Trypocølliphora Peusr 1960 (Diptera: Calliphoridae)

Revision of the bird-parasitic blowfly genus Trypocølliphora Peusr 1960 (Diptera: Calliphoridae)

KNUT ROGNES

Rognes, K. : Revision of the bird-parasitic blowfly genus Trypocalliphora Peus, 1960 (Diptera: Ent. scand. Calliphoridae). Ent. scand. 15: 371-382. Copenhagen, Denmark 31 January 1985. ISSN 0013-8711.

European material of the bird-parasic blowfly genus Trypocalliphora Peus, 1960, incfyding type material of. Avihospita braieri Hendel, 1901 and Trypocalliphora lindneri Peus, L960, has been examined. The genus is redefined on the basis of the male phallosome structure and its systematic position ind status discussed. The genus contains a single Holarctic species, Trypocalliphora braueri (Hendel, 1901) ; Protocalliphora hirudo Shannon & Dobroscky, 1924, f . Iindneri Peus , 1960 and 7. compaclø Grunin,, L966 are new synonyms. Knowledge of the biology is summarised and a host list for the Palaearctic Region is given.

Knut Rognes ,HavØrnbrauteneTa, N-4040 Madla, Norway.

The genus Trypocalliphora Peus, 1960 was torisches Museum, Vienna (NMW), Staatliches erected for blowflies of Phormiinae (Phormiini Museum ftir Naturkunde, Stuttgart (SMN), sensu Zumpt 1956) with larvae living obligatory as Zoologisk Museum, Bergen (ZI|{B) and Zoologi- subcutaneous parasites on birds (nestlings and cal Museum of the University, Helsinki(ZldUH). juveniles). Peus removed these flies from the Adults and some puparia of Protocalliphora genus Protocalliphora Hough, 1899 where they azurea (Fall6n), P. chrysorrhoea (Meigen), P. had been placed previously (e.g. by Zumpt 1956). nuortevai Grunin, P. proxima Grunin , Boreellus Larvae of Protocalliphora are external blood- atriceps (Zetterstedt), Protophormia terraenovae sucking parasites in birds' nests that feed intermit- (Robineau-Desvoidy), Phormia reginø (Meigen) tently on the nestlings (Sabrosky & Bennett 1958; (Phormiinae) and Chrysomya albiceps (Wiede- Rausch l97L). 3 nominal species were originally mann) , C. megacephala (Fabricius) (Chry- included in Trypocalliphora: the Palaearctic somyinae) have been available for comparison in braueri (Hende[, 1901) and lindneri Peus, 1960 Norwegian museum collections (Bergen, Oslo, and the Nearctic hirudo (Shannon & Dobroscky, TromsØ) and in my own collection. 1924). Most European workers have accepted Peus' genus (except Zumpt 1965) but North Genus Trypocalliphora Peus , L960 American ones have not. The purpose of this Peus 1960: 199 . Type-species: Avihos- paper is to redescribe Trypocalliphora with em- Trypocalliphora-ptto , braueri Hendel, 1901, by original designation. phasis on the male phallosome (not considered by (Sabrosky (1967) erroneously cited Trypocalliphora Peus), discuss its status and its position within the lindneri Peus, 1960 as type-species). Phormiinae, establish some new specific Diagnosis. Phormiine genus with male frons at narrow- synonymies and summarise existing knowledge on est point 0.05 x head width, parafrontals subcontiguous; its biology and host range. female frons at vertex 0.20x head width, parafrontal The study is based on European material half as wide as interfrontal stripe and without a preverti- (adults and puparia) in the following institutions: cal seta; female parafacial on upper part in certain views triangular dark spot; arista with long London showing large, British Museum (Natural History), hairs a6ove and below; basicosta, squamae, subcostal (BMNH), Museum ftir Naturkunde der Hum- sclerite, anterior spiracle and palp yellow to light brow- boldt-Universitåt, Berlin (MNHU), Naturhis- nish yellow; subcostal sclerite with black setulae; haltere

@ Entomologica scandinavica (DIPT 098) 372 Knut Rognes ENT. SCAND. VOL. 1s (1e84)

yellow; posterior spiracle brown; squamae and postalar frontals in male; the second group by mesonotal wall bare; body colour metallic greenish blue in male, flattening, dark colour of above mentioned struc- brassy green with white pruinosity in female; male distiphallus (Fig. 1) with very wide apical opening nearly of tures and separated parafrontals in male . Zumpt same calibre as base of phallus; hypophallic lobes small, (1965) did not accept Peus' genus and treated its pointing laterally, not ventrally; ventral surface flat, nominal species in Protocalliphora. He admitted evenly sclerotised, with spinules widely distributed but that the absence of the crown of setae on the most concentrated in two subapical ventral fields; a complete ring-shaped sclerotisation around apex; a distinct pseudocephalon and other features of the third dorsal sclerotisation extends towards base from ring stage larva might justify Trypocalliphora as a structure; no midventral sclerotisation (meso- separate genus but considered this premature in hypophallus) developed; ventral surface with stripe of view of the overall insufficient knowledge of the hyaline membrane along midline; male surstylus as in Figs . 9-20. Female with telescopic ovipositor with well larvae of bird-parasitic blowflies. Neither Peus developed sclerites; probably oviparous. Third stage nor Zumpt considered the structure of the phallo- larva without a crown of setae on pseudocephalon; an- some. Sabrosky (1967), in a catalogue entry, rec- terior spiracle with 2-l branches. ognised Trypocalliphora as a subgenus of Pro- tocalliphora and thus, by implication, viewed Trypocalliphora and Protocalliphora (s.str.) as Systematic position sister-groups. However, I have not been able to Zumpt (1956) and Schumann (1973a; 1973b; find synapomorphic characters of these two taxa 197 4) used the structure of the male distiphallus to (apart from the bird parasitic habit) to support split Onesia Robineau-Desvoidy, 1830 (s. lat.) this. Alternatively, Trypocalliphora could be the into the genera Onesia (s. str.) , Melinda Ro- sister-group of Protocalliphora + Boreellus + bineau-Desvoidy, 1830 and Bellardia Robineau- Protophormia * Phormiata Grunin ,197I. Trypo- Desvoidy, 1863. Similarly, the structure of the dis- calliphora has at least two good autapomorphic tiphallus readily defines Trypocalliphora as a valid characters: (1) absence of a prevertical seta in genus.Apparently, this taxon differs from all female , (2) male distiphallus with dorsal longitu- species of Protocalliphora by the absence of a dinal sclerotisation apically. The monophyly of mesohypophallus (a torpedo-shaped ventral the sister-group is suggested by two assumed aut- sclerotisation on distiphallus) . Further, in Pro- apomorphies: (1) a mesohypophallus of charac- tocalliphora the ventral surface of distiphallus teristic shape present in male distiphallus , (2) informs a deep groove in which the mesohypophal- terfrontal stripe in male frons usually broader than lus lies, the hypophallic lobes point ventrally, one parafrontal. The group Trypocalliphora + spinules are confined to two sub-lateral fields api- (Protocalliphora * Boreellus * Protophormia * cally, no apical ring-shaped or dorsal longitudinal Phormiata) is probably monophyletic, its sclerotisations are developed, and the apical aper- monophyly suggested by two assumed aut- ture is narrow (compare Figs. 1 and 2; see also apomorphies: (1) presence of 2 apical clusters of Hall 1948 (as Apaulina); Gregor & Povolny 1959; spinules on male distiphallus , (2) conspicuous flat- Grunin 1966; 1970; l97lb; 1972; Kano 8. tening of mesonotum between suture and scutel- Shinonaga 1968). In the Phormiinae a meso- lum (variable in Trypocalliphora, indistinct in hypophallus is also present in Protophormia ter- Boreellus, condition unknown in Phormiata). The raenovae (Fig. 3) and Boreellus atriceps (personal group Boreellus + Protophormia is probably observations) and in Phormiata phormiata Grunin monophyletic, ås evidenced by at least six good (Grunin 1971a). The distiphallus in Phormia re- autapomorphies: (1) both squamae dark brown, gina is very different, and without a midventral (2) black hairs on disc of upper squama, (3) hind sclerotisation representing the mesohypophallus. coxa with hairs laterally on posterior surface (an Peus (1960) divided the Phormiinae in two overlooked character, also present in Chryso- groups: (1) Trypocalliphora + Phormia Ro- myinae, but occasional specimens may lack the bineau-Desvoidy, 1830 and (2) Boreellus Aldrich hairs) , (4) prst acr absent, (5) body dark metallic & Shannon, 1923 + Protophormia Townsend, bluish-green, non-pollinose, (6) hypophallic lobes 1908 + Protocalliphora. The first group he de- with spinules distally. Collin (1925) and Seguy fined by alleged absence of flattening of (1928) even regarded the constituent species (ar- mesonotum, yellow colour of anterior spiracle, riceps Zetterstedt - boganidae Erichson and ter- basicosta and wing veins, and contiguous para- raenovae Robineau-Desvoidy - groenlandica ENT. SCAND. VOL. 1s (1984) Revision of the bird-parasitic Trypocalliphora 373

Zetterstedt) as congeneric . Phormia has 3 equivocally belong to the syntypic series and have the in common: (1) "Brauer apomorphic characters which elsewhere are pre- foilowing white labels [print]; Austria ink]" (2) "Braueri [farl_ed ilk];det, B. B. sent only within the Chrysomyinae (personal ob- [taOeO [print]" (3) "Avihospita; braueri H. [lfl; det. H.endel servations, see also S6guy 1928; Patton & Cushing [irint]" (4) "Trypocåtlipnora d [ot ?]; braueri (.Hen- 7934a;1934b; Thomas 1951; Emden 1954): (1) in- iiel) ; F. Peus det. 1959 [ink]". The lecto_typ e d cylies an "Paratypus" label It is lacking the conspicuous white hairs laterally on disc of upper additional red [print]. abdomen and I believe this to be the specimen dissected squama , (2) phallosome with dense cover of short by Hennig, as removal of the whole abdomen was his hairs on ventral surface proximal to hypophallic way of måting genital preparations (A. C. Pont in litt. lobes, (3) hypophallic lobes circular. It may there- from J. P. Dear 24.ii.1981). His reference to the "Typus von braueri Hend." whose hypopygium he examined fore be more closely related to the Chrysomyinae (Hennig 1939:361) I choose to interpret as the first valid than to the other phormiine genera. lectotype designation. Hennig's preparation was illus- According to the above analysis Protocal- trated 6y Peuslte6O: 223,,224, fig. 18), but could not be (Frankfurt, liphora is a member of a monophyletic group not traced in NVIW, MNHU, DEI, or coll. Peus The paralectotype C carries an additional red including Trypocalliphora. This means that these Geneva). "Type" iabel and a red låbel reading "braueri" [p9ryiU two taxa entered the bird niche independently at This^ must be the "Holotypus" mentioned by Peus (1960: different times which was also suggested by Peus 226, note page number). The abdomen appears intact (1960). This is perhaps reflected in the different but is without terminalia (dissected?). A paralectotype with an additional red "Type" label is the'Allotypus" larval habits of the two genera, and in the fact that Q of Peus (1960: 226). A second paralectotype ! carries an Protocalliphora contains about 50 species (Grunin additional red "Paratypus" label. 1,977a) (many still undescribed) which are gener- The additional twospecimens (1d 1?) in NMW have ally either Nearctic or Palaearctic, whereas the following labels in common: (1) "Coll. Bgst. [print]" (2) "azure{ [faded ink]; det. Bergenst. [print]" (3) Trypocalliphora (like the remaining four phor- ;Paratypus (4) "Trypoca]figlor1C ^braueri[print, red låbel]" miine genera) is monotypic. In view of these con- det. 1952 [or ?];-carries (Hendel); F. Peus [in11"' Th" siderations it appears preferable to retain a male the following additional labels: (5) "Cal- label]" (q) "Calliphora generic status for Trypocalliphora. liphora azurea Fall. [ink, yellow azvrea aus Sperling [pencil]". The pin also carries an empty puparium. As these two specimens are without Hendel-'s betermination labels, Peus may be wrong in as syntypes. I assume, Trypocalliphora braueri (Hendel, 1901) having labelled them authentic however, that they were bred from the same young spar- Figs. L, +23, Tables 1-5. row as the syntypes (cf . Hendel 1901: 31). Brauer (1883: not state the number of specimens bred. Avihospita braueriHendel, 1901:29. Lectotype C, Au- 74) did after Hennig's valid indication of a lectotype stria, eX nestling of Passer domesticus, date and Note that a lectotype for braueri twice. The further locality unknown, Brauer leg. et cult. Peus (1960) designated p.224 tig. tS, where he figures the preparation (NMW), by designation of Hennig (1939: 361) first is on made by Hennig (see above) and in the legend says it to Iexamined]. the "Hblotypus", which means that the speci- Protocalliphora hirudo Shannon & Dobroscky, 1924: be from by Hennig and illustrated by Peus by 252. Holotype C, U.S.A.: Colorado, ex "nestling men dissected implication is designated as lectotyPe (ICZN, Art .74 (b), warbler", 10.vii .1911, Palmer leg. et cult. (U.S. Na- besignation by means of a figure) The second designa- tional Museum, Washington, No . 26862) [examined made on p .226 where he identifies a specimen car- on my behalf by C. W: Sabrosky]. Syn. n. tion ii rying certain labels as the "Holotypus". He is clearly re- Trypocalliphora lindneri Peus, 1960: 227 . Holotype d , the carrying the "Type" label, dealt with West Germany: S. Baden, Kreis Kehl, Diersheim, fårring to d above. These designations, which do not apply to the ex. Alauda arvensis 1 . viii . 1951 , Schlorer leg. , Lind- , specimen, are invalid and unnecessary because of ner cult. (SMN) Syn. n. Same [exanlined]. previous action. Trypocalliphora compacla Grunin, 1966:902 (as ssp. of Hennig's lindneri Peus). Syntypes 5 12 U.S.S.R.: Lenin- d ?, Trypocalliphora lindneri Peus. In sMN 3d 3-Q, includ- grad Prov., Luga district ,, ex Phylloscopus trochilus; ing^trototype, all examined;2C 1?, MNHU, ld Tatar A.S.S.R. district, Raifa , ex Stur- 1? ,Zelenodol'sk including genital preparations. Peus (1960) nus vulgaris; Okeanskaya railway station near Vla- exåmined,- gives information on labels. For details on the types and divostok, eX Passer sp. (Institute of Zoology , further specimens bred from the same lark, see Heinz U . S. S. R. Academy of Sciences) examined] . [not (1954) Zumpt (1956), T indner (L957), Gregor &- Syn. n. , Povolny (1959), Peus (1960). Terminalia of the holotype dissected and kept in glycerol in micro-vial on separate pin labelled: (1) ;'Holotype c Trypocalliphora lindneri Notes on type material Peus , 1960" (2) "Genitalprep. 23 Knut Rognes prep. Avihospita braueri Hendel. In NMW 3C_ 3?, all 1980 C, (3) "Trypocalliphora braueri (Hendel) K. examined. All are more or less teneral. 2d 2? un- Rognes det. 1981". 374 Knut Rognes ENT. SCAND. VOL. 1s (1984)

mhph

5 uffiff

{igl l{, Male genitalia. - 1. Trypocalliphora braueri; distiphallus, left and ventral view. - 2. Protocalliphora azurea; distiphallus,rightandventralview.-3. Protophormiaterrabnovae;distiphallus,ventralview.-4-5. T.'braueri;phallapodeme, profile and surface view. - 6. T. braueri; ejaculatory sclerite. - mhph: mesohypophallus.

O ther Trypocalliphora material examined rer l"g.; Beckenwort, ld, ex Emberiza schoeniclus, 29.vi.1963, Isenmann leg., dissected. SMN. FINLAND. Lapponia kemensis: Sodankylå, Seipåjoki, lC ,, with empty puparium, ex Oenanthe oenonthb (Gru- Diagnosls. See above diagnosis for the genus. nin, Nuorteva & Raj ala L969), terminalia glued to piece of celluloid with the specimen; Regio kuusamoensis: Salla, Vårriotunturi ,, 2d, eX Anthus pratensis (Haka- Description nen, Grunin 8L Nuorteva 1974), dissected; Karelia ladogensis: Parikkala, Siikalahti, ld, with empty Male. Length: 6.I-9.3 mm. Head. Black; frontal puparium, ex Luscinia luscinia, L7.vi.1970, pupated stripe blackish brown to reddish brown; lunula 20.vi., emerged29.vi., J...S...leg. (illegible); Ostrobott- bare, nia borealis: Pello 2Q, one with intact puparium, the brown to brownish black, shining; face and other with empty puparium and semipupated larva, ex mouth-edge black; facial ridge yellowish below; Motacilla alba, 24.vi.1973, J. flalonen leg. AIIZldUH. genae reddish or brown; occipital dilation black; - NORWAY. Buskerud, Ål: Å1, Zd 7Q,-wittr 5 puparia 1.ant.segm. black or reddish; 2.ant segm. reddish of which 4 empty, ex Ficedula hypoleucø (nestlings in in apical half; 3.ant.segm. reddish basally and nest-box), 6.vii .1944,, L. O. Odden l"g. , one male dissected; Sogn og Fjordane, Balestrand: BrekkestØlen, along hind edge; arista black, yellow in middle; Brekka 15d 3!, with 3 empty puparia, ex Oenanthe hairs of occiput whitish with slight yellowish oenanth€, 8.,9.,14.vii.1935 1Q as larvae 23.vifrom ,2d sheen ., I-3 rows of black setulae behind postocular an approx. 7 days old nestling, pupated 24-25 .vi, setae; emerged 14.vii, O. Meidell leg. et cult.,4 males dissec- row of frons at narrowest 0.034-0.064x ted. ZMB and coll. Rognes. U.S.S.R. Karelian head width (Fig .7 ,Table 1); parafrontal with 12- A.S.S.R. : near Kivach, Ld, with empty puparium "s de- 14 inclinate frontal setae, foremost at lunula, reven. lastochki" [ex Hirundo rusticaf, 15.vii .1974,1?, hindmost separated from anterior ocellus by with empy puparium, ex "penochka-vesnich" fPhyllo- scopus trochilus], 10.vii.1974, both A. Pokrovskiy leg., length of ocellar triangle; iu strong, no ev; para- male dissected. BMNH. - WEST GERMANY. No lo- frontals with black setulae in front half ; cality, lC , ex Emberiza schoeniclus,28.vii .1961, Schlo- parafacialia with parallel edges and numerous ENT. SCAND. VOL. 1s (1984) Revision of the bird-parasitic Trypocalliphora 375 setulae arranged in 2-3 irregular rows in upper Il2 or 213; large vibrissa above mouth-edge by distance about ll2-113x width of 3.ant.segm.; facial ridges with 10-15 hairs in double or triple rows to slightly below middle, reaching level of lowermost parafacial hairs; 5-8 subfacials; mouth-edge just visible in profile; arista with longest hairs ll2-215x length of arista. Thorax. Sometimes flattened on dorsum between suture and scutellum; 3+3-4 acr,3+3-4 dc; 5h; l*3 ia; 2 strong + 1-3 weak npl; 5-8 weak ptpl distinguishable among hairs; postalar wall bare, rarely with a few short hairs visible at middle; propleuron, prosternum and basi- sternum setose; posterior spiracle brown to yellowish brown with a few minute setulae near ventral edge; subapical scutellars strong , 7.5-2x length of scutellum; apical scutellars hair-like, well in front of apex; 2-3 latbral scutellars 1l2x Figs. 7, 8. Trypocalliphora braueri; frons . - 7. Male. - length of subapicals; basal scutellars slightly shor- 8. Female. ter than subapicals; lateral slopes of scutellum hairy, underside with small whitish hairs. Legs. Black with some reddish transparency affecting all ocelli (Table 4); parafrontal with setulae for whole / and p surfa ce of. fi; claws slightly shorter than api- length; parafacialia becoming narrower down- cal tarsal segment; r, with a single pv at apical ll3; wards. Wings. Veins more yellow than in male. 12 with stronge st ad as strong as v, both at same Terminalia. Described by Gregor &. Povolny level at apical 113; f, without pv ctenidium api- (1959) and Lobanov (1976). T8 with complete cally ) cxt bare behind . Wings. Veins normally yel- marginal row of setae, not interrupted in middle. lowish brown, more brownish apically, sometimes Eggs.Unknown. all yellow; 1-4 small hairs sometimes present at junction of h and sc on underside of wing) rq+s Larvae. 1st and Znd stage larvae unknown. 3rd above with setulae halfway to r-m; node at base of stage larva: Length Glz mm; width up to 6 mm. rq+s hairy below; costal spine at exit of sc at most No crown of setae on pseudocephalon. Mouth- 2x length of neighbouring spinules; csz bare parts as in Fig. 2l (Rodendorf 1957, ås "Genus below ) cs+with spinules in bas al ll2; bend of m1 as species"; Gregor &, Povolny 1959; Cais 1965; distant from hind margin of wing as from m-cu; Zumpt 7965; Hakanen, Grunin & Nuorteva apical crossvein2.5x length of ml between bend 1e7 4). and m-cu, slightly concave near bend . Terminalia. Surstylus variable (Figs. 9-20); phallosome (Fig. Puparium. Length 5.3-7.6 mm, maximum dorso- 1) normally straight, no angle between basal and ventral depth 2.+3.4 mm (n-10). Brown, trun- apical parts of distiphallus in profile view; cate at both ends, maximum depth a little behind paraphallic processes projecting ventrally slightly middle and with bands of rather small spinules. 4 below ventral surface; phallapodeme as in Figs . + Anterior stigma usually with 3, rarely 2 or 5; ejaculatory sclerite as in Fig. 6. branches (Fig . 22). Posterior stigmata as in Fig. 23. Female. Length: 6.0-8.7 mm . Head. Frons at vertex 0.18-0 22x head width (Fig. 8, TableZ); para- Variation and notes on synonymy frontal with 9-ll inclinate frontal setae, hindmost at level with anterior ocellus, proclinate orbitals, The status of Trypocalliphorø lindneri Peus without prevertical seta, iu and ey strong (Fig. 8); interfrontal stripe below anterior ocellus 1.63- Peus (1960) described a number of supposed mor- 2.28x width of parafrontal (Table 3) and 1.0- phological differences between Trypocalliphora 1.5 x distance between outer rims of posterior lindneri and T. braueri: ENT. SCAND. VOL. 1s (1984)

15 ENT. SCAND. VOL. 1s (1e84) Revision of the bird-parasitic Trypocalliphora 377

Figs. g-20. Trypocalliphora braueri;cerci and surstyli, right (upper) and left (lower) profile views. -9. Holotype of. lindneri (SMN). - L0. Paratype of. lindnerl (MNHU). - 11. West Germany. - 12. U.S.S.R. - 13 , L4. Finland, Salla. - 15. Finland, Sodankylå. - L6. Norway, Al. - 17_20. Norway, Balestrand.

Male surstylus. The type materialof lindneri (Figs. Female frons. The interfrontal stripe was de- 9-10) and other material from the same series scribed as scarcely broader than a parafrontal in (Heinz 1954; Gregor & Povolny 1959) show a va- braueri, and twice as broad in lindneri. The ratio riability of the same range as in Figs . 9-20. The d (Table 3) actually shows a continuous range of paratype in MNHU, oo which Peus based his de- variability. The interfrontal stripe was also description of the surstylus in lindneri, is unusual in scribed as exactly equal to the distance between having a distinct "pråapikale Einschntirung". Evi- the outer rims of the posterior ocelli in braueri, dently the surstylus of the lectotype d of braueri, and distinctly broader in lindneri. This ratio figured by Peus (1960:224, fig. 18), falls within the (Table 4) also shows a continuous range of varia- variational range of that structure in lindneri. bility.

Male frons. The parafrontals were described as Colour of female abdomen. I do not agree with narrowly separated in lindneri, and contiguous in Peus (1960: 224-225) as to colour differences be- braueri. The holotype of lindneri agrees with Peus' tween the type material of braueri and lindneri. description, but one of the paratypes in SMN d Colour of posterior spiracle. Peus gives "hell- parafrontals. Most males have a has contiguous braun" for braueri and "braun" for lindneri. I frons as in Fig. 7. Frons/head ratios (Table 1) show agree with this with regard to the type material, the type material of braueri to lie in the lower part but the difference is slight and Norwegian material and that of. lindneri in the upper part of the varia- shows intermediate colours. bility range. The Norwegian material shows intermediate values. Colour of wing veins.These were described as yel- 378 Knut Rognes ENT. SCAND. VOL. 15 (1e84)

Table 1. Trypocalliphora braueri; males. Ratio frons width (narrowest point)/head width. Specimens with a teneral frons have been omitted.

Origin of material Ratio Austria (type material 0.034, 0.049,0.050 of. braueri W. Germany (type ma- 0.056, 0.060, 0.062,0.064 terial of lindneri) Finland 0.038, 0.045, 0.046,0.049 W. Germany 0.046, 0.050 Norway, Ål 0.050, 0.058 Norway, Balestrand 0.045 ,0.047 ,0.048 ,0.049, 0.051, 0.051, 0.053, 0.054, 0.057,0.060 U.S.S.R. 0.056 Range: 0.034 - 0.064 Mean:0.051 N:26

Table 2. Trypocalliphora braueri; females. Ratio frons width (at vertex)/head width. Specimen from Finland with a teneral head has been omitted.

Origin of material Ratio Austria (type material 0.19, 0.19, 0.19 of.braueri) W. Germany (type ma- a.20,0.21,0.21,0.21 terial of lindneri) Finland 0.22 Norway, Ål 0.19, 0.19, 0.19, 0.19, 0.19 0.20,0.20 Norway, Balestrand 0.18,0.20,4.21 U.S.S.R. 0.20

Range:0.18-10.22 Mean:0.20 N:19

The status of Trypocalliphora lindneri compactø Grunin Grunin (1966) described a subspecies compacta which differed from the nominate form in having Figs. 2L-23. Trypocalliphorø braueri; puparium. - 21. "less curved and thicker harpes not protruding on Mouthparts of 3rd stage larva . - 22. Anterior spiracles. the ventral side of the phallosome" (cited from the 23. Posterior spiracles. - Ent. Rev. Wash. translation). None of the 12 phallosomes I have seen has the paraphallic processes as slender and strongly curved as in Gru- nin's tig.25 which represents the nominate form. low or brownish yellow in braueri and as becoming All have the paraphallic processes shaped as in his darker towards the wing margin in lindneri. Most tig. 26 which represe nts compacta. In the latter fig- specimens examined agree with the description ure, however, the acrophallus is distinctly bent re- applying to lindneri, but a few Norwegian speci- lative to the base of the distiphallus, the un- mens are exactly like the type material of braueri. sclerotised membrane is less distended than in his ENT. SCAND. VOL. 1s (1984) Revision of the bird-parasitic Trypocalliphora 379

Table 3. Trypocalliphora braueri;females. Ratio inter- tion was verified by C. W. Sabrosky who had ac- frontal stripe width/parafrontal width (measured at tip cess to the type material of hirudo in the of ocellar triangle). Parafrontal width defined as half the Despite of difference between frons width and interfrontal stripe U. S. National"Museum, Washington. width. Specimens with a teneral frons have been omit- this Peus (1960) concluded, after comparison with ted. specimens of hirudo (1d 19, alcohol preserved, Sabrosky det.) from Wyoming, U.S.A., and Origin of material Ratio Shannon & Dobroscky's (L924) description, that Austria (typ" material 1.63,1.92,2.00 the Schlorer material from West Germany was of. braueri sufficiently different to require status of a new W. Germany (type ma- 1.81,, 1,.8',7,, 1.87,2.L4 species , lindneri. Apart from minor details con- terial of lindneri) cerning the male and female frons and the male Finland r.77 surstylus which are of no taxonomic value (see above) the differences between lindneri and Norway, Ål 1 .93, 2.00, 2.07, 2.14,, 2.15 hirudo mentioned by Peus concern the colour of Norway, Balestrand 1..92,2.00,2.28 the wing base and squamae and (female only) the U.S.S.R. 2.00 amount of pollinosity on the ocellar triangle. He Range:1.63 -2.28 Mean:1.97 N:17 found additional support in the fact that hirudo in North America apparently infests a wider range of birds of different ecology. Sabrosky (in litt. Table 4. Trypocalliphora braueri; females. Ratio inter- 2.iii.1981) informs me that the colour of the frontal stripe width (at tip of ocellar triangle)/distance squamae of hirudo in the U.S. National Museum posterior with a ten- between ocelli inclusive. Specimens "varies somewhat with the condition of the speci- eral frons have been omitted. mens, and possibly with latitude. I would describe Origin of material Ratio those of the holotype male as light brownish yellow, but I have seen males that would be called (type 1.09, 1.18 Austria material L.00, brown. Females are brighter than of.braueri) brown to dark males, and could often be called yellow, even W. Germany (typ" ma- 1.22,1,.26,1.27, 1.35 the same reared series with dark brown terial of lindneri) females in males. " I regard this information together with the Finland 1.10 present knowledge about morphology and host Norway, Ål I .17 ,1 . L 8, | .26, 1 .26 ,1 .30 range of the Palaearctic species as sufficient evi- Norway, Balestrand 7 .04, 1 .26 ,1 .50 dence for regarding hirudo as identical with U.S.S.R. 7.41 braueri. Range: 1.0-1.50 Mean: 1.23 N:17 Number of branches of the anterior stigma of 3rd stage larva and puparium tig. 25, and the paraphallic processes do not pro- trude on the ventral side. In these respects com- Rodendorf (L957) first described the anterior pacta disagrees with all material I have examined spiracle of the T. braueri 3rd stage larva as 3- and also with material from Poland (Cais 1965). branched. Sabrosky reports (in litt .2.iii.1981) that However, I believe that these differences are the few puparia present in the U. S. National merely artifacts. Museum also have a low number Qa) of spiracu- lar branches. Species of Protocalliphora and other Phormiinae have a higher number (5-15) as far as The status of Protocalliphora hirudo Shannon & this is known (Eng el 1920; S6guy 1941; Hall L948; Dobroscky Rodendorf 1957; Zumpt 7965; Ishijim a 1967).

Specimens of the type series of lindneri and other specimens bred from the same lark were originally Distribution identified by Heinz (1954) as Protocalliphora hirudo Shannon & Dobroscky, a species until then Europe: Austria, Czechoslovakia, East Germany, only known from North America. This identifica- Finland, Hungzty, Norway, Poland, LJ.S.S.R., 380 Knut Rognes ENT. SCAND. VOL. 1s (1984) West Germany. - Asia: U.S.S.R. (east to Vla- Hirundo rustica, Luscinia luscinia and Ficedula divostok). - North America: Alaska to Quebec, hypoleuca are recorded for the first time. south to New Mexico and Georgia. (Rodendorf Acknowledgements. I want to thank the following per- 1957; Peus 1960; Cais 1965; Hall 1965; Grunin sons for loan or gift of material, and other assistance: A. 1966; Draber-Morik o 1971 ; Raus ch 1971 ; Bedard Bernhoft-Osa, A. Brinkmann jr., R. Contreras- & McNeal 1979; Cepelak 1982, and present Lichtenberg, J. P. Dear, L. Greve Jensen, W. Hack- study. man, B. Herting, H. Holgersen, B. Lindeberg, A. ) Løken, J. F. McAlpine, L. O. Odden, A. C. Pont, C. W. Sabrosky, H. Tambs-Lyche and H. Schumann.

Biology and hosts The information given below is compiled from labels on examined specimens, letters from A. References Løken (10.x.1980) and L. O. Odden (7.xii.1980) Bedard, J. & McNeal, J. N. 1979 . Protocalliphora concerning Norwegian material, and from Zumpt hirudo (Diptera: Calliphoridae) infesting Savanna (1956 , 1965),, Rodendorf (1957), Gregor & Sparrow, Passerculus sandwichensis (Aves, Fringil- Povolng 0959), Peus (1960), Cais (1965), Grunin lidae), in Eastern Quebec. -Can. Ent. 111: 111-112. (1966), Grunin, Nuorteva 8L Rajala (1969) Brauer, F. 1883. Die Zweifligler des kaiserlichen , Museums zu Wien. III. Systematischen Studien auf Rausch (1971), Hakanen, Grunin 8t" Nuorteva Grundlage der Dipteren-Larven nebst einer Zusam- (1974), Itåmies & Ojanen (1975) and Bedard & menstellung von Beispielen aus der Literatur tiber McNeal (1979). dieselben und Beschreibung neuer Formen. - Denkschr. Akad. Wiss., Wien 47: l-100. Eggs. Probably laid in clusters directly upon nest- Brauer, F. & Bergenstamm, J. E. von 1894. Die Zwei- lings of various ages, apparently also on newly fltigler des kaiserlichen Museums zu Wien. VII. Vor- arbeiten ztJ einer Monographie der Muscaria hatched ones. Schizometopa (exclusive Anthomyidae). Pars IV. - Ibidem 6l: 537-624. Larvae. These make permanent subcutaneous Cais , L. 1965. Protocalliphora hirudo Shannon & Dob- burrows (up to 12 mm deep and 4.1 mm broad) on roscky, 1924 (Diptera, Calliphoridae) w Polsce. - various parts of the body (from nostrils to feet) but Fragm. faun . 12: 183-191. most frequently on the wings. Extensive tissue de- Cepelak , J . 1982. Some groups of higher flies of the Mal6 Karpaty (Diptera, Brachycera) . II. - Biologia struction may occur. The digestive tract of the lar- Bratisl . 37: 599407. (In Czech, with English sum- vae mostly contains blood. A single infested nest- mary). ling usually supports 9-20 larvae but numbers vary Collin, J. E. t925. Diptera (Cyclorrhapha) from Spits- bergen. Results of the Oxford University Expedition from 1 to 57. Not all nestlings of an attacked brood to Spitsbergen 1924. - Ann. Mag. nat. Hist. (9) 16: may be affected although this is usually the case. 332-337. Duration of the larval stage is 8-9 days. Mature Draber-Moriko , A. 1971. Niekt6re Calyptrata (Diptera) larvae leave the host at death or at fledgling stage Bieszczad6w. - Fragm. faun . 17: 483-543. (In Polish, (thus outside nest). Pupation takes place in the with Russian and German summaries). Emden, F. I. van 1954. Diptera Cyclorrhapha Calyp- ground or in the nest. Infestation weakens the host trata (I) Section (a) Tachinidae and Caltiphoridae. - and is frequently lethal. Nestlings may recover if Handbk Ident. Br. Insects 10 (+) (u): 133 pp. the larvae are removed artificially. Engel, E. O . 1920. Dipteren, die nicht Pupiparen sind, als Vogelparasiten. - Z. Wiss. InsektBiol. 15: 249- Pupal stage. DurationT-19 days. 258. Gregor, F. & Povolny, D. L959. Kritischer Beitragzur Adults. Rarely captured in the open. My§licka Kenntnis der Tribus Phormiini (Diptera, Calli- phoridae). Cas. 26-51. (1968) and Cepelak (1982) reported captures from - dsl. Spol. ent. 56: Grunin,, K. Ya. 1966. New and little known Calli- 8.viii and 3.ix. Bred material has emerged on 10.vi phoridae (Diptera), mainly bloodsucking or sub- - 10.viii. Adults hibernate in unknown locations. cutaneous parasites of birds. - Ent. Obozr. 45: 897- 903. (In Russian, English translation in Ent. Rev., Host range. A wide range of passeriform birds Wash . 45: 503-506). with different nesting habits, even box-nesting 1970. New species of Calliphoridae (Diptera: Calli- - phoridae). Ibidem 49: 471483. (In Russian, En- ones are attacked, but the fly may prefer ground - , glish translation in Ent. Rev., Wash . 49:282-289). breeders in open environments. Known hosts in l97la. Phormiata Grunin, gen. n. - A sixth genus of the Palaearctic Region are listed in Table 5. - the tribe Phormiini (Diptera, Calliphoridae). * Ent. ENT. SCAND. VOL. 1s (1984) Revision of the bird-parasitic Trypocalliphora 381

Table 5. Host list for Trypocalliphora braueri; Palaearctic Region only. The classification and nomenclature of the birds follow Voous (as in Haftorn 1979), vernacular names Peterson, Mountfort & Hollom (1963). For sources, see text.

Host Country PASSERIFORMES Alaudidae Alauda arvensis Sky Lark Austria, Czechoslovakia, West Germany Hirundinidae Hirundo rustica Swallow U.S.S.R. Motacillidae Anthus pratensis Meadow Pipit Finland, U.S.S.R. Anthus trivialis Tree Pipit Poland, U.S.S.R. Motacilla alba White Wagtail Finland, Hungary Motacilla flava Yellow Wagtail ? (Zumpt 1965) Turdidae Luscinia luscinia Thrush Nightingale Finland Oenanthe oenanthe Wheatear Finland, Norway, U. S. S. R. Oenanthe sp. U.S.S.R. Sylviidae Sylvia communis Whitethroat West Germany Phylloscopus bonelli Bonelli's Warbler Austria P hy llo s c o p us tr o chilus Willow Warbler U.S.S.R. Muscicapidae Ficedula hypoleuca Pied Flycatcher Norway Paridae Parus major Great Tit Finland Corvidae Cyanopica cyanus Eastern Blue Magpie U.S.S.R. Sturnidae Sturnus vulgaris Starling U.S.S.R. Passeridae Passer domesticus House Sparrow Austria, E,ast Germany, Czechoslovakia Passer sp. U.S.S.R. Fringillidae Carduelis flammea Redpoll Finland Emberi zidae Emberiza schoeniclus Reed Bunting West Germany Miliaria calandra Corn Bunting Norway

Rev., Wash. 50: 253. (Russian original in Ent. as a subcutaneous parasite of the Wheatear in north- Obozr. not seen). ern Finland. - Ibidem 35: 56-57 . 1971b . Ergebnisse der zoologischen Forschungen Haftorn, S. 1979. Våre fugler. En felthåndbok. - Oslo. - von Dr. Z. Kaszab in der Mongolei. 277. Calli- Hakanen, R., Grunin, K. Ya. & Nuorteva, P.1974.Lar- phoridae (Diptera) II. - Acta zooL hung. 17:255- vae of Trypocalliphora lindneri Peus (Dipt., Cal- 2s9. liphoridae) as subcutaneous pathogens on nestlings 1972. Beschreibung einer ornithoparasitischen of the Meadow Pipit and Common Redpoll in the - Fliege, Protocalliphora nuortevai Grunin, SP. n. subarctic. - Suom. hyont. Aikak . 40: 15-18. (Dipt., Calliphoridae) aus Nord-Finland. - Suom. Hall, D. G. 1948. The blowflies of North America. - hyont. Aikak. 38: 156-158. [6 + ] 477 pp. , 51 pls. , Lafayette, Ind. Grunin, K. Ya., Nuorteva, P. & Rajala, P. 1969. 1965. Family Calliphoridae. - Pp . 922-932, in: Stone, Trypocalliphora lindneri Peus (Dipt., Calliphoridae) - Sabrosky, Wirth, Foote & Coulson, A catalog of the 382 Knut Rognes ENT. SCAND. VOL. ls (1e84)

Diptera of America north of Mexico. - Agric. larval Protocalliphora h. hirudo Shannon and Dob- Handb. 276: iv*1696 pp., Washington D. C. roscky , 1924 (Diptera: Calliphoridae) . - Aquilo Heinz, H. J. 1954. Weitere Vogelblutfliegen (Gattung (ZooL ) 13: 1,4. Protocalliphora, Dipt.) in Deutschland und Oster- Rodendorf, B. B. 1957. On the parasite flies, injurious reich. - Vogelwarte 17: 3942. to nestling singing birds. - Ent. Obozr.36: 11G724. Hendel, F. 1901. Beitrag zur Kenntnis der Calliphorinen (In Russian with English summary. English transla- (Dipt.) - Wien. e.!t. Ztg20:28-33. tion in the British Library also seen). Hennig, W. 1939. Uber Namen und Artenzahl der Sabrosky, C. W. 1967. Coriections to A catalog of the deutschen "Vogelblutfliegen". Arb. physiol. Diptera of America north of Mexico. - Bull. ent. Soc. angew. Ent. Berl . 6:359-364. Am. 13: 115-125. Ishijima, H. 1967. Revision of the third stage larvae of Sabrosky, C. W. & Bennett, G. F. 1958. The utilization synanthropic flies of Japan (Diptera: Anthomyiidae, of morphological, ecological, and life history evi- Muscidae, Calliphoridae and Sarcophagidae). - Jap. dence in the classification of Protocalliphora (Dipt- J. sanit. ZooL 18: 47-100. era: Calliphoridae). - Proc. 10. Int. Congr. Ent. Itåmies, J. & Ojanen ,M. 1975. Trypocalliphora lindneri (19s6) t:163-164. Peus (Dipt. Calliphoridae) infesting a great tit nest- Schumann, H. 1973a. Bemerkungen zum Status der ling(Parus majorL.). - Suom. hyont. Aikak . 41: 42. Gattungen Onesia, Melinda und Bellardia (Diptera, Kano, R. & Shinonaga, S. 1968. Calliphoridae (Insecta: Calliphoridae). -Mitt . zooL Mus. Berl .49:333-344. Diptera). - Fauna japon., 166 pp, 23 pls., Tokyo. 1973b. Revision der palaearktischen Melinda-Arten Lindner, E. 1957. Vogelblutfliegen (Protocalliphora, - (Diptera: Calliphoridae). - Dt. ent. Z. (N. F.) 20: Calliphorinae, Diptera). Ein.Beitrag zur Kenntnis 293-3t4. ihrer Systematik und Biologie. * Vogelwarte 79:8F 1974. Revision der palaearktischen Bellardia-Arten 90. - (Diptera, Calliphoridae). - Ibid em 27: 237-299. Lobanov, A. M. 1976. Opredelitel' samok sem. Cal- S6guy, 8.1928. Etudes sur les mouches parasites. Tome lophoridae (Diptera) fauny SSSR po yaitsekladam. - I Conopides, Oestrides et Calliphorines de l'Europe Opred. Faune SSSR 109: 1-48. occidentale. Recherches sur la morphologie et la dis- My§lick d,, Z. 1968. Much6wki podro dziny Calliphorinae tribution g6ographique des Diptåres å larves para- (Diptera) G6r Swigtietokrzyskich. - Zesz. nauk. sites. - Encycl. ent. (A) 9: 1-251. Uniw. L6dz. (2)28:125*136. (In Polish). 1941. Etudes sur les mouches parasites. Tome II Cal- Patton, W. S. & Cushing, E.C. 1934a. Studies on the - liphorides. Calliphorines (suite), Sarcophagines et higher Diptera of medical and veterinary impor- Rhinophorines de l'Europe occidentale et tance. A revision of the genera of the subfamily Cal- m6ridionale. Recherches sur la morphologie et la dis- liphorinae based on a comparative study of the male tribution g6ographique des Diptåres å larves para- and female terminalia. The genus Chrysomyia sites. - Ibidem 2l: 1436. Robineau-Desvoidy (sens. lat.). * Ann. trop. Med. Shannon, R. C. & Dobroscky, I. D. 1924. The North Parasit . 28: 123-130. American bird parasites of the genus Protocalliphora 1934b. Studies on the higher Diptera of medical and (Calliphoridae, Diptera). - J. Wash. Acad. Sci . 14: - veterinary importance. A revision of the genera of 247-253. the subfamily Calliphorinae based on a comparative Thomas, H. T. 1951. Some species of the blow-fly gen- study of the male and female terminalia. The genus era Chrysomyia R.-D., Lucilia R.-D., Hemipyrellia Phormiø Robineau-Desvoidy (sens. lat.). - Ibidem Tnsd. and Calliphora R.-D. from South-Eastern 28:305-314. Szechuan, China. - Proc. zooL Soc. Lond.721:147- Peterson, R. T., Mountfort, G. & Holloffi, P. A. D. 200. 1963. A field guide to the birds of Britain and Zumpt, F. 1956.64i. Calliphorinae. - In: Lindner, Europe . - 318 pp., London. Fliegen palaearkt. Reg. 1l: l-140, Stuttgart. Peus, F. 1960. Zur Kenntnis der ornithoparasitischen 1965. Myiasis in man and animals in the Old World. Phormiinen (Diptera, Calliphoridae). - Dt. ent. Z. - A textbook for physicians, veterinarians and (N. F.) 7: r93-23s. zoologists. - xv*267 pp., London. Rausch, R. L. 7972. Cutaneous myiasis in a bird by the

Manuscript received Septernber 19Bl . Revised February 1984.

Journal and subscription to reprints of particular groups to be ordered from: Publishing House of the Swedish Research Councils, P.O. Box 6710, S-113 85 Stockholm, Sweden.