SEXUALDIFFERENCESINLEVELSOFBLOODCAROTENOIDSIN CIRLBUNTINGSEMBERIZACIRLUS

JORDIFIGUEROLA 1RICARDGUTIERREZ2

FiguerolaJ.R.Gutidrrez1998.Sexualdifferencesinlevelsofbloodca­

rotenoidsinCirlBuntingsEmberizacirlus..

Carotenoidsareresponsiblefor thebrightredandyellowplumageof birds. Thesesubstancescannotbesynthesisedbybirdsandmustbeobtained fromthedietandtransportedthrough thebloodtothefeathers.Thecarot­ enoidcontentofbloodwasmeasuredinCirlBuntingsEmberizacirlus, a specieswithsexuallydimorphicyellowplumage.Carotenoidconcentra­ tion,estimatedfromthecolorationofthe plasma,washigherinmalesthan infemalesandjuveniles.Thesedifferenceswereunrelatedtothegreaterin­ cidenceoffeathermoultinmales.Althoughthereasonsforthedifferences arenotunderstood,theresultsofthisandtwopreviousstudiessuggestthat, inspecieswithsexuallydichromaticplumage, the quantityofcarotenoids transportedin the bloodishigherinthemorebrightlycolouredsex.

Keywords:Emberizacirlus -carotenoids-passerines-plumagebrightness

-sexualdimorphism-sexualattractiveness

‘DepartamentdeBiologiaAnimal(Vertebrats),FacultatdeBiologia,Avda. Diagonal645,E-08028Barcelona,Spain; 2Departamentd’Agricultura, RamaderiaiPesca,GranVia612-614,E-08007Barcelona,Spain; Present address:DepartmentofAppliedBiology,EstaciónBiológicadeDoflana,

CSIC,Avda.MariaLuisas/n,41013,Spain,

Manyofthesexualdifferencesinplumagecolor­ ationofbirdsresult fromtheoccurrenceofcarot­ enoid-derivedcoloursinmales (Gray1996). Brightyellow,orange,redandvioletcolorations areproducedbytheaccumulationof carotenoids infeathers(Brush1978).Unlikeotherplumage pigmentssuchasmelanin,carotenoids cannotbe synthesisedby birds,andhaveto beobtained fromthebird’s diet(Brush1978,1990;Goodwin

1984).Afteringestion,carotenoids aretransported

fromthedigestivetract tothedevelopingfeathers viatheblood(Fox1962).Differentstudieshave shownthatfemalesprefertomatewithbrightly plumagedmales(i.e.Hill1991).Arelationship betweenthe quantityofcarotenoidsintheblood (estimatedfromthecolorationoftheplasma)and thebrightnessofthedevelopingplumage,aswell assexualdifferences incarotenoidconcentra­ tions,havebeenreportedintheHouse FinchCar-

podacusmexicanus(Hillet al. 1994). Asimilar sex-specificpatternofcarotenoidcontentinthe bloodoccursintheNorthernCardinalCardinalis cardinalis,alsoaspecieswitharedcarotenoid­ derivedandasexuallydimorphicplumage(Hill

1995a).Redandyellowplumagesseemtobepro­

ducedbydifferentkind ofcarotenoids(i.e.Brush

Power1976;HudonBrush1992),butupto nownostudyhasexaminedthesexualdifferences inbloodcarotenoidcontentinayellow-plumaged sexuallydimorphicspecies.Inthispaperwecom­ paretheplasmacolorationofmale,femaleand juvenileCirlBuntingEmberizacirlus,andpro­ videthefirstevidenceofsexualdifferencesin plasmacolorationinaspecieswithsexuallydi­ morphicyellow plumage.

TheCirlBuntingisa15.5cmlongpasserine

weighingabout25g.Itiswidely distributed throughout southern Europe and Africa

titles);thecontinuationofWieken Sneb Watervogels(1976-1981,vol.1-6;147titles);se­ riesdiscontinued

DeGraspieper (1982-1998, vol. 2-18(1); 556

titles)

Inall,thefilelists31,867titlesof17journals. Furtherinformationisavailablebythecompilers ofthisdiskette:

H.J.Lichtenbeld,Valparaisodreef47.3563VL Utrecfit,TheNetherlands

G. Speek,Zonegge16-16,6903GNZevenaar,T

heNetherlands

EDITORIALACKNOWLEDGEMENTS

Theeditorialteamwishestothankthe following colleagueswhorefereedmanuscriptsorotherwise providedhelpand advice:

M.Arts,A.V.Badyaev, P.Bednekoff,A.J.Bein­ tema,A.Berg,R.M.Bevan,E.Bignal,R.G.Bijl­ sma,C.Both,H.Boyd,J.S.Bradley,R.Brandl,

M.Brinkhof,M.E.Brown,B.Bruderer, L.W. Bruinzeel, C. Carey,J.Carroll,A.J.Cave,B.Eb­ binge, M. Engelmoer,Y.Ezaki, J.R. Faaborg, A.D.Fox,J.A.vanFraneker,W.R.Fraser,H.Gal­ braith,A.Gardarsson,S.M.Gatesy,J.Gonzalez- Soils,G.E.Hill,H.J.Hill,P.A.Hockey,H.Hoi, H.Hdtker,A.I.Houston,J.Hudson,B.Kempe­

naers,E.Korpimaki,A.Kvist,A.Lindstrom,M.

Loonen,E.Meelis,T.Meijer,P.Monaghan, J.J. Negro, G.Nehis,j.A.Nillson, B.Nolet,K.Nor­

ris,M.Owen,W.J.Peach,I.Pen,T.Piersma,J. Prop,K.Reid,G.Ruxton,H.Schekkerman,T. Slagsvold,A.L.Spaans,R.A.Stiliman,I.Tinber­ gen,I. vander Veen,S.Verhulst,H.Visser,I.Wa­ nink,P.Wiersma,B.Zonfrillo.Inaddition,we wantto thankGeryEngelmoer-Hoekstra (layout; Computekst,Groningen),DickVisser(artwork; GroningenUniversity),andJosZwarts(artwork) fortheverypleasantco-operation andhighqua­ lityoftheirworkforArdea. ArieSpaans(IBN­ DLO,Wageningen)kindlycorrectedthe‘Samen­ vattingen’.RobDapper(NIOZ)wasinstrumental inthemaintenanceandupdatesoftheNOUhome page.

(SnowPerrins1997).Maleshavehighlyvari­ ableplumages,withvariousextents ofyellow featheringonbreast,bellyandhead.Theplumage offemales,andespeciallyjuvenilesofbothsexes, islessconspicuous,withmainlylightgreyand brownunderparts,andsomelightyellowpartsin adultfemales(Gutiérrez 1997).

CirlBuntingsweretrappedwithmist-netsat

Juncosa (Lleida, northeast Spain, 41°23N,

00°46’E)inJuly-August1996andAugust1997. Individual4swereringed,wing-lengthandbody-

masswiremeasured,and moultstageand differ­

entplumagecharacteristicsofthemaleswerere­ corded.Bloodwasextractedbypuncturingthe brachIalvein,andrecoveredinaheparinizedcap­ illarytubeandonamicroscopeslide.Aftercen­ trifugationofthecapillarytubes,thethreebasic characteristics of colour, were later estimated (light,chromaandhue)oftheplasmaweremeas­ uredusing a Minolta CR200 colorimeter.The colorimetersendsastandardisedlightflashover thetableusedtoscoretheplasmaandanalyses thereflectedlight(seeSenaretal.1998).Huerep­ resentsthewavelengthofacolourandismeas­ uredindegreesofacirclewith redat0degrees, yellowat90degrees,greenat180degrees,blue at270degreesandcompletingthecirclewith red at 360 degrees. Lightness corresponds to the physicallightintensity,tothecolour sensation producedoverascaleofgreys.Lightnesswas measuredonaslidingscalewith0forblackto

100forwhite.Chromaispositively correlatedto colourmonochromatism(Kuppers1996),andis

measuredaspercentagesaturationrangingfrom0 forwhiteto100forpurecolour.

Bloodsampleswereobtainedfrom93indi­

viduals(Table1).Nodifferencesweredetectedin lightnessandhuevaluesofplasma between young,adultmales andadult females(Kruskall­ Wallistest:light,H393=2.61,P=0.27;hue,H393

=0.03,P=0.93).Highlysignificantdifferences

werefoundinthechromavaluesofplasma(H393

=24.02,P0.0001). Thesedifferencesweredue

toahigherchromavaluesofmales comparedto females (Mann-WhitneyU-test,Z229=-2.99,P

0.003)andjuveniles(Z229 -4.80,P0.0001),

but no differencesoccurred betweenjuveniles andfemales(Z229=-0.42,P=0.68).Ahigher proportion ofmalesthanfemaleswereundergo­ ingbodymoult(F-Fisher,P=0.001).Nodiffer­ enceswerefoundinplasmacoloration inmoult­ ingandnon-moultingmales(chroma:Z175=-

1.14,P=0.25;light:Z175 -0.51,P=0.61;hue:

Z175=-0.12, P=0.91). Non-moultingmales

showedhigherchromascoresthannon-moulting

females(Z58=-2.49,P=0.01),althoughnodif­ ferenceswerefoundfortheothertwovariables (light:Z58=-0.44,P=0.66;hue:Z58 -0.44,P

0.66).Comparisonsformoultingandnon-moult­

ingfemales,andformoultingmalesandfemales werenotcomputedduetothesmallnumberof moultingfemalescaptured.

The basicassumptionof thisstudyis that

plasmacolorationreflectscarotenoidcontent(see

BrushJohnson1976;Brush1990;Hilletal.

1994,foradiscussionofitsvalidity).Onthisba­

Table1. Aspectofplasma colour(seetext)inunsexedjuvenile, andadultmaleandfemale,CirlBuntings (ave­

rages±l SD).

n / Lightness / Hue / Chroma
Juveniles / 62 / 83.4±3.9 / 87.7± / 7.0 / 47.2±6.2
Males / 22 / 82.5±3.3 / 87.9± / 4.4 / 57.1±8.2
non-moulting / 5 / 82.7 ±3.5 / 88.5 ± / 5.0 / 61.3±8.5
moulting / 17 / 82.4±3.4 / 87.8 ± / 4.3 / 55.9±7.9
Females / 9 / 81.5±7.3 / 83.5 ± / 15.3 / 47.1±8.1
nonmoulting / 8 / 80.9±7.5 / 82.2± / 15.7 / 48.7±7.0
moulting / 1 / 86.1 / 94.3 / 34.4

cludedonthemailinglistforbrochuresandfor informationonthe23rlCongressshouldbesent totheSecretary-GeneralortheAssistantSecre­ tary-General.

Allinquiriesaboutthescientificprogramof the23Mcongress,aswellascommentsandsug­ gestionsforthe generalprogram,plenarylec­ tures,andsymposia shouldbesenttoDrFer­ nandoSpina,Chair,ScientificProgramCommit­ tee,23’ Congress,InstitutoNazionaleperIa FaunaSelvatica,ViaCa’Fomacetta9,1-40064

OzzanoEniila (BO),Italy,phone+39516512

111,fax+3951796628,e-mail<infsioc@iper­

bole.bologna.it>.

General questionsandcommentsshouldbe senttoDrWalterJ.Bock,Presidentofthe23td Congress(addressseebelow).Inquiriesaboutthe International Ornithological Committee should besenttoDrDominiqueG.Homberger,Secre­ taryoftheInternationalOrnithologicalCommit­ tee, DepartmentofBiologicalSciences,508Life SciencesBuilding,Louisiana StateUniversity, BatonRouge,LA70803-1715,USA,phone+1

504388 1747, fax+15043882597,e-mail <zod­ >.Informationaboutthe IOCcan also be obtained from a new home page at

<

whichiscurrentlyunderconstruction.

DrWalterJ. Bock,Presidentofthe23Congress, Department ofBiologicalSciences,Columbia University,l200AmsterdamAvenue,MailBox

5521,NewYork,NY10027-7004,USA,phone

+12128544487,fax12128658246,e-mail

<>.

COMPLETETABLEOFCONTENTSOF

ARDEA AVAILABLEONDISKETTE

AcompletetableofcontentsofARDEAisnow availableondiskette(DbaseIV format),inclu­ dingtitlesofarticles,shortnotes,notesandnoti­ ces,bookreviews,necrologiesandintroductions,

3455titlesinall:1912-1998,vols. 1(1)to86(1). ARDEAwasthecontinuationof‘Jaarboekjesder NederlandscheOrnithologischeVereeniging’ (An­ nualreportsofthe NetherlandsOrnithologists’ Organisation),whichwasissuedbetween 1904 and1911,andthetableofcontentsofwhichisin­ cludedon thediskette(152titles).

Theindexfilewasissuedforthe first time in December1998,andwillbefollowed byupdates, whichcanbeorderedatanymoment,convenient fortheuser.Thedisketteisissuedatapriceof Dfl.50,=percopy(Dii.45,=formembersofthe NOU,institutesandlibrariesDfl.250,=).Updates areavailableforDfl.25,=(Dfi.22,50/Dfl.100,=) percopy.Thediskettedoesnotincludeasearch engine,foritisassumedthatthedatawillbeim­ portedintoreferencesystemsofone’s own choice.BesidestheindexofARDEAand ofJaar­ boekjesderN.O.V.,thediskettecontainsacom­ pleteindexof:

DrentseVogels(1985-1998, vol.1-11;151titles) DutchBirding(1979-1998,vol.1-20(5);2290titles) Jaarberichtder ‘Club’ (1911-1928,vol.1-17;

684titles)

Orgaander‘Club’(1928-1933,vol.1-6,868titles) Limosa(1934-1998, vol.7-71(3);3845titles);the continuationofOrgaander‘Club’

RingingMigration(1975-1998,vol.1-19(1);

469titles)

SlechtvalkNieuwsbrief(1995-1998,vol.1-4(1);

56titles)

Sula(1987-1998,vol.1-12(2);797titles)

DeTakkeling(1993-1998, vol.1-6(3);326titles)

Vanellus(1948-1998, vol.1-51(5);6621titles)

OphetVinkentouw(1963-1998,vol.1-87;1311 titles)

WiekenSneb(1951-1956,vol.1-4;317titles)

Het Vogeljaar(1957-1998, vol. 5-46(5); 9688

sis,the carotenoidcontentofmalep’asmawas higherthanthatoffemalesandjuveniles.These differencescould berelatedto thehighercaroten­ oidcontentofmaleplumage,andtheneedfor highercarotenoidlevelstoproducemorebrightly colouredplumage(Hilleta!.1994).Thatthepro­ portionofmoulting malesinoursamplewashig­ herthanthatoffemalescouldhavebiasedour es­ timatesofplasma carotenoidsiftheingestionof carotenoids,orthedegreetowhichtheywerere­

tainedintl1e blood,increasedduringthemoult

(seeHill1995b).However,therewasnodiffer­

enceinplasmacolorationbetweenmoulting and non-moultingmales.Sexualdifferencesinplas­ ma hroma remained statistically significant whenonlythedatafromnonmoultingbirdswere considered.Additionally,juvenilebirdsthatwere stillgrowingtheirfirstfeathersinsomepartsof thebodyhadlowercarotenoid levelsthanmales. Thissuggeststhatthepattemsfoundarenotan artifactofsexualdifferencesinmoultinitiation date.

Differencesin plasmacoloration havebeen

alsoreportedin theHouseFinchand theNorthern Cardinal(Hill1995a).Thesestudiessuggestthat therearesexualdifferencesinthequantityofca­ rotenoidstransportedintheblood.Thecausesof thesedifferencesarenotunderstood,butseveral non-mutually exclusive hypotheses have been proposed.Forexample,malesandfemalescould differinthedensityofcarotenoid-carryingpro­ teinsintheblood (seeTrams1969).Sexualdiffer­ ences in foraging behaviour or diet selection couldalsoresultinahighercarotenoidintakein males(Hill1992).Ourdatadonotallowusto de­ terminethereasonsforthispattern,whichhasso farbeeninvestigatedinveryfewspecies.Investi­ gationofcarotenoidlevelsin non-sexuallydimor­ phicspeciescould provideadditionalinforma­ tion.Ifthereporteddifferenceswerereallyre­ latedtoplumageneedsandnottootherecological factorsdifferingbetweensexes,wewouldexpect tofindnodifferencesintheplasma colorationof non-dimorphicspecies.However,Bortolottietat (1996)haverecentlyreportedtheexistenceofsex­ ualdifferences in theconcentration of plasma

caretonoidsinLoggerheadShrikesLaniusludovi­ cianus,aspecieswithoutcarotenoid derived colorations.

DrJuanCarlosSenar(MuseudeZoologiade

Barcelona)kindlyallowedustousethecolorime­ ter.ManelPomarolprovidedessentialsupportfor thework.TheArbonesfamilyprovidedlodging facilitiesandgreatlyimproved ourforagingsuc­ cess.FranciscoCerdàandMontserrat Panyella collaboratedinthefieldwork.Thecommentsof GeoffHill,JocelynHudonand TheunisPiersma inprovedanearlierversionofthismanuscript.

REFERENCES

BortotottiG.R.,1.1.Negro,J.L.Telia,TA.Marchant D.M.Bird1996.Sexualdichromatisminbirdsin­ dependentof diet,parasitesandandrogens.Proc. R.Soc.Lond.B263:1171-1176.

BrushA.H.1978.Avianpigmentation.In:BrushD.E. (ed.).Chemicalzoology,10:141-161.Academic

Press,NewYork.

BrushA.H.1990.Metabolismofcarotenoidpigments inbirds.Fed.Am.Soc.Exp.Biol.J.4:2969-2977.

BrushA.H.N.K.Johnson1976.Theevolutionofco­

lordifferencesbetweenNashvilleandVirginia’s

warblers.Condor78:412-414.

FoxD.L.1962.Metabolicfractionation,storage,and

displayofcarotenoid pigments byflamingoes. Comp.Biochem.Physiol.6:1-40.

GoodwinT.W.1984.Thebiochemistryofcarotenoids.

Vol.2,Animals,2nded.ChapmanandHall,New

York.

GrayD.A.1996.Carotenoidsand sexualdichromatism

inNorthAmericanpasserinebirds.Am.Nat.148:

453-480.

Gutidrrez R.1997.Identification ofCirlBuntings.

Alula3:174-180.

HillG.E.1991.Plumagecolorationisasexuallyse­

lectedindicator ofmalequality.Nature350:337-

339.

HillG.E.1992.Proximatebasisofvariationincaroten­

oid pigmentationinmaleHouseFinches.Auk109:

1-12.

HillG.E.1995a.Interspecificvariationinplasmahue

inrelationtocarotenoidplumagepigmentation.

Auk112:1054-1057.

HillG.E.1995b.Seasonalvariationincirculatingca­

rotenoidpigmentsintheHouseFinch.Auk112:

1057-1061.

HillG.E., R.Montgomerie,C.Y.Inouye3. Dale

sitism,parentage,extra-paircopulations(includ­ ingnecrophilia),ectoparasites,predators,crèches, informationcentres, kieptoparasitism,andnon- breeding.However,itisabookfullofanecdotes rather than insightful scientificobservations, a book of how a fairly self-confident academic trainsandleadshisratheranonymousassistants andstudentsinthefield.Thereisnodoubt,how­ ever,thattheCliffSwallows,intheirtightlypac­ kedcoloniesinculvertsformanidealstudyob­ jectandthatthemanyyearsofstudyinNebraska havebeanexceptionallyproductive.Goodnews isthe factthatthereisan extensivesubjectindex inthebackofthispaperback,sothatinteresting subjectscanbeeasily(re-)located. (CJC)

SECONDMEETINGEUROPEAN ORNITHOLOGISTSUNION GDANSK(POLAND),

15-18SEPTEMBER1999

TheEuropeanOrnithologistsUnionwas estab­ lishedin1997.TheaimoftheUnionisthead­ vancementofornithologyandthepromotionof thescientificstudyofbirds amongornithologists withinEurope.ThefirstEOUMeetingwasheld inBologna,Italy.Therewasnearly250partici­ pantsfrom28countries(mostlyfrom thewest partofEurope).The aimoftheSecond Meetingis tocreateanopportunityforareallylargenumber ofornithologistsfromthewholeofEuropeto ex­ changethemostrecent resultsof theirworkin differenttopics,tomakeitpossibletodiscussdif­ ferentaspects oftheirresearch,tomake contacts between centralleasternscientistsandwestern onesaseffectiveaspossible. Thusitwasagreed toholdthis conferenceinPolandtoenablemany scientiststocomefromcentralandeasternpartof Europe.Theconferencewillincludethreedaysof meetingsandwillconsistof:-plenarysessions (withinvitedkeyspeakers)- symposia/work­ shops -postersessionswithspecialtimeforpre­ sentationanddiscussion. Therewillbepost-con­ ferenceexcursions.The conferencelanguagewill beEnglish,and theproceedingswillbepublished

inEnglish.

Thesecondconferencewilltakeplace in Gdansk,an oldtownsituatedonthecoastofBal­ ticSeainthenorthpartofPoland.Itwillbeheld attheUniversityofGdansk.Thevenue offers goodfacilitiesformeetings,isnotfarfromthe centreofthetown,easilyaccessiblebycar,tram orsubway.Theconferencefee($120),includes conferencedocumentsandproceedings.

Organisingcommittee:University ofGdansk, BirdMigrationResearchStation,Przebendowo

84-210Choczewo,Poland,phone+4858676 32

20, fax +4858676 32 65, e-mail <eou.meet­

>.

23RDINTERNATIONALORNITHO­ LOGICALCONGRESSINBEUING, AUGUST2002

Atthe22ndInternationalOrnithologicalCon­

gressheldinDurban,SouthAfrica,16-23August

1998,theInternationalOrnithologicalCommittee votedtoaccepttheinvitationfromtheChinese ornithologiststohostthe23rdInternationalOrni­ thologicalCongressinBeijing,China,on11-17

August2002.Informationcanbeobtainedviaem­

ail<>,viatheinternetat

< the22uidcongressat <

Itwillbepossibletoregisterandtosubmitab­

stractsviatheinternet.Thefollowingarecontact addressesofpeopleresponsibleforthe23Con­ gressinBeijing.ProfessorXuWeishu,Secretary- Generalofthe23’’ Congress,BeijingNatural HistoryMuseum,1-1-302,BeijingScienceand TechnologyCommissionApt.,Balizhuang, Hai­ dianDistrict,Beijing100037,China,phonefax:

+861068465605,e-mail<>.The HonorableLiuFeng,AssistantSecretary-General ofthe23k’Congress,ChinaInternationalConfer­ enceCenterforScienceandTechnology,Xuey­ uanNanRoad,Beijing100081,China,phone+

861062174952;fax+861062180142,e-mail

<>.Requeststobein-

248ARDEA86(2),1998

1994.Influence ofdietarycarotenoids onplasma andplumagecolourinthehousefinch:intra-and intersexualvariation.Funct. Ecol.8: 343-350.

HudonJ. &A.H.Brush1992.Identificationofcaroten­

oid pigmentsin birds.MethodsEnzymol. 213:

312-321.

KUppersH.1996.Atlasdeloscolores.Blume,Barce­

lona.

SenarJ.C.,J.Domênech &M.J.Conroy1998. Sexing Serinfledgingsbyplumagecolourandmorpho­ metric variables.OrnisSvecica 8:inpress.

Snow D.W.C.M.Perrins1997.TheBirdsofthe

Wesern Palearctic.ConciseEdition.OxfordUn­

sivitdyiPress,Oxford.

TramsE.1969.Carotenoidtransportintheplasmaof theScarletIbis(Eudocimusruber).Comp.Bio­ chem.Physiol.28:1177-1184.

SAMENVATTING

Carotenenzijnplantaardigebiologische kleurstoffen diemeestalverantwoordelijkzijnvoordefellerode, oranjeengelekleurenvanvogels.Vogelskunnencaro­ teennietselfaanmaken,endestoffenmoetendusmet hetvoedselwordenopgenomen. Caroteenwordtnaar groeiendeverengetransporteerdviadebloedbaan. In dezestudiewerdencaroteenconcentratiesinhetbloed geschat aandehandvandematevankleuringvanhet bloedplasmabij deinZuid-Europaalgemeenvoorko­ mende CirlgorsEmberizacirlus.Bijdezegorzensoort heefthetmannetjeeengelerverenkleeddanhetvrouw­ tje.Inderdaadwas hetbloedplasma vanmannetjesge­ lerdandatvande vaal-gekleurdevrouwtjesenjuvenie­ len.DeresultatenvoordegeelgekleurdeCirlgorszijn inovereenstemmingmeteerderestudiesaanroodge­ kleurdezangvogels:bijdichromatische vogelsoorten wordt inhetbloedvandefelgekleurdesexehetmeeste caroteen getransporteerd. (TP)

Received5January1998, accepted14July1998

Correspondingeditor:TheunisPiersma

Browsingthroughthis fascinatingbookgives the impressionthatmostbirderswerebearded men.Thereis someexceptions,likeEvelynBax­ ler (1879-1959) and Leonora Rintoul (1875-

1953),EmilieSnethiage(1868-1929),Elizabeth Kozlova(1892-1975) anda few other women whohaveaprominentplacein thisbook.All thesepeoplearedescribedandwhenpossiblede­ pictedinabookthatreadslikeanovel.Thisisa bookaboutthenameswe so oftenseeattachedto birdnames:Linnaeus,Hutton,Pleske,Temminck, Steller, MtiGillivray, Bulwer,Franklin,and Gouldtonameafew.Chaptersaredevotedto the birdartists(e.g.JohnGould),armyofficers (e.g. RichardMeinertzhagen),clergymenand mission­ aries(e.g.ReverendFrancis Jourdain)andthe professionalfieldcollecters(e.g.AlfredRussell Wallace).Itdescribestheirmethodsofkilling, skinningandpreparing,labellingandnote taking, andinparticularthedifficultiesinthefieldand the severeproblemsinvolvedwithgettingthe specimenshome.Ifallwentfine, therewere still themuseumdisasterslikefireorinsectsruining vastcollections.

Thewriters,Barbara andRichardMearns, mustbecongratulatedwiththeirwork.Onthe dustcoverwereadthat“theystrongly believethat theimportantroleofthescientific collectors shouldnotbeforgottenandthattheir collections shouldnotbeundervalued.”Thisbookisaseri­ ouscontributiontowardsachievingjustthat.

Kees(C.].)Camphuysen

NetherlandsInstituteforSeaResearch, P0.Box59,1790ABDenBurg,Texel, The Netherlands

ALSORECEIVED

ERRITZOEJ.H.B. ERRITZOE1998.-Pittas oftheWorld.AMonographonthePittaFamily- TheLutterworthPress,Cambridge.ISBN07188

29611.Hardback,2O’lpp,32colourplates,some

linedrawingsand B&Wphotographs, numerous maps. Price: £30.=(f94=).

Onlytwoyearsafterthepublicationofanidentifi­ cationguideonPittas,Broadbillsand Asities (LambertWoodcock1996, PicaPress,Sussex), thismonographonthePittafamilyhas beenpub­ lished.Afteraverydetailed chapteronthePitta familiyanditsplacewithin thePasseriformes thereare30speciesaccounts,akeytothesyno­ nymsandnewproposed names,aglossary, 5ap­ pendices and an impressive bibliography(ca.

1300references). AnyoneinterestedinPittas shouldhavethisbook.Thetextsareverywellor­ ganised,withclearheadingsandsubdivisions,in­ formationonrecentrecords (after1975),habitat, vocalisations,foodandfeedingbehaviour,breed­ ingbiology, moult,biometrics,andworldwide museumholdingsofskins.Perhapsnot expected inabooklikethisaresectionsonPittasincaptiv­ ity,whichincludeincredibledetailssuchasthe survivalofindividualsonadietofWayme dog foodwithvitaminsandmixedwithfinelydiced three-day-old mice, apparentlybecause“Pittas arenotbirdsforthebeginnerinaviculture.”This bookisamust,because ifitisn’tinthebook,or cannotbefoundusingthespectacular bibliogra­ phy, itiseitherunknownornotworthbeing known. (CJC)

BROWNC.R.1998.-SwallowSummer-Univer­

sityofNebraskaPress,Lincoln.ISBN0-8032-

6145-4. Paperback, 371pp, 26 B&W photo­

graphs.2maps.Price$16.95(f31.50).

Inthisbook,bestdescribedasanovelindiary format,CharlesBrown,oneoftheco-authorsof ColonialilyintheCliffSwallow(Universityof ChicagoPress,1996),leadsthereaderthrough onefieldseason,infactthe15thconsecutivesea­ sonofstudy.ThebookstartsonMay8, 1995(the startofanewfieldseason)andendsonJuly27, whenthe expedition packs up to movehome again.Itdescribesinaratherentertainingmanner thefieldworkatcoloniesofCliffSwallowsPet­ rochelidonpyrrhonotanear Ogallala insouth­ westernNebraska,USA.Althoughadescription offieldworkratherthanstudyresults,several interestingbiologicalaspectsareaddressed,such ascolonyformation,sitefaithfulness,broodpara

249

ESSAYSONRECENTPUBLICATIONS

(ANINVITATION)

In addition to the ‘ordinary’book reviews,in

1999,Ardea hasinauguratedanew‘bookreview’ sectioncalled‘Essays on recent publications’,to publishin-depthcriticalreviews ofimportant booksinornithology.Newornithologicalbooks tendtoreceiverather cursorytreatmentinthe bookreviewsectionsofthemajorjournalsand, inspiredby,theextensivebookreviewspublished byEvolution,Ardea nowaimstoprovideoppor­ tunitiesforcriticalreaderstopresentlastingeval­ uationsofnewpublicationsintheirfieldofinter­ estandexpertise.

Weinvitereviewerstosubmitessaysonchal­ lengingbookspublishedinrecentyears.Essays shouldbeformattedasshortpublications andin­ cludea title,alist ofreferences andthenameand addressoftheauthor.The titleandpublicationde­ tails ofthebookunder reviewwillbe givenasa footnotetotheessay.Ardea aimstopublish‘Es­ says on recent publications’assoonaspossible aftersubmission.Manuscriptswillbereviewed bymembersoftheeditorialteam,with outside helpifnecessary.Twenty-fivereprintswillbeof­ feredfreeofcharge.

BOOKREVIEWS

BookreviewsinArdea arewritten oninvitation. Booksreviewedbelowwere receivedfromthe publishersandwillbekeptinthelibraryofthe NetherlandsOrnithologists’Unionforconsulta­ tion byNOUmembers.Thereviewsaremeantto alert readerstonewbooksandtogiveanidea whether ornotthebookisworthreadingorpur­ chasing.Reviewsshouldthereforebedescriptive, butalsohighlightboththe strengthsandweak­ nesses ofthebook.Reviewsshouldbeinforma­ tive,interestingandpertinent.Technicalinforma­ tion onthebooksunderreviewincludespub­ lisher, ISBNnumber(ifavailable),anindication oftheprice(andf),thenumberofpages,fig­ ures,tablesandphotographs.Prices,ifnotmdi-

catedbythepublishers,arebasedonrecentcata­ loguesofmajorbooksellers.Englishpounds () havebeenconvertedintoDutchguilders(f)ata rateof1:3.13; 1$=f1.86,1DM=f1.13.

BYRKJEDALI.D.B.A.THOMPSON1998- Tundra Plovers:theEurasian,PacificandAmeri­ canGoldenPloversandGrey Plover -T.A.D. Poyser,London.ISBN0-85661-109-3.Hardback,

422pp.,onecolourplate,57blackwhite pla­ tes, 89 figures, and 16 tables. Price £ 27.95 (f87.50).

“Mostofallthisisabookbyandforpeoplewho lovethebirdsandtheirhaunts.Youmayhave beenon alargecoastal mudflat, in themidstofan agriculturallandscape.orevenwanderingacross hugetundraplainsclosetotheNorthPole.Itdoes notreally matter,forsolongas youhavewatched atundraplover, andthoughtaboutitsbehaviour, ecologyandappearance,youarewithus.” This quotefromthebeginningofIngvarByrkjedaland DesThompson’s impressivebookonthefour speciesofPluvialis,thetundraplovers,isspoton. Itisadeeplyinspiredbook,thelabouroflove fromagiftedNorwegianlScottishduo.Thebook isbeautifulforits manyoriginaldrawingsbyIng­ varByrkjedalandforitstext,anditprovidesan incredible,worldwide,compilationofdataonthe birds’morphology,distribution,populationsizes, breedingschedules,breedingbehaviour,migra­ tionpatternsandschedulesanddiets.Browsing thebookgivesoneafeelfortheecological and behaviouraltraitsthatunitethetundraploversas agenus,butalsoafeelforwhat setseachofthe speciesapartfromtheothers;itimpressesone withtheaccumulatedamountofknowledgeon tundraploverswhilstatthesametimeconveying thehugegapsinourunderstandingofthisworld­ wideandenigmaticgroupofbirds.

Thesceneis setintheforewordbyDerekRat­ cliffe,thedescriptionsoftheauthors’plover-la­ denpersonalhistories,asurveyofthestudyareas, andareviewoftheploverandlapwingfamily Charadriidae.Itisclearthatthetundraploversare

ARDEAt(2),I99

ershipofover50,000skinswasnotunusual.Wil­ hamHenry Phelps(1875-1965),withhelpofhis sonBillyandhisdaughterinlawKathleencol­ lected76,000skins,mainlyfromVenezuela(Co­

.lecciónOrnitologiaPhelpsinSabena Grande). LouisBishopofConnecticut(1865-1950)person­ allyshotabout40%ofthe53,000skinsinhiscol­ lection,manyofthemwhileonvacation.Count HansvonBerlepschfromHessen(1850-1915),a hummingbird specialist, had a collection of

55,000s4inswithnofewerthan6000ofthesebe­

inghuirimingbirds.Thebirdsandbutterfliesthat

WalterRothschildinitiallyhousedin hisbedroom andagardenshedgraduallyswelledto280,000 bird skins,2400mountedbirds,200,000birds’ eggs,3400mammals(mounts,skinsandskulls),

2,250,000lepidoptera,300,000beetlesand other invertebrates,reptilesandfish(laterleading tothe establishmentofTringMuseuminwhichtohouse them).Therewereseveralmoreofthose‘veryac­ tive’collectors,convenientlysummarizedinthe chapteron‘TheGreatAccumulators’.

Obviously, therewasnotjustkillingforcol­ lectionsandthisbookexplorestheharvests of birdsand theireggs insimilardetail.Birdsasver­ min,birdsbeingdecorative anduseful,birdskil­ ledforsport,birdsasfood.Asonedramaticillus­ trationthereisthispictureoftheLaysanAlba­ trosses(p.14),beingrobbedoftheireggsonLay- san(Hawaiianislands)in1906forcompaniesthat usedalbumeninthemanufactureofphotographic paper. Thebirdsstandaroundintheircolony whichhasjustbeenclearedofeggs.Heapsof eggsarevisible,piledupinwagonsandwheel­ barrows,withthecollectorstakingarest.Among thecollectorsisasmallgirlvisible,TillieSchiem­ mer.Tillielivedontheislandwithherfatherand she,despitethelocalpopulationofafew millions ofseabirds,demandedpets. Father introduced Englishrabbits,GuineaPigs andBelgianHares. Tillie’spetsbredsoproficallythatherfatherre­ leasedthem.Threeyearslater,Japanesefeather hunterslanded(illegally)ontheislandandkilled

200,000seabirds,mainlyalbatrosses.Thecolo­

nieswouldprobablyhaverecoveredfromtherob­

bery,buttheintroducedherbivores,Tillie’spets,

hadcausedecologicaldisaster(erosion).Withthe exceptionofSchlemmer’stobaccopatch,every greenleafontheislandwasconsumed. mci­ dently,theendemicMillerbirds,LaysanHoney- creepersandLaysanRailswerewipedout.Thisis justone‘little’drama,orratherafewincidents leadingtodisaster,ofwhichthisbookgivesso manyexamples.

Intheappendixofthisbook,69museumsare

listed, housing atleast7,362,574 skins.Since

1600,90speciesand60subspeciesofbirdshave vanishedfromthe faceoftheearth,thevastma­ jorityofthembeingislandendemics. Collectors aresometimesblamedforthe extinctionofbirds, mostnotablysoincaseoftheDodoandtheGreat Auk.However,thiswasalsotheerawherethe rapidlyadvancingcivilizationtransformed the worldandwherethelossofnatural habitat broughtsomanyspeciestoextinction.Habitatde­ structionissomethingatwhichwe arestillvery good.Whilewenowgenerallyfind itmorally wrong toharmor killbirds,wecannotstopclear­ ingawaytropicalrainforests.Wenowtry andre­ habilitatetheoddoiledseabirdwashed ashore, shockedasweare,butatthesametimewefailto stopchronicoilpollutionofourseasandwecon­ tinueoverfishing.So,beforethe fingerispointed totheold collectors,we shouldlookat ourselves firstandperhapstryandappreciatetheknowl­ edgethesepeoplebrought forwardwiththeir work.

Theimportanceof‘old’and‘new’bird collec­

tionsisdiscussedinthebook,aswellas current methodsofaddingtomuseum bird collections (mainly‘passive’collecting).Thecollectionsare very valuableindeed. Zoologistsdependonspec­ imencollectionsasthebasisofsystemsforclas­ sification.Anobviousfunctionofskincollections istheircontributiontotheidentificationprocess. Thebrilliantpaintingsinmodernhandbooksand fieldguides,whichwetendtotakeforgranted, areoftenlargelybasedon carefulstudiesofskins inthemuseum.Furtherreasonsinthebookaregi­ ven astowhyweshouldappreciate theskincol­ lectionsworldwide,althoughsomeargumentsare ratherfarfetched.

ARDEAO5(2),1998

agroupanthrirown,staudiugquitoapartfrom boththrChar-ads’iua-ptovrrsasthrVanellun-lap­ wiugs.Thryarruuiqurforthrirnorthrrnbrood­ inghabitsauditisthought thatthrapprarancrof icragrs,2mittiouyawsago,mayhavrhad much todowiththrorigins ofPtacialis.Thrrarty chaptrrs contain rxcrrdingty dataitad dasccip­ tiousofthaptumagrsandmouttsofrachatthr fourspacirs(audadiscoursaouthrrrcrutdis­ tiuctionbrtwrrnthrAmrricanaudPacificGot­

druPtos’rs P.dominicaaudfulca).ttatsacon­

taiusanorigtnatctadisticanatytis ofthagroup

basad00ptumagacharactarisfics.Tharaisa totto sayforthacouctusionthatthaGrayPtovar istha siftarspaciastotha gotdeuptovars,audthattha EurasianGotdanPtovaaisthasistarspaciatto tha tassargotdanptoveas,butmotecutardataoaruow uaadadtocoaroboratathis.Thaauthorsdooat comaupwithadafinisivastatamantanthamis­ tancaoftwodistinctsuhspacias ofEurasianGot- danPtovars(thasauthamapsicas-iaandthanorth- amatttjrons).ThairdatafarNorwayshowactina inthaaxtentofcantrasfingbraadingptumaga,and thara appaarstohaintriguingassociationsha­ twaanhabitatcharactaaisfics,timingofsnowwatt andptumagatraits.Thisisjustanaofthamany topicsthatthabookindantifiasasbaingripafar fuatharandfmitfutanquicy!

Tharaisabigbtackoffourchaptarsthatra­ viaw diffarantaspactsofbraading:distribution andpopulationsiaa,baaadingschadutas,andso­ ctatandparantatbahavioucAlt ofthatopicsma basadonnawsumsoariasofaxistingdataandan unpubtishad absarvatiansbythaauthors,aspa­ ciattybyEyrkjadatwhohasaccumutatad axpan­ anca withattfour ptas’aasspaciasin diffarant partsofNorway,anthaYamatPaninsutatnEus­ siaandinthaCanadianArctic.

Thachapsar andistributioncomasupwith shockingcanctusiousaboutprobabtapopulation siaasofaachofsha spacias.Parallfourspacias Iharaviawsofbaaadingdansitsasincombination withastimalasofavailablahabitattaadtopopula­ tionastimalasofImillionpairsormara,twoto

19limasthacusrantlyavailablapopulationsiza astimatas.Thisdoasnotnacassaritymaanthattha

populations arefaring watt!AmaricanGatdaa Plavarssaamtohas’asuccassfuttyracos’aradfrom IhaonslaughtsofthamarkathuntinginUnitad fitatasintha19thCantury,andoarstiltsarmingly incraasing thairranga tn the CanadianArctic. EurasianGaldanPloverscontinuasariouslysoda­ dmainthahaathar moorsofwastamEuropa,but hasamanagad toanpandIhairaangadnato dafo raslalianinthabaraatforastzana,aspaciallyin Pinland.ThawidalyraporladincraasaofGray PlovarsinwaslamEuropa(f-foldinsha Eritish Islassincatha197fs(isunlikatytohaduasosaul populationincraasasbut,accordingtoEyrkjadal andThompson,mustaaflacldistributionalchan­ gasinwinlacIamnotsocanvincadthatthaam­ partad disappaarancaof an apparantEurapaan wintaringpopulationofPacificGaldanPlavars fhighlyanigmaticinitsawnrighticouldhava brancausadbythaGulch‘wilslawallars’,aspa­ cialtyasIhahistoricalrapartsofthisspactasmdi­ catathat(incangmancawishihaiehabitatchaica inAsia-Pacifc( thayusadinsarsidatratharthan grasslandaraas.Farthisraasan,Ihayshouldhava bran ralativatyimmunatothisfann ofmartcas hunting.

Thansharaarm wandarfulsammariasofsha

hraadingschadulasshowingshaimprasstoalongi­ tudinalvariafianinplavarbraadingsaasanalily, aspaciatlyfarEurasian GaldanandGrayPlavars. Thasongfight andgrounddisplaysofthafont tundraplavarsarmbroughtalivabyIha many drawingsandschamasofaarialmavamants,as wallasbythasanogramsandIhairanalysas;allof thisbringoriginal workagain!

Thatthaauthorswaranotafraidtowork vary

hardansynthasasofinfas’mafian,comasoutin thachaptaronmigration. Easadontha datagai­ nadfrom 4dfifimusaum-skinstha warid aver, backad-upbythoroughtisarasuraravsaws,savaral diagramsare campitadtoshowthaworldwide phanatagyofsouth-andnorthwardmigrationof Ihafaurplavars.Gaagraphicallinasofsimilardo- lasofaccuwanca (‘isaphanas’(,basadonthama­ diandatasforctustarsofmuseumspacimans,pro­ videaninnovativewaytodocument migratory advanceinsuchwidelyrangingspecies.Eming

interestedinage-differencesinmigrationsched­ ules andbeingfrustratedwiththelackofageing informationinthemuseumrecords,rather than tryingtoborrow somanyskins theauthorsasked forskinstobephotocopied:“soonphotocopied specimenspiledup, andvirtuallyallofthemwere sufficientlycleartoenableustoagethebirdscon­ fidently!”.Quitestriking,andnotunderstood,dif­ ferencesinthe autumndistributionofjuveniles andadultsinespeciallyAmericanGoldenPlovers comeoutasaresult.

Afterdealingwithmigrationandnonbreeding distributions, thebookiscompletedwith reviews of foragingandfood,interactionsbetween p1o- versandothershorebirdspecies,andaspectsof conservation.Inthelatterchapteritispointedout howlittleweknow aboutpopulation processesin anyofthetundraplovers, inspiteofthefactthat suchknowledgeisapparentlyconsideredmanda­ toryforEurasianGoldenand GreyPlovers inthe contextoftheECBirdDirective. Gosh,doesthis reallymeanthatthereare‘societalneeds’forde­ tailedstudiesonthe demographyofourtundra plovers?Let’sgetgoing!

No seriouscriticismsatall,then?Well,maybe

afew.To dojusticetotheimportantdata-compi­ lationsondistributionandbreeding-andmigrat­ ion-phenology,therecouldhavebeen greaterat­ tentionto theclarityofthelay-outoftheresulting maps,whichcouldalsohavebeenprintedina biggerformat.Theencyclopaedictreatmentof topics,andthechapterarrangements,aresome­ timesatoddswiththedevelopmentofastrong theme.Forexample,thechapterontheevolution ofthegenusPluvialiswouldprobablyhaveseen greater development,andbeenmoresatisfying,if ithadbeenwrittenasthegrandsynthesisofthe currentknowledgeontundraploversand their habitats.Theseparationoftext+figuresfromthe tables(afterReferencesattheendofthebook) andtheextensiveAppendices(beforeReferences) is sometimes a little cumbersome. The Index wouldhavebeenmore effectiveifthebiological subjectshadbeengivenseparateentriesforeach ofthespecies,ratherthansimilarlistsoftopics beinggivenforeachofthem.Butthesearereally

minor quibbles,andprobablyrather personal ones.Ifyoulove,orevenjustlike,ploversand thehabitatsinwhichtheyoccur,thisisabookfor you.

TheunisPiersma,

NetherlandsInstituteforSeaResearch(NIOZ), P0.Box59,1790ABDenBurg,Texel,

TheNetherlands

MEARNSB.R.MEARNS1998.-TheBird

Collectors-AcademicPress,SanDiego.ISBN0-

12-487440-1.Hardback, 472pp.,manyunique B&Wphotographsandportraits,3tables,4maps. Price:£29.95(f93.50)

TheBirdCollectorsisanamazingbook.Weall knowthatourgrandfathersand great-grandfa­ thers(butonlyonrareoccasionsour grandmoth­ ers)wentbirdingwithagunratherthanwitha pairofbinoculars.Weallatleastfaintlyknow aboutthefashiondemandingternsonwomen’s hats,aboutthelargescaleeggingonislandsand inseabirdcolonies, aboutthelasttwoGreatAuks killedonEldey(Iceland),andtheextinctionof the DodoonMauritiusandthePassengerPigeon inNorthAmerica. Butthisbookgoesfarbeyond thatfaintknowledgeandshowsusindetailwhat isabsolutelyworthknowingaboutourorntiho­ logicalhistory!Prismbinoculars werenotin­ venteduntilthebeginningofthe1900sandthe notionthatanexperiencedbirdwatcher could confidentlyidentifyeveryspecieshesaw,using binocularsandafieldguide,wasonlyslowlyac­ ceptedby‘theoldshotgunschoolofornitholo­ gists’.Thebeliefthat“What’s hitishistory, what’smissedismystery”wasfirmlyentrenched intheirminds.So,theVictorianbirderwaswork­ ingonhispersonalbirdcollection. Therefore, earlybirdbooksconcentratedontheidentifica­ tionofbirdsinthehand,notin thefield,depicting wingformulae,tarsi,websandtoes,beaks,and thepatternonindividualfeathers.

Thesepersonalcollectionswereoftenhuge

andbytheendofthenineteenthcenturytheown-