©EntomologicaFennica. 25August2010 Pre-imaginal stages of the blowfly Protocalliphora falcozi in nests of the tree sparrow (Passermontanus) VieraJanoskova,IvanOrszagh,JanJamriska&MartinKopani Janoskova,V.,Orszagh,1.,Jamriska,J.&Kopani,M.2010:Pre-imaginalstages of the blowfly Protocalliphorafalcozi in nests of the tree sparrow (Passer montanus). —Entomol. Fennica21: 107—116. ThispaperofferstheinitialdescriptionofthefirstinstarlarvaofProtocalliphora falcoziSéguy, 1928(Diptera: Calliphoridae)andredescriptionofthesecondand thirdinstarlarvaeandpuparium, aswellasdistinctionbetweenP.falcoziandits relativeP. azureaFallen(Diptera: Calliphoridae). Thematerialwas sampledin theNationalNatureReserve Sur (SW Slovakia) fromnests ofthetree sparrow (Passer montanus), which has not previously been classified as a host of P.falcozi. Knowledge ofdiagnosticallyrelevantcharacterics to determinebird blowfly larvae andpupariawillundoubtedly contributeto the currentaccepted phylogenyandclassificationofthegenusProtocalliphora. V.Janoskova,I. Orszaglz&J.Jamriska,DepartmentofZoology,FacultyofNat— ural Sciences, Comenius University, Mlynska dolina B], 842 15 Bratislava, Slovakia; E—mails.‘ [email protected], [email protected], [email protected] M. Kopani, Institute ofPathology, SchoolofMedicine, Sasinkova 4, Comenius University, 81108Bratislava,Slovakia;E—mail.‘[email protected] Received10July2008, acceptedI1January2010 1. Introduction including P. rognesi and T. braueriwhich have Holarctic distribution (Sabrosky et al. 1989, Birdblowflies(ProtocalliphoraHoughspp.)be- Whitworth2003a, b). long to the family of Calliphoridae (subfamily Very little data has been published on the Chrysomyinae) togetherwith some othergenera identification ofpre-imaginal stages ofthis ge- (Chloroprocta Wulp, Chrysomya Robineau— nus.Whenthirdinstarsareavailable,theyshould Desvoidy, Chrysopyrellia Séguy, Cocnliomyia be allowed to pupate for easier identification. Townsend, Compsomyiops Townsend, Hemilu— However,emptypupariaareusuallytheonlyevi- ciliaBrauer,ParaluciliaBrauer&Bergenstamn, denceoflarvaeinbirds’ nests. Phormia Robinneau—Desvoidy, Protoplzormiata Various determination keys from the Palae- Grunin, Protoplzormia Townsend and Trypo— arctic region have been published to identify calliplzora Peus). Theirlarvaearebloodsucking Protocalliphora larvae, and these are predomi- parasites on altricial nestlings. 15 species of nantlyforthirdinstarlarvae,whichpossessmore Protocalliphora and 1 ofTrypocalliplzora have diagnostic characters than the first and second been recorded from the Palaearct (Schumann instar larvae: Protocalliphora azurea (Engel 1986,Kurahashi 1993), 10fromEurope(Rognes 1920,Rohdendorf1957, Iwasa&Hori 1990),P. 2004), and29 species fromtheNearctic region, falcozi(Séguy 1941, Zumpt 1965, Lehrer 1972), 108 Janos/(ova etal. ° ENTOMOL. FENNICAVol. 21 P. maruyamensis (Iwasa & Hori 1990) and P. Slovakia on the SE fringe ofthe Male Karpaty rognesi(Engel 1920). Sabroskyetal.(1989)pro- Mts. (48° 42’ N, 17° 16’ E). The conservatory videdthekeyforidentificationofthirdinstarlar- areaisformedbyrarealderwetlandsinaflooded vae of 16 species in the Nearctic region. Older terraindepressioncombinedwiththermophilous referencesincludedescriptionsofthirdinstarsof oak woods. The collected specimens belong to P.avium(Coutant1915,Dobroscky 1925)andP. firstinstarlarvae (11), secondinstarlarvae (17), metallica (Hall 1948). thirdinstarlarvae(32)andpuparia(20),andwere Second instar larvae were described for allobtaineddirectlyfromnestlings ornestmate- Protocalliphora avium (Coutant 1915) fromthe rial duringthe 15t—3relbroods ofthetree sparrow. Nearctic region and P. azurea (Draber—Monko Thematerialwas sampledintheperiods ofMay 2002),P.falcozi(Zumpt1965),P.maruyamensis 20—June 26, 2006 and May 2—June 27, 2007. (Iwasa&Hori 1990),P. rognesi(Draber—Monko Some of the larvae were kept to develop to 2002)fromthePalaearcticregion.Theonlypaper pupariaandtoadults,andthisallowedspeciesde- concerning description ofthe first instar larvae termination according to Draber-Monko (2004). was published by Iwasa & Hori (1990) for P. The pre-imaginal stages were identified from maruyamensis. nests where only specimens ofP.falcozz' adults PupariaofProtocalliphoraazurea,P.falcozi, occurred. MixedinfestationbyP.falcozz'andP. P. rognesiandTrypocalliphorabrauerl'werede- azurea appearedonlytwice during ourresearch, scribed by Cais et al. (2001), and Whitworth and in these cases the pre-imaginal stages were (2003a) published his key for the determination thus not identified. The puparia ofProtocalli— ofpupariaof27speciesintheNearcticregion.P. phoraazureawereobtainedfromnests ofParas falcozihasbeenrecordedonlyfromthePalaearct, majorintheBotanic Gardens inBratislava. and mainly as adults (Gregor & Povolny 1959, Larvaewerekilledbysoakinginhotwater(to Peus 1960, Lehrer 1972, Mihalyi 1979, Rognes avoiddeformation) andtheywere storedin76% 1997). P.falcozz'has been reported as aparasite alcohol. Larvae and puparia were mounted in onParasmajor(Gregor&Povolny 1959,Lehrer Canadabalsam. AdigitalLeicaDFC195 camera 1972),Paruspalustris(Szpila1999,Wesolowski mountedon aLeicaDM 1000 stereomicroscope 2001),Parascaeruleus(Rognes 1997),Ficedula was used for microphotography. Samples for albicollz's(Gregor&Povolny 1959,Lehrer1972) SEManalysisweredehydratedingradedalcohol, and Phoenicurus phoenicurus (Gregor & subjected to the critical drying point of CO2 Povolny 1959, Lehrer 1972). The tree sparrow (CPD 030, BAL-TEC, BGPRUFZERT). Speci- (Passermontanus) was declaredto be an exclu- mens were mounted on carbon stubs andcoated sive host of Protocalliphora azurea (Rognes withalayerofgoldinanionsputteringapparatus 1991) (SCD 050, BALZERS, Lichtenstein). SEMpic- Thispaperaimstodescribethefirstinstarand tures weretakenusing aJXA 840 A (JEOL, Ja- to re-describe the second and third instar and pan) scanningmicroscope. pupariumofP.falcozisinceallavailablerelevant ThestudymaterialisdepositedattheDepart- publisheddatahasbeeninsufficientandtoogen- ment ofZoology, Faculty ofNatural Sciences, erally diagnostic for the whole genus. Here, we ComeniusUniversity. emphasize the characteristics to distinguish the pre-imaginal stages ofP.falcozi andP. azurea, since they belong to the most common species 3. Terminology foundinnests ofcavemicolousnestersincentral Europe. Withtheirtapering anteriorandtruncatedposte- riorendsProtocalliphoralarvaegenerallydonot much differ from other genera ofCalliphoridae. 2. Material and methods Their body consists of 12 segments, pseudo- cephalon, 3 thoracic segments (T l—T III), 7 ab- Sampling ofthe material was conducted in the dominal segments (Al—AVII) andthe analdivi- National Nature Reserve Sur situated in SW sion,completelycoveredwithspines. TheTIan- ENTOMOL. FENNICAVol. 21 ° Pre-imaginalProtocallzphomfalcozi 109 H1125w mesoI%n5)’:'-.""'SE"'TWm'.oe“s'a:Z;T'-''"I:‘I.I Fig.2. SEMofanteriorendofbodyofthefirstinstarof Protoca/Iiphorafalcozi,ventralview;a:antenna,as: additionalsenseorgan,0:cirri,or:oral ridges, pf: pro- thoracicfringe, mh: mouthhooks, mp: maxillarypal- pus. instars. The posterior spiracle region is divided intoahyperspiracularregion(abovetheposterior Fig. 1. FirstinstarofProtoca/Iiphorafalcozi.—a. spiracles), amesospiracularregion (betweenthe Wholebody, lateralview.—b. Cephaloskeleton,ven- posterior spiracles) and a hypospiracular region tralview;dc:dorsalcornua, is: intermediatesclerite, (undertheposteriorspiracles). mh: mouthhooks, pb: parastomal bar,vc:ventral Apart from a well-sclerotized and darkened cornua.—c. Posteriorspiracularregion;ao:anal cuticle, the puparium also differs from larvae, opening,ap:anal papillae, p1:dorsal papillae, p3: havingaless distinctprothoracic fringewhichis supralateral papillae, p4: infralateral papillae, p5: inverted during pupation and also poorly distin- subventral papillae, p6:ventral papillae. guishable papillae. Externally the puparia have varyinglydevelopedcuticularfolds (closetothe teriorspinosebandisexceptional,sinceitismod- posteriorspiracles)andcuticularridges(dorsally ified to a crown of setae (prothoracic fringe), andventrallyoneachsegment),andthesearenot which most likely adheres the larva to the host visibleonlarvalinstars.Theterminologyusedfor during blood-sucking. Each abdominal segment thedescriptionofpre-imaginalstageswasthatof hasnormallythreedistinctventralspinalbands— Courtneyera1. (2000) andWhitworth(2003a). ventralbandsboundedby8ventralpads(Fig. l). Thepseudocephalonhas antennae, maxillary palpus andmouthhooks. The cephaloskeleton is 4. Descriptions oflarvae composedofbasalandintermediatescleritesand and puparium mouthhooks. Intermediate sclerite ofeach instar proceeds to the basal sclerite which consists of 4.1. Firstinstarlarva dorsal andventral comua. Paired labial sclerites aresituatedbetweenthemouthhooksandtheme- Thebodylengthis09—225m(n:4). Spinesof dianbeamofthe intermediate sclerite. Alabrum each segment are onlyjust visible and lack dis- andaccessory sclerite are absent. tinctdifference intheir shape and size. Thepro- Theanaldivisionhas analpapillaeand7 dis- thoracic fringe is very short comparedto that in tinct pairs ofpapillae (dorsal, subdorsal, supra- other instars (Fig. la), and the ventral pads are lateral, supraventral, infralateral, subventral and clearly distinguishable on each abdominal seg- ventralpapillae), butthese arenotdistinctonall ment. Three ventralbands are distinct andunin- llO Jdn0§k0vdetal. ° ENTOMOL. FENNICAVol. 21 Table 1. Second instarlarvalcharactersofProtoca/IiphorafalcoziandF’. azurea. Reference Length (mm) min—max Numberoflobes mean 1r SD. (n) ofanteriorspiracles (n) P. falcozi Presentstudy 1.6—5.6* 3.65 i 1.36 (13) 8 (10) P. azurea Draber-Monko (1996) 6.0—6.5— 6—7 (occasionally8) *Fullydevelopedsecondinstarlarvae,theonlydifferencewasinlength. terrupted. Anteriorspiracles areabsent. 3a, 3b). Theparastomalbaris fusedWiththe in- Each lobe ofthe pseudocephalon bears an- termediate sclerite (their border is marked by a tenna, maxillary palpus (cluster of several dashedline, Fig. lb). sensilla)andaccessorysensillum. Onthesurface ofthe pseudocephalon there are numerous cirri andoralridges, Whilethe labial lobe andventral organareabsentorindiscernible (Fig. 2). Cephaloskeleton is small, mouthhooks are more strongly sclerotized than the other parts. The intermediate sclerite is elongated (When comparedtothesecondandthirdinstars),andthe dorsalcomuaislongerthantheventralone(Fig. fa.“-d‘ c 0.01mm Fig.4. Second instarofProtoca/Iiphorafalcozi.—a. Wholebody, lateralview.—b. Cephaloskeleton,ven- tralview:dc:dorsalcornua, is: intermediatesclerite, ls: labialsclerites, mh: mouthhooks, pb: parastomal bar,vc:ventralcornua.—c. Posteriorspiracularre- gion;ao:analopening,ap:anal papillae, p1:dorsal papillae, p2:subdorsal papillae, p3:supralateral Fig.3. CephaloskeletonoffirstinstarofProtoca/Ii- papillae, p4: infralateral papillae, p5:subventral pa- phorafalcozi.—a. Lateralview.—b.Ventralview. pillae, p6:ventral papillae, p7:supraventral papillae. ENTOMOL. FENNICAVol. 21 - Pre-imaginalProtocalliphorafalcozi l11 Fig. 5. SEMofanteriorendofbodyofthesecond instarofProtocalliphorafalcozi. They have anal papillae and only 5 pairs of tiny papillae (dorsal, supralateral, infralateral, subventralandventral)developedontheanaldi- vision, With the supralateral, infralateral, sub- ventral, ventral and anal papillae clearly distin- guishable. Theanalpapillae areroundedandthe anal opening is situated slightly anterior to the anal papillae. The posterior spiracles are small andbilobed(Withoutagapbetweentwoforming slits) and they are poorly sclerotized. The peri- treme is incomplete, With an absent spiracular Fig.6. Cephaloskeletonofsecond instarofProtocalli— scar(button).Spinesinthemesospiracularregion are also absent(Fig. 1c). phorafalcozi.—a. Lateralview.—b.Ventralview. sensillumindiscernibleorabsent.Numerouscirri 4.2. Secondinstarlarva and oral ridges occur on the surface of the pseudocephalon. The labial lobe andventral or- The body length is given in Table l. Spines of ganare absentorindiscernible (Fig. 5). each segment andprothoracic fringe are distinct Thecephaloskeletonisstout,mouthhooksare (Fig. 4a). On T II—A VII the central spines are more strongly sclerotized than the other parts. slightly longer than the marginal ones. Ventral TheparastomalbarisnotfusedWithintermediate pads are easily distinguishable on each abdomi- sclerite, and the dorsal cornua is longer than the nalsegment,andtherearethreedistinctandunin- ventralone(Fig. 4b, 6a, 6b). terrupted ventral bands. Anterior spiracles are Analpapillaeandall7pairsofpapillaearede- present, number ofdeveloped lobes is given in veloped on the anal division, including distinct Table 1. supralateral, infralateral, subventral and ventral Each lobe ofthe pseudocephalon bears an- papillae. The subdorsal papilla is poorly devel- tenna and maxillary palpus, with an accessory oped, and anal papillae are rounded, While the ll2 Jdn0§k0vderal. a Table2.Third instarlarvalcharactersofProtocalliphorafaicoziandP azurea Reference Length (mm) min-“max Numberoflobes mean i 8.0. (n) ofanteriorspiracles (n) P. faicozi Thisstudy 11.3404" 7.98 i 1.91 (15) SW11 (15) Zumpt(1965) 1243“- 5;; (occasionally 10) P. azurea Draberw Monko (2004) 13~15~ 8 lwasa & Horl (1990) 1345“- 5-40 *Fullydevelopedsecondinstarlarvae,theonlydifferencewasinlength. anal opening is situated slightly anterior to the analpapillae.Posteriorspiracleshave2slitswith adistinctgapbetweenthem. Theperitremeisin:- complete but more sclerotized in comparison withthe firstinstar. A spiracular scar(button) is developed although it is not clearly distinguish- able. and. spines are developed in the meson- spiracularregion(Fig. 4c). 4.3. n mmlarva The body length is given in Table 2. Spines of each. segment andprothoracic fringe are distinct (Fig. 7a). On T HmA VH central spines are pnsaalidglsshetalgyrmeleoennatgs.iealyrnddthtishateninretghuaeirsemhtaharbregleeindoainslteoinnacecthsa.naVbdd.eoumnnit_irn—au-l \g\h1“2(‘Mo‘‘f“i-‘‘3l1'l‘s:_ll9ni.'g5Ii..‘11{rI901n“:,:Ix1H}‘{,l!15‘‘1,l'’‘If!u;dfii?uiHI’(fpgHlIfirIg‘fr"fr”fr';i'l.sfnIiI9l,?""9"‘a'n'‘9’?pf'W?'f7F.'’Hf5riI3«‘arMbc:fll'Vfi3Egfaf!fl'll'1ih‘iggl5f“,,d'frFf1i‘.3!!”f'-"hf!3'f«!1!-ai4’f{“1’fd:‘I’i1I.p;":-”/‘rf,Il,'d‘;.IIf;lf‘0}JIJ'r. tempted ventral bands. Anterior spiracles are A. 'b r-I. /-p3 ptTserearemnbsEsleaefanllc2aut.;hn.mndluoinmmbdeabisxeocirleflaomtrhfyiebdlpeepavsoleeprluuoadspb.oescdweenipltoth.hbaOetlhsonenitshabecgecasiveuressrfnsaaocrirrney- .yHwfl:.w..‘ua1"u";’.nm.m.t1ie'y-wIr-.i.wIwfll“rfV.iv,w.»r;f;(ff’““rfiI“v,(jIJfalli'd;Id“;“wr,9irf,‘!-a'1,o“I,:lir’I“«y;tIi,o''r“v‘5’a:if"’;”?1.’’g4!"ff1r!"w'fa1,¢/!lVo0y,I,arr'Ir,4.;(.‘j'”f.’'iI#”IIl‘.lIfe9Nlfl’iy’.11vlfgglJ§,a’;'"NJ,:,fI’u”3';l'1fJ"f1a3"It3jgt("1#5f[J‘"j3#.Jn‘"li"“l411:3.{I3“‘fI9'.'”31i9',J’t1l“l'3r‘"’,a3}l’fl;1rf"f‘ii4'3ifsIfu'‘H"{i't«5J(‘{i1t91‘’“anP"3i;~B‘,c}“..Ia,t‘113f"“ia1f‘I11i‘;'v“1.‘“341lsh4l)'4r.a51.t““f“Amn“.“‘‘I}n‘\“i:*‘12t1mh$mtWrk”q“.h.“-u.§.“E‘.; ...M--’ew-'7”p4 ofthe pseudocephalon there are numerous cirri and.oralridges.and.thelabiallobeand.ventral0r- ganare also indiscemible orabsent(Fig. 8}. The cephaloskeleton is thick. with the mouthhooks moresclerotizedthanotherparts.Theparastomal barisnotstedwiththeintermediatescleriteand the dorsal comma is longer than the ventral one (Fig. 7b. 9a. 9b). Fig. 7.Third instarofProtocaliiphorafalcozi-—-a. Theanalpapillaeandall7pairsofpapillaeare Wholebody, lateralView.--—-b. Cephaloskeleton,venm developedontheanaldivision.includingdistinct tralView;dc:dorsalcomma, is: intermediatesclerite, ls: labialsclerltes, mh: mouthhooks,vc:ventral supralateral. infralateral. subventral and ventral comma.---c. Posteriorspiracularregion;a0:anal papillae. The subdorsal papilla is poorly devel- opening,ap:anal papillae, p1:dorsal papillae, p2: oped.Whiletheanalpapillaeareroundedwiththe subdorsal papillae, p3:supralateral papillae, p4: analopeningsituatedslightlyanteriortotheanal infralateral papillae, p5:subventral papillae, p6:ventral papillae.Thereareposteriorspiracleswith3 slits. papillae, p7:supraventral papillae. ENTOMOL. FENNICAVol. 21 - Pre-imaginalProtocalliphorafalcozi 113 Fig.8. SEMofanteriorendofthethirdinstarof Protocalliphorafalcozi. withadistinctgapbetweenthem. Theperitreme iscompleteandaspiracularscar(button)isdevel- opedbutseldomeasilydistinguishable.Thereare alsomesospiracularspines (Fig. 7c). 4.4.Puparium Thebodylengthis7.11—8.34mm(meaninSD.= 7.95i0.42, n=9),withall3ventralspinebands easilydistinguishable.Therearecuticularfoldsin the hypospiracular and mesospiracular region. However, there are only small ones in the ' if {é hyperspiracularregion. Additionally, thecuticu- .n - .. . larridgesareasdistinctasthecuticularfolds.The Fig.9. CephaloskeletonofthirdinstarofProtocalli— puparia found in nests were sometimes covered phor‘afalcozi.—a. Lateralview.—b.Ventralview. bynestmaterial. tocompletetheirdevelopmentsuccessfully(Ben- net & Whitworth 1991). Additionally, enough 5. Discussion hostnestswithnestlings arerequired inorderto collect sufficientmaterial oflarvae andpuparia. Identification of pre-imaginal stages of bird Wehavenotyetacquiredanapproachtokeeplar- blowflies accordingtotheirmorphologicalchar- vae artificially, dueto analternativehostusedto acteristics appears indisputably more difficult nourish bird blowfiy larval instars (larvae pre- than for adults. In some cases this seems almost sumably cannot complete their development on impossible. presocial nestlings). Another problem occurs in Lack of available data on determination of obtainingeggssincefemaleskeptunderartificial Protocalliphora larvae demands arduous samp- conditionscannotlaytheireggsdespiteapparent lingsincetheseobligatebirdparasitesareneces- mating(Bennet&Whitworth 1991). Suchinade- sarilyconnectedtotheirhost, andthisisessential quate materialresults in insufficientopportunity 114 Jdn0§kovd etal. ° ENTOMOL. FENNICAVol. 21 to compare more species in their pre-imaginal ofnestlings and spend their period in hosts, so stages.Therefore, someauthorsdescribedProto— thatthey do nothave segments completely cov- calliphoralarvaeentirelyaccordingtocharacter- ered with spines, and this can play a defensive istics distinctive ofadditional or even all genus role in a host’s body (also e.g. Auchmeromyia species (e.g. Zumpt, 1965: descriptions of the Brauer andBergenstamm andthe third instar of secondandthirdinstars ofP.falcozi). Cordylobia Gruenberg). Sincelarvaeofthesameinstarcanhavevary- ThemainfeaturesofP.falcozilarvaeinclude ing nutrition their characteristics can vary, and enlargedantennae,numerouscirriandoralridges this makes identificationmore complicated(e.g. onthepseudocephalon.Enlargementofantennae distance between spines, prothoracic fringe di- andmaxillarypalpiinthefirstinstarsofparasitic ameterandthe distancebetweenposteriorspira- or predatory species from the genera Bellardia cles). Also, during food deficiency, the body Robineau-Desvoidy, Onesia Robineau—Desvoi- lengthofamore advancedinstarmaybe shorter dyandPollem'aRobineau—Desvoidywasalsoob- thanthatofayoungerone. Moreover, lengthof- served(Szpila2003, 2004). Thisphenomenonis tenvarieswithinthe same instardueto differing possiblyassociatedwithanecessityforbetterori- food sources and conditions. Certainbody parts entation in nature. Females ofCalliphora Robi- such as the cephaloskeleton andposterior spira- neau-Desvoidy, Phormia Robineau-Desvoidy clesalsohaveasurprisingdegreeofvariabilityin and Lucilia Robineau—Desvoidy, which do not theirsclerotizationinthe sameinstar. have enlarged antennae andmaxillary palpi, lay In comparison with the Nearctic region, re- their eggs directly onto a suitable food source, centresearchonbirdblowfliesinthePalaearctic whereas parasitoids do so when actively search- area has not been intensive enough. Larvae of ing fortheirhostin soil (Szpilaetal. 2008). Fe- Protocalliphora distincta, P. isochroa, P. Iii, P. males ofProtocalliphora spp. lay their eggs di- peusi,P. nuortevaiandP.proxima, allPalaearc- rectly on nestlings or on nearby nest material. ticspecies,havenotbeendescribed.Mostdataal- Even though they can lay on a suitable food ready distinguishes only Trypocalliphora larvae source(nestlings),larvaeareintermittentfeeders, from Protocalliphora and Protocalliphora leaving their hosts facultatively after sucking rognesifromP. azurea andP.falcozi. enough blood, but they soon search for this LarvaeofP. azureaandP.falcozz'ofteninter- sourceagain. actwiththesamehostandtheirhostsbuildsimi- There is a lack ofdetailed information con- larnestsorremaininnestsforapproximatelythe cerningtheanalpadofProtocalliphomlarvalin- sameperiod, therefore identificationoftheirlar- stars. The analpad, whichhasbeendescribedin valinstarsstillseemsdoubtful. Originalmorpho- the second instar ofP. rognesi (Draber—Monko metric analysis supported by other references 2002)issimilartothatofP.falcozi.Theposition showedshortersecondandthirdlarvalinstarsfor oftheanalopeningofthefirstinstarofP.falcozi P.falcozithanforP. azurea.Thenumberofante- corresponds with that of Phormia regina rior spiracle lobes may be the same inboth spe- (Meigen, 1826) and Lucilia illustris (Meigen, cies, andhencethis does notrepresentareliable 1826) and this was described by Szpila et al. characteristic foridentification(Tables 1 and2). (2008). Differences in the shape ofcephaloskeletons in Therearemorphologicaldifferencesforiden- these2speciesappearalsoinsignificant(Draber— tificationinpupariaandadultsbetweenP.falcozi Monko2004). However, thereis somedegreeof andP. azurea (Cais et al. 2001, Draber—Monko discrepancy in the intermediate sclerite region, 2004). In our analyses, some additional charac- including theparastomalbar. Detailed synthesis teristicsusedindeterminationofNearcticspecies ofcephaloskeletons requires additional material (Whitworth2003a)havealsobeentakenintoac- foranalysis. count. The cuticle surface of fully developed Segments ofProtocalliphora species, which pupariaofP.falcoziisdistinctlydifferentthanin arecompletelycoveredwithspines, exhibitrela- P. azurea. Onthe surface they arewrapped in a tively rare features for dipteran larvae. The Try— hair cocoon, and this is usually absent in P. pocalliphoralarvaeburrowdirectlyintotheflesh azurea. ENTOMOL. FENNICAVol. 21 ° Pre—imaginalProtocalliphorafalcozi 115 Weproposethatdataobtainedinouranalyses ticonbirdsinPoland.7AnnalesZoologici(Warsza- will be useful in complex phylogenetic simula- wa)52: 3337337. Draber-Monko, A. 2004: Calliphoridae. Plujki (lnsecta: tions. IwasaandHori (1990) declaredthat some Diptera).FaunaPolski.Tom23.7Muzeumilnstitut characteristics,whichareabsentinotherChryso- ZoologiiPAN,Warszawa. 662pp. myinae genera, presumably corresponded with Dobroscky,I.D. 1925:Externalparasitesofbirdsandthe specializedbloodsuckinghabitsuchasinProto— fauna ofbirdnests. 7Biological Bulletin48: 2747 phormia,PhormiaandChrysomya.Forexample, 281. Engel,E. O. 1920:Dipteren,dienichtPupiparensind,als these include undeveloped tubercles at abdomi- Vogelparasiten. 7 Zeitschrift fiir Wissen—schaftlich nal segment VIII, prothoracic fringe, relatively Insektenbiologie 15: 2497258. smallmouthhooks, smallposteriorspiracleswith Gregor, F. & Povolny, D. 1959: Kritischer Beitrag zur a long distance between them. Except in Kenntniss der Tribus Phormiini (Diptera, Calli- Protocalliphora, obligatory haematophagy is phoridae). 7Acta societatis entomologicae Cecho— seen only in rare forms ofparasitism ofCalli- sloveniae56: 26750. Hall, D. G. 1948: The blowflies ofNorth America.7 phoridaelarvae.Morphologyofbirdblowflylar- ThomasSayFoundation,Washington.477pp. vaemaybebeneficialto comprehendthe evolu- Iwasa,M.&Hori,J. 1990:Thecalliphoridlarvaeparasitic tionofCalliphoridaewhenlarvalhaematophagy onbirdsinJapan(Diptera:Calliphoridae).7Medical originating in saprophagy is considered. How- andVeterinaryEntomology4: 1417146. ever, thedataofferedinthispaperisproposedto Kurahashi, H. 1993: Japanese Species of the Ornitho— become essential forcomplexphylogenetic syn- parasitic Blow Flies: Protocalliphora and T73490- theses. calliphora(Diptera:Calliphoridae).7NewEntomol- ogist42: 8715. 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