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Absence of Blood Parasites in the Red-Necked Nightjar

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Absence of Blood Parasites in the Red-Necked Nightjar J. Field Ornithol., 68(4):575-579 ABSENCE OF BLOOD PARASITES IN THE REI•-NECKFJ• NIGHTJAR M•uEI• G. FO•RO •I• Jose L. TEI•I• Estacitn Bioltgicade Do•ana, C.S.I.C., Avda. M a Luisa s/n, Pabelltndel Perd, 41013 Sevilla,Spain ALVAROGAJON Serviciode Diagntsticode Fauna Silvestre,Facultad de Veterinaria, Universidadde Zaragoza,Miguel Servet177 Zaragoza,Spain Abstract.--One-hundredand six Red-NeckedNighjars (Caprimulgusruficollis) were sampled in southern Spain to determine the incidence of haematozoainfection in relation to their expressionof sexual ornaments (i.e., white wing and tail patches). Birds were sampledjust after breeding and during molt, when blood parasitescould affect plumagecoloration, but none of these birds was infected by haematozoa.The low prevalenceof blood parasitesin this and other nightjarspecies cannot be satisfactorilyexplained at present. AUSENCIA DE PAR•ITOS SANGUiNEOS EN CAPRIMULGUS RUFICOLLIS Sinopsis.--Semuestrearon 106 individuosde Caprimulgusruficollis con el fin de determinar la posiblerelacitn entre la presenciade parfisitossangulneos y la expresi0nde ornamentos sexuales(manchas blancas en el ala y la cola) en estaespecie. Las avesfueron capturadas en Dofiana (sur de Espafia),coincidiendo con el final de la reproducci6ny la muda, finico momento en que podria verseafectada la coloraci6ndel plumaje. Sin embargo,ninguna de las avesestuvo parasitada. La bajaprevalencia de parfisitossanguineos en •sta y otrasespecies de chotacabrasno puede ser explicadaen basea la informacitn disponibleen la actualidad. In recent years, the incidence of avian blood parasitismhas received considerableattention from both evolutionistsand behavioral ecologists. Severalstudies have focusedon the possibledetrimental effectson host fitness (Allander and Bennett 1995, KorpimSki et al. 1993, RStti et al. 1993, Tella et al. 1996). Hamilton and Zuk (1982) have suggesteda re- lationship between a heritable resistanceto parasitesand bright colora- tion in birds. Individuals could signal differences in parasite loads through the expressionof bright colorations,and this would enable fe- males to choosemales of high genetic quality. As a part of a study on secondarysexual charactersof Red-Necked Nightjars (Caprimulgusruficollis) (Forero et al. 1995, Forero and Tella 1997), our aim wasto testif the expressionof ornaments(wing and tail patches) reflects the parasite burden of their hosts.We examined blood samplesfrom 106 nightjars captured from June-October in 1994 and 1995. This period coincideswith the molt (Forero and Tella, unpubl. data; Gargallo 1994), when parasitescould influence plumage coloration (Lozano 1994). The study area was Dofiana National Park in southern Spain, a wetland region of marshes,mediterranean forests, and coastal scrub.Birds were caughtduring nocturnal car transects,using a flashlight and a hand net (Forero et al. 1995). We sexed and aged the birds (through capture-recapturedata and molt patterns) into three age classes (juveniles,one-year-old birds [SY], and older birds [ASY]; Table 1). See 575 576] M. G. Foreroet al. j. FieldOrnithol. Autumn 1997 T•LF. 1. Number of Caprimulgusruficollis sampled at Dofiana National Park (1994-1995) by year, sex, and age. Sex Year Juvenile SY ASY Male 1994 7 18 10 1995 3 3 14 Female 1994 14 7 10 1995 3 5 12 Total 27 33 46 Gargallo(1994) and Forero et al. (1995) for more details.A drop of blood wasextracted from the brachialvein, smearedon an individuallylabelled slide, and air dried. We fixed smears with absolute methanol and stained them with Giemsa stain. One hundred oil microscopefields (1000x) were chosen from each smear in a line from one to the other border of the slide to avoid biases related to the thickness of the smear (Weatherhead and Bennett 1991). None of the 106 examinedbirds wasinfected by blood parasites.This cannot be attributed to samplingbias due to season,sex, or age differ- encesin parasitizationrates (Weatherheadand Bennett 1991, Tella et al. 1996). All birds were caught during and just after breeding (when blood parasitismis expectedto be the highestdue to breeding effort; Ots and Horak 1996, Richner et al. 1995), adults were molting and juveniles had recentlygrown feathers,and sampleswere equallydistributed with regard to sexand age. Our samplesizes were sufficientto establishthat the high variable expressionof sexualornaments in Red-NeckedNightjars (Forero et al. 1995, Forero and Tella 1997) is not correlatedwith prevalenceof blood parasites,at least in this population. Studiesshowing low blood parasitismin birds are scarceand the results are usuallyattributed to the lack of suitableornithophilic vectorsin some habitats (Bennett et al. 1992a, Little and Earl• 1995, Tella et al. 1996), an insufficienttime for the coevolutionof host, vector,and parasitein a biogeographicalregion (Earl• and Underhill 1993), or host behavior (Bennett et al. 1992b). Ours is the first surveyfor blood parasitesin the Red-NeckedNighjar, so we have not been able to test these hypotheses intraspecifically.Several species of blood parasiteshave been reported to parasitizeother nightjars (Bennett et al. 1982), but at a low rate. Out of 23 speciesexamined, only sevenwere infected,and the prevalences(num- ber of infected birds/sampled birds) were low (Table 2). Furthermore, Bennett et al. (1982) reported additional recordson the lack of parasit- ization in nine of ten nightjar species,but they did not mention the sample sizes. Overall, theseresults could be influenced by their generallysmall sam- ple sizes (Table 2). However, the variety of habitats and broad biogeo- graphical range of these samplednightjars (Table 2) make unlikely the first two hypothesesexplaining the absenceof blood parasites.Taking into Vol.68, No. 4 BloodParasites in Nightjars [577 oo••oo• 578] M. G. bbreroet al. j. FieldOrnithol. Autumn 1997 account the generalized absenceof blood parasitesin nightjars, two fac- tors could preclude thesespecies from parasitization(Combes 1991, Hart 1994). The physiologicalor immunological characteristicsof nightjars could constitutea barrier to parasites.On the other hand, the generally nocturnal and/or ground-dwellingbehavior of this group of birds could minimize encounterswith appropriate vectors. In fact, Greiner et al. (1975) found that ground-nestingspecies tend to have low frequencies of hematozoaninfections, presumably related to the preferenceof vectors to feeding in higher vegetationstrata. However, as Tarof et al. (1997) pointed out, parasitelife cicles,feeding activityof vectors,and their in- teraction with hostsare largely unknown for most avian species.A con- siderable research effort is required to find a clear explanation for the high variabilityof blood parasitizationrates shownby birds. ACKNOWLEDGMENTS We thank L. Garcia and personnelof Estaci6nBio16gica de Dofiana for their help in the field work, to G. E Bennett for his first guidance, and M. Brigham,J. A. Donfizar,J.J. Negro, G. E. Hill, C. R. Chandler, and an anonymousreferee for constructivecomments on this manuscript.I. de Bustamantehelped with the Englishtraslation. LITERATURE CITED ALLANDER,K., •tNr)G. E BENNETT.1995. Retardationof breeding onsetin Great Tits (Parus major)by blood parasites.Func. Ecol. 9:677-682. BENNETF,G. E, M. WHITEWAY,AND C. B. WOODWORTH-LYNAS.1982. Host-parasitecatalogue of the avian haematozoa.Occasional Papers in Biology,5. Memorial Univ. of Newfound- land, Newfoundland.243 pp. , g. MONTGOMERIE,AND G. SEUTIN.1992a. Scarcityof haematozoain birds breeding on the arctic tundra of North America. Condor 94:289-292. •, R. A. EARLg,AND M. A. PEIRCE.1992b. The Leucocytozoidaeof South AfYicanbirds: Caprimulgidae,Columbidae, Gruidae and Spheniscidae.OnderstepoortJ. Vet. Res.59: 229-234. •, R. A. EARLP•,H. Du TOIT, ANDE W. HUCHZERMEYER.1992c. A host-parasitecatalogue of the haematozoaof the sub-Saharanbirds. OnderstepoortJ. Vet. Res.59:1-73. COMBES,C. 1991. Evolutionof parasitelife cycles.Pp. 62-82, in C. A. Toft, A. Aeschlimann, and L. Bolis, eds. Parasite-hostassociations. Oxford University Press,Oxford, United Kingdom. 389 pp. EARLf•,R. A., G. E BENNETT,H. Du TOIT, D. DE SWARDT,ANr)J.J. HERHOLDT. 1991. Regional and seasonaldistribution of avian blood parasitesfrom northern South Ai?ica.S. Ai?.J. Wild. Res. 21:47-53. , AND L. G. UNDERHILL. 1993. Absence of haematozoa in some Charadriiformes breeding in the Taimyr Peninsula,Russia. Ardea 81:21-24. FORERO,M. G., J. L. TELLA,AND L. GARCIA.1995. Age related evolution of sexual dimor- phisinin the Red-neckedNightjar Caprimulgntsruficollis. J. orn. 136:447-451. , ANDJ. L. TELLA.1997. Sexualdimorphism, plumage variability and speciesdeter- mination in Nightjars: the need for further examination of the Nechisar Nightjar Ca- prirnulffussolala. Ibis 139:407-409. GARGALLO,G. 1994. Flight feather moult in the Red-neckedNightjar Caprimulgntsruficollis. J. Avian Biol. 25:119-124. GREINER,E. C., G. F. BENNETT, E. M. WHITE, AND R. F. COOMBS. 1975. Distribution of the avian hematozoaof North America.Can. J. Zool. 53:1762-1787. HAMILTON,W. D., ANDM. ZUK. 1982. Heritable true fitnessand bright birds:A role for parasites?Science 218:384-386. Vol.68, I•o. 4 BloodParasites in Nightjars [579 HART, B. L. 1994. Behaviouraldefense against parasites: interaction with parasiteinvasive- hess.Parasitology 109:139-151. KORPIM'AKI,E., H. HAKKARAINEN,AND G. F. BENNETT.1993. Blood parasitesand reproductive successof Tengmaln'Owls: Detrimental effects on femalesbut not on males?Func. Ecol. 7:420-426. LITTLE,R. M., ANDR. A. EARLI•. 1995. Sandgrouse(Pteroclidae) and
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