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24 PAGEET AL. [Auk, Vol. 100 Bird Observatory. Ralph Widrig, Steven Cellman, tain Plover social systems.Living Bird 12: 69- Barbara Swarth, Susan Peaslee, Erica Buhrman, Don- 94. na Vaiano, Paul Neal, Sharon Goldwasser, Dennis HAYS, H., & M. LECI•o¾. 1971. Field criteria for Parker, John Harris, David Shuford, Stuart Johnson, determining incubation stage of the Common John and Ricky Warriner, Stephanie Zeiler, Elliot Tern. Wilson Bull. 83: 425-429. Burch, Brett Engstrom,Leslie Smith, BernadetteAl- HOLMES,R.T. 1966. Breeding ecologyand annual len, Greg Calllet, Tom Pogson,Tom Cassidy, Geoff cycle adaptationsof the Red-backedSandpiper Geupel, Doug Burgess,Judy Wagner, Kim Sullivan, (Calidrisalpina) in northern Alaska. Condor 68: JanetKjelmyr, and Tad Theimerhelped with the field 3-46. work and Philip Henderson, Carolyn Frederiksen, 1972. Ecologicalfactors influencing the and Frances Bidstrup provided valuable observa- breeding season schedule of Western Sandpi- tions. John and Ricky Warriner generouslyshared pers (Calidrisrnauri) in subarcticAlaska. Amer. with us their knowledge of Snowy Plovers at Mon- Midi. Natur. 87: 472-491. terey Bay, and CharlesSimis provided weather data. MAYFIELr•,H. F. 1961. Nesting successcalculated Jan Simis allowed us to camp on her land and ex- from exposure.Wilson Bull. 73: 255-261. tended to us very warm hospitality while we were --. 1975. Suggestionsfor calculatingnest suc- there. David Ainley, Peter Connors,David DeSante, cess. Wilson Bull. 87: 456-466. Marshall Howe, Harriet Huber, Frank Pitelka, and OI•INO, L. W., & M. L. KNOrrSON. 1972. Monogamy Chris Ribic provided helpful comments on the and polyandry in the SpottedSandpiper. Living manuscript.We thank theseorganizations and peo- Bird 11: 59-72. ple for their generosity.This is ContributionNo. 189 PITELKA, F. A., R. T. HOLMES, & S. F. MACLEAN. of Point ReyesBird Observatory. 1974. Ecologyand evolution of social organi- zation in arctic sandpipers.Amer. Zool. 14: 185- LITERATURE CITED 204. RITIINGHAUS,H. 1975. Seeregenpfeifer,Charadrius ANDEI•SSOtW,M., & C. G. WI•<LUtWD.1978. Clump- alexandrinusalexandrinus. Pp. 205246 in Hand- ing versusspacing out: experimentson nest pre- buch der Vogel Mitteleuropas,vol. 6 (U. N. Glutz dation in Fieldfares(Turdus pilaris). Anim. Be- von Blotzheim, Ed.). Wiesbaden, Akademische hav. 26: 1207-1212. Verlagsgesellschaft. GORANSSON,G., J. KARLSSON,S.G. NILSSON, & S. SAFI•IEL,U. N. 1975. On the significanceof clutch Ulfstrand. 1975. Predation on birds' nests in size in nidifugous birds. Ecology56: 703-708. relation to antipredator aggression and nest TINBERGEN, N., M. IMPEKOVEN, & D. FRANCK. 1967. density: an experimentalstudy. Oikos 26: 117- An experiment on spacing-out as a defence 120. against predafion. Behaviour 28: 307-321. GOTTFI•IEr•,B. M. 1978. An experimental analysis WILSON,R.A. 1980. Snowy Plover nesting ecology of the interrelationshipbetween nest density and on the Oregon coast.Unpublished M.S. thesis. predation in old field habitats. Wilson Bull. 90: 643646. Corvallis, Oregon, Oregon State Univ. GRAUL,W. D. 1973. Adaptiveaspects of the Moun- The American Ornithologists'Union offers several Marcia Brady Tucker Travel Awards to help defray expensesof outstanding students wishing to present researchpapers at the society'sCentennial Meeting in New York (26 September to 1 October, 1983). To apply, send the following materials to Douglas Mock, Departmentof Zoology,University of Oklahoma,Norman, Oklahoma73019 by 1 May 1983:(i) ex- panded abstractof the paper (max. 3 pp, typed, double-spaced;to include methods,major results,and scientificsignificance), (ii) curriculumvita, (iii) your anticipatedbudget of expenses,and (iv) a letter of support from the academicadvisor supervisingyour research(which should be mailed by itself). Please note that you must provide seven(7) copiesof everything,including the letter of support,because each application is reviewed by the AOU Student Awards Committee's seven members. Also, be advised that applying for a MBT Travel Award does not guaranteea place on the Scientific Program--each student must make arrangementsfor that as per the instructions given with the Call for Papers announcement. INTRASEASONAL REPRODUCTIVE COSTS FOR THE HOUSE SPARROW (PASSER DOMESTICUS) W. BRUCE McGILLIVRAY • Museumof NaturalHistory, University of Kansas,Lawrence, Kansas 66045 USA ABSTP,•CT.--HouseSparrows (Passer domesticus) near Calgary, Alberta begin breeding in earlyspring and continuethrough to late summer.High productivityfrom previousbroods is negativelycorrelated with fledglingproduction from secondand third broods.Although fat reservesmay limit the ability of femalesto raiseyoung, there is no concomitantdrop in clutchsize or in the probabilityof renesting.Pairs that fledgemany young in a yearspace fledgling productionevenly over the breeding seasonbut are most productivein mid- season.The interval between fledging and the initiation of the next clutchincreases with the numberfledged. This delay,an indicationof the physiologicalstrain involved in rearing young,is greaterfor laterbroods and for femalesnesting in trees.Measures of reproductive effort(dutch size, numberfledged, length of the nestlingperiod) vary seasonallybut give no indicationof peakingfor last broods.Thus, reproductiveeffort is not adjustedto parallel changesin the probability of surviving to the next breeding attempt. Received3 November 1981, accepted24 April 1982. A COMMONassumption underlying theoret- ship of double-broodedfemale House Martins ical discussionsof life-history phenomena is (Delichonurbica), and male Pied Flycatchers that trade-offsin life-history characteristicsoc- (Ficedulahypoleuca) feeding many young were cur (Williams 1966, Chamov and Krebs 1974, lesslikely to return to the study area than those Steams 1976, Snell and King 1977). For exam- feeding fewer young (Askenmo 1979). Lack ple, Pianka and Parker (1975)partition the re- (1966) and Kluyver (1970) present evidence productive value of an individual of age X in suggesting an inverse relationship between a stablenongrowing population into fecundity brood size and parental survivorship. De Ste- at age X and expected fecundity conditioned ven (1980), however, did not find a reduction by survival to ages X + 1, X + 2.... , X + in the survivorship of female Tree Swallows N. Clearly if survivorshipis lowered by current (Iridoprocnebicolor) as a result of brood en- reproduction,future fecundity is reduced. largement. Smith (1981), after demonstratinga Many researchersof life-historyphenomena positive associationbetween reproductive ef- in birds have investigated whether or not the fort and survivorship, concludedthat a trade- modal clutch size is the most productive, as off between fecundity and survivorshipshould predicted by Lack (1947, 1966). If productivity not be assumedfor passerinebirds. is measured as the number of young fledged The longevity and mobility of birds usually from a brood, then clutch sizes larger than prevents a precise determination of mortality modal appear to be optimal (Klomp 1970, von rates for individuals with a known reproduc- Haartman 1971,Jones and Ward 1976, Murphy tive history. If dispersers are a nonrandom 1978). If parental survivorship is reduced by sampleof the population (Lowther 1979),then the effort of raising a large brood, however, estimatesof survivorshipbased on individuals then the optimal clutch size might be lower staying at or returning to the study area will than the most productive (Fisher 1930, Char- be biased. As a consequence,the effect of re- nov and Krebs 1974). Studies testing the rela- production on survivorship is often estimated tionship between brood size and parental sur- from physiologicalevidence. Weight lossesfor vivorshipshow equivocal results. Bryant (1979) femalesthrough the breeding seasonsuggest a found a reduction in the overwinter survivor- physiologicalstrain that may reduce survivor- ship (Newton 1966,Hussell 1972, Bryant 1979). For femaleHouse Sparrows(Passer domesticus), Pinowska (1979) has shown that levels of fat • Present address: Water ResourcesBranch, On- influence clutchsize, while lean dry weight (an tario Ministry of the Environment 135 St. Clair Av- indicator of protein level) may determine the enue W., Toronto, Ontario M4V 1P5, Canada. number of broods raised in a season.Norberg 25 The Auk 100: 25-32. January 1983 26 w. BRUCEMcGxLLXVRA¾ [Auk, Vol. 100 (1981) presented the hypothesis that adult countedand numbered. Nestlings 5•5 days old were weight lossesduring breedingmay be adap- banded with a U.S. Fish and Wildlife Service alu- tive. This would be true if the reduced cost of minum leg band and color marked with leg bands collecting food for the nestlings increases after reachinga 20-g weight. Data analysis.--Becausethe nestswere not checked fledgling numbersmore than it reducessur- every day, exact clutch-initiation data and nestling vivorship and future fecundity. ages were not known for all nests. For these cases, Birds that raise more than one brood in a estimateswere used. Nestling age when first found seasonmay be expectedto show differential was estimatedby comparingthe weight of the largest reproductiveeffort and productivitythrough nestling in a brood to averagevalues obtained from the breedingseason. There is a higher proba- broods of known age. Clutch-initiation date was es- bility of survivingfrom one broodto the next timated by assuming an average incubation period ct-uringthe breedingseason than from the last of 11 days and a laying rate of one egg