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Effects of Geographic Location and Habitat on Breeding Parameters of Great Tits

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Effects of Geographic Location and Habitat on Breeding Parameters of Great Tits The Auk 115(4):1034-1051, 1998 EFFECTS OF GEOGRAPHIC LOCATION AND HABITAT ON BREEDING PARAMETERS OF GREAT TITS JUAN Jos12SANZ • ZoologicalLaboratory, University of Groningen,P.O. Box 14, 9750 AA Haren,The Netherlands ABSTRACT.--Istudied variation in laying date,clutch size, and numberof fledglingsin Great Tits (Parusmajor) in relationto habitat,elevation, and latitudeusing data from 137 breedingareas. Laying date was not affectedby habitattype but increasedwith elevation andshowed a significantquadratic relationship with latitude.Food availability, ambient tem- perature,and photoperiodtogether can explainwhy laying date increaseswith latitude. However,more information is neededto understandwhy birdsin northernAfrica start lay- ing later than in nearbysouthwestern Europe. Variation in clutchsize and numberof fledg- lingswas significantly affected by habitattype, with lowervalues in coniferousforests. Mean clutchsize decreased with elevation.Mean clutch size and mean number of fledglingsof first and secondbroods showed a significantquadratic relationship with latitude,with thehigh- est valuesat about 55 to 60øN.The resultssuggest that latitudinal variationin life-history parametersis relatedto variationacross marginal and centralparts of the species'distri- butionand is influencedby factorssuch as daylength,temperature, and/or seasonalityof food resources.Birds at southernlatitudes are probablyunder time constraints,whereas birds at northernlatitudes are probablyunder energyconstraints. Alternatively, Great Tits at northernlatitudes might be less well adapted to thesehabitats because they invaded north- erly latitudesduring the last decades.The proportionof pairs laying a secondclutch de- creasedwith latitude.The observed pattern of a latestart of laying,lower variability in laying date,and lower frequencyof secondclutches in northernpopulations likely was due to the late and shortgrowing season for the GreatTit's main prey,caterpillars. Received 13 January 1997, accepted22 April 1998. THE GEOGRAPHICTRENDS in reproductivepa- sivelycooler climates, the onsetof breedingis rameters of birds have been well known for de- expectedto be delayedowing to low tempera- cades(Lack 1947, Klomp 1970).The ultimate turesand slow developmentof the vegetation regulatorof the timing of breedingis the re- (Lack 1950, Slagsvoid1976, Orell and Ojanen quirementthat reproductiontake place during 1983a). the seasonin whichbirds can raise their young The evolution of clutch size is one of the most most efficiently (Lack 1950, Drent and Daan activelystudied of life-historytraits in birds 1980,Martin 1987).The availabilityof food for (Stearns1992). Clutch size is constrainedby layingfemales has been considered to be both severalfactors, the mostcommonly discussed an ultimate factor (Lack 1950, Perrins and beinglatitude and habitat (Klomp 1970, Perrins McCleery 1989,Daan et al. 1990) and a proxi- and Birkhead 1983, Murphy and Haukioja mate factor (Perrins 1970, Kallander 1974, 1987).For example,within temperateregions Drent and Daan 1980, Martin 1987) in the tim- clutchesof passerinesare larger in deciduous ing of theonset of laying.However, factors such woodlandsthan in coniferousforests (Klomp as habitat type (Klomp 1970, Blondel et al. 1970,van Balen 1973,Zang 1980,Blondel et al. 1993), temperature (Svensson and Nilsson 1987, Sanz 1995). The trend for clutch size of 1995), and photoperiod(Meijer 1989;Silverin passerinesto increasewith latitudeis oneof the 1995;Lambrechts et al. 1996,1997) also may in- moststriking patterns within and amongspe- fluencethe onsetof breeding.In areasat high cies of birds (Perrins and Birkhead 1983). latitudesand elevations,which haveprogres- Clutch size of passerinestends to be two or three eggsin tropicallatitudes (Moreau 1944, • Presentaddress: Departamento de EcologfaEv- Skutch1985), four or five eggsin middle lati- olutiva, Museo Nacional de Ciencias Naturales tudes,and five to seveneggs in arcticlatitudes (CSIC), Jos6Gutierrez Abascal 2, E-28006 Madrid, (Ricklefs 1969). This trend occursin both the Spain.E-mail: mcnsl 1 [email protected] Old World (Lack 1968) and the New World 1034 October1998] BreedingParameters of Great Tits 1035 (Cody 1971) in the Northern Hemisphere, riod or more energy to be investedin a single whereasthe trendin the SouthernHemisphere nestingattempt rather than in severalattempts is weak (Moreau 1944, Yom-Tov1987, Yom-Tov (Slagsvoid1982). The degreeof nestpredation et al. 1994, Martin 1996). seems to decrease with increasing latitude Threemain hypotheseshave been presented (Ricklefs 1969, Jehl 1971, Pedroli 1978, Kulesza to explainthe latitudinal trend in clutchsize in 1990;but see Martin 1996). passerines.First, Lack (1947) suggestedthat The precedinghypotheses predict a positive the increase in clutch size with latitude could relationshipbetween clutch size and latitude, be attributedto the increasein daylighthours but this trend may not be linear A linear in- available for parents to collect food for their crease in clutch size with latitude has been nestlings.Royarea (1969) extendedLack's hy- found in many studies(e.g. Lack 1947, Ash- pothesisto incorporatethe energy require- mole 1961, Cody 1966, Slagsvoid1975a, Kule- mentsof the youngrelative to ambienttemper- sza 1990, Young 1994), but exceptionsare not atureand the time availablefor foraging.Owen rare (e.g. Owen 1979, Orell and Ojanen 1983b, (1979)accepted Lack's daylength hypothesis as Mailer 1984, Isenmann 1987, Soler and Soler a partial explanationand introducedthe diver- 1992, Sanz 1997). Ecologicalfactors can influ- sity of potential food as an additional factor. ence to a different degree the breeding deci- Lack's(1947) hypothesis has been reformulated sions at different latitudes. At low latitudes, in a life-historycontext to encompassrepro- predationor the availabilityof daylightcan be ductive costs in terms of adult survival and/or the mostimportant factors (Sanz 1999), where- future reproductioninvolved in raising a cer- as at high latitudes,variability or unpredicta- tain number of young (Charnov and Krebs bility of food and weathermay be more impor- 1974), and with a focus on individual differ- tant (J•irvinen1983). Northern breeding popu- encesin quality (Drent and Daan 1980, Pettifor lations of passerines are more unstable, and et al. 1988,Slagsvoid and Lifjeld 1990). their environmentis more unpredictable,than Second,Ashmole (1961) suggestedthat bird are southern populations (O. J•irvinen 1979, populationsin temperateareas are regulatedin Slagsvoid1981, A. J•irvinen1983). Stability is the nonbreedingseason by foodresources. The defined as year-to-yearpersistence of commu- breedingdensity thus tendsto be low in north- nity structure(i.e. total density,number of spe- ern latitudes, and competition for food in cies, diversity and frequenciesof species;J•ir- spring and summer is relaxed. Reduced com- vinen 1979).J•irvinen (1979) showedthat pop- petition favors large clutches(Ashmole 1961, ulationstability of birds is greatestin the areas Ricklefs 1980, Koenig 1984). Studies typically of maximum density (usually in the southern relate clutch size to differences between winter or centralparts of the range).At high latitudes, and summer evapotranspiration(AE), a mea- birds under severe climatic conditions are sure that is tightly correlatedwith primary probablynear their ecologicallimit in termsof productivity (Rosenzweig1968) and, presum- acquiringenough energy for maintenanceand ably, with food availability.Correlations be- reproduction (J•irvinen 1983). Moreover, for tween clutchsize and seasonalityof AE have somepasserines clutch size tends to be lowerin been found in some studies (Ricklefs1980; Ko- marginalhabitats than in nearbyoptimal hab- enig 1984,1986) but not in others(Mailer 1984, itats (Klomp 1970, van Balen 1973, K•illander Dunn and Macinnes 1987,Young 1994). It may 1975,Alatalo et al. 1985,Sanz 1995). Peripheral be questionedwhether so crudean estimateof and central parts of the geographicranges of resourcelevels can justify rejectionor accep- speciesare associatedwith marginal and op- tanceof this hypothesis(Mailer 1984). timal habitats,respectively (Slagsvoid 1981). Third, Skutch(1949) invoked predation as a The existenceof a distributionallimit may re- possiblefactor in the evolutionof clutchsize. flect that the conditionsfor reproductionare The classicinterpretation has been that re- poor and larger clutchescannot be selectedfor duced nest predation favors larger clutches (Slagsvoid1981). Therefore,a nonlinear rela- (Skutch1949, Cody 1966, Perrins1977, Slags- tionshipof clutchsize or brood size with lati- void 1982, Ekman and Askenmo 1986, Lima tude may exist, with the maximum valuesoc- 1987, Kulesza 1990, Martin 1992) by allowing curring in the centralparts of the distribution more young to be fed over a longernestling pe- (Sanz 1997, 1999). 1036 JUANJose 5ANZ [Auk, Vol. 115 The GreatTit (Parusmajor) is oneof the most factorsaffecting the evolutionof breedingtime well-studied bird species.Great Tits breed and clutch size in Great Tits. from about10øS to 71øN(Gosler 1993), and they The aims of my study are to: (1) document are resident even at the northern limit of their latitudinalvariation in layingdate, clutch size, breeding distribution (Cramp and Perrins number of fledglings,and proportionof second 1993,Silverin 1995). In a pioneeringstudy, Lack clutchesin the GreatTit usinga large dataset (1950) showedthat clutch size increaseswith collectedover an extensivegeographic range; latitude for different races
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