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Population Ecology of the Long-Billed Curlew (Numenius Americanus) in Western Idaho

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Population Ecology of the Long-Billed Curlew (Numenius Americanus) in Western Idaho POPULATION ECOLOGY OF THE LONG-BILLED CURLEW (NUMENIUS AMERICANUS) IN WESTERN IDAHO ROLAND L. REDMOND AND DONALD A. JENNI Departmentof Zoology,University of Montana,Missoula, Montana 59812 USA ABSTRACT.--IIlwestern Idaho, a breeding population of Long-billed Curlews (Numenius americanus)remained relatively stable over a 7-yr period.Productivity was monitored closely from 1977through 1979. Clutch mortality rates did not differ significantlyamong years, and meanclutch survival for a 32-daynesting period (4 daysegg laying and 28 daysincubation) was40% for all 3 yr combined.Females laid justone clutch(usually of 4 eggs)each season, and meanclutch size was significantlysmaller in 1979than in 1977and 1978.Estimated fledgingsuccess ranged from 0.40in 1977to 0.17in 1978and wasconsistently greater for femalesthat nestedearly eachseason. Mean annualadult survivalwas estimatedat 85% basedon resightingsof color-markedindividuals. Limited data for subadultsurvival pre- cludedcomplete demographic analysis. Nevertheless, given the estimatesof productivity and adult survivalpooled for 3 yr, survivalof subadultsfrom fledginguntil first breeding as3-year-olds needed to beonly 58% to maintainthe stablepopulation size observed through 1983. Received10 June1985, accepted 7 April 1986. THE population ecology of shorebirds (Cha- land et al. 1982), and Common Redshank (Yates radrii) is not well known, probably because 1982). Larger speciestend to be better studied many speciesare wary, breed in remote areas, during the nonbreeding season,and survival or both. Most thoroughly studied are species estimatesbased on annual returns to wintering that nest at temperate latitudes such as the Eur- grounds exist for Bar-tailed Godwits (Limosa asianOystercatcher (Haematopus ostralegus; Goss- lapponica)and Eurasian Curlews (Numeniusar- Custard et al. 1982, Safriel et al. 1984), Common quata) (Evans and Pienkowski 1984). Ringed Plover (Charadriushiaticula; Laven 1940, We studied breeding density, clutch size, Bub 1962, Pienkowski 1984a), and Common nesting mortality, reproductive success,and Redshank (Tringatotanus; Grosskopf 1959, 1964; adult survival in a population of Long-billed Yates 1982). Many aspectsof breeding biology Curlews (Numenius americanus). These demo- are known for other shorebirds, especially in graphic data are integrated by life-table anal- Europe and Scandinavia (see Cramp and Sim- yses,and the longer-term reproductive perfor- mons 1982, Evans and Pienkowski 1984). But mance of the population is evaluated. even among calidridine sandpipers(Scolopaci- dae), a subfamily with diverse social adapta- STUDY AREA AND METHODS tions and mating systems(Pitelka et al. 1974, Studyarea.--The study area was an upland strip of Myers 1981), detailed demographicparameters shortgrassrangeland (•21,600 ha) in western Idaho have been reported only for the Dunlin (Cali- (Redmond and Jenni 1982). We concentrated on a dris alpina; Soikkeli 1967, 1970a, b) and Tem- portion of the area (• 1,600 ha) that supportedhigh minck's Stint (C. temminckii;Hild•n 1978). Trin- breeding densities of Long-billed Curlews. This gine sandpipersexhibit a wider range of body smaller plot was grazed intensively by large bands sizes and tend to breed at lower latitudes than of sheepfrom late March through early May each do calidridines (Johnsgard1981). Hence, they year;other portions of the studyarea were grazedby should be more accessiblefor study, and also cattle from November to June. Methods.--Thenesting cycle was divided into pre- more accessiblefor comparative demographic laying, laying, incubation,and prefledgingperiods. analyses.Long-term reproductive performance A standardclutch of 4 eggswas laid in 4-7 days,and and population dynamics,however, are known incubation began after the last egg was laid. Time for only a few smaller speciessuch as the Spot- from laying to hatchingof the last egg was consid- ted Sandpiper (Actitis macularia;Oring et al. ered the incubation period; it ranged from 28 to 30 1983), Common Sandpiper (A. hypoleucos;Hol- days(•? = 28.4 days). 755 The Auk 103: 755-767. October 1986 756 REDMONDAND JENNI [Auk, Vol. 103 Population densitieswere estimatedeach year by fornicus;Ring-billed Gulls, L. delawarensis;and Frank- a modified Finnish line-transect method (Redmond lin's Gulls, L. pipixcan)occasionally passed over the et al. 1981),and nestswere locatedduring egg laying study area during April and May. Only ravens were or incubation according to methods described by mobbed by adult curlews. Redmond (1986). When found, each nest was marked Grazing livestock also destroyed curlew nests. with a numbered wooden stake (2.5 x 5 x 40 cm) Characteristically,one or more eggsdisappeared from placed 10 m upslope from the nest cup. To deter a nest coincidentwith the presenceof sheepor cattle mammalianpredators from following our scenttrails, in the vicinity. Sometimes a crushed egg remained paradichlorobenzene(PDB) crystalswere sprinkled in the cup or an egg was dislodgeda short distance. at a 1-m radiusaround each nest cup. On subsequent Gophersnakes (Pituophis melanoleucus) were the only visits, a nest was always approached directly from reptiles capableof preying on curlew eggsor chicks. the marking stake;when departing, a line of PDB Thesesnakes were encounteredregularly during late crystalswas spreadacross this path. Becausethe crys- spring, but individuals were always relatively small tals were small and very volatile, they evaporated and probably unable to swallow a curlew egg whole. quickly and probably had only a short-term effect. The existence of individual gopher snakes large Whereasthe efficacyof this method for discouraging enough to ingest whole curlew eggs was document- mammalian predatorshas never been tested,Lehner ed in Nebraska (Tremaine 1975), but we doubt that et al. (1976) showed that similar chemicals deterred such predation was frequent on this study area. coyotes(Canis latrans) and dogs(Canis familiaris) from Daily mortality rates were calculatedfor all clutch- food rewards. Similarly, Knight (1983) found PDB es, for individual eggs within successfulnests, and and napthaleneto be useful in repelling skunksfrom for chicks before fledging according to Mayfield's specificareas. (1975) method as modified by Willis (1981). When Nests found during incubation usually were the day of nest destructionwas not known, it was checkedevery 2-5 daysuntil the eggsfirst starredor assumed to have occurred at 40% of the interval since pipped, and then every day until hatching was com- the last visit (Johnson 1979). Variance estimates of pleted. Nests located during egg laying were not thesemortality ratesfollow Johnson(1979), and rates checked for the first 2-3 weeks of incubation. Clutch were compared by G-testsof independence (Willis size was estimatedby the maximum number of eggs 1981). Capture and marking techniques were de- found in a given nest. scribedby Redmond and Jenni (1982), and Redmond The diversityof potentialpredators on and around (1984) provided details about radio-tracking meth- the studyarea made it difficult to interpret every act ods. of predation and to identify responsiblespecies. Resightingsof natally philopatric curlews (indi- Mammalian carnivores that hunted regularly over viduals color-bandedas chicksin the study area and portionsof the studyarea included coyotes, fetal dogs, resightedthere in later years) were made opportun- feral cats(Felis domesticus), badgers (Taxidea taxus), and istically along survey routes from 1979 through 1981. long-tailedweasels (Mustela frenata). Red foxes(Vulpes By themselves,these resightings represent minimum fulva), striped skunks (Mephitis mephitis),and rac- survival rates for subadults.Because of a significant coons(Procyon lotor) occurred on adjacentagricultur- male bias in natal philopatty (Redmond and Jenni al lands but were never seen hunting in upland hab- 1982), however, subadult survival for males was es- itat. timated as one half the number of color-marked chicks Nest losswas attributed to a badger if the nest cup known to fledge and later resighted. Estimatesof an- was damaged and buried under several centimeters nual adult survival utilize resighting data from 1978 of soil. Large, crushedshell fragmentsoften lay be- through 1981 and are basedon the maximum-likeli- neath this soil or in the immediate vicinity of the hood method of Cormack (1964). To obtain resight- destroyednest. Canids were presumedto be respon- ings of color-marked adults, their previous nesting sible for an empty, undamagednest cup whenever territoriesand adjacentterritories were searchedreg- the previous visit had revealed an intact clutch. In ularly during the prelaying and incubationperiods. several casesalso attributed to canids, the nest cup was damaged but not buried, and large shell frag- RESULTS ments lay nearby. Avian predators usually did not destroy entire Breedingdensity.--Because of high variances, clutches;instead, they opened and consumedpor- there were no significant differences among tions of 1 or 2 eggs.Partially destroyedclutches were abandoned by adult Long-billed Curlews, and re- annual density estimates of territorial male maining eggs were subsequently consumed. Black- Long-billed Curlews on the 1,600-hastudy plot billed Magpies (Pica pica) were the most abundant (Fig. 1). Thus, for the duration of the study, we avian egg predator. Common Ravens (Corvuscorax) assumedthat the breeding population was rel- and severalgull species(California Gulls, Laruscali- atively stable in size, even though a gradual
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