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ShortCommunications andCommentaries

the rate of mate incubation feeding in the Pied FTycatcherFicedulahypoleuca. Behav.Ecol.SociobloT. 24:417-420.

SOK•L,R. R., ANDF. J. ROHLF. 1981. Biometry. W. H. Freemanand company,New York. TINBERGEN, J.M.,ANDM. C. BOERLIJST. 1990. Nestling weight and survivalin individual GreatTit (Parus major).J. Anita. Ecol.59:1113-1127. WALSBERG, G. E. 1983. Arian ecologicalenergetics. Pages161-220 in Arian biology, vol. 7 (D. S. Far-


net, J. R. King, and K. C. Parkes,Eds.).Academic Press, New York. W•THœRS, W. W., ANt• K. A. St•LLIVAN. 1989. Juve-

nile foragingproficiency,parentaleffort and avian reproductivesuccess.Ecology59:223-246. WœBB,D. R.

1987. Thermal tolerance of avian em-

bryos:A review. Condor 89:874-898. Received27 February1995,accepted20 June1995.

The Auk 113(1):253-256, 1996

Absenceof Blood-parasitization Effects on LesserKestrel Fitness Jos• L. TELLA,• MANUELAG. FORERO, t ALvAROGAJ(•N,2 FERNANDOHIRALDO,• AND Jos• A. DON.•ZAR•

•Estacidn Bioldgica de Do•ana,C.S.I.C.,Avda.M Luisas/n, Pabelldndel Perg,41013Sevilla,Spain;and 2Centrode Diagn•sticode FaunaSilvestre,Facultadde Veterinaria,Universidadde Zaragoza, Miguel Servet17, Zaragoza,Spain BToodparasiteswere thought to be benign to their avian hosts,but recentreviewsuncoveredimportant alterationsin birds infected with blood parasites(Atkinsonand van Riper 1991,Bennettet aT.1993).Hamilton and Zuk (1982) proposedthat secondarysexual traits evolved assignalsof parasiteresistancethat are used in mate choice. This hypothesis has been the focus of recent research and reviews (Gibson 1990, Pruett-Joneset aT. 1990, Weatherhead 1990, Clayton 1991, Weatherhead et al. 1991, Weatherhead and Bennett 1991, 1992, Lozano 1994, Seutin 1994). Recent

studiesalso have focusedon possibledetrimental effectsof haematozoaninfection on reproductiveeffort (Apanius 1993, Norris et al. 1994), breeding success (Davidar and Morton 1993, Korpim•iki et aT. 1993, Allander and Bennett1995),malespring arrival (R•itti et al. 1993), dominance (Weatherhead et al. 1995), and bird survival (Davidar and Morton 1993). Weatherhead (1990) failed to find any fitnesscostcausedby blood parasites. The LesserKestrel (Falconaumanni)is a migratory colonial falcon with strong sexual dimorphism in plumage.Adult malesare brightly colored,whereas femalesand juveniles of both sexesare dull (Cramp and Simmons1980). We report on levels of parasitization by haematozoain a LesserKestrelpopulation and relate these to hosts'reproductiveeffort, clutch size, and survival.

Methods.--Our study was conducted in Los Monegros(northeasternSpain;41ø25'N,0ø1I'E), where a large population of LesserKestrel breeds in abandoned farm houses(Tella et al. in press).Adult birds were caught while roostingor attending nests;cap-

turesoccurredfrom spring arrival (March) to the end of the breedingseason(July) in 1993and 1994.Nestlings were sampledin 1993.We took 498 blood sampTesfrom the brachial vein of as many hosts.Thin blood smears were individually labelled, air dried, fixed with 100% methanol, and stained with Giemsa

(Bennett 1970). A 100 x oil-inmersion lens was used

to count blood parasitesin 100 microscopefields on each smear.




in a line



end of the slide to the other to compensatefor differences in the thickness of the smear (Weatherhead and Bennett 1991).Haemoparasiteprevalencewas defined as the percentageof infected individuals in a sample,and intensity as the number of parasitesper infected bird per 100 microscopefields. The identity of parasite specieswas determined at the International Reference Centre for Avian Haematozoa (Memorial University of Newfoundland, Canada). We were able to age mostof the birds asthey were bandedwhen young. Clutch size was determined in focalpairs,after successive visitsto estimateegg losses due to predation (Tella et al. in press).A two-factor ANOVA


no differences

in clutch

size be-

tween years (F•,•07= 1.774, P = 0.18), but significant differencesbetweenfirst-yearand older females = 4.257,P = 0.016). Thus, we only analyzedclutches from after-first-year(AFY) femalesand pooledyears. Laying and hatchlingdateswere estimatedaccording to the length of the eighth primary feather of the largestchick in eachbrood (Negro et al. 1992).Based on these data, we grouped the known parents into the prelaying (March to beginning of May), incubation (most birds until beginning of June), and nest-


ShortCommunications andCommentaries

TABLE1. Number and percentage (in parentheses) of infected and uninfected LesserKestrelsdepending on sex and year.

= 4.47, P < 0.05). However, of 64 adults and 4 nest-

lings taken in 1993and resampledin 1994(3 of them parasitizedin 1993),only 1 adult shifted statusfrom infected




1994 Total

variations, no trends in either sex were found in the

1 (1.28) 1 (1.11) 2 (1.19)

percentageof parasitizedbirds throughout the sampling period (males,X 2 = 1.81, P = 0.40; females,X2 = 0.16, P = 0.92; Table 2). Furthermore, only 1 of the 28 birds resampled in successiveperiods in 1994 changedfrom unparasitizedto parasitizedstatus. Clutch size of AFY parasitized females (œ= 4.66 + 0.70, n = 9) did not differ from that of unparasitized ones of the sameage group (œ= 4.61 + 0.70, n = 65; Mann-Whitney U-test, z = -0.046, P = 0.96). Blood parasitizationapparentlydid not affectLesserKestrel survival, although the sample size of infected birds was very small for a meaningful comparison;2 of 6

77 (98.72) 89 (98.89) 166 (98.81)

Females 1993 1994 Total

to uninfected.

Regarding the reproductive effort and seasonal

Males 1993

[Auk,Vol. 113

5 (4.59) 5 (4.95) 10 (4.76)

104 (95.41) 96 (95.05) 200 (95.24)


12 (3.17)

366 (96.83)

infected birds, and 90 of 179 uninfected

ling (until middle July) periods. We resampled several birds in differents periods of the 1994 breeding seasonto measuretheir reproductiveeffort. To avoid pseudoreplication,we randomly selected one blood samplefrom birds that were caughtmore than oncein the sameyear. However, we considered smearstaken from the samebird in different yearsas independent samples, as blood parasitization can change depending on year and bird's age (Gibson 1990, Weatherhead and Bennett 1991, Davidar and Morton 1993, Allander and Bennett 1994, Norris et

al. 1994, Seutin 1994). We comparedprevalencesbetween years using chi-squaretests,with Yates' correction when the expected values were lower than five (Zar 1984).

Results.--Theonly bloodparasitefound in our Lesser Kestrel population was Haemoproteus tinnunculi.It was detected in 3.17% of adult birds (n = 378; Table

1), and there were no differencesin prevalence between the two yearsof study(1993,3.20%;1994,3.14%; X 2 = 0.0, P = 0.97). Intensity ranged from 1 to 95 infected erythrocytesper 100 inspectedmicroscope fields (median = 21.5, lower quartile = 9, upper quartile = 53.5, n = 12). None of the sampled nestlings (n

birds re-

turned from 1993 to 1994 (X 2 = 0.16, P = 0.68).

Discussion.--Ourstudy is the first to report Haemoproteustinnunculiparasitizing the Lesser Kestrel. This parasiteis broadlydistributedin falconidspecies from the Old and the New World (Peirce et al. 1990,

Bennettet al. 1992a).However, its prevalencein the studied LesserKestrel population (3.17%) is much lower than that shownby this or otherbloodparasites in most of well studied avian hosts (28-100%; see Gibson 1990, Pruett-Jones et al. 1990, Weatherhead

and Bennett 1991, 1992, Apanius 1993,Davidar and Morton 1993,Korpim•iki et al. 1993,R•itti et al. 1993, Allander and Bennett 1994).The causesof the scarcity of haematozoain our LesserKestrel population are not clear, but might be due to the local absenceof suitablevectors(ornitophilic ceratopogonids).Lesser Kestrelsbreedin open,arid regions(typicallysteppes). In the tundra, another treelesshabitat, both sedentary and migratory birds are almost free from blood parasites,while they are heavily parasitizedin forested boreal environments (Bennett et al. 1992b, Earl• and Underhill 1993). In forested areas of North America, 85% of American Kestrels (Falcosparverius)are para-

sitizedby H. tinnunculi (Apanius1993).Furtherstudies

= 85) were infected.

are needed to evaluate these differences (Bennett et

Prevalencewashigher in femalesthan males(Table 1; X 2 = 3.87, P < 0.05 for pooled data). All infected birds were more than two yearsold, this age-related trend being only significantin the caseof females(X2

al. 1992b).A low densityof vectorsmight alsoexplain that the risk of infection increaseswith the age of the bird, due to the longer time of exposure,at a lower rate than in highly parasitizedspecies(Weatherhead

TABLE2. Prevalence(number with percent in parentheses)of Haemoproteus tinnunculiin adult male and female LesserKestrelsin relation to reproductivestatus. Breeding period








2 (2.33)

84 (97.67)

5 (4.72)

101 (95.28)


0 (0.00)

49 (100.00)

4 (5.33)

71 (94.67)

Chick rearing

0 (0.00)

28 (100.00)

1 (3.45)

28 (96.55)


ShortCommunications andCommentaries

and Bennett 1991, Davidar and Morton 1993, Allander

and Bennett 1994, Seutin 1994).

As in other species(e.g. Korpim•iki et al. 1993,Norris et al. 1993), more female than male Lesser Kestrels

were parasitized. This fact may be related to reproductive effort decreasingthe host'sability to control chronic infections and/or a greater exposure of females to vectors (Norris et al. 1994). Our results do

not supportthe first hypothesis,sinceprevalencedid not increase intrasexually during the breeding season. Furthermore, males feed the females during the prelaying period to improve the female'sbody con-

dition (Don•zaret al. 1992),andstudiesusingdoubly labelledwater showedthat malestend to spendmore energy than femalesduring the chick-rearingperiod (J.L. Tella,J.A. Don•zar,andF. Hiraldo unpubl.data). Taking into accountthat LesserKestrel femalesinvest more time in nesting activities than males (Negro 1991),sexdifferencesin blood parasitizationcouldbe related to the spatiotemporalpattern of activity of vectors.

Blood parasitization does not appear to affect individual fitnessof LesserKestrelsgiven that there is no apparent reduction of clutch size or adult survival. Variation in male spring arrival (up to two months) alsocannotbe attributablein this populationto bloodparasite loads (R•tti et al. 1993, but see Davidar and Morton 1993).Our resultsalsohave implications concerning the relation between blood parasites and


ulation of Great Tits Parusmajorfrom Gotland, Sweden. J. Avian Biol. 25:69-74. ALLANDER,K., AND G. F. BENNETT. 1995. Retardation

of breeding onset in Great Tits (Parusmajor)by blood parasites.Funct. Ecol. 9:677-682. APANIUS,V. 1993. Blood parasitism,immunity and reproduction in American Kestrels (Falcosparverius).Pages117-125 in Biologyand conservation of small falcons (M. K. Nicholls and R. Clarke, Eds.). The Hawk and Owl Trust, London. ATKINSON, C. T., AND C. VAN RIPER. 1991.


genicity and epizootiology of avian haematozoa: Plasmodium,Leucozytozoon, and Haemoproteus. Pages 19-48 in Bird-parasite interactions: Ecology, evolution, and behaviour (J. E. Loye and M. Zuk, Eds.). Oxford Univ. Press, Oxford. BENNETT, G.F. 1970. Simple techniquesfor making avian blood smears. Can. J. Zool. 48:585-586. BENNETT,G. F., R. A. EARLR,H. DU TOIT, AND F. W.

HUCHZERMEYER.1992a. A host-parasite catalogueof the haematozoaof the sub-Saharanbirds. OnderstepoortJ. Vet. Res. 59:1-73. BENNETT,G. F., R. MONTGOMERIE, AND G. SEUTIN. 1992b.

Scarcityof haematozoain birds breeding on the Arctic tundra of North






1993. Avian haematozoa:Mortality and patho-

genicity.J. Nat. Hist. 27:993-1001.

plumagebrightnessin birds(Hamilton and Zuk 1982). CLAYTON,D. H. 1991. The influence of parasiteson host sexualselection.Parasitol.Today 7:329-334. Variability in the highly colored plumage of male LesserKestrelscan hardly be explained by haema- CRAMP, S., AND K. L. E. SIMMONS. 1980. Birds of Europe, the Middle East,and North Africa, vol. tozoainfestations,unlessthe population that we stud2. Oxford Univ. Press, Oxford. ied is exceptionalin terms of its low parasite prevaDAVIDAR,P., AND E. S. MORTON. 1993. Living with lence. Thus, we provide additional evidence that, at parasites:Prevalence of a blood parasite and its least in large populations of some species,processes of mate choice and sexual selection are not mediated effecton survivorship in the Purple Martin. Auk 110:119-116. by blood-parasite infestation (Bennett et al. 1992b, DONb,ZAR, J. A., J. J. NEGRO,AND F. HIRALDO. 1992. Earl• and Underhill 1993,Seutin 1994). Functionalanalysisof mate-feedingin the Lesser Acknowledgments.--We thank R. L6pez and I. S•nKestrel Falco naumanni. Ornis Scand. 23:190-194. chezfor their help during the fieldwork, and V. ApanEARLf•, R. A., AND L. G. UNDERHILL. 1993. Absence ius for his first guidancein haematozoanstudies.G. of haematozoa in some Charadriiformes breedF. Bennett kindly confirmed the blood-parasitespeing in the Taimyr Peninsula, Russia.Ardea 81: cies.G. F. Bennett,G. R. Bortolotti,J. J. Negro, and two anonymous reviewers provided constructive

commentson the manuscript.I. de Bustamantehelped with the Englishtranslation.Finnancialsupportwas

providedfromthe CICYTprojectPB90-1021, andpartially from the Servicio de Conservaci6ndel Medio


GIBSON,R. M. 1990. Relationships between blood parasites,mating successand phenotypic cues in male Sage Grouse Centrocercus urophasianus. Am. Zool. 30:271-278.

Natural, Diputaci6nGeneralde Arag6n. J.L.T.benefitted from a FPI predoctoral fellowship from the Ministerio de Educaci6ny Ciencia,and a stayat the

HAMILTON, W. D., AND M. ZUK. 1982. Heritable true

Avian Scienceand Conservation Centre (McGill Uni-


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LOZANO,G.A. 1994. Carotenoids, parasites,and sexALL•NDER, K., AND G. F. BENNETT. 1994. Prevalence

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ShortCommunications andCommentaries nlcaloprimilia Falconaumanni. Ph.D. thesis,Univ. Sevilla, Sevilla, Spain.


Copulatorybehaviourin a colony of LesserKestrels:Sperm competitionand mixed reproductive strategies.Anim. Behav.43:921-930. NORMS, K., M. ANWAR, AND A. F. READ. 1994. Re-

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Received24 April 1995,accepted21 August1995.

The Auk 113(1):256-257, 1996

An Associationof Habitat with Color Dimorphism in Finches TREVOR PRICE

Biology Department 0116,Universityof Californiaat SanDiego,La Jolla,California92093,USA Sexual dimorphism in the color of birds is often attributed to sexual selection (Moller and Birkhead

1994),althoughthereare alternativeexplanations(e.g. Baker and Parker 1979). One of the more puzzling

observations isthatmanycloselyrelatedspeciesdiffer in degree of dimorphism. For example, the House Sparrow(Passer domesticus) is dimorphicand the Tree Sparrow (P. montanus)monomorphic. Understanding differences

such as these would

be aided

if environ-

mental associationswith dimorphism could be detected. In one of the few examplesof such an association, Crook (1964a, b) showed that the forest-dwell-

ing weaver fincheshave dispersedterritoriesand are monogamousand monomorphic,whereassavannah speciesare colonial, polygynous,and dimorphic. In this note ! demonstrate

an association

of habitat


dimorphismacrossfinches on five different continents.

Schluter(1986) presentedlists of finch speciesoccurringin similarhabitatsin five differentregionsof the world (North America, South America, Europe, Africa, Australia). Finches come from four different

families (the Emberizidae,Frigillidae, Estrildidae,and Ploceidae),and no speciesare held in commonacross all five regions investigated (for detailed discussion of dataset, see Schluter 1986). Distributions of mono-

morphicand dimorphicfinchesin different habitats are shown in Figure 1. Following Schluter(1986)and Schluterand Ricklefs(1993), I useda two-way ANOVA (region x habitat) to test for differencesbetween habitatsin the proportion of finch speciesthat are monomorphic. Since there are no replicatesper cell, the interaction



be tested

and is used as

the error term in the ANOVA. There is a significant differenceamonghabitats(Fs,•o = 4.8,P < 0.05).There is no significantdifferenceamongregion(F4,•o = 1.1, P > 0.4). Testsweighting by samplesize in eachhabitat and after arcsin-transformingthe data gave similar significancevalues. Reasonsfor the associationof sexualdimorphism with


are unclear.


an association


tween habitat and mating systemdoes seem to be generallyupheld acrossbird species(Vehrencampand Bradbury1984), the associationbetween mating sys-