Contributions ro a

MAI{TIAL OF PALAEARCTIC DIPTERA (with special reference to flies of economic importance)

Volume 3

Higher Brachycera

Låszl6 Prpp and B6la Darvas (Editors)

Published by

Science Herald, Budapest

l.ggg Issued November 30, 1,998

Original illustrations by Albert Szappanos (Kecskemdt) Jånos Pål (Budapest) and the authors

Cover design: Jånos Lengyel (Budapest)

Illustrations editor: Zsuzsanm E. Prpp (Budapest)

Authorised distributor: E. §ø. Classey Ltd Oxford House l\'larlborough Street Faringdon, Oxfordshire SN7 TP, U.K.

Price subject to change without notice

ISBN 963 04 8835 1, (Series) ISBN 963 04 8 83 8 8 (Volume 3)

O Foundation for the Publicity of the Hungarian Science Baross u. 13, Budapest, PO Box 137, H-1431, Hungary

Åll rights reserved. No part of this book mav be reproduced in any form or by any means' particilarly involving elictronic processing, rvithout written permission from the publisher. Educarional use is permirted for universiry and high school courses.

Type-setting: PARS Ltd, Budapest, Hungary Printing: Dabas-Jegyzer Kft, Dirbas, Hungary

Cover figures: Chrysotoxum elegans Loew and Neomyia cornicina (Fabricius), del. Jånos Pål Contents

3.0. Introduction - L. Papp and B. Darvas 7 3.L. Family Lonchopteridae - M. Bartåk 13 3.2. Family Opetiidae - P. J. Chandler 17 3.3. Family Platypezidae - P. J. Chandler and A. I. Shatalkin 27 3.4. Family Phoridae - R. H. L. Disney 51 3.5. Family Syrphidae - F. C. Thompson and G. Rotheray 81 3.6. Family Pipunculidae - M. Kozånek, M. De Meyer and A. Albrecht 1,4L 3.7. Family Cypselosomatidae - D. K. McAlpine 151 3.g. Family Pseudopomyzidae - D. K. McAlpine and A. I. shatalkin 155 3.9. Family Tanypezidae - J. RohåEek 165 3.10. Family Strongylophthalmyiidae - M. Iwasa 173 3.1,1. Family Psilidae - M. Iwasa 177 3.72. Family Otitidae - L. Greve 185 3.13. Family Platystomatidae - D. K. McAlpine L93 3.74. Family Pallopteridae - B. Merz 201. 3.15. Family Carnidae - L. PaPP 211, 3.16. Family Clusiidae - M. Sasakawa 21,9 3.17. Family Acartophthalmidae - L. Papp and A. L. Ozerov 227 3.18. Family Odiniidae - L. PaPP 233 3.19. Family Tethinidae - L. Munari 243 3.20. Family Canacidae - §f. N. Mathis 251, 3.21. Family Opomyzidae - E. Brunel 259 3.22. Family Anthomyzidae -J. Rohådek 267 3.23. Family Aulacigastridae - L. Papp 279 3.24. Family Periscelididae - §0. N. Mathis and L. Papp 285 3.25. Family Asteiidae - L. Papp 295 3.26. Family Xenasteiidae - L. Papp 305 3.27. Family Nannodastiidae - L. Papp and'W. N. Mathis 309 3.28. Family Milichiidae - L. Papp and T. A. §Theeler 315 3.29. Family Braulidae - L. Papp 325 3.30. Family Teratomyzidae - D. K. McAlpine 331 3.31. Family Coelopidae - D. K. McAlpine 335 3.32. Family Helcomyzidae - D. K. McAlpine 341 3.33. Family Heterocheilidae - D. K. McAlpine 345 3.34. Family Dryomyzidae - A. L. Ozerov 349 3.35. Family Sciomyzidae - R. Rozko§nyi 357 3.36. Family Phaeomyiidae - R. Rozko§nf 377 3.37. Family Lauxaniidae - L. Papp and A. t. Shatalkin 383 3.38. Family Celyphidae - L. Papp 401 3.39. Family Cremifaniidae - L. Papp 409 3.40. Family Chamaemyiidae - I. F. G. Mclean 41,5 3.41. Families of Heleomyzoidea - L. Papp 425 3.42. Family Chyromyidae - T. A. 'Wheeler 457 3.43. Family Sphaeroceridae - J. Rohååek 453 3.44. Family Curtonotidae - L. Papp 497 3.45. Family Drosophilidae - G. Båchli 503 3.46. Family Campichoetidae - P. J. Chandler 515 3.47. Family Diastatidae - P. J. Chandler 523 3.48. Family Camillidae - L. Prpp 531 3.49. Family Ephydridae - '§(/'. N. Mathis and T. Zattwarnicki 537 3.50. Family Anthomyiidae - M. Suwa and B. Darvas 571, 3.5 1.. Family Calliphoridae - K. Rognes 61,7 3.52. Family Sarcophagidae - T. Pape 649 3.53. Family Rhinophoridae - T. Pape 679 3.54. Family Tachinidae - H.-P. Tschorsnig and V. A. Richter 691, 3.55. Family Nycteribiidae - K. Hfirka 829 3.56. Family Streblidae - K. Hfrrka 839 Index - L. Peregovits and L. Prpp 849 3.51 . Family CALLIPHORIDAE

Knut ROCNES

Small to very large robust flies. (Fig. 1) of terior ocellus, but occasionally sexes more or less black or yellow ground-colour, with mostly sil- similar in this respect. Length 4-1'5 mm. very, yellowish or golden microtomentum in a variable paftern, though sometimes undusted; Adult. Head. usually much shorter than high non-metallic or metallic blue, bluish-black, blu- or deepr ås wide as thorax. Frons sloping, anten- ish-green, brassy or green. Some species all yel- nae usually inserted at middle of eye as seen in low. Male frons usually much narrower than in profile. Eyes bare, occasionally hairy. Facets oc- the female , at narrowest even narrower than an- casionally enlarged towards frons and in upper

Fig. 51.1 . Lucilia sericata (Meigen), male. # 11, u u { å 618 Knut ROGNES f

half. Lunula exposed, more or less shinin g, rarely groups siruated much higher than lower facial setose. Frontal vitta of varying width, withour margin (e.g. , Pollenia). small vibrissae above cruciate setae (except in some Mesembrinellinae large one ascending for varying distances ro- - a Neotropical group). Male frons usually only wards lunula. These ascendirrg setae very strong with inner and outer vertical setae, but occasion- in Catapicephala. Scape short, sometimes more ally with a lateroclinate (prevertical) seta, and or less erect. First flagellomere of varying length, even with proclinate orbital setae in some spe- from as long as pedicel (Fig. 2) to 7-B times as cies. Female frons usually with inner and outer long. fuista long- (Fig. 9) or short-plumose (Fig. verricals, a single lateroclinate orbital (preverti- 8), Lare (Figs i-l) or pectinate [is. I).. t# | f cal) and 2 proclinate orbitals, but in Villeneu- with or without a carina. Gena with genal dila- ' uiella with only inner or inner and outer vertical tion below eye. Gena very narrow (e.g., in Ben- setae retained (Fig. 3). Frontal setae reaching galia, Fig. 7) to very broad (Fig. 9), depending pedicel, these setae reduced in Villeneuuiella on size of eye. Occiput concave or convex, more (Figs 24). Fronto-orbital plate usually with nu- or less microtomentose with numerous black or merous setulae beside stronger frontal setae. white setulae, also in upper part, but in Rhiniinae Paraf.acial setose for longer or shorter distances the occiput lacks setulae in upper parr and is al- or bare; in one genus very strong setae in lower most always shining black. Paravertical setae pre- part (Eurychaeta) (Fig. 5). Lower facial margin sent, absent in Rhiniinae. Intrapostocular cilia vertical in profile or more or less projecting as often present benveen posterior eye margin and seen in profile; in Rhiniinae it is often strongly the row of postocular cilia. Proboscis short, but protruding (Fig. 4). Vibrissae srrong, in some lengthened e.g., in some Bellardia and Rhiniinae,

I

I J

Figs 512-9. Hcads of Calliphoridae.2-3: Villeæutiella stackelberyf Grunin. anterior view: 2: male, 3: female. 4-9: male heads in profile: 4z Stomortina lunata (Fabriøts);5: Phormia regina (Meigen);6: Eurychaeta pølpalis (Robineau-Dewoidy); 7z Bengølia lyneborgt James (not Pdaearctic, paratype); 8: An§oneurø acerba (Meigen);9: Calliphorø uralensis Villeneuve (2-3 afær Grunin 1957,4-9 after Rognes l99ta). 3.51. Family CALLTPHORTDAE 619

and very stout palpus in Bengalia (Fig. 7). cla- Polleniinae (Neso dexia, Morinia, and also the vate, moderate to long, sornetimes flattened; Nearctic Melanodexia) and many Rhiniinae the often yellow. anterior lappet is a narrow fringe, whereas the Thorax. Humeral (- posrpronotal) chaeto- posterior one is a larger almost circular flap (Fig. taxy variable: usually 3 serae in a gently curved 15). In other groups the merarhoracic spiracle is line, or in the corner of a right-angled triangle. much larger, and particularly the anterior lapper often a single or several additional setae in front is greatly enlarged, sometimes much larger than of these. The number of humeral setae reduced the posterior lappet (Figs 1s, 18). In some to 2 in e.g. Dexopollenia , and Bengalia. usually Melanomyinae the spiracle is very much re- 34+34 acrostichal serae and 34 + 34 dor- duced. In some Polleniinae and in Catapicephala socentral setae, but often these series are re- there are a few or several yellow or black setae duced. Presutural intraalar setae (- posterior on the anrerior lappet. In Protophormia aticeps presutural intraalar seta) absent or present. No- (Zeæerstedt) the lappets of the rnerarhoracic spir- topleuron with 2 setae, occasionally with supple- acle (and the fringes bordering all other spir"ll.t mental notopleural setae in Protocalliphora and on body) are greatly thickened. In Chrysornyinae Trypocalliphora. Proepisrernal and proepimeral there is a row of long serulae along the an- setae usually present. Meron with a row of teroventral edge of the metathoracic spiracle pro- strong setae below metathoracic spiracle. (Fig. 15). subscutellum somerimes somewhat sternum, proepisternal depression, postalar wall swollen, but never as much as in the Tachinidae. and metasternal '§7ings. area setose or bare in various srem vein with a row of slender setae combinations in the various subfamilies. In- on dorsal surface of wing in Rhiniinae and Chry- frasquamal hairs always present. Coxopleural somyinae (also in Lucilia sinensis Aubertin and streak absent or present (Figs 15-18). Katatergite Pollenia atramentaria (Meigen), Figs zz-23). (supraspiracular convexiry) usually bare, but Costa setulose below ar least ro iunction with hu- with long thin erect hairs in Hemipyrellia. Sup- meral crossvein (Fig. 2{), but for varying dis- rasquamal ridge bare or serose, in Luciliinae a tances beyond this poini accordirrg ro subgroup [ø separate green metallic sclerite is present towards (Figs 23-25). Subcostal sclerite usually bare, but the hind end of the ridge which has a bundle of with setae in some Luciliinae and Polleniinae. strong setae (Fig. 20). Tympanic membrane brr. vein M with a bend that varies from very sharp setose (Figs or 19-21). Ourer posthumeral seta to gently rounded. Often a small appendix is pre- (: anterior presutural supraalar seta) outside of sent at bend. Bend usually rather close to wing line through presutural seta (- posrerior presu- margin (never as widely separated as occurs in tural supraalar seta), but often in line with this some Sarcophagidae). Base of vein R++s usually seta (several Polleniinae) or absent. Inner pos- with setulae at node or for some distance ro- thumeral seta (anterior presurural intraalar seta) wards R-M crossvein. Cell r++5 usually open, but absent, single or double. Anepisternum with a sometimes closed in margin or petiolate. Lower rnore or less complete row of setae along poste- calypter mostly broad with inner edge converg- rior edge. Anepimeron (Figs 10-1 1) usually bare irg with long-aris of fly (Figs 20-21), but nar- in anterior half, but often invaded with hairs in row and rongue-like in Rhiniinae (except in ventral half of anterior parr (e.g., Calliphora), or some Oriental groups) and in some gene ra of the wholly setose (e.g. , chrysomya). scutellum usu- Chrysomyinae, Melanomyinae and Polleniinae ally with 2-S marginal and 1-2 discal pairs of se- (also the orienml genus of Helicoboscinae) (Fig. tae (Figs 12-1,4). Katepisrernal setae usually ar- 19). upp.r calyprer usually bare, but hairy on the ranged in a 2+ 1, fashion, but other pafterns oc- upper surface (as seen when wing is in resting po- cur (1+ 1, 1+ L + 1, 2+ 1+ 1). Merakatepisternum sition) in a few groups (Phormia, Protophormia, bare or setulose. Metathoracic spiracle variable chrysomya) (Fig. 26). Lower calypter often in size, openirg closed by anterior and posrerior more or less hairy on rhe upper surface in all Cal- lappets of varying size. In Helicoboscinae, sorne liphorinae, a few Melanomyinae (Tricycleopsis,

L-__ 620 Knut ROGNES

Angioneura xiniiangensis, and Chrysomyinae the male front tibia is armed with a comb of (Chrysomya) (Fig. 21). short stout spinules on the upper half of inner Legs. Legs moder ately robust. Male claws side, a similar ctenidium is presenr apically on sometimes elongated. Other sexual differences the posteroventral side of the middle femur (Fig. most often found in vestiture of middle dbia, al- 27); the tibiae sometimes also have a tuft of long though in Pollenia some species may show sexual hairs in some species; in the female these fearure dimorphism in other legs also. ln Bengalia sexual are lacking, but the hind femur is often provided differences in l.g chaetotaxy are pronounced: with strong ventral seta which is replaced by

Figs 51.1G-21. Adult morphology of Calliphoridae. 1G-11: anepimeron: 10: Lucilia caesar (Linnaeus); 11: Calliphora uralensis Villeneuve. l2-l4z scutellum: 12: Eurychaeta palpølis (Robineau-Desvoidy); !3: Melanomya ndna (Meigeni; 14: Egisops pecchiolii Rondani. 15-18: area around metathoracic spiracle: 15: Phormia regina (Meigen); 16: Euryciaeta pal- pølis (Robineau-Desvoidy); 17 Pollenia uagabunda (Meigen); 18: Cynomya n ortuonul, Linnaeus. 19-21: area Lehind wing base: 19: Stomorhina lunau (Fabricius); 20: Løcilia caesar (Linnaeus); 21: Cølliphora gendflim (Zetterstedt) (after Rognei 199ta). 3.51 . Family CALLIPHORIDAE 621

hairs in the male. Fore femur with rows of pos- usually erect in the male and more or less decum- terodorsal and posteroventral setae, with partial bent in the female. Sometimes the ventral part of second posterodorsal row. Mid femur usually the tergites has a ground vesrirure that is very with anteroventral and posteroventral rows of dense compared to that of the dorsal surface. rwo to six serae in basal half, rarely with pos- Spiracles 1-5 enclosed in tergites near ventral teroventral row to apex, anteriorly with short margin. Alpha-setae usually present on anterior row at middle and sometimes a preapical sera, margins of sternites, but absent in some groups. posteriorly with diagonal row of three serae near Female terminalia. The females are oviparous apex. Hind femur with complete anterodorsal (e.g., Calliphora), multilarviparous (e.g. , Onesia) and incomplete or uneven anteroventral rows, or unilarviparous (macrolarviparous) (e.g., Eury- with posterovenrral rows confined to basal half. chaeta). ovipositor wirh varying sclerite lengrh. Front tibia with more or less complete antero- In Luciliinae, Chrysomyinae, and mosr Callipho- dorsal row, usually with one or rwo setae poste- rinae (Fig. 34) they are long and tergite 8 is sepa- riorly. Mid tibia usually with one ro several an- rate from the epiproct. In some genera of the terodorsal setae, a ventral seta regularly presenr calliphorinae they are rather short (e.g., Bel- as well as a few in posterodorsal and posterior lardia, Onesia, Polleniop.srs, Tainanina) (Fig. 3T). position. Hind dbia with antero- and posrero- In Melanomyinae they are very long and rergite dorsal rows, in addition to a varying number or 8 is fused with the epiproct (Fig. 3s). In the anteroventral seme of varying size. In a few Pol- Rhiniinae the sclerites are of moderare length, lenia there ate also numerous anteroventral elon- and the marginal serae often extend laterally be- gate hairs aparr from the setae (Figs ZB-29). yond the lateral edge of rhe sclerite ro which they Hind tibia also with anrerodorsal and dorsal pre- belong (Fig. 33). srernite 8 is of full lengrh ex- apical setae, a posterodorsal preapical seta well cept in Chrysomyinae and Nesodexia where it is developed in some genera (Fig. 30). Hind coxa reduced in anterior half, and in Melanomyinae bare or setulose on posterior surface. Tarsi vari- where it is very short or absent. The apex of ster- able in length, most often longer than tibia, but nite 8 is bifid in Calliphorinae (Fig. 3B), but en- in som e Pollenia species males have the hind rar- tire in other groups. In mosr Calliphorinae, all sus shorter than the hind tibia. Luciliinae, and all Chrysomyinae the epiproct Abdomen. Tergite 1,-2 more or less excavated and the cerci are shining black wirhout micro- 'the at base; excavation may or may not reach trichiae. The cercus carries some strong setae hind end of the tergite. Tergite 5 in female some- that tend to be concentrated towards the exter- times cleft mid-dorsally near hind end (Fig. 3Z), nal margin and its upper sur face is provided with tergite 4 or tergite 5 somerimes also provided numerous sensillae (Fig. 34). Such an ovipositor with a comb of exceptionally strong marginal se- tip seems to be characteristic of the saprosarco- tae (Fig. 32). Sternite 1 very short. Sterni te 2 usu- phagous species. In some Calliphorinae (e.g., ally overlapping margins of tergite 1-2, bur other Onesia, Bellardia) the cerci are oval, lack sensil- arrangements occur. sternite 5 in male consisting lae, are provided with long serae and are micro- of a basal piece with rwo posterior more (Fig. 3 1) trichiose (Fig. 37). Such an ovipositor tip seems or less well developed lobes. In one species (Cal- to be characteristic of earthworm parasites or liphora stylifera (Pokorny) from the European predators. In the highly specialised ovipositor of Alps) the basal piece of the male sternire 5 carries the Melanomyinae the cerci are reduced to small a single median long shining posteroventrally di- warts (Fig. 36). In Rhiniinae the ovipositor tip is rected process in addition to the ordinary lobes. often provided with strong spines probably used rn Bengalia the male is provided with a more or for digging. In Helicoboscinae the ovipositor is less square flap-like appendage benveen the srer- very short. In Polleniinae the pleural membranes nite 5 lobes. There are usually some discal setae are extensively microtrichose, but the cerci and on the posterior tergites, but some groups lack epiproct may or may not be microtrichiose. In discal setae altogether. The ground vesrirure is some species of Pollenia the ovipositor tip is pro- 622 Knut ROGNES vided with slender stiff spines. In Bengalia the articulates through its " Gelenkfortsatz" (of sternites of the ovipositor are very broad. Spir- Salzer, "basal apodeme of the gonostylus" of Pape) acles 5 and 7 are located in tergite 6. with the base of the aedeagus. It usually has one Male terminalia. Tergit e 6 a separate sclerite, strong seta, at about middle, and numerous small shaped like a short transverse band. The com- sensillae in its apical half. The aede aga| apoderne pound syntergosternite 7 +8 is large, enclosirrg (phallapodeme) is elongate and flattened dor- spiracles 5 and 7, sometimes carrying peculiar soventrally, in some Rhiniinae it is flattened appendages and projections (Callipl'tora calli- from side to side (stomorbina). Aedeagus (Figs phoroides Rohdendorf). The epandrium is rather 39-50) with a basiphallus and a distiphallus. small. The bacilliform sclerites are usually in the Basiphallus usually short and carries a dorsal shape of rwo parallel widely separate rods, but process, the epiphallus, projecting from its base often they are close to each other, or even fused or near its junction with the distiphallus. The dis- into a single piece (Stomorhina). The pregonite tiphallus has a basal sclerotization - the ventral is a low, more or less triangular sclerite carrying plate - most often encircling the base of the dis- setae. The postgonite (gonosrylus) is slender and tiphallus. A mesohypophallic sclerotization pro-

26

Figs 5122-26. Vings of the Calliphoridae. 22: Stomorhina lunata (Fabricius), from below; 23: Protophormia øtriceps (Zet- terstedt), from above; 24: Lucilia caesør (Linnaeus), from above; 25: An§oneura cyrtoneuina (Zetterstedt), from below; 26: Protophormia atriceps (Zenerstedt), wing base with upper calypter (after Rognes l99la). *l

I I 3.51 . Family CALLIPHORIDAE 623

ceeds forward rnid-ventrally from the basal ring- sclerites through long and slender parastomal like sclerotizatton in most species. Sclerottzatton sclerites (Calliphora, Cynomyo, Protophormia, along dorsal surface of distiphallus proceeds dis- Lucilia) (Figs 51-52) or separated from tentoro- tally and is most often split in rwo paraphallic pharyngeal sclerites (Eurychaeta, Pollenia), processes, which have various shapes: long, very sometimes dorsally concave (i.e., upturned) and slender (Calliphora, Pollenia, Pbormia) (Figs 41, firmly fused to tentoropharyngeal sclerites (Beltardia\ (Fig. rods (Melanomyinae). Segments with varying nurn- {"s 49h hooks 4n; straight (Polleniopsis); large rounded plates serrate along bers of complete or incomplete spine bands or ft edge (Eurychaeta) (Fig. 4?). The aedeagus of additional spines and tubercles. Anterior spiracle Bengalia is of very complex build, whose parts absent, posterior spiracle bilobed. The first instar are not easily homologized with those of other of the rhiniine genus Villeneuuiella has peculiar calliphorids. In Rhiniinae the dorsal surface of appendages on several body segments. the distiphallus is provided with a strong shield- Second instar 2-9 ffiffi, sirnilar to third but like sclerotization (Fig. 39). The ventral surface with weaker mouthparts and rwo slits in poste- of the distiphallus is most often provided with rior spiracles. posteriorly directed denticles, but these may be Third instar 9-22 mm long. Body shape grub- entirely absent in several groups (Melinda, like and rounded at both ends (Boopottus, Try- Pollenia) (Figs 41, 46). The ejaculatory apodeme pocalliphora) or pointed at anterior end and (ejaculatory sclerite) is of varying size, but usu- truncate posteriorly (Fig. 57). Most segments ally small, and more or less triangular (Fig. 39). with more or less complete anterior bands of posteriorly-directed spines and posterior bands Egg. Eggs shining white, in oviparous species of anteriorly-directed spines. Rarely, segments 0.9-L.5 mm long and 0.3-0.4 mm wide. In completely covered with spines (Protocallipho- macrolarviparous species the egg is very large rd), or most segments without spines (Angio- and rich in yolk (Eurychaeta).In lateral view the neur*, Pollenia). The third instar larvae of Chry- egg is flat or slightly concave dorsally and convex somya albiceps §Tiedemann) and C. rufifacies ventrally. Chorion with a reticulate pattern. A (Macquart) are unique in having several fleshy narrow median area mid-dorsally, bounded by protuberances on the body segments. Anterior parallel carinae which unite posteriorly. The me- end in Protocalliphora with a crown of setae on dian area functions as a plastron, enabling the the cephalic segment formin g a kind of "sucker" egg to respire when occasionally submerged in through the centre of which the mouthparts pro- water. trude. Mouthparts well-developed (Fig. 59). Mandibles often large, sometimes with a single Larvae. The larvae of most Calliphoridae are accessory oral sclerite berween them apically. rypical pale maggots with a tapering anterior and Tentoropharyngeal sclerite varying in shape, a truncate posterior end. usually without an "incision" of weak scle rotrza- First instar less than 2 mm long. Mouthparts tion in dorsal arm (except Angioneura and Sro- with well-developed labrum and mandibles (me- morhina). Anterior spiracle variously shaped, dian and lateral hooks) (Calliphora, Cynomld, with numerous openings in a regular row (Fig. Ph ormia, Protopb ormia, Lucilia, Angioneura, Eg- 58) or at the end of an irregularly branched tra- gisops, Melinda) (Figs 51-52), with very srrong cheal tube (Eurychaeta, Booponus). Posterior labrum and smaller, toothed mandibles (Eury- truncate end of maggot with a spiracular field cbaeta), with well-developed labrum and anteri- surrounded by 7 pairs of papillae (Fig. 50), åtr orly reduced mandibles (soine Pollenia) (Figs additional pair of papillae present next to the 53-54), or with strong mandibles and the labrum anus. The spiracular field is usually flat, but vestigial or absent (Bellardia, Onesia, Protocal- sometimes it can be closed by n ffansverse folding liphora, Booponus) (Figs 55-55). Labrum dor- and bringing together of dorsal and ventral sally convex and connected to rentoropharyngeal halves. Posterior spiracles usually flush with sur- s ræ.-H!;s ffi, 624 Knut ROGNES ffi ;.,.nIs

då- rounding area. Ecdysial scar median and more or Puparium. Reddish-brown to black, retaining l# : j ' rl.r less distinct. Peritreme complete (Fig. 61), in- most characters ..;-; of the third instar larva. Puparial :;,.,:r, complete (Fig. 52), or entire spiracular plate horns protruding to various degrees. ',til

'!,, irr. completely sclero tized (Fig. 53). Each spiracle 3::'li. with 3 usually straight, more or less horizontal Biology and behaviour.'§Øhereas rhe adults in slits, angle benveen upper and lower slit much general visit flowers, faeces, and dead animal less than a right angle. occasionally lower slit bodies to obtain nourishmenr, the larval habits vertical and angle more than a right angle are extremely diverse. A survey is given below of (Angioneura, Pollenia, Stomorhina) (Fig. 55). the different kinds encountered among Pa- Sometimes slits slanting strongly downwards lat- laearctic species. The biology of several genera is erally (Booponus) (Fig. 54). not known (Xøntbotryxus, Dexopollenia, Mor-

preap preap

i'l N t§ illil{

Figs 51.27-32.27: Bengølia lyneborgi James (not Palaearctic, paratype), tip of male left middle femur and rrbia.28-29: Pollenia spp., left hind tibia: 28: P. rudis (Fabricus); 29t P, pediculatø Macqaart. 3U Melanodexia grandis Shannon (not Palaearctic), apex of left hind tibia. 31: Calliphora subalpina (Ringdahl), male abdomen. 32: Cøltiphora steluiana (Brauer et Bergenstamm), female tergite 5 with protruding ovipositor (abbreviations: ad: anterodorsal, d: dorsal, pd: posterodorsal, preap: preapical) (28-29,31-32 after Rognes l99la,30 after Shewell 1987). 3.51 . Family CALLIPHORIDAE 625 inia, NesodexiA, Polleniopsis, most Rhiniinae). In least for a certain period, oo the host's dead or Arctic or high alpine areas calliphorids are often living tissue, liquid body-substances, or ingested associated with rodent burrows, which provide food. This phenomenon is called myiasis and is shelter in extreme temperatures and a source of an important aspect of the biology of the Cal- dung and dead bodies. liphoridae. Myiasis gener ally leads to extensive Necrophagy, coProphagy, saprophagy, mli- tissue destruction and in serious cases ultimately asis. Many blowflies are oviparous and l^y their to the death of the victim. eggs on the exposed dead bodies of various ani- In the Palaearctic Region a few blowflies are mals, especially vertebrates, irrespective of size. obligatory myiasis producer s. Lttcilia bufoniuora Being very numerous, their larvae play an ex- Moniez oviposits on living amphibians and its tremely important part in narural communities as larvae cause their eventual death. The subcuta- decomposers and scavengers. It has been esti- neous larvae of Trypocalliphora braueri (Hendel) mated that the offspring of 2000 blowflies need make permanent burrows in the bodies of nest- 3 00 kg of meat in 3 days to mature (Schumann ling and juvenile birds. Booponus larvae are skin L955).In the Palaearctic Region the genera Cal- parasites of bovids, cervids and elephants. B. bo- liphora, Chrysornya, Cynomya, Phormia, Proto- realis larvae are skin parasites of bovids, cervids phormiø, Hemipyrellia and Lucilia belong to this and elephants. B. borealis Rohdendorf develops group, and they make up "1.90/o of the Palaearctic in the soft skin covering the young antlers of the calliphorids (61 of 321, species). Chrysomya al- maral (Centus elaphus Linnaeus, subsp. sibiicus biceps (Wiedemann) may attack and feed on Sev.) and less frequently beneath its skin or be- other blowfly larvae. The time spent in the dif- neath the skin of the elk (Alces Linnaeus). One ferent stages of development varies with species, case is reported in which the larvae even parasit- temperature and other factors. rzed man, a four year old boy (Gornoyunova et Blowflies pose a threat to all stored meat- al. 1973). Booponus inerpectatus (Grunin) de- products, even vacuum-packed ham. It should be velops beneath the skin of the musk deer noted that direct access to the meat is not a con- (Moschus moschiferus Linnaeus) causing subcuta- dition for successful infestation as the female is neous warbles, mainly on the back. able to force the ovipositor through the meshes The Neotropical obligatory myiasis producing of a fly net or through minute holes in plastic chrysomyine blowfly, Cochliomyia hominiuorax wrappirrg covering dishes containing meat and to (Coquerel), the New §forld Screwworm, which drop eggs on to the substrate. affacks livestock, has recently been introduced Species of Calliphora, Chrysomya, Lucilia, into Libya, but swift action with a sterile male and Protophormia are among the first insects to technique seems to have succeeded in eradicating arrive on dead bodies, including humans, and are this livestock pest from the Old World therefore of great significance in forensic m.edi- (Lindquist et al. 1,992). It somewhat resembles cine (cf. Smith 1986). Chrysomya but the greater ampulla lacks long The habit of feeding on faeces and visiting bristly hairs and the palpus is short and filiform. fresh or cooked meat, fish or dairy products, or Facultadve myiasis agents consist of maggots even wounds, make many species of Calliphora, (e.g., of Calliphora, Chrysomya, Cynomya, Pro- Chrysomyd, Cynoffild, Lucilia, Phormia and Pro- tophormia, Lucilia) normally developing in car- tophormia potent vectors of bacteria, viruses, casses or other decayirrg organic matter, which protozoans and helminths causihg various en- occasionally gain access to the living bodies of teric and other diseases in man (.f. Greenberg wild or domesticated animals through oviposi- 1.971). Such species are therefore of the greatest tion in wounds, lesions, natural body openings hygienic importance and pose a permanent or wool, and there cause myiasis. Myiasis has threat to human health and well being. been reported in birds, various mammals, hu- Dipterous larvae may infest the bodies of liv- måns, and even fish (Bristow et al. §9q. In irg human or vertebrate animals and feed, at sheep myiasis, blowflies such as Lucilia sericata 626 Knut ROGNES

År 34 6

§b Gl o*-r Co

Figs 51.33-38. Female terminalia of Calliphoridae. ,33-.35: complete ovipositor: 33: Stomorhina lunata (Fabricius); 34: Protophormia øticeps (Zetterstedt); 35: Melinda gezfiIli Robineau-Desvoidy. 36l. Melinda geztilis Robineau-Desvoidy, ster- nite 8, hypoproct and cerci. 37-38: Bellardiapubicornis-(Zetterstedt): 37: ovipositor tergites, epiproct and cerci 38: sternite 8 and hypoproct (after Rognes t99la). 3.51 . Family CALLIPHORIDAE 627

and L. cuprina are often the primary agents, rype of biology may thus occur in as rnany as whilst other species such as Callipbora uicina Ro- 20.2a/o of the Palaearctic calliphorids (55 of 321, bineau-Desvoidy and C. uomitoria (Linnaeus) species). The adults are commonly found on may oviposit in the lesions caused by the primary flowerirg plants. A number of species are at- attack (secondary and tertiary strike), and if un- tracted to freshly-opened termite mounds in Af- attended the attacks are fatal within a short time. rica and Australia, and females may l^y eggs on Sheep-strike is a serious problem in New Zea- the mound material or in the disrurbed earth land, Australia, South Africa, England, Scotland near the excavation. The egg hatches within a and other sheep-rearing countries. few minutes of deposition into a small acdve first Blood-sucking species. The larvae of Protocal. instar larva (Zumpt 19581 Ferrar 1,987:90). The liphora feed intermittently on the blood of nest- adults of Rhiniinae do not visit filth, carrion or ling birds. This type of biology occurs in 4.70/o food and seem to have no hygienic importance. of the Palaearctic calliphorids (15 of 321, species). Stomorhina lunata (Fabricius) is a predator on Exceptionally a rhiniine species Isomyia tri- locust egg-masses (schistocerca gregaia). The partita (Bigot) is reported to have larval stages Bengaliini are aggressive predators as adults, living as external bloodsuckers of domestic pigs feeding on termites (workers, dealared adults), in China (Chen and Zhi-zi, cited in Ferrar 1,987: lepidopterous larvae, ants and ant pupae or 91). snatching prey from the ants themselves. They Association with gastropods. So far as their bi- are oviparous, and oviposition in loose soil ology is known, all members of the Melanomyi- freshly excavated by dealated termites has been nae appear to be parasitic upon living snails. observed. The eggs hatched almost immediately They are oviparous or viviparous. Melinda spe- and there was s(,me evidence that the larvae were bies place their eggs in the pallial caviry of snails. predacious on termites in the soil. The larva lives as an internal parasite, ultimately causing the death of the snail and devouring it. Classification and distribution. It is by no Members of Helicoboscinae are viviparous and means certain that the Calliphoridae is a valid place a single very large first-instar larva on dy- monophyletic taxon. Hennig (1,973) and Grif- ing or dead Helicidae, which are completely de- fiths (1982) suggested that the family as pres- voured. Catapicephala is probably also a snail ently conceived rnay be paraphyletic. Others (Le- scavenger. It is multilarviparous. Among Palae- hrer 1,970; Rognes L986, l99La; McAlpine arctic calliphorids reproductive dependence on 1989; Pape 1,992) have maintained the view, snails occurs in 10.5o/o of the species (in 34 of based on various characters, that the "Calliphori- 321, species). dae" is a natural group. Parasites or predators of earthworms. At least In a recent attempt at elucidating rhe phylo- three genera are biologically associated with geny of the Calliphoridae by means of cladistic earthworms: Bellardia, Onesia and Pollenia. Pol- analyses of a number of "old" and often-used lenia females lny their eggs on the soil and the characters as well as new ones, Rognes (1997b) newly-hatched larvae find their way ro their concluded that there is no convincing basis on hosts or prey by following the natural spaces in which to maintain it as a monophyletic tæron in the soil. Biological dependence on earthworms its currently accepted sense within the Oestroi- possibly occurs in 3 L.5 a/o of the Palaearcic cal- dea (- Tachinidae family group : Callipho- liphorids (101 of 321 speciesi. roidea). However, I propose that no forrnal ac- Association with termites and other insect lar- tion is taken for the time being to dismember this ude. Very little is known about rhe biology of the apparently polyphyletic group, or ro rearrange Rhiniinae, but all of them seem ro be associated the oestroid tæra accordingly. The srudy of the with the developing stages of inse*s, especi ally phylogeny of the group is still in its infancy and termites, hymenoptera and orthoptera, on which no premature action will be taken in a Manual the larvae feed as predators or parasites. This of this kind. a

628 Knut ROGNES

On the other hand there is good reason to ac- of these groups has been discussed in detail by cept a number of sub taxa, which are given status Rognes (I99'1,a, 1,997b) and by Pape (1992). as subfamilies in this treatment, as monophyletic Note that this classification differs considerably groups. In the Palaearctic Region these are: from the one presented in the Palaearctic cata- Chrysomyinae, Calliphorinae, Luciliinae, Mela- logue (Schumann 1985) or by recent Chinese nomyinae, Helicoboscinae, Bengaliinae (with the treatments (Fan 1"992, 1997). The Chrysomyi- tribes Auchmeromyiini and Bengaliini), Pollenii- nae, Luciliinae and Calliphorinae contain all the nae, Rhiniinae and Ameniinae. The monophyly sarcosaprophagous, mostly oviparous blowflies

.,,tt., d :,r. , il il g 'i ,å t. l

I ,ilf ' 'ii ',t,,';'r- i

:r1lr

i"." I

,,,..ii,.

- -ti,.. .{ .", r,'t .-. ,,; 14 . .i Figs 51.39-50. Aedeagus of the male Calliphoridae. 3}{iU right lateral view: 39: Stomorhina lanata (Fabricius); 40: Pro- .'$ .,'i,å tophormia drtceps (Zetterstedt). 4142l- Polleniø bkolor Robineau-Desvoidy: 41: right lateral view,42: posterior view. 43: .:,-, - 1 Eurychaeta palpalis (Robineau-Desvoidy), right lateral view.4{45t Attgionerra acerba (Meigen)t 44 right lateral view, 45: '; ":,'.',r:tl i posterior view. 46-50: right lateral view: 46: Melinda uiidicyønea (Robineau-Dewoidy); 47: Lucilia seicata (Meigen); 48: 'i'+# Bellørdiø uiantm (Robineau-Desvoidy); 49 Calliphora loeui Errderlein; 50: Cynottrya ,floltuonot, Linnaeus (after Rognes 'ffil1,r*3:1..; ,:.-L i t991,a). :+F. ;.1L : n?le :f i'l ].-- ^t . i;li: 'isilg y'' . i:!i-l: I .;;_ti;" i :{i-}s- 1 {-& * tlxt' $ 3.51 . Family CALLIPHORIDAE 629

which act as decomposers of dead animals in na- structure in taxonomic studies at various levels ture, and which, feeding on faeces, food and lags behind. It will no doubt prove very fruitful flowers, have an enorrnous impact on human hy- and bring about significant progress. The delimi- giene. The chrysomyines are distributed all over tation of genera is problematic within several the tropics and occurs in the southern parts of groups, but especi ally within Calliphorinae and the Palaearctic, whereas Luciliinae mostly occur Melanomyinae. In some cases I have diverged in tempe rate areas. The Calliphorinae occur in considerably from the generic concepts used in all zoogeographical regions. The Melanomyinae the Palaearctic Catalogue (Schumann 1986). are oviparous (e.g. , Melinda) or multilarviparous Boreellus is sunk as a synonyrn of Protophormia (e.g. , Eggisops, Gymnadichosia pusillø Ville- in accordance with Rognes (1991a), but I have neuve [dissected for this work]), and encompass kept Trypocalliphora generically distinct from mostly snail-parasitizing forms. The subfamily is Protocalliphora. Pollenia is most likely a para- widespread in the Palaearctic, Nearctic, Oriental phyletic taxon in its present sense. Regarding Lu- and Australasian regions. The Helicoboscinae is cilia I have followed the Old World tradition and a small group of probably rather ancient origin have not split it up. The monophyly of the genus which is macrolarviparous (unilarviparous), d.- is not firmly established (Rognes l99la: L50). positing a single large lawa on dying or dead Stevens and §7all (1,996) have analysed its phy- snails. It is represented by one genus each in the logeny and found that L. bufoniuorø Moni ez and '§Testern Palaearctic (Eurychaeta) and North- L. siluarum (Meigen) (usually assigned to Bufolu- Eastern Oriental Region (Gulmargia). The Ben- cilia by North American authors) always grouped galiinae is mostly an Afrotropical and Oriental among the species classified by North American group, but the genus Bengalia penetrates into the authors in the genus Phøenicia, giving a further Palaearctic Region in China, and the genus argument against splitting. The concept of sev- Booponus in the east of Russia. Bengølia is asso- eral East Palaearctic genera of Melanomyinae is ciated with termites in their development, adopted from Fan (1992). Thus Paradichosia, whereas Booponus is an obligatory parasite of Gymnadichosia and Pollenomyia, often united in mamrnals in the larval stages. Polleniinae is well Paradichosia in the pffit, are treated as separate represented with several genera in the Palaearctic taxa. Interestingly both the aedeagus, pre- and Region. Among these, Pollenia is very rich in spe- postgonites and spermathecae of Gymnadichosia cies which all seem to be parasites or predators pusilla Villeneuve share peculiarities (possibly of earthworms. This genus is also represented in apomorphies) with those of Angioneura. Melindn the Oriental and Ausrralasian Regions. The Rhini- is treated as separate from these three genera, al- inae is also mosdy a tropical and subtropical though Kurahashi (1970') merged them all. In group, absent from the New Vorld (except for Calliphorinae the nominal genera Abago, Acro- a single introduced species in Bermuda). A few nesia, Aldichitrn, Steringomyia and Ticerato- species penetrate into more temperate areas. Wgd are sunk as synonyms of Calliphora (as sug- Their biology is mostly associated with the de- gested in Rognes 1,99La; see also Schumann and veloping stages of other insects, mostly termites. Ozerov 1" 992). The generic limits of Polleniopsis, The Australasian and Oriental group Ameniinae Onesia, and Bellardia and even Calliphora are is possibly represented by the genus much more problematic than is usually appreci- Catapicephala in the very close neighbourhood ated. A delimitation valid on a wbrld scale must of the Palaearctic region (recorded in the Pa- await a comprehensive cladistic analysis based on laearctic catalogue by Schumann 1985). many more characters than has hitherto been The taxonomy at the generic level in Cal- used. Note that Kurahashi in papers on the Ori- liphoridae is based to a large extent only on ex- ental fauna lumps all species of. Bellardia into a ternal adult characters. The structrrre of the male broadly based Onesia. terminalia is also exploited to a certain degree, " In the k.y below I have included some extra- but not consistently. The use of the ovipositor Palaearctic genera e.g., Catapicephala (Amenii- 630 Knut ROGNES

nae), Tainanina (Calliphorinae), Gulmargia (Heli- Becker, 1,91,0 (Rhiniinae) and Pericallimyiø Vil- coboscinae), since they occur very close to the leneuve, l9l5 (Calliphorinae) in the k.y, even formal, but perhaps unnafural, eastern bounda- though Deeming's (1996) records of species of ries of the Palaearctic Region. I have not in- these genera from Oman (Dhofar region) might cluded the Afrotropical genera Pararhyncomyia be regarded as first records from the Palaearctic

md hy(?+lab)

)) ) )

perit spr op

60 61

Figs 51.51-61. 51-55: anrerior segments and mouthparts of first instar larvjre: 5l-52t Protophormia aticeps (Zetterstedt); 53-54: Polleniø sp.; 55-56: Bellardia wlgais (Robineau-Desvoidy). [-6-5f : Lucilia seicata (Meigen), third instar larva:

6r/ 62 9r A a Er ffi t6e 65 Figs 51.62-65. Postcrior spiracles of third instar Lx,ræ.622 Ptotophormia teriderrouae (Robineau-Desvoidy); 63: Ewychæu palpølis (Robineau-Desvoidy); 64: Booponts borealk Rohdendorf; 652 Pollenia orgtlrtigeu Vainwright (64 after RoMen- dorf 1959, 6243,55 after Rognes l99la).

Region (as this is defined in the Afrotropical and equals Phyto discrepans Pandelld (new synon- Palearctic catalogues). Neither have I included ymy) (holotype and genital slide llo. 249 of D. any of the Ærotropical or Oriental species re- lecheualieri in Paris, examined). corded by him from Oman in the number cf spe- For each genus the k.y below gives the cies given for the various genera in the key. number of species and their distribution in the The genus Diprodexia S6S,ry, 1935, cata- Palaearctic Region, in addition to literanrre for logued in Calliphoridae by Schumann (1985), is the identification of adult specimens. A prelimi- a junior synonym of. Phyto Robineau-Desvoidy nary check-list of Palaearctic species, on which and thus belongs in the Rhinophoridae (new syn- these figures are based, is available from the onymy). Its rype species D. lecheualieri Sig',ry author.

KEY TO GENERA Adults

L. Stem-vein setose above (Figs 22-23)

Stem-vein bare above llucilia sinensis Aubertin (China), bright metallic green species with bare parafacial, andPollenia atramentaria (Meigen) (Europe), non-metallic species with golden yellow crinkly hairs on thorax, have setulae on the stem-vein" but are keyed out here] 19

2. Occiput almost always without setulae in upper half, this area often without microtomentum and therefore shining. Lower facial margin often protnrding well in front of vibrissal corners. Costa hairy below only to the costal break iust beyond the humeral crossvein (Fig. 22), bare below beyond this point. Subcostal sclerite bare RHIMINAE 3 T at * i i I

1 632 Knut ROGNES

.

t

. Occiput with setulae upper part. Lower facial in margin usually not protruding much. Costa I i hairy below as far as junction with Sc, bare below beyond this point Gig. 23). Subcostal sclerite I + .,

setose CHRYSOIvIYINAE 1,4 :

3. Mouthparts strongly reduced (Figs 2-3): mentum shorter than pedicel, palpus shorrer than first flagellomere" which is at most twice as long as wide. Body all yellow. Setosity reduced. Arista bare. Cell t4+S r;yith a long stalk Villeneuviella Austen 8 spp.; North Ærica, Iraq Kazzkhstarq Saudi Arabia, Turkmenistan, Yemen, Oman; Grunin 1957,Ferrar 1987. Mouthparts normal: mentum and first flagellomere longer than antenna. Body usually at least partly with dark ground colour 4

4. fuista pectinate (Fig. a)

Arista bare, pubescent or plumose 9

5. Proepisternal depression setose. Cell r4+5 open Arrhinidia Brauer et Bergensramm 1 tp., A. øbørazs (Schiner); China (Shanghaih Peris 1,952b, Fan L992, 1997.

Proepisternal depression bare. Presunrral acrostichal setae not distinguishable 6

5. Outer posthumeral seta absent. fuista short-pectinate, longest hairs less than half the width of first flagellomere. Lower facial margin not protruding much in front of vibrissal corner. Thorax and abdomen totally glosqr metallic green or purple without distinct microtomentum. Cell r++s long petiol ate Chlororhinia Townsend I sp., Ch. erernpu (Walker); China; Peris 1952b, Dear 1977,Pan 1992, 1997. Outer posthumeral seta present. fuista long-pectinate, longest hairs longer than half the width of the first flagellomere 7

7. Cell r++s with long stalk. Katepisternum glossy in upper half, without microtomenrum. Anepisternum without shining bare spots around base of setae. Abdomen wholly or at least mostly yellow in ground colour.'Proepimeral seta absent Rhinia Robineau-Desvoidy 1 sp., Åå. apicølis (Wiedemann); widespread; Zumpt 1956a.

Cell r++s open or closed in margin. If with a stalk (Stonorhina uibrata (Bigot) and S. mnthogas- ter (Wiedemann)), then pleura with conspicuous shining bare spots around base of setae. Abdo- men usually black with yellow markings (stomorbina xanthogaster has an orange yellow abdo- men, sometimes with a black pattern; S. discolor (Fabricius) also has the abdominal ground col- our yellow)

8. Hind tibia without conspicuous row of anterodorsal setae, but with 2-3 anterodorsal setae as long as or longer than tibial diameter Idiella Brauer et Bergenstamm 3 tpp.; China; Peris 1952b, 7-umpt 19564Far,l, 1992,1997.

Hind tibia with a row.of more or less similar-sized anterodorsal setae, sometimes 2-3 of these longer than the rest stomorhina Rondani 5 spp., widespread; Peris 1952b; Zumpt 1956a.

9. fuista pubescent or bare 10

Arista plumose 11 3.51. Family CALLIPHORIDAE 633

10. Proepisternal depression hairy Metallea Wulp 4 spp.; Egypt, China; Peris 1952b; Fan t99Z (inMetalliopsis). Proepisternal depression usually bare Rhyncomya Robineau-Desvoidy 18 spp', widespread; Peris 1951, 1952b, Rohdendorf l93O,Zumpt 1956a,1958, Zumpt and Tsacas !976, L97å, Gonzales-Mora and Peris 1988, Fan 1992,Deeming 1996,Fan 1997. 1,1,. Outer posthumeral seta absenr Alikangiella Villeneuve 1 sp., Å. uittata (Peris); reaching the Palaearctic in China; Peris 1952a, 1952b, Fan L992, 1997.

Outer posthumeral seta present 1,2

1'2. Presutural acrostichal setae not distinguishable from the ground vestirure Cosmina Robineau-Desvoidy 9 rpp.; North Africa, Middle East, Italy; peris 1952b, zumpt 1956a, Deeming1996.

Presutural acrosrichal setae distinct. Proepisternal depression bare 13

13. Anepisternum with a group of black setae on its upper parr nexr to anterior notopleural seta. Male and female postabdomen of normal size Isomyia Valker 10 spp.; Egypt, China, Japan; Peris 79SZb, Zumpt l9S6a, Zumpt and Tsacas 1,978, Fang and Fan 1985, Fan 1992,1997.

Anepisternum without setae on its upper part next to anterior notopleural seta. Male: postab- domen very strongly developed, almost the same size as the rest of ihe abdomen. Female: last sternite widely uncovered by the corresponding tergites, its posterior border projecting outrvards Strongyloneura Bigot 3 rpp.; Japan, Korea, China; Far. 1992, Chen, Fan and Fang 1992, Fan 1997.

14. No acrostichal, dorsocentral or intraalar setae distinguishable among the thoracic ground vesti- ture. Thorax and abdomen shiny black phor*i"t" Grunin I sp., Ph. phonniata Grunin; Tadzhikistan, China; Grunin l977ai Xiang and Ye 1986; Fn 1992, 1997. - Acrostichal, dorsocentral and intraalar setae well differentiated among dorsal vestiture, at least behind surure 15

15. Lower calypter hairy above (as in Fig. 21), broad with inner edge converging backwards with longitudinal axis of fly. Upper calypter usually with white hairs on lateral fari of upper surface (viewed when wings are in resting position) (as in Fig. 25). §7hole anterior half of anepimeron, including greater ampulla, clothed with erect hairs. Hind coxa with hairs on posrerioi surface. Outer posthumeral seta absent Chrysomya Robineau-Desvoidy 7 spp., widespread; Senior-White, Aubertin and Smart, l94},Zumpt l956a,Zumpt 1956b, Kano and Shinonaga t966.

Lower calypter bare above, narrow with inner edge diverging from longitudinal axis of fly (Fig. 19). Outer posthumeral seta present 16

1,6. Both calypters dark brown. Upper calypter with black hairs on upper surface (viewed when wings are in resting position) (Fig.26). Hind coxa usually with a few hairs laterally on posterior surface. Presutural acrostichal setae absent. 2 postsutural intraalar setae. Very dark bluiih-reen metallic body without microtomenrllm protophormia Townsend 2 rpp., widespread; Rognes 1991a. Both calypters white or yellow, at most weakly infuscated. Upper calypter bare or with white hairs laterally on upper surface (not to be confused with hairs along margin). Presutural acros-

L 634 Knut ROGNES

at least tichal setae conspicuous. Body blue or green metallic with conspicuous microtomentum, on anterior slope of mesonotum L7

lZ. Upper calypter with white hairs on lateral half of upper surface. Presutural acrostichal setae Postalar frååfy *i." long as ground-vestiture (thorax viewed in profile). 1+2 intraalar setae. wall with no-.ro,rl", lorig black hairs. Basicosta and anterior thoracic spiracle yellow. Hind coxa with a few hairs or bare on posterior surface Phormia Robineau-Desvoidy I sp., Ph. re§na (Meigen); widesprcad; Rognes l99la' Upper calypter bare on upper surface. Presutural acrostichal setae longer, about 4 times as long grourrd-restirure GhoÅ viewed in profile). 1*3 intraalar setae. Hind coxa always bare be- ",hind 18 practically 1g. Basicosta, both thoracic spiracles and both calypters golden yellow. Postalar wall bare (a few short inconspicuous hairs sometimes present). Male: fronto

Postalar wall setose (bare in a few small black non-metallic Melanomyinae, which have the coxopleural srreak absent). No very strong setae in lower part of par af.acial 2l

ZO. Eyes bare. Very strong serae in lower half of parafacial. fuista strongly pubescent in basal half. nålpus yellow. Outerlosthumeral seta in line with presutural seta. 2+l+l katepisternal seta9. 5-i pairs of marginal scutellar setae. Lower calypter with inner edge converging backwards with longitudinal axis of fly Eurychaeø B-rauer et Bergenstamm 3 rpp.; §gest Palaearctig incl. North Ærica; Patton 1939, Rognes 1986, l99le, 1993. Eyes with sparse short hairs. At most weak setae in lower part of parafacial. fuista plumose in more than basal trno thirds. Palpus black. Outer posthumeral seta outside line through presutural seta. 1+1+1 katepisternal setae.3 pairs of marginal scutellar setae. Lower calypter with inner edge diverging from longinidinal axis of fly (as in Fig. 19) Gulmargia Rognes 1 .p, G. øngrctisquaaø Rognes, Oriental Region: Northem India and Kashmir; Rognes 1993. 21. Ground colour yellow. Anal vein reaching wing margin. Costa setulose below far beyond iunc- tion with Rr, usually almost to junction with Rz+s. Coxopleural streak absent. Prosternum se- tose. Proepisternal depression bare. Hind coxa bare or setulose on posterior surface. Male some- times with an apical pott.ror"ntral ctenidium on mid femur (Fig.27) BENGALIINAE 22 3.51 . Family CALLIPHORIDAE

Ground colour all black, more or less yellow or all yellow. Anal vein not reaching wing margin. Hind coxa bare behind. Male with no ctenidium apically on posteroventral surface of mid femur 23

22. Arista long plumose. Gena very narrow (Fig. 7), about one ninth of eye height in profile view. Hind coxa hairy on posterior surface. Metakatepisternum setulose. 2 humeral setae. Male ster- nite 5 with median flap-like lobe BENGALIINI Bengalia Robineau-Desvoidy 5 spp.: 4 species possibly reaching the Palaearctic from the Oriental Region in China; 1 Afrotropical species in Oman; Senior-§fihite, Aubertin and Smart 1940, Kano and Shinonaga 1968, Fan 1992, Deeming 1996, Fan 1997. Arista very short pubescent, longest hairs at most as long as width of arista. fuista thickened at least in basal half. Gena broad, 0.4-0.5 times eye height in profile view. Hind coxa bare on posterior surface. Metakatepisternum bare. 3 humeral setae. Male sternite 5 without median flap-like lobe AUCHMEROIvffiINI BooPonus Aldrich 2 spp.; Russia: §fest and East Siberia, Far East; Mongolia; Grunin 7947,1949, Rohdendorf 1959, Gomoyunova, Grunin, Evstigne'eva and Kovalev 1973. A ZwnTt 065 23. Prosrernum and proepisternal depression bare. Non-metallic species, with black or yellow ground colour POLLENIINAE 24

Prosrernum serose (bare in a few small black non-metallic Melanomyinae, which have the coxopleural streak absent). Usually metallic species with black ground colour (a[lyellow inTri- cycleopsis pøradoxa Villeneuve of Melanomyinae) 28

24. Thorax with numerous long golden curly hairs in addition to setae and ground setulae. Metathoracic spiracle large, with anterior lappet about the same size as the posterior one (Fig. L5). Lower calypter usually broad [some species of. Dexopollenia may have a rather narrow lower calypterl 25

Thorax without golden crinkly hairs..Metathoracic spiracle small, with anterior lappet narrow and much smaller than posterior one (as in Fig. 15). Coxopleural streak present. Parafacial bare (except in Morinia skufyini Khitsova from Caucasus and M. piliparafacia Fan from China, with serulae in upper part). Subcostal sclerite without setae among microscopic pubescence 27

25. 1+2 intraalar setae. (1-)2 posthumeral setae, outer very rarely absent, on line with presutural seta. Coxopleural streak almost always present (absent e.g., in P. iaponica Kano et Shinonaga). Hind tibia with posterodorsal preapical setae not differentiated or very much smaller than the anrerodorsal and the dorsal preapicals. Parafacial setulose for whole length. Subcostal sclerite with a bundle of long black or yellow setae among the layer of microscopic pubescence. 3(-5) humerals. Ground colour of abdomen and legs usually black (except Pollenia bicolor Robineau- '§Testern Desvoidy and P. ruficrura Rondani in Palaearctic) Pollenia Robineau-Desvoidy 42 spp.; widespread, but few species in Eastern Palaearctic; Kano and Shinonaga 1966b,1968; Rognes 1987a,

L987b, 1 98 8, 1 99 1 a, 1991b, 1991c, 1992a, 1992b, 1'992c, 1998.

0+2 intraalar setae. 1 posthumeral seta, outer lacking. Coxopleural streak absent or Present. Hind tibia with preapical anterodorsal, dorsal and posterodorsal setae of equal size. Parafacial quite bare or almosr (a few setulae in extreme upper part). Postscutellum slightly swollen. Sub- costal sclerite with or without setae among velvery pubescence 25 §I !rj

636 Knut ROGNES

26. Large species with dense cover of curly golden vestiture. Ground colour black (thoughX bazini (Sdguy) holorype male has some yellow transparency at base of abdomen, especially ventrally). Abdomen with white microtomentose cheiquered pattern . Parafacial bare except for extreme up- permost part where a few short setae usually are present. Subcostal sclerite with numerous black or yellow setae among the microscopic pubescence. 1"-2 presutural acrostichals. 24 humeral setae. Coxopleural streak absent. Lappets of metathoracic spiracle dark brown Xanthotryxus Aldrich 7 tpp.; China, Japan, Korea; Aldrich 1930, Kano and Shinonaga L968, Fan L992, Chen et al. 1992,Fan 1997. Small species. Curly golden vestiture sparse. Ground colour often extensively yellow. Paraf.acial absolutely bare. Subcostal sclerite with only a few pale setae or wholly without setae among microscopic pubescence. 1 presutural acrostichal seta. 2 humeral setae. Coxopleural streak ab- sent or present. Lappets of metathoracic spiracle yellow or dark Dexopollenia Townsend 5 .pp.; China, Japan; Aldrich 1930, Kano and Shinonaga 1968,Far. 1992; Chen et al. L992; Kurahashi 1992; Fan et al. 1993, Kurahashi 1995, Fan 1997. 27. Lower calypter narrow, inner edging diverging backwards from longitudinal axis of fly. Node at base of R++s bare on both surfaces of wing. 0+2 intraalar setae. 1+1 acrostichals. Outer posthumeral absent. Hind tibia with anterodorsal, dorsal and posterodorsal preapical setae of same size. 2 pairs of scutellar marginals. Small dark brownish microtomentose species with un- striped thorax and abdomen without chequered pattern Morinia Robineau-Desvoidy 5 tpp.; Europe, Caucasus, China, Japan; Villeneuve 1936, Herting 1951, Khitsova 1983, Rognes 1991a, Fan 1997. Lower calypter broad, inner edge converging backwards with longitudinal axis of fly. Node at base of R++s setulose on both sides of wing. 1t3 intraalar setae. 3+4 acrostichal setae. Outer posthumeral seta present, on line with presutural seta. Hind tibia with posterodorsal preapical setae about half as long dorsal preapical. 3-5 pairs of scutellar marginals. Large robust species with white microtomentose striped thorax and white microtomentose chequered abdomen Nesodexia Villeneuve 1 tp., N. corsicana Villeneuve; Corsica; Villeneuve l9l1a, b, S€guy 1,928, Zuntpt 19 5 6a.

28. Parafacial bare. Coxopleural streak present. Bright metallic green specles 29

Paraf.acial usually setose at least in upper part (often requires quite careful observation) (but bare in e.g., Angioneura cyrtoneurina (Zetterstedt)). Coxopleural streak absent 32

29. Proepisternal depression ,.toi.. Suprasquamal ridge posteriorly with separate shining green sclerite bearing setae (Fig. 21). Anterior lappet of metathoracic spiracle without vestirure. Facial ridge with very small and inconspicuous setulae ascending at most halfway to lunula LUCIUINAE 3O

Proepisternal depression setose or bare. No separate sclerite at the posterior end of the su- Prasquamal ridge, this ridge bare. Upper corner of anterior lappet of metathoracic spiracle with bundle of long black setae directed backwards AMENIINAE 31

30. Katatergite (sclerite immediately above metarhoracic spiracle) with long upstanding setae. Sub- co§tal sclerite setose. Basicosta black. Hind coxa hairy on posterior surface Hemipyrellia Townsend 2 .pp.; Egypt, China, Japan, Korea; Aubertin 1931, Fan 1997. 637 3.51 . Family CALLIPHORIDAE

Katatergite with short pubescence only, no long upstanding setae. Subcostal sclerite setose or bare. Basicosta black or yellow. Hind coxa bare on posterior surface Lucilia Robineau-Desvoidy 24 spp.; widespread; Aubertin 1,933, Zumpt 1955a, Grunin 1969, Mihålyi 1977, Holloway 1'991, Rognes l99la, "1.994, Fan 1992, L997. 3L. Large species. Ground colour black, brilliant metallic green or blue. Facial carina absent. Strong on facial ridges more rhan halfway to lunula. More than 5-6 humeral setae. erect setae ascending"proepisternal prosternum setose. depression bare. Metasternal area setose. Coxopleural streak present. Outer posthumeral seta outside line through presutural seta. Anterior half of anepi- L.ron invaded irom behind by long setulae ventrally. Subscutellum strongly convex. 3 +4 dor- socentral setae, 3+4 acrostichal setae (anteriormost postsutural acrostichal seta weak) Catapicephala Macquart region are: C. pøttoni Aubertin and Smart 8 spp; Oriental region; species closest to Palaearctic -senior-§(hite, Villeneuve (Taiwan) 1tnåi"r Uttar PradJsh, Mussooriel, C. sinica Fan (China: Guangdong), C. dasyop_hthaåøa Kano and C. taficomri Villeneuve (Taiwan); Townsend L916, 1931, Senior-§7hite et al. L940, Tumrasvin and 1977, Fan 1997, Kurahashi et al. 1997 . Other combination of characters OtheT AMENIINAE Australasian and Oriental region; Crossk.y 1,955, Colless l'998. 32. Lower calypter almost always bare above (hairy in Ticycleopsis from l1p"l - T- tibialis Kura- para- hashi with tlack, T. paradoxn Villeneuve with yellow ground colour of body and legs, T. 6oxah*sthe abdominal tergite 5 long with a transverse row of 8 discal setae, and strong marginal setae on tergite 4 and, tergiie 5; hairs in a small patch also reported f.ot Angioneura xiniiangensis junction Fan, Chen .tY"o from China). Body non-metallic or metallic. Costa hairy below beyond with Rr (only to junction with Rr inT. paradoxa). Aedeagus without denticles on ventral surface of distipirallus. 6vipositor tip with T8 and epiproct fused into a shovel-shaped structure with small wart-like cerci MELANOTMNAE 33

Lower calypter almost always hairy above (a few Bellardia may have a bare lower calypter)- Hairs on lower ."lypt.r usually covering whole disc, but often confined to a smaller area or even oc- casionally Costa hairy below usually only to junction with Rr. Aedeagus with numerous "brånt. denticles on ventral surface of acrophallus and most of distiphallus. In ovipositor tergite 8 and epiproct separate. Epiproct and cerci shining black, but microtrichiose in several genera CALLIPHORINAE 40

33. proepisternal depression bare. Notopleuron bare or with 7-2 odd hairs. Scutellum with apicals *""k o, lacking, usually only a singie additional very much stronger pair in subapical or lateral position (Fig. 13). Body black, non-metallic, grey or brownish microtomentose 34

Proepisternal depression setose. Notopleuron setose. Scutellum with 3-5 strong marginal pairs, apicals strong (Fig. fa). Prealar seta strong, as strong as or stronger than first postsutural dor- såc..rtral r.tr. Lo*"r calypter broad, inner edge converging with longitudinal axis of fly 35

34. Prealar sera srrong, much itronger than hind notopleural seta. fuista long plumose, its width (hairing included)1.5-2 times rhat of first antennal flagellomere. L-Z presutural acrostichal setae conspicuous. 2 postsutural intraalar setael anterior strong postsutural intraalar seta on a line connecting pr""i", and first postsutural dorsocentral seta. Lower calypter narrow' subcircular, T I !

638 Knut ROGNES

inner margin diverging from scutellum, bare above. Posterior spiracle very narrow, compressed dorsoventrally. t1 with 1 anterodorsal seta Melanomya Rondani 1 sp., M. nana (Meigen); Europe, Japan; Rognes 7997a. Prealar seta short and weak, at most as strong as hind notopleural seta. Arista almost bare or short plumose. 0-L presutural acrostichal seta conspicuous. L-2 postsutural intraalar setae; an- terior strong postsutural intraalar seta well behind a line connecting prealar and first postsutural dorsocentral setae. Lower calypter narrow and subcircular or broad, usually bare above. Poste- rior spiracle broader. tr with 1,4 anterodorsal setae Angioneura Brauer et Bergenstamm 4 rpp.; Europe, China (Xinjiang); Herting 1961, Rognes l99la,FanL997.

35. Lunula setulose (very inconspicuous hairs on lunula in Eggisops petiolata (Pape) from Japan and Korea). l+2-3 intraalar setae; if only 2 postsutural intraalars, first postsutural intraalar seta closer to suture than to posterior postsutural intraalar. First postsutural intraalar seta on a line connecting prealar and first postsutural dorsocentral seta. Cell r++s open or closed with a long stalk (E. petiolata), opening or stalk in front of apex of wing. tr with 2-5 anterodorsal setae. Lower calypter bare above Eggisops Rondani 2 spp.; Europe, Korea and Japan; Pape 1,988, Rognes 1991a. Lunul a bare. Cell r4+5 not long-petiolate 36

36, Lower calypter with some black hairs on part of upper surface, often indistinct and easily rubbed off. 0-1*2 intraalar setae. Thorax and abdomen subshining bluish black or restaceous yellow. Legs yellow or black Tricycleopsis villeneuve 2 rpp.; Japan, [China: Taiwan]; Kurahashi 1972, Kurahashi and Suenaga 1991. Lowe r calyptir bare above 37

37. In males some or all of the abdominal sternites 24 with specialised vestiture in the form of bundles or brushes of very strong or densely-set setae mostly in posterior part. '1,-2+1 katepister- nal setae. t+2-3 intraalar setae. Abdomen black or partly testaceous basally. Legs sometimes yellow or brown and in males often with very long setae, very fine towards tip. Male: paraphallic process slender, distally with or without a small hook. Female: sternire 7 sclerotized for its full length Paradichosia Senior-§fhite 7 .pp.; China, Japan; Zumpt 1955a, Kano 1962, Kano and Shinonaga, 1968, Kurahashi 1970, Fan t992; Chen et al, 1992, Fan 1997.

No specialised vestiture on the male abdominal sternites24. Male tibial serae normal 38

38. Legs and base of abdomen yellow. 2*L katepisternal setae. L *2 intraalar setae. Male: aedeagus with very long and strongly curved basiphallus and epiphallus. Distiphallus with broad shield-like paraphallic tips, serrated along their ventral edge. Pre- and postgonites very long and slender; Female: sternite 7 sclerotized for its full length Gymnadichosia Villeneuve 2 spp.; Russia: Maritime Territories, Japan, China; Hori 1951, Kano and Shinonaga 1958, Grunin 1970c, Kura- hashi 1970, Fan 1992, 1997.

Ground colour of thorax and abdomen black. Aedeagus and gonites otherwise 39

39. 1*2 intraalar setae. Abdomen microtomentose with chequered panern. Legs yellowish. Male: aedeagus with a distally massive hook-shaped paraphallic tip and ventral plate strong. Female: sternite 7 sclerotized for its full length pollenomyia S6guy 3 tPp.; Russia: Maritime territories, China, Japan; Kano and Shinonaga 1958, Grunin 1970c, Fan tggZ, t§gi. 3.51 . Family CALLIPHORIDAE 639

0-1+2:3 intraalar setae. Abdomen microtomentose with metallic black, blue or coppery col- ouring. Male: aedeagus distally with slender paraphallic tips. Female: sternite 7 only sclerotized towards base (known f.or M. gentilis Robineau-Desvoidy, M. uiridicyanea Robineau-Desvoidy and M. septentrionis Xue) Melinda Robineau-Desvoidy 10 spp.; widespread; Kano and Shinonaga 1958, Schumann t973, Rognes 199La,Fan 7992,1997. 40. Small size 3-5 mm. First flagellomere very long, 5-5 times as long as broad in male, not quite as long in female. Body blackish with greyish white microtomentum. Abdomen with shifting chequered markings. Arista plumose. Facial carina absent. 2*3 acrostichal setae. 0+3 intraalar setae. Costa hairy below from wing base almost to junction with Rz+r. Lower calypter pale with a small patch of hairs on upper surface, usually only on basal half, sometimes only a few setulae. Lateral margins of abdominal sternites 2-5 overlapping ventral margins of abdominal tergites 2-5. Male: frons very broad, 0.20-0.30 times head width, sometimes with a proclinate orbital seta Tainanina Villeneuve 2 spp. possibly reaching the Palaearctic region in Japan; Kurahashi 1978, L989A i k^ First flagellomere much shorter. Abdominal sternites 3-4 not overlapping margins of abdominal tergites 3-4 41

41". 1(-2)+0 acrostichal setae. Black, partly metallic blue, microtomentose. Sizeg-L1 mm. Parafacial and fronto-orbital plate plate very wide, silvery microtomentose, with black hairs. fuista with very short hairs. Palpus brownish black. 0+2 intraalar setae. §7ing base and calypters pale yel- lowish. Lower calypter with black setae basally on upper surface. Outer posthumeral seta present (the only posthumeral seta). 2 humeral setae. Costa hairy below to junction with Rr. Male: frons at narrowest part 0.30 times head width. A bundle of strong moderately long setae present on tip of lobes of sternite 5. Female: frons 0.41 times head width. Sternite 5 of ovipositor 2 times as broad as long Onesiomima Rohdendorf 1 sp., O. pamiica Rohdendorf; Tadzhikistan, Kirgizia, China; Rohdendorf 1962, Sychevskaya 1968,Fan 1992, 1997.

At least 1 pair of postsutural acrosrichal setae 42

42. 0-1 presutural acrostichal setae. Facial carina usually well-developed. Body dark grey, micro- tomentose. Arista plumose. Outer posthumeral seta absent. 1-3 postsutural acrostichal setae. 0+2 intraalar setae. Lower calypter with a small patch of setulae, usually only on basal half, sometimes with only a few setulae. Male: paraphallic processes short, straight, without distal hook. Mesohypophallic rod long Polleniopsis Townsend 16 spp.; China, Mongolia, Japan, Korea; Villenewe L942, Kano and Shinonaga 1958, Fan 1992, 1997, Chen et al. 1992.

1-2 presutural acrostichal setae. Outer posthumeral seta usually present. 1-4 postsutural acros- tichals. Body metallic dark blue, green or olive green. Lower calypter with hairs on about basal half or more of upper surface 43

43. 3 postsutural intraalar setae..Costa hairy below to junction with Rr (at least European species) or hairy below even beyond this point (at least some East Palaearctic species). First flagellomere much more than 2x as long as pedicel (at least European species). Lower calypter pale with dark hairs on upper surface except on outer and posterior third or half (at least European species). Abdomen olive green metaiiic wrth a shifting pattern. Male: aedeagus with paraphallic processes fused mid-dorsally and apically straight (European and some East Palaearctic species) or distally Knut ROGNES

curved (East Palaearctic species). Mesohypophallic rod long, twice as long as paraphallic proc- esses Onesia Robineau-Desvoidy 19 spp.; widespread; Schumann 1964, 1974, Kano and Shinonaga 1968, Kurahashi and Park 1972, Rognes 1991a, Rudzinski L992,Fan 7992, 7997, Chen et al. 7992, Rognes 1997a. 2 postsutural intraalar setae. Costa bare below beyond junction with Rr 44

44. Costa setulose below to junction with Sc, usually bare beyond this point (thus second costal sector bare below). First flagellomere about 2x as long as pedicel. Lower calypter pale or infus- cate with hairs usually confined to anterior and median half, but bare e.g., in B. mascariensis (Villeneuve), and occasionally bare in B. pubicomis (Zetterstedt). Abdomen usually olive green with shifting pattern. Male: aedeagus with hook-shaped paraphallic tips, mesohypophallic rod much shorter than paraphallic processes (Fig. 48) Bellardia Robineau-Desvoidy 40 spp.; widespread; Schumann 1974a, 1974b, Kurahashi 1985, Rognes l99la, §7u et al. 1991,Far.r.1992, 1997,Fan et al.1993. Costa setulose below betrneen junctions with Sc and Rr (second costal sector setulose below). First flagellomere much more than 2x as long as pedicel 45

45. 3 postsutural acrostichal setae. 0-1 presutural intraalar setae. Lower calypter pale or dark, with hairs on most of upper surface (Fig. 21\. Abdomen blue metallic, more or less microtomentose, with or without a shifting pattern Calliphora Robineau-Desvoidy 23 spp.; widespread; Zumpt 1956a, Kano and Shinonaga 1,958, Rogne s 1991a, Fan 1,992, L997 , Rognes , 1997 a. 1-2 postsutural acrostichal setae. 0 presutural intraalar seta. Lower calypter white 46

46. Male: cerci normal, about as long as surstyli, apex hooked. Aedeagus with complicated distal half. Female: tergite 4 with strong discal and marginal setae; tergite 5 peculiar, very much smaller than tergite 4; cleft mid-dorsally and almost invisible from above. Tergite 6 completely divided into rwo sclerites. Sternite 7 very broad proximally Cynomyiomima Rohdendorf 1 sp., C. stackelbergi Rohdendorf; Russia: East Siberia, Mongolia, China; Rohdendorf 1924, 1962. Male: cerci rudimentary, very short compared to the long shining surstyli. Female: tergite 4 without discal setae; tergite 5 normal in size, well visible and with numerous strong erect discal setae. Tergite 5 a single sclerite Cynomya Robineau-Desvoidy 2 spp.; widespread; Rognes 1991a.

Identification of larvae. A general account of (1,972), Greenberg and Szyska (1984), Erzinclio- the larval morphology of the Diptera and a key glu (1985, 1988), Rognes (1985, 1,985), Smith to families are given by Teskey (1981a, b, I99L), (1985), Shewell (1987), Liu and Greenberg and Ferrar and Smith in Volume 1 of the present (1989), Tantawi and Greenberg (1993). Manual. A detailed survey of the breeding habits and immature stages of cyclorrhaphan families of Species of economic importance Diptera, includi.g the Calliphoridae, has re- Calliphora uicina Robineau-Desvoidy cently been published by Ferrar (1987). Descrip- This is a common blue-bottle fly widely dis- tions of larvae of many Palaearctic species of Cal- tributed throughout the Palaearctic Region and liphoridae are given by Nielsen (1917), Keilin has followed man into most parts of the §7orld. (1919), Hall (L948), Kano and Sato (1,952), It is the cornmonest fly in urban areas in North- Schumann (1954), Rohdendorf (1957), Zumpt, ern Europe. It is attracted to faeces, meat and (1,965), Ishijima (1967), C.pelak and Rozko§ny fruits. The larvae develop in carrion. It may be (1958), Du§ek (L971), Yahnke and George associated with sheep-strike in Britain, but is not 3.51 . FamilY CALLIPHORIDAE 641 able to act as primary invader (Zumpt 1965). It yunova et al. 1,973). Booponus inexpectatus is involved intestinal and urinary myiasis (Zumpt (Grunin) develops beneath the skin of the Musk 1,965). It is the species most commonly found on Deer (Moschus moschiferus Linnaeus) (Musk human corpses in forensic cases. Reiter (1984) Deer Skin Maggot) causing subcutaneous war- has constructed an "isornegalendiagfam" by which bles, mainly on the back, an average of 70FS00 one rnay estimate the age of a larvae from knowl- larvae occur per animal (up to 2000 per animal edge of lengfh and temperature during develop- recorded for some individuals) (Grunin 1949). ment. In the Sikhote-Alin Mountains every musk deer Lucilia sericata (Meigen) and Lucilia cupittø is reported to be infected, even the young born (§Tiedemann) in early summer. The development of the larvae L. sericata is a very common green-bonle fly lasts approximately 2 months. The fully grown in the temperate zone of the Nonhern hemi- larvae drop to the ground where they pupariate. sphere. It has often been confused with Lucilia They hibernate in this stage. B. borealk causes cuprina both of which have great practical sig- economic loss to maral-rearing farms. nificance as primary agents of myiasis in sheep. Pollenø tp. In some sheep farming areas of the Southern The biolory is known only for single members hemisphere it has become, in addition to indige- of the rudis, dasypoda and uiatiø species-groups nous blowflies, a more or less important pro- of Pollenia and published records should be ducer of sheep-strike, as in South Africq Austra- treated with caution as to the identity of the spe- lia and New Zealand. L. seicata is a facultative cies snrdied. Members of the first group overwin- wound parasite also in animals other than sheep. ter as adultr, are on the *it g from early spring It has been recorded in cases of human myiasis to late autumn, have several generations during (Greenberg 1984, Rognes 1,991a), and is fre- the surlmer, and their larvae are parasitic or quently involved in forensic cases. predatory on lumbricid earthworlns. Members Booponus borealis Rohdendorf and Booponus of the uiatica species group seem to have only a inexpectatus (Grunin) single generation each yetr, to overwinter as first The adults of these species have totally yellow instar larvae within an eafthworm host, and to ground colour, and a broad frons in both sexes. devour the host and emerge as adults during the The arista is strongly thickened in its basal 0.7 5 summer. Eggs are laid singly or in batches in and provided with very short hair, whose length small crevices in the upper parts of the soil. The does not exceed the width of the arista itself. The first instar larva hatches in 3 days and locates the gena is about half as high as an eye. The outer host by random locomotion through the soil in posthumeral seta is lacking. The anal vein nanrrally occurring spaces. Larvae penetirate the reaches the wing margin. Third instar larvae are body of the worm and feed on it with only the unique among Palaearctic species in having the posterior spiracles exposed on the surface of the spiracular slits very srrongly slanting downwards worm. The following species have been reported externally (Fig. 53) . Booponus larvae are obliga- as hoss: Eisenia rosea (Savignyl, Nlolobophora tory skin parasites of bovids, cervids and ele- chlorotica (Savigny), Å. caltginosa (Savigny) and phants. The Palaearctic B. borealis Rohdendorf Lumbrians tetrestris Linnaeus. develops in the soft skin coverirrg the young ant- lers of the maral (Centus ehph*s Linnaeus, Accordirrg to Shewell (1987), adult cluster- subsp. sibiicus Sev.) (Red Deer,Skin Maggot, flies may enter buildings in the autumn in huge The Antler-Bud Fly) and less frequently beneath numbers to hibernate. Most of them cannot es- its skin or beneath the skin of the elk (Alces alces øpe from the building the following spring and Linnaeus). There are probably two generations either die by the hundreds in the attics or else annually. The puparial period lasts about 3 find their way into the lower rooms. The accu- weeks" One case is reported in which the larvae mulations of dead flies attract dermestid beedes parasitized man, a four year old boy (Gomo- or other household pests. 642 Knut ROGNES

REFERENCES Aldrich, J. M. 1930. New rwo-winged flies of Deeming, J. c. 199d. The calriphoridae (Di- proc the family Calliphoridae from China . (J. ptera: cyclorrhapha) of oman . FAuna of S. natn. Mus (1): 78 l-S. Saudi-Arabia 1 5 : 254-ZT g . Aubertin, '1,971. D. 1931. Revision of the genus Hemi- Du§ek, J. K.y to larvae. Pages 1,GZ-190, pyrellia proc. Tns. (Diptera, calliphoridae). in Greenberg, B. (ed.): Flies and disease. vol. Zool. Soc.i Lond. 1931: 497-509. 7. Ecology, classification and biotic associa- Aubertin, D. 1933). Revision of the genus Lu- tions. Princeton Universiry press, princeton, cilia (Diptera, R.-D. calliphoridae). Jr. Linn. New Jersey. 855 pp. Soc. Lond. (Zaol.) 38: 389436. Erzinclioglu, Y. z. 198s. Immaftrre stages of Bennett, G. F. and whinporth, T. L. rggl. British calliphora and cynomya, with re- snrdies on the life history of some species of evaluation of the tæ(onomic characrers of Protocalliphora (Diptera: calliphoridae). larval calliphoridae (Diptera). I. nat. Hist. Can. t. hol. 69: Z04B-20S8. 19: 69-96. Bristow, G. A., Berland, B. and Fosså, S. A. Erzinclioglu, Y. Z. 1988. The larvae of the spe- \990. A tirsr case of rnyiasis in fish. /. Parasi- cies of Phormia and Boreellus: Northern, tol. 76: 256-257. cold-adapted blowflies (Diptera: Calliphori- eepelak, J. and Rozko§ry, R. lg68. zur Biono- dae) /. nat. Hist 22: '!,'!,-16. mie der Art Arsoneura cyrtoneurina zet- Fan, z. 1992. Key to comtt on flies of china. terstedg (Rhinophorinae, 1859 Diptera). Acta second edition. shanghai Institute of Ento- zootechnica, gl. Nitra lT: 181-1 mology, Academia Sinica, i-xlviii + ggT pp. Chen, 2., Fan, Z. and Fang, J. lgg?,. Diptera: + 40 plates. lin chinese, with English nores calliphoridae. vol. 2, pages 1183-1 219, in: on new tæraJ Insects of the Hengduan Mountains Reson Fan, z. '1,997. Fauna sinica. Insecta vol. 6. Di- (Dec. 1992). [in chinese, with English sum- ptera: calliphoidae. science press, Beijing. i- maryJ xii + 707 pp. [in chinese with English sum- colless, D. H. lg99. Morphometrics in the ge- mary of keys and new tæ

Gomoyunova, N. P., Grunin, K. Ya-, Evstig- Grunin, K. Ya. 1,970a. 108. Sern. Calliphoridae ne'eva, N. S. and Kovalev, V. G. 1,973. A case Kalliforidy. Vol. 512, pages 607424, in of subcutaneous myiasis in the child caused Bei-Bienko, G. Ya. (ed.): Opredelitel' nasek- by larvae of Booponus borealis Rohdendorf omykh anopeiskoi chasti SSSR. Akad. Nauk (Diptera, Calliphoridae). EnL Obozr. 52: SSSR, Zool. Inst., Leningrad [in Russian] 944-947. [English translation in Entomologi- Grunin, K. Ya. 197Ab. Flies of the family Cal- cal Review, Washington 52: 61,141,3 1'973)j liphoridae (Diptera) new to the USSR. Eltt. Gonzales-Mora, D. and Peris, S. V. 1"988. Los Obozr. 49: 471,483 [in Russian; English Calliphoridae de Espana: L: Rhiniinae y Chry- translation rn Ent. Reu., Wash. 49: 282-2891. Pa- somyinae (Diptera). EOS, l'lødid 64: 9l-139 - Grunin, K. Ya. 1,970c. Species of the genus Greenberg, B. 1984. Two cases of human myi- radichosia S.-'§U'. (Diptera, Calliphoridae) new asis caused by Phaenicia sericata (Diptera: to rhe USSR. Ettt. Reu., Wash. 49: 139-140 Calliphoridae) in Chicago area hospitals. I- [Russian original in Ent. Obozr. not seen] med. EnL 21: 615. Grunin, K. Ya. 1971a. Phormiata Grunin gen. Greenberg, B. and Szyska, M. L. 1984. Imma- n. A sixth genus of the tribe Phormiini ture stages and biology of fifteen species of (Diptera, Calliphoridae). Ent. Obozr. 50: Peruvian Calliphoridae (Diptera). Ann. ento- 4 I I 145 [in Russian; English translation in mol Soc. Am. 77: 488-5 17. Ent. Reu., Wash.50: 2531. Gregor, F. and PovolnY, D. 1959. Kritischer Grunin, K. Ya. l97lb. Ergebnisse der zoologi- Beitrag zur Kenntnis der Tribus Phormiini schen Forschungen von Dr. Kaszab in der (Diptera, Calliphoridae). eat. åsl. Spol. ent- Mongolei. 277. Calliphoridae (Diptera) II. 55: 26-SL. Acta zool. hung. 17: 255-259. Griffiths, G. C. D. 1982. On the systematic po- Grunin, K. Ya. 1972. Beschreibung einer neuen sition of Mystacinobia (Diptera: Calliphori- ornithoparasitischen Fliege, Protocalliphora dae) . Mem. ent. Soc. Wash. 10: 70-77. nuortevai Grunin, sp. n. (Dipt., Calliphori- Grunin, K. Ya. 7947. Podkozhnyi paræit kabargi dae) aus Nord-Finland. Suomen lryhnt. Ai- Pavlovskiomyia inexpectata gen. r., sp. 11. knk. 38: 155-158. (Diptera, Calliphoridae) . ParAzitol. sborn. hol. Grunin, K. Ya. I97 5. Gasterophilidae, Cal- Aknd. Nauk §SSR 9: 183-190. liphoridae, Oestridae, Hypodermatidae (Di- Grunin, K. Ya. L949. Predstavitel' roda Cordy- ptera) of the Soviet-Mongolian expedition, lobia Grtinb. (Diptera, Cdliphoridae) v SSSR. 1969-1971. Vol. 3, pages 620427, in: Nø- Ent. Obozr.30: 440442. sekonye Mongolii. Akad. Nauk SSSR, Zool. Grunin, K. Ya. 1957. Neue Arten der Gatnrng Inst., Leningrad [in RussianJ Villeneuviella Austen (Rhynchoestrus S6grry) Hall, D. G. 1948. The blouflies of North Aner- aus UdSSR und Iran (Diptera, Calliphoridae). iø. Thomas Say Foundation 5 + 477 PP., 51 Ent. Obozr. 36: 538-545. pls, l,.afayette, Indiana. Grunir, K. Ya. 1966. New and little known Cal- Hollowty, B. 1991. Morphological characters liphoridae (Diptera), mainly bloodsucking or to identify adult Lucilia sericata (Meigen, 1826) subcutaneous parasites of birds. Ent. Obozr. and L. cuprina flfiedemann, 1830) (Diptera: 45: 897-903. Russian; English translation Cdliphoridae). N. Z. hol. 18: 415-420. [in 'W'. I. tn Ent. Reu., Wash.45: 503-506I Hennig, 1973. 31. Diptera (Zweifliigler). Grunin, K. Ya. 1969. 172. Calliphoridae, Gas- Hattdb. tuL aQ)A3l:1-337 (Ueferung 20). terophilidae, Oestridae. Ergebnisse der zoo- Herting, B. 1961. 64e. Rhinophorinae. Vol. 9, logischen Forschungen von Dr. Z. Kaszab in pages l-36, in: Lindner, E. (ed.): Die Fliegen der Mongolei (Diptera). Faun. Abh. Staatl. d$ palaearhtischen Re§on. Schweizerbart, Mus. Naturkde Dresden 3: 5-1 1. Snrngart. 644 Knut ROGNES

Hori, K. 1961. Taxonomical snrdies on the Kurahashi, H. 1,967. Snrdies on the calypterate muscoid flies in Japan. IX. A new species of muscoid flies from Japan VI. Revision of the the genus Paradichosia (Dipt., Calliphoridae) tribes Bengaliini and Polleniini of the subfam- from Japan. Scienæ Reports of the l(attazau)a ily Calliphorinae and the subfamilies Ch.y- Uniuersity 7: 103-105. somyinae and Rhiniinae (Dipterq Calliphori- Ishijima, H. 1967. Revision of the third stage d".). kient. W. IØtazøuaUniu. 12:255-302. larvae of synanthropic flies of Japan (Dip- Kurahashi, H. 1970f Tribe Calliphorini from tera: Anthomyiidå€, Muscidae, Calliphoridae Australian and Oriental regions, I. Melinda- and Sarcophagdae). Iap. I. sanit. hol. 18: group. Pacific Insects 12: 519-542. 47-100. Kurahashi, H. 1972. A new species of the genus James, M. T. 1977. Family Calliphoridae. Vol. Tricycleopsis from J"pan (Diptera: Cal- 3, pages 526-556, in Delfinado, M. D. and liphoridae) . Kontyfr 40: 2+26. Hardy, D. E. (eds): A Catalog of the Diptæa KurahæH, H. L978. Tribe Calliphorini from of the Oriental Re§on. Subordcr Cyclor- Australian and Oriental regions. V. Onesia- rhapha (excluding Diuision Aschiu). The Uni- group: Genus Tainanina (Diptera: Callipho- versity Press of Hawaii, Honolulu, 854 pp. ridae) . Pacific lræects 18: L-8. trGno, R. 1962. Notes on flies of medical impor- KurahæH, H. L986. The genus Onesia (Diptera, tance in Japan. Part XVI. Three new species Calliphoridae) from Tsushima, J"pan, with of the genus Melinda (Calliphoridae, Dip- description of a new species. Kontyfi 54: tera) from J"p"n . Iap. J. sanit. hol. 13: 14. 429432. Kano, R. and Sato, K. 1952. Notes on the flies Kurahæh, H. 198 9a. 109. Family Calliphori- of medical importance in Jap.n. (Part VI) dae. Pages 702:718, in Evenhuis, N. L. Larvae of Luciliini in Japan. Iap. I. bp. Med. (ed.): Catalog the Diptera of the Aus- 22:3342. tralasian attd Oceanian"f Regions. Bishop Mu- Kano, R. and Shinonaga, S. 1966a. Notes on seum Press, Honolulu, and E. J. Brill, Leiden, flies of medical importance in Jap"n. Part 1155 pp. XXV. Description of two species belongng Kurahashi, H. 1989b. New record of Tainanina to the genus Protocalliphora (Diptera: Cal- sarcophagoides (Malloch) (Diptera, Catlipho- liphoridae). IaP. I. sanit. hol. 17: 16+168. - ridae) from Japan. Iap. I. Entomol. 57:515. I(ano, R and Shinonagq S. 1966b. Notes on,flies Kurahashi, H. 1992. A new species of Dexo- of rnedical importance in Japan. Part XXV[. pollenia from Sabah, Malaysiq with a key to the Description of a new species belons.ng to the Oriental species. Iap. I. snit. hol. 43: ?3-27. genus Pollenia in Japan (Diptera, Calliphori- Kurahashi, H. 1993. Japanese species of the dae). Iap. I. sanit. hol. 17: 223-225. ornithoparasitic blow flies: Protocalliphora Kano, R and Shinonaga, S. 1968. C,alliphoridae and Trypocalliphora (Diptera: Calliphoridae). (Insecta: Diptera). Fauna Iaponica. Bio- Neut Entomol. 42: 8-15. geographical Society of Japarl. 181 pp., 23 pls. Kurahashi, H. 1995. Two new species of Keilin, D. 1919. On the life history and larval Dexopollenia from Thailand, with a k.y to anatomy of Melinda cognata Meigen (Dip- the Oriental species (Diptera: Calliphoridae). tera Calliphoridae) parasite in the snail He- Iap. I. sanit. Zool. 46: 139-1 44. licella virgata DaCosB, with an account of Kurahashi, H., Benjaphong, N. and Omar, B. the other Diptera living upon r.nolluscs. Para- 1997. Blow flies (Insecta: Calliphoridae) of sitology 11: 430455. Malaysia and Singapore. Raffies Bulletin .of l(Iritsova, L N. 1983. A new species of the genus Zoology 30: 1-88. Morinia (Diptera, Rhiniphoridae [sicJ from Kurahashi, H. and Park, S. H. 1972. A new the Caucasus. Zool. Zh. 62: 1588-1590. [in species of the genus Onesia from Korea (Dip- Russian with English summaryl tera: Calliphoridae) . Kontyfi 40: 22-24. 3.51 . Family CALLIPHORIDAE 645

Kurahashi, H. and Suenaga, O. 199'1,. A new Peris, s. v. '1,956. Nuevas notas sobre Rhiniini record of Tricycleopsis paradoxa Villeneuve con descripcion de formas nuevas. Eos, IvIød- from Japan (Diptera: Calliphorida e). Tropical rid 32: 231-254. Medicine 33: L-5. Peus, F. L950. Zur Kenntnis der ornithoparas- Lehrer, A. 1,97 0. Consi d€,rations phylo gend- tischen Phormiinen (Diptera, Calliphoridae). tiques et taxonornique sur la famille Calli- DL ent. Z. (N. F.) 7: L93-235. phoridae (Diptera). Annotnes zool.-bot., Bra- Ponr, A. c. 1980. 90. Family calliphoridae. tisl. 61: L-5 1. Pages 779-800, in Crossk.y, R. (ed.): Lindquist, D. A., Abusova, M. and Hall, M. J. catalogue of the Diptera of the Afrotropical R. L992. The New §Øorld Screwworm fly in Region British Museum §atural Histo ry), Libya: a review of its introducrion and eradi- London, 1,437 pp. cation . Med. uet. Entomol. 6: 2-8. Reiter, C. 1984. Zum §Tachsnrmsverhalten der Liu, D. and Greenberg, B. !989. Immature Maden der blauen Schmeissfliege Calliphora stages of some flies of forensic importance. vicina. Z. Rechtsmed. 91: 295-308. Ann. ent. Soc. Am. 82: 80-93. Richards, P. G. and Morrison, F. O. lg72. The McAlpine, J. F. 1,989. Phylogeny and classifi ca- egg and chorion of Pollenia rudis (Fabricius) tion of the Muscomorpha. Vol. 3, pages (Diptera: Calliphoridae). Cdn. I. Zool. 50: 1397-15 1 8 , in McAlpine, J. F. er al. (eds): 167 6-1679. Manual of Nearctic Diptera. Research Branch Rognes, K. L985. Revision of the bird-parasitic Agriculrura Canada, Ottawa. Agric. Can. blowfly genus TrFpocalliphora Peus, 1,960 (Di- Monogr. No. 32, vi + 1333-1581 pp. ptera: calliphoridae). Ent. scAnd. 15: 371-392. Mihålyi, F. 1977. A new k.y to Hungarian Lu- Rognes, K. 1986. The systematic position of the cilia species (Diptera, calliphoridae). Annls genus Helicobosca Bezzi with a discussion of hist.-nat. Mus. natn. hung. 59: 181-1 84. the monophyly of the calyptrate families cal- '1,9'1,7. Nielsen, J. C. [§Tithout title, in minutes liphoridae, Rhinophoridae, sarcophagidae and of meeting]. vidensk. Meddr dansk naturh. Tachinidae (Diptera). Ent. scand. 17: 7s-gz. Foren. 68: xix-xxi. Rognes, K. 1,987a. A new species in the interme- Pape, T. L988. First record of Melanomya Ron- dia-group and a new synonymy in the genus dani (Diptera, Calliphoridae) from Japan. Pollenia Robineau-Desvoidy, 1830 (Diptera: Kontyfi 56: 641,-644. Calliphoridae). Sysf. ent. 12: 381-388. Pape, T. L992. Phylogeny of the Tachinidae Rognes, K. 1987b. The tæ(onomy of the pol- family-group (Diptera: calypffarae). Tiidscbr. lenia rudis species-group in the Holarctic Re- Ent. 135: 43-86. gion (Diptera: Calliphoridae). Sys/. ent. t2: Patton, w. s. L939 . The termin aha of the genus 47 5-502. §Tohlfahrtia B. & 8., and those of some al- Rognes, K. 1988. The tæ(onomy and phyloge- lied genera, together with notes on the natu- netic relationship of the Pollenia semicinerea ral grouping of the species of the subfamilies specieryroup (Diptera: calliphoridae). sysf. sarcophaginae and Miltogrammatinae. Bull. ent. 1.3: 315-3 45. §oc. Fouad f' Entomol. ZZ: 67-109. Rognes, K. '1,991a. Blowflies (Diptera, callipho- Peris, '!,9s1. s. v. Descripciones preliminares de ridae) of Fennoscandia and Denmark. Fauna nuevos Rhiniini (Dip. calliphoridae). Eos, ent. scand. 24: !-272. Rognes, K. l991b. Revision of the species of Peris, s. v. 1,9s2a. Notas sobre Rhiniini con de- Pollenia Robineau-Dewoidy described by ca- scripcion de nuevas forma s. Anales de la Es- millo Rondani (Diptera: Calliphoridae). Ent. tacion Experimental de Aula Dei z: zz4-233. scAnd. 22: 355-3 67. Peris, s. v. L9szb. La,subfamilia Rhiniinae (Dipt., Rognes, K. l99lc. Revision of the cluster-flies Calliphorida e). Anales de la Estacion Expiri- of the Pollenia viatica species-group (Diptera: mental de Aula Dei 3: l-224. Calliphoridae). Sysf. ent. 16: 439498. 646 Knut ROGNES

Rognes, K. 1"992a. Revision of the cluster-flies 4'1,: 931,-941 [in Russian; English translation of the Pollenia vagabunda species-group (Di- in Ent. Reu., Wash. 4l:S7G-Sg1]. ptera: calliphoridae). Ent. scand. gs-1,14. 23: Rohdendorf, E. B. 1"9sr. on the parasite flies, in- Rognes, K. lggzb. Revision of the clusrer-flies jurious to nestling singing bir ds. Ent. oborr. 36: of the Pollenia venrurii species-group, with a 1,15-1'24. [in Russian, with English summaryJ cladistic analysis palaearctic of species of Rudzinski, H. G. '1,992. zw unterscheidung d;; Pollenia Robineau-Desvoidy (Diptera: cal- rTeibchen von onesia austri aca villeneuve liphoridae). Ent. scAnd. 233-24g. 23: 1920 und onesia kowarzi villeneuve 1,920 Rognes, K. 1992c. pollenia First record of fulvi- (Diptera: Calliphoridae). Entomologische kit- palpis Macquart, 1835 from spain (Diptera, schrift mit Insektenborse l0z: gd-90. Calliphoridae). Eos, '§(/'., L,Iadid d8: Z0l. sabrosky, c. Bennefr, G. F. and §Thinvorth, Rognes, K. 1993. A new genus of the Heli- T. L. L989. Bird blout (protocailiphora) coboscinae flies from the Himalayas (Diptera: in North America (Diptera: cailiphoridae) calliphoridae, with emended genus sub- with notes 07, the palaearctic speciis. smith- family concepts. Ent. scand. ".å '§Tashington 21: 391,404. sonian Institurion. 3lz pp. Rognes, '/-,994. , K. First record of the sheep Schumann, H. 1954. Morphologisch-systemati- green bonle fly Lucilia cuprina (§Tiedemann) sche srudien an Larven von hygienisch wich- from Europe (Diptera: calliphoridae) with tigen mitteleuropåischen Dipteren der farni- additional spanish records of.calliphoridae, lien Calliphoridae Muscidae. wr'ss. z. [Jniu. :f" Muscidae and sarcophagid".l. Eos, Madrid Greifauald III, 1.9 s 3 s 69:4144. ]alngang I 4 lvIathænatisch- naturutisæn*c.ha{tliche Reihe Nr. 4l s : z4s-27 4. Rognes, K. 1997a. Addirions to the swiss fauna Schumann, H. 1964. Revision der Gatrung one- of blowflies with an analysis of the systematic sia Robineau-Desvoidy, 1g30 (Diptera: position cal- of calliphora srylifera (pokortry, liphoridae). Beitr. Ent. 14: gli-g3g. 1889) includirg a description of the female. schumann, H. 1973. Revision der palaearktis- Mitt. schueiz. ent. Ges. T0: 63-76. chen Melinda-Arren (Diptera: caliiphoridae). Rognes, K. 1997b. The calliphoridae (Blow- Dt. ent. Z OL F.) Z0: Zg3-314. flies) (Diptera: oestroidea) are not a mono- schumann, H. 1974a. Bemerkungen zum Stanrs phyletic group. Cladktics 13: 2246. der Gatnrngen onesia, Melinda und Bellar- Rognes, K. 1.999. 103. calliphoridae. pages dia (Diptera, calliphorida e). Mitt. zool. Mus. 323-325, in: Merz, B., Båchli, G., Haenni; J. Berlin 49 (1973): 333-3 44. G. and Gonserh, Y. (eds) : Fauna Heluetica i, schumann, H. 1,974b. Revision der palaearkti- Diptera checklist, cscF und schweizeri- schen Bellardia-furen (Diptera, Calliphori- sche Entomologis.che Gesellschaft, 369 pp. dae). Da ent. Z. fAL F.) Zl: Z3l-Zgg. Rohdendorf, B. B. 1924. Calliphorinen-studien Schumann, H. 198G. Farnily Calliphoridae. Vol. r. (Dipt., Tachinidae). ht. Mitt. 13: zgl-zgs. 12, pages 11-58, in So6s, Å. and p"pp, Rohdendorf, L. B. B. I 930. calliphorinen-studien (eds): catalogue of Palaearctic Diptetd, m (Dipt.). cal- Zool. Anz.88: 176-1,7g. liphoidae-,sarcophagidae. Akadirniai Rohdendorf, Kiad6, B. B. lgsg. Booponus borealis sp. Budapest, zGS pp. n. (Diprera, calliphoridae), a new parasite of Schumann, H. and Ozerov, A. L. '!,g92. Zum Siberian Red Deer (Cervus elaphus sibiricus systematischen stanrs von Abago rohdendorfi sev.) in southern g: siberia. E?t. obozr. 3 Grunin, 1966 (Diptera, calliphoridae). 583-589. Dt. [English translation in Ent. Reu., ent. Z. AL F.) 39: 403408. Wash. 38: 52 $ S ZBl. s6guy, E. lgzg. Enrdes sur les mouches para- Rohdendorf, B. B. 1,962. Neue und wenig be- sites. Tome I. conopides, oesrrides et Cal- kannte calliphorinen und sarcophaginen (nip- liphorines de I'Europe occidentale. Recher- tera, I-arvaevoridae) aus Asien. Ent. oboir. - ches sur la morphologie er la distribution 647 3.51 . FamilY CALLIPHORIDAE

g€ographique des Diptåres larves parasites. Foote, B. A. (coord.): Order Diptera. Pages 'W'. Encycl. ent. (N 9: 'l'-251,- 590-919, in: Stehr, F. (ed.) : Immature Senior-§7hite, R., Aubertin, D. and Smart, J. Insects.. Kendall/Hunt Publishing CompåtrY, 1940. Diptera Vol. VI. Family Calliphoridae. Dubuqu€, Iowa, xvi + 1'-97 5 pp. In: Fauna of British India. xiii + 288 pp. Townsend, C. H. T. 1915. New genera and sPe- Shewell, G. E. 1987. 106 Calliphoridae. Vol. 2, cies of muscoid flies. Proc. U. S. nat. Mus. pages 1133-1 145 , in McAlpine, J. F- et al. 51: 299- 323. (eds) : Manual of Nearctic Diptera. Research Townsend, C. H. T. 1.931. Notes on Old Vorld Branch, Agriculture Canada, Ottawa. Agric- Oestromuscoid types. Part 1. Annk Møg. nat. Can. Monograph No. 28, vi + 675-L332- Hist. (10) 8: 369-39'1,. '§(/'. Smith, K. G. V. 198 6. A mnnual ,f forensic en- Tumrasvin, and Kano, R. 1977. Two new tomology. British Museum §atural History). species of Catapicephala Macquart from London, 205 pp. Thailand (Diptera: Calliphoridae). lap. J. sanit. '§7a[[, ,1-,996. Stevens, J. and R. Classification of Zool. 28: 127-132. the genus Lucilia (Diptera: Calliphoridae): a Villeneuve, J. 1,9'l.La. Dipterologische Sammel- prelirninary parsimony analysit. .,[. nat. Hist. reise nach Korsika. Tachinidae. Dt. ettt. Z. 30:1087-1094. 7911: 62-1,00. ''1,968. Sychevskaya, V. I. Contributions to biol- Villeneuve, J. L9! 1b. Dipterologische Sammel- ogy and morphology of Onesiomima pami- reise nach Korsika. (Dipt.) (Schluss). Tachini- rica Rohd. (Diptera, Calliphoridae). Ent. dae. Dt. ent. Z. 19IL: L1,7-130. Obozr. 47: 815-823 [in Russian; English Villeneuve, J. 1933. Myodaires sup6rieurs asia- translation rn Ent. Ret)., Wash. 47: 497-501]. tiques nouveaux. Bull. et Annls Soc. r. Ent. Tantawi, T. I. and Greenberg, B. 1,993. Chry- Belg. 73: 195-1 99 . somya albiceps and Chrysomya rufifacies Villeneuve, J. L936. 52. Dipteren, '1,6. Musci- (Diptera: Calliphoridae): Contribution to an dae. In Schwedisch-chinesische wissenschaft- ongoing taxonomic problern. I. med. Ento- liche Expedition nach den nordwestlichen mol. 30: 646-648. Provinzen Chinas, unter Leinrng von Dr. Sven Tawfik, M. F. S. and El-Husseini, M. M. 'l'972. Hedin und Prof. Si.i Ping-chang. Ark Zaol. Life-history of Pollenia dasypoda Portochisky 27l. (34): 1-1 3. [sic], a parasite of the earth worm, Allolobo- Villeneuve, J. 1942. Descriptions de myodaires phora caliginosa (Sav.). Bull. Soc. ent. Egypte supdrieurs nouveaux (Dipt. Tachinidae) . Bull. 55: 275-287. Soc. ent. Fr. 47:50-55. Teskey, H . J. 1981.a.3. Morphology and termi- §ilu, Y., Chen, Z. and Fan, Z. 199'l-.. A new nology larvae. Vol. 1, pages 55-88, in species of the genus Bellardia R.-D. (Diptera: McAlpine, J. F. et al. (eds): Iulanual of Ne- Calliphoridae) from Shaanxi, China). Ento' arctic Diptera. Research Branch, Agriculrure motaxonomia 13: 307-309 [in Chinese with Canada, Ottawa. Agric. Can. Monograph English summaryJ. No. 27, vi + I-674. Xiang, C. and Ye, Z. 1973. Description of the Teskey, H. J. 1981b. Key to families larvae. female of Phormiata phormiata Grunin, 1,971 Vol. 1., pages 125-147, in McAlpine, J. F. et (Diptera: Calliphoridae) . Acta zootaxon. sin. al. (eds): Manual of Nearctic Diptera. Re- 1t: 111-1 72. '1,972. search Branch, Agriculture Canada, Ottawa. Yahnke, §7. and George, J. A. Rearing and Agric. Can. Monograph No. 27, vi + l-674. immature stages of the cluster fly (Pollenia Teskey, H. J. '1,991. Introduction + K.y to rudis) (Diptera: Calliphoridae) in Ontario. families of larvae. Vol. 2, pages 690-730, in CAn. Ent 104: 567-576. Knut ROGNES

Zumpt, F. 1955a. 64i. Calliphorinae. Vol. 11, Zumpt, F. 1965. Myiasis in man and animals of pages 1-14A, in Lindner, E. (ed.): Die Fliegen tbe old world. A textbook fo, physicians, uet- dcr palaearktischen Regioa. Schwe izerbart, Snrtt- erinarians and zoologisls London, iv + 26T pp. gatt. Zampq F. and Tsacas, L. 1976. The Rhyncho- Zumpt, F. 1,955b. Calliphoridae (Diptera Cy.- mya callopis-group sensu S6Sry (Diptera: Cal- lorrhapha) . fa* I: Calliphorini and Chryso- liphoridae). /. nat. Hist. 10: 347-349. myiini. Explor. Parc. natn. Alb;ert. Miss. G. F. Zumpt, F. and Tsacas, L. 1,978. Ta>