryhallux. Auk 89:457. ALLEN,A. A., AND H. KYLLINGSTAD. 1949. of long- of the bristle-thighedcurlew. . 1973. Growthand development eggs and young Auk90:435A38. Auk66:343-3S0. billedcurlew chicks. 1957. Check- GOSS-CUSTARD,J.D.? ANDR. E. JONES.1976. The AME:RICANORNITHOLOCISTS UNION. cllrlew.Bird Study ofNorth American birds. Fifth Ed. PortCity diets of redshankand list 23:233-243. Press BaltimoreMd. Observationsat a long-billed BAILEY, F. M. 1904. Handbook of birds of the GRAUL,W.D. 1971. and curlewnest. Awlk 88:182-184. westernUnited States.Houghton, Mifflin the Pp. 101-102. GSNNELL,J. 1921. Concerningthe status of Co., Boston7Mass. ofthe long-billed curlew. C. 1907. Summerbirds of southwestern supposedtwo races BENT,A. Condor23:21-27. Saskatchewan.Auk 24:407430 Notes011 water hirds . 1962. Life historiesof Northkmerican > ANDR. HUNT. 1929. Inc., New at MorroBay7 California Condor 31:62-73. shorebirds. Dover Publications, ctlrlewin Massa- York,N.Y. Pp. 97-109. GRISCOM)L. 1939. Long-billed Ameriean chusetts.Auk 56:332-333. BISHOP, L. B. 1910. Subspeciesof North mam- 27:59-63. HALL,E. R., ANDK. R. KELSON. 1959.The birds.Auk RonaldPress Co., New BROOKS,A. 1920. Notesonthe Limicsolaeof south- malsof North America. Columbia.Condor 22:26-32. York7N.Y. ern British Comparativebehavior of BURTON7J. K. 1974. Feeding and the feedingap- HAMILTON,R. B. 1975. and ad- theAmerican avocet and the black-necked stilt paratusin waders:A studyof anatomy Monogr.17:1-98. aptationsin theCharadrii. Brit. Mus. Nat.Hixt., (Recurvirestridae).Ornithol. HIBBERT-WARE7A. ANDR. F. RUTTLEDGE.1944. London,Eng. foodhabits of the common BYSTRAK,D. (ED.). 1974. Winteringareas of bird A studyof the inland aircraft.Natl. curlew.Brit. Birds 38:22-27. species potentiallyhazardous to Flora New York,N.Y. HITCHCOCK7C. L.>AND A. CRoNQvIsT. 1973. AudubonSoc. Inc.7 Univ.of Washington CAMERON, E. S. 1907. The birds of Custer and ofthe Pacific Northwest. Mantana. Auk 24:241-270. Press7Seattle, Wash. Dawson counties, thebrink of extinction. Tex- CAMPBELL, R. W. 1972. Cc)astalrecords of the IVERSENE. 1976.Qn long-billedcurlew br BritishGolumbia Can. as ParksWild. March.24-26. LAFAv L. D. 1954. Breedingof northern long- Field-Nat.86:167-168. Lake. Murrelet CLINE, J. F.) D. W. URESK, AND W H RICKARD. billed cllrlewsat Sprague 53:48. 1975. Characterizationof plant communities on C. Cribs controlleclarea and LITTLLFIELD,C. D. 1973. Birdarrival dates adjacentto the B. Refuge Oregon. Redox Pond areas on the 200 Area plateau MalheurNatic)nal Wildlife EENWL-1916,Battelle, Pacific Northwest Lab- WesternBirds 4:83-88. Richland,Wash MANOLIS)T. AND G. V. TANGREN.1975. Shore- oratories, Valley,Calibrnia. COLEX T.7 AND R. S. SHARPE 1976. The effeetsof birdxof the Sacramento grazing managementon a sandhills prairie WesternBirds 6:45-54. densityand di- MCCALLUM,D. A.SW. D. GRAULAND R ZACCAG- community.III. Breedingbird cur- versity.Proc. Nebr. Acad. Sci. Affll.Soc. 86:12. NINI. 1977. The statusof the long-billed DAUBENMIRE,R. 1959. A canopy-coveragemethod lew in Montana.Auk 94:599-601. analysis.Northwest Sci. 33:43-64. MCCAQ,KIE,G. 1970. Shorebirdand waterbird use ofregetation FishGame 56.87-95. DAWSON W. L. 1909. The birds of Washington. ofthe Salton Sea Calif. Co. Seattle Wash. Pp. 680- MILLER?R. G., JR.1966. Silnultaneous statistical OceidentalPublX Co.>New York, 682. inirence.McCraw-Hill Book DICE, L. R. 1918. The birds of Walla Walla and N.Y. Washington 013ERHOLSER,H.C. 1918.Notes on the subspecies Columbia counties,southeastern Auk Auk35:40-51. of Numeni7ls utnericanus Bechstein. observedin Saskatchewan 35:188-195. FERRY,J. F. 1910. Birds of North the summerof 1909. Auk 27:185-204* PALMER7R. S. 1967. The shorebirds during In G. D. Stout R. E. 1977. Time-lapsephotography br AmericaVol. III. Pp. 183-184* FITZNER Anlerica.Viking fieldstudies of raptorialbirds. In J. T. Kitch- (Ed.). The shorebirdsof North Tarr(Eds.). Natl.Environmental Press,New York,N.Y. ingsand N. E. gath- ResearchPark Symp.:natural resource inven- PETTINGILL,O. S.7 JR.1968. An llncommon Alldubon70:6 tory,charactcrization, and analysis. ORNL- ering of statesideshorebirds. Oak Ridge Natl. Lab., Oak Ridge,Tenn. 17. 5304, and Middle D. M. 1967. Egg teethand hatching RIDGWAY,R. 1919. The hirdsof North FonsYTHEt 50:390-395. meth£dsof the long-billed curlew. Wilson Bull. America.U.S. Natl. Mus. Bull. MAHER.1976. Notes 79:340-341. SADLER,D. A. R. ANDW. J. . 1970. Vocalizationsof the long-hilled£:ur- on the long-billedcurlew in Saskatchewan. lew. Condor72:213-224. Auk93:382484.

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SAUER, R. H., AND S. L. OWZARSKI. 1979. Collec- APPENDIX tionand analysis of vegetative cover data. PNL- Ltstof Commonand ScientificNames of Plants 3068,Pacific Northwest Laboratory, Richland, Mentionedin theText Wash. SHUFELDT, R. W. 1913. An introductionto the GRAMINEAE studyof the eggs of the North American Limi- colae.Condor 15:138-151. Cheatgrass Bromustectorum L. SILLOWAY,P. M. 1902.Some experiences of 1901. Sandberg's bluegrass Poa sandbergiiVasey Condor4:31-34. Bluebunch wheatgrass Agropyronspicatum . 1909.A problemin indeterminates. Condor (Pursh) Scribn. & Smith 11:86-90. Sand dropseed Sporoboluscryptandrus (Torr.) SIMMONS,K. E. L. 1955. Thenature of the predator Gray reactionsof waders toward humans; with spe- POLYGONACEAE cialreference to the role of the aggressive-, es- Buckwheat Eriogonumsp. Michx. cape-,and brooding-drives.Behaviour 8:130- 173. CHENOPODIACEAE . 1957. The taxonomicsignificance of the Russian thistle Salsolakali L. head-scratchingmethods of birds. Ibis 99:178- Spiny hopsage Atriplexspinosa (Hook.) Collotzi 181. 1961.Problems of head scratching inbirds. CRUCIFERAE Ibis 103a:37-49. Jim Hill mustard Sisymbriumaltissimum L. STEBBINS, R. C. 1966.A fieldguide to western rep- Wallflower Erysimumasperum (Nutt ) DC. tilesand amphibians.Houghton Mifflin Co., LEGUMINOSAE Boston,Mass. Lupine Lupinussp. L. STENZEL, L. E., H. R. HUBER, AND G. W. PAGE. 1976. Feedingbehavior and dietof the long- POLEMONIACEAE billedcurlew and willet. Wilson Bull. 88:314- Long-leaf phlox Phloxlongifolia Nutt. 332. BORAGINACEAE STONE, W. 1901. Reportof the committee on the protectionofNorth American birds for the year Tarweed fiddleneckAmsinckia Iycopsoides Lehm. 1900.Auk 18:68-76. ONAGRACEAE STONE, W. A., D. E. JENNE, AND J. M. THORP. Pale evening primrose Oenotherapallida Lindl. 1972. Climatographyof the Hanfordarea. BNWL-1605,Battelle, Pacific Northwest Lab- MALVACEAE oratories,Richland, Wash. Munro globe-mallow Sphaeralcea munroana SUCKLEY, G. 1860. Waterbirds. Pp. 245-246.In (Dougl.) Spach The naturalhistory of WashingtonTerritory andOregon. Bailliere Brothers, New York, N.Y. LILIACEAE SUGDEN, J. W. 1933.Range restriction ofthe long- Asparagus Asparagusofficinalis L. billedcurlew. Condor 35:3-9. Brodiaea Brodiaeadouglasii Wats. TAVERNER,P. A. 1934.Birds of Canada. Natl. Mus. UMBELLIFERAE Can.Bull. 72:189-190. Desert parsley Cymopterusterebinthinus (Hook.) TIMKEN, R. L. 1969.Notes on thelong-billed cur- T. & G. lew.Auk 86:750-751. TREMAINE,M. M. 1975.Life at an avocetnest. Au- CACTACEAE dubon77:68-75. Plains prickly-pear cactus Opuntiapolyacantha U . S . DEPARTMENT OF THE INTERIOR. 1973. Haw. Threatenedwildlife of the United States. U.S. FishWildl. Serv. Res. Publ. 114:204. ROSACEAE WATSON, D. 1972. Birdsof moorand mountain. Antelope bitterbrushPurshia tridentata (Pursh) ScottishAcademic Press, London, Eng. Pp. 88- DC. 91. COMPOSITAE WICKERSHAM,C. W. 1902. Sickle-billedcurlew. Big sagebrush Artemisiatridentata Nutt. Auk19:353-356. Gray rabbit-brush Chrysothamnusnauseosus WIENS, J. 1969. Anapproach to thestudy of eco- (Pall.) Britt. logicalrelationships among grassland birds. Green rabbit-brushChrysothamnus viscidifiRorus Ornithol.Monogr. 8:1-93. (Hook.) Nutt. WILSON, A.,AND C. L. BONAPARTE. 1831.Ameri- Fleabane Erigeronsp. L. can ornithology.Porter and Coates,Philadel- Balsamroot Balsamorhizacareyana Gray phia,Pa. Pp. 320-324. Yarrow Achillealanulosa (Nutt.) Piper WOLF, L. R. 1931. The breedingLimicolae of Utah.Condor 33:49-59.

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