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Fauna ofNew Zealand


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Editorial Advisory GroupMem bers at Entomology DivisionDepartment of Scientific and Industrial ResearchMount Albert Research Centre

Private Bag, Auckland, New ZealandEx officioDirector - Mr J. F. LongworthGroup leader,* Systematics Section - Dr G. W. Ramsay

Co-opted from w ithin Sy stematics SectionDr T. K. Crosby, Dr B. A. HollowayUniversities representative*

Dr G. W. GibbsZoology Department, Victoria University of WellingtonPrivate Bag, Wellington, New ZealandMuseums representative*

Dr J. C. YaldwynDirector, National Museum of New ZealandPrivate Bag, Wellington, New ZealandOverseas representative*

Dr J. F. LawrenceCSIRO Division of EntomologyP.O. Box 1700, Canberra City, ACT 2601, Australia*on a rotational basis

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Series EditorMr C. T. DuvalSystematics Section, Entomology DivisionDepartment of Scientific and Industrial ResearchMount Albert Research CentrePrivate Bag, Auckland, New Zealand


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Fauna of New ZealandNumber 8

Calliphoridae( I n s ec ta : D ip te ra )

James P . Dear*Department of Entomology

British Museum (Natural History)Cromwel l Road, London SW7 5BD, England

with three contributed sections:Immature stages of New Zealand Calliphoridae

by . . Holloway1Fly strike in New Zealandby . C. G. Heath 2

R e a r i n g C a l l ip h o r i d a eby Pritam Singh1

* Present address: Devonport Cottages, Main Road, Stickney, Lincs 22 8 , England Contribution seen through press by A . C. Pont, of this address, to who m correspon denceconcerning it may be addressed

1 Entomology Division, DSIR, Mt Albert Research Centre, Private Bag, Auckland, New Zealand2 Wallaceville Animal Research Centre, Ministry of Agriculture and Fisheries, Private Bag,

Upper Hutt, New Zealand


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Frontispiece Calliphora quadrimaculata resembles Northern Hem isphere members of its genus andyet is confined to New Zealand. It epitomises the biogeographical complexities of the subregion.Artist: D. W. Helmore.


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Cataloguing-in-publication citationDEAR, James P.Calliphoridae (Insecta: Diptera) / James P. Dear.- Wellington : DSIR, 1985.(Fauna of New Z ealand, ISSN 0111-5383; 7)ISBN 0-477-06764-6I. TitleII. SeriesUDC 595.773.4

Date of publication: see back cov er of N um ber 9

Suggested form of citationDear, J.P. 1985: Calliphoridae (Insecta: Diptera). Fauna of N ew Zealand[number] 8 .

This publication was produced by offset lithography. The author's textwas supplied as typescript, and after editing was keyboarded on to aword processor. Style coding was added, and the de finitive text wasphototypeset at the N.Z. Government Printing Office. Times New Romantype is used for most of the text; major headings and figure labels areset in Univers; Garamond and Geneva are used in the titles.The E ditorial Advisory Grou p and the Series Editor acknowledgethe following co-operation.DS IR, M t A lbert Research Centre:Mr D.W. Helmore - insect habitus drawings, figure labellingMrs L.S. Howey - word processor inputMr M.B.B. Irwin - photoreduction of line figuresMrs H.A. Whelan - computer operation and file managementScience Information Publishing Centre, DSIR :Dr N. Hawcroft - style coding and phototypesetting

-Front cover: The insect depicted is Calliphora quadrimaculata(Swederus), female

Crown Copyright

Published by Science Information P ublishing Centre, DSIRP.O. Box 9741, Wellington, New Zealand


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ABSTRACTNew Zealand's Calliphoridae are revised. Keys are given to genera and species, andcharacters of diagnostic and system atic importance are illustrated. All taxa are fullydescribed, and (where known) reference is made to their biology. The 7 generaincluded contain 54 species, of which 31 27 in Pollenia and 4 in X enocalliphora are new. Huttonophasia Curran, 1927 is synonymised under genus Pollenia Robi-neau-Desvoidy, 1830. Pollenia demissa var. minor Malloch, 1930 is absorbed intoP. demissa (Hutton, 1901). Calliphora ruf ipalpis Macquart, 1851 becomes a juniorsynonym of Xenocalliphora hortona (Walker, 1849). Two new combinations areproposed: Gymnophania pernix Hutton, 1901 is transferred to Pollenia, and Cal-liphora flavipes Lamb, 1909 is moved into X enocalliphora. Three names Polleniaatrifemur Malloch, 1930, Pollenia demissa var. cuprea Malloch, 1930, and Calli-phora nothocalliphoralis Miller, 1939 are considered to be of uncertain identity.Lectotypes are designated for 10 nominal species: Calliphora hilli Patton, 1925;Calliphora dasyphthalma Le Guillou, 1842 (and Macquart, 1843); Pollenia auro-notata Macquart, 1855; Sepimentum fumosum Hutton, 1901; Calliphora antipodeaHutton, 1902; Calliphora eudy pti Hutton, 1902; Calliphora rufipalpis Macquart, 185 1;and Calliphora aureopunctata Macquart, 1855. Sections by contributing authors coverthree topics: immature stages and life history (B.A. Holloway); fly strike (A.C.G.Heath); and rearing (Pritam Singh). A note by K. Rognes reports the occurrence oftwo Pollenia species in the rudis-group, apparently recently introduced from theNorthern Hemisphere.

CHECKLIST OF TAXA(Note: the many names in synonymy are listed in the Taxonomic Index, p. 82)

PageSubfamily CALLIPHORINAE21Genus Calliphora Robineau-Desvoidy, 18 3021hilli Patton, 192522quadrimaculata (Swederus, 1787 )23stygia (Fabricius, 178 1)24vicina Robineau-Desvoidy, 18 3026Genus Hemipyrellia Townsend, 191 827ligurriens (Wiedemann, 18 30)27Genus Lucilia Robineau-Desvoidy, 18 3027cuprina (Wiedemann, 1830)27sericata (Meigen, 1826)27Genus Pollenia Robineau-Desvoidy, 18 3028advena new species32aerosa new species33antipodea new species34astrictifrons new species34atricoma new species34commensurata new species35consanguinea new species35consectata new species35demissa (Hutton, 1901 )36dysaethria new species37dyscheres new species37enetera new species38eurybregma new species38fulviantenna new species38 Pagefumosa (Hutton, 1901 )39hispida new species39immanis new species40insularis new species40lativertex new species41limpida new species41nigripalpis new species41nigripes Malloch, 193042nigrisquama Malloch, 193042notialis new species43opalina new species43oreia new species43pernix (Hutton, 1901) newcombination44primaeva new species44pulverea new species45sandaraca new species45scalena new species46uniseta new species46New Zealand Pollenia of uncertain identity46Two Ho larctic species of Pollenia recentlyintroduced to N ew Zealand (note by K.Rognes)47Genus Ptilonesia Bezzi, 192747auronotata (Macquart, 1855)47

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Genus X enocalliphora Malloch, 192448antipodea (Hutton, 1902)49clara new species50divaricata new species51eudyptis (Hutton, 1902)51flavipes (Lamb, 1909) newcombination52hortona (Walker, 1849)53neozealandica (Murray, 1954)54solitaria new species55vetusta new species56viridiventris Malloch, 193056"Calliphora nothocalliphoralis" Miller56Subfamily CHRYSOMYINAE57Genus Chrysomya Robineau-Desvoidy, 183057megacephala (Fabricius, 1794)57rufifacies (Macquart, 1843)58CONTENTS

PageAcknowledgments7Introduction7Genus Calliphora8Genus Hemipyrellia9Genus Lucilia9Genus Pollenia9Genus Ptilonesia10Genus X enocalliphora1 1Genus Chrysomya1 1Immature stages of New Zealand Calliphor-idae, by B. A. Holloway12Fly-strike in New Zealand, by A. C. G. Heath15Rearing Calliphoridae, by Pritam Singh19Text conventions: abbreviations21Key to genera of Calliphoridae knownfrom New Zealand21Descriptions (see Checklist of taxa)21References59Illustrations62Taxonomic index82ACKNOWLEDGMENTSI am very grateful to the following colleagues whohave lent me material for study from the collec-tions in their care: Dr R. J. Gagn (USNM ); Dr R.A. Harrison (LCNZ); Dr B. A. Holloway (NZAC);Dr R. Contreras-Lichtenberg (NHMW); Dr L.Lyneborg (ZMKD); L. Matile (MNHN); Dr K.R. Norris (ANIC); and Mr R. A. Savill (CMNZ).I am also grateful to Mr Adrian Pont (BM NH) andLoic Matile for their help with historical facts aboutthe Bigot and Macquart collections, and to RoyHarrison for the useful suggestions he suppliedwhile the work was in progress.

This work was originally conceived as a straight-forward systematic revision for publication as ajournal paper. I welcome the opportunity to haveit remodelled as a Fauna contribution, and amindebted to the authors of the new text sections fortheir willingness to contribute. Adrian Pont haskindly acted on my behalf to see the publicationthrough its processing stages into print, and withBeverley Holloway has read the MS. and proofs;their help has been of exceptional value.

INTRODUCTIONThe dipteran fam ily Calliphoridae, to which belongthe familiar blowflies, bluebottles, and greenbot-tles, is represented in New Zealand by 52 species.These are contained in 2 subfamilies comprising 7genera. Six of the genera Calliphora, Hemipy-rellia, Lucilia, Pollenia, Ptilonesia, and X enocalli-phora belong to the subfamily Calliphorinae. Theseventh, Chrysomya, belongs to the Chrysom yinae,a group represented here by only two species. Tw oother subfamilies, the Ameniinae and Rhiniinae,have been recorded from New Zealand by earlyworkers, but these records are based on erroneouslocality data (see Miller 1950, pp. 138, 139, and142).In general appearance the New Zealand speciesof Calliphoridae are very much like the othermembers of this large and cosmopolitan family,which are mostly stout, large to moderate in size,and often metallic in appearance, at least on theabdomen. T hey can be distinguished from all othercalyptrate f l ies in New Zealand by the presence ofa row of hypopleural setae, an undeveloped post-scutellum, and a haired propleuron (if the prop leu-ron is bare then the outer posthumeral sets isabsent). However, in structural details some of thespecies differ from the norm al calliphorid pattern.These anomalies are discussed below under thegenera Pollenia, Ptilonesia, and X enocalliphora.There is a long history of work on N ew Zealand'scalliphorids. The first species was described bySwederus (1787) from material collected duringCook's first voyage to New Zealand. By the earlytwentieth century a few species had been describedby European workers such as Macquart and Walker,and Hutton had recognised species endemic to theoffshore islands. Subsequent workers such as M al-loch (1924, 1927, 1930), Miller (1939b), and Mur-ray (1954) continued to describe new species, andin general they had a good eye for valid species,although they often confused the nomenclaturebecause of the inaccessibility of much type materialin European museums at that time. Miller (1939b,p. 11) gave a very interesting account of the history

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of blowflies in New Zealand, including their sig-nificance in M aori lore. Harrison (1976, p. 147) hasgiven a useful revision of the species from thesouthern islands.Calliphorids are generally ubiquitous. They flymainly by day, though a number of specimens Ihave examined were collected at light traps. Mostspecies of Pollenia appear to be restricted to higheraltitudes, whereas Ptilonesia an d X enocalliphora are

often found at the seashore. Adults are oftenattracted to sweet liquids, and can be collected atflowers, where they feed on nectar. They also feedon the liquid products of organic decomposition,which provide the proteins essential to the femalefor egg m aturation. M ost species breed in carrion,but Pollenia may parasitise earthworms as it doesin other geographical regions. Some of the speciesof Calliphora, Chrysomya, and Lucilia are of sig-nificant veterinary importance, causing the cuta-neous myiasis of livestock known as `fly strike', butthe life history details of most New Zealand Cal-liphoridae are unknown.This revision is based on almost 1400 speci-mens, from numerous sources. Virtually all typematerial has been examined.

G e n u s Call iphoraThe genus Calliphora in New Zealand contains fourspecies, two introduced from Australia and onefrom Europe, and the fourth endemic. The Aust-ralian species, Calliphora stygia and C. hilli, aretwo of the well known g olden blowflies, or golden-haired blowflies. The first record of their appear-ance in New Zealand dates from 1841, when DrA. Sinclair collected specimens of both (used in thetype series of laemica W hite) in the Bay of Islands.In 1845 he made further collections of both spe-cies, and these are preserved in the BMNH collec-tions. Colonel Bolton collected stygia in 1845 atthe Auckland Islands. French expeditions visitedNew Zealand in 1824, 1827, and 1831, but whenin Paris (MNHN) I found no golden blowflies col-lected on these expeditions. In his narrative ofCook's first voyage, Hawkesworth refers to "fleshflies" being like those of Europ e (see Miller 1939b,p. 58); this probably refers to the endemic species,which has the appearance of European blowflies.Joseph Banks collected this species during Cook'svoyage of 1768, and would surely have taken agolden blowfly had he seen one (he did while inAustralia). During this voyage Cook was mappingthe North and South islands, and Banks wen t ashoreat every opportunity. It seems unlikely, then, thatgolden blowflies were in New Zealand at that time;or at best they were present only in very smallnumbers. It seems most probable that they wereintroduced from A ustralia between 177 9 and 1841.

J.S. Pollack in 1838 mentions a "gad fly" or oestrusdepositing larvae on meat (see M iller 1939b, p. 61);this may refer to a golden blowfly. Hennig (1966,p. 10) accepts a passive dispersal from A ustralia inrecent geological time, saying that man may nothave been the agent.The two golden blowflies C. hilli and C. stygiahave been confused tax onomically by most authors.Hutton (1901, p. 64) mentions neither of them inhis New Zealand synopsis. Malloch, in his Aust-ralian paper (1927, p. 309), discusses the confusion,keys the flies successfully, and also mentions thathe has females of hilli from New Zealand, but hedoes not refer to them in his New Zealand work(1930, p. 313). Hardy (1937, p. 21) gives a ratherconfusing list of species and synonym ies, includingthe names hilli , laemica, miller, rufipes, an d stygia.He does not mention stygia as occurring in NewZealand, but gives laemica and miller as the twogolden blow flies of New Zealand. He alludes to hisnew species milleri as the common New Zealandblowfly, and gives hilli as a synonym of rufipes(milleri is here referred to hilli). Miller (1939b, pp.30 and 32) and Murray (1954, p. 719) both referto hilli , as recognised here, as rufipes and to stygiaas laemica. The name rufipes has in the past causedgreat confusion, as Macquart described both a

ollenia rufipes and a Calliphora rufipes, and bothnames have been applied to blowflies in NewZealand. Dr K .R. Norris (pers. comm.) has studiedthe type specimens of these species in conjunctionwith his work on Australian blowflies, and hasfound that the syntypes of P. rufipes ar e Calliphorastygia and that the holotype of C . rufipes is an Aus-tro-Oriental Calliphora species. Kurahashi (1971,p. 158) placed C. stygia in his subgenus Neocalli-(see below) and referred to C. hilli as C.(Paracallipho ra) rufipes ssp. milleri, but I can findno justification for subspecific status for this species.Calliphora quadrim aculata is an endemic speciesof striking appearance, being large, robust, andmetallic. It is mentioned in M aori folklore, and wasrecorded by all the early authors, having beendescribed by Sw ederus from m aterial collected byBanks during Cook's first voyage. Malloch (1930,p. 316) unaccountably referred to this species asCalliphora sacra (Fabricius). Kurahashi (1971, p.155) placed this species in his subgenus Neocalli-phora along with five others, of which I find thatonly two, namely C. nigrithorax from Tasmaniaand C. ochracea from Australia, are closely relatedto quadrimaculata. These three species constitutea distinct group, and are probably living exam plesof something near to the an cestral stock of present-day Calliphora species. They have in commondensely haired eyes, holoptic head in the m ale, verylarge thoracic spiracles, unicolorous undusted

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abdomen, and curved paralobes in the male geni-talia. Calliphora testaceifacies is closely related tothis group, but in having bare eyes and a dustedfifth tergite is reminiscent of the Australian Calli-phora robusta group. It may be a relatively recentoffshoot from the ancestral line of the numerousspecies of the robusta group found in Australiatoday.Similarly, C. stygia is closely related to the nigri-thorax - ochracea group but has a golden-dustedabdomen, sparsely haired eyes, and straight para-lobes. The thoracic spiracles are very large, and thisis a characteristic feature of the ancestral group;enlargement is found to some degree in other goldenblowfly species, but less strikingly so. It is probablethat stygia is still quite close in its characters to theancestral stock, and marks the first step in a mor-phological progression to the numerous goldenblowfly species found in Australia today.C. quadrimaculata has been geographically andgenetically isolated in New Zealand, and may formthe stock for future speciation. Indeed, on Camp-bell Island and the A uckland Islands it differs fromthe mainland form in having the abdomen blue-green an d me tallic rather than violet. This subant-arctic form was described by H utton (1904, p. 155)as a distinct species, but I do not believe that itdeserves this status. However, speciation appearsto be rapid in the New Zealand subregion, with itsmany islands.Calliphora vicina, accidentally introduced fromEurope, was first recorded in New Zealand by G .V.Hudson in 1889, but under the name C. vomitoria,a misidentification. This species has now reachedall geographical regions of the world, most prob-ably dispersed passively by human agency in thedays of sail.G e n u s HemipyrelliaThis genus is very closely allied to Lucilia, and mayat some time be considered synonymous. LikeLucilia, Hemipyrellia species breed in decayinganimal matter, but adults are attracted to sweet-smelling substances.In material received from Lincoln College I founda single specimen of Hem ipyrellia ligurriens (Wie-demann), possibly incorrectly labelled. As withChrysomy a m egacephala, this species is tropical andwould probably not become established in NewZealand, but have identified it amongst materialcollected at airports in New Zealand. The genushas been included in the key so that it may be iden-tified should it turn up in imported cargo.

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Genus LuciliaThe genus Lucilia is represented here by the wellknown, cosmopolitan sheep blowfly L. sericataMeigen, which was first recorded from New Zealandin 1872 by Hutton (1901, p. 63). Miller (1939b, p.52) gave a key to three species, one sericata andthe others called simply "A" and "B". His speciesB can be eliminated as it obviously did not origi-nate in New Z ealand, having a yellow third tergite;it fits the description of, and most likely is, theAustralian species Hemipyrellia fergusoni Patton.Miller separates his species A from sericata usinga poorly diagnostic colour character, but his geni-talia drawings show quite marked differences.However, these may be artefacts due to distortionduring slide mounting and to over-clearing of theaedeagus. I have not found a second species in thematerial I have examined. Material recentlyreceived from New Zealand airport authoritiesincluded a specimen of the tropical species L.cuprina Wiedemann. This is unlikely to becomeestablished in New Z ealand, but I have included itin the key on p. 27 for the benefit of those iden-tifying insects found in imported cargo.Genus PolleniaPollenia has previously been considered a predom -inantly Palearctic genus, with one or two migrantspecies reaching N orth Africa, America, and north-western India. It is now clear that this is erroneous,and that Pollenia is substantially represented in theSouthern Hem isphere. Many species await descrip-tion from southern Australia, and new species aredescribed here from New Zealand.Apart from two recent introductions from theNorthern Hem isphere, all New Zealand species areendemic. Pollenia is by far the largest as wellas the least known genus of Calliphoridae in thiscountry.A few members of the tribe Polleniini weredescribed from the Oriental and Austro-Orientalregions by early authors such as Francis Walker,but these have subsequently been transferred toother genera within the Calliphorinae and Rhini-inae. No Polleniini have been recorded from theAfrotropical and Neotropical regions. In NorthAmerica the tribe is represented by two introducedspecies and the genus Melanodexia Williston, whichis confined to river g ullies in California and Idaho.There are, then, four distinct geog raphical groupsof Polleniini: Palearctic Pollenia (30 + species);Nearctic Melanodexia (8 + species); Oriental andAustro-Oriental Pollenia (8 + species, includingX anthotryx us); and the Australian and New ZealandPollenia (60 + species, including A nthracomyia).Basic external and genital morphology is identical

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in the four groups, head shap e (Figure 12) and thebare prosternum and propleuron in particular dis-tinguishing them from other Calliphoridae. TheOriental - Australasian genus Polleniopsis Towns-end has a Pollenia-like head but a haired propleu-ron and prosternum, short, larviparous ovipositor,and Onesia-like aedeagus.The question of origins and phylogenetic rela-tionships of the Po lleniini within the Calliphoridaeis especially difficult, and its detailed resolution isoutside the scope of this study. H owever, the exist-ence of a large numb er of very similar species indi-cates that the Polleniini are a comparatively recentgroup, possibly originating in and dispersing fromthe Palearctic region. It is likely that this dispersaltook place across the Bering land-bridge and alongthe island chains of South-west Asia to Australiaand New Zealand. Whatever their origin, onceestablished the groups would have been isolated bychanges in sea level during Pleistocene interglacialsand the post-Pleistocene period, and rapid specia-tion apparently took place in the more temperateareas. The distinct Oriental and Austro-Orientalgroup consists of the few species that were able toadapt to a subtropical climate.In basic structure and general appearance theNew Zealand Pollenia species resemble thePalearctic representatives, with only a few minordifferences, and I regard the two groups as con gen-eric. New Zealand Pollenia do not have the facialcarina that is present in most Palearctic species,nor such lo ng, crinkled thoracic ground setulae. Insome of them the abdomen is metallic blue or green,without distinct dusting, and often with pale groundcolour to the thorax.A few Palearctic species placed by some authorsin a separate genus, Nitellia Robineau-Desvoidy,have shorter ground setulae and an undusted,metallic black abdomen. These are much moresimilar to the New Zealand species than they areto the rest of the Pa learctic Pollenia (the rudis-group;see note on p. 47 ), which have long, crinkled, goldenthoracic hairing and the abdomen densely dustedand olivaceous. The male genitalia of the NewZealand and Palearctic species are of very similarconstruction, but unlike any of the New Zealandspecies the rudis-group has a strongly sclerotisedrod on the basal half of the hypophallus. In theNitellia-group the marginal spines are confined tothe dorsal portion of the hypophallus, and the lowerportion is not produced forwards into a spine-likeprojection as in many of the New Zealand species.The aedeagus of Melanodexia resembles that of theNitellia-group but has a broader, blunter para-phallus.The Australian species are generally darker, andoften have the tho racic hairing typical of the rudis-

group. In structure of the aedeagus they are iden-tical with the New Zealand species.I wholeheartedly endorse the view of Mihlyi(1976) concerning the H ungarian Pollenia species:"Pollenia is the most difficult genus of the Calli-phoridae". I have experienced considerable diffi-culty in producing useful taxonomic con clusions tothe problems encountered w hile preparing a key tothe New Zealand species of Pollenia, as with thePalearctic species. These problems have arisenmostly because of infraspecific variability, thegeneral similarity of the species as regards externalmorphology, and the relative paucity of materialavailable. Although infraspec ific variability has beenaccounted for in the key, the figures of the malegenitalia should always be used to confirm anidentification.A separate key to females has been given, andagain variability has been accounted for, but asmore material becomes available this key mayprove unsatisfactory. Except w here I have had reli-ably associated sexes, the females have not beenassigned to species but left unnamed, and insteadreferred to as "species a" to "species q" .H utton (1901 ) was the first to describe any NewZealand Pollenia species, but placed his new spe-cies demissum and fumosum in the genus Sepi-mentum. Malloch (1924) synonymised this genuswith Pollenia and considered H utton's two speciesto be conspecific. In a subsequent paper (1930),published after examination of more material,Malloch reversed his opinion and restored them asdistinct species. He also described three new spe-cies, and two new varieties of demissa; these latterare discussed under P. demissa (p. 36).Although unhappy about the validity of thegeneric characters of Huttonophasia as distinct fromPollenia, Malloch (1930) considered it a good genus.I consider it to be synonymous with Pollenia, as Ican find no characters important enough to war-rant its distinction (see Remarks under P. pernix).Genus PtilonesiaPtilonesia is a very unusual calliphorid genus foundonly in New Zealand and two localities in Austra-lia. It is monotypic, and in general appearance isvery Onesia-like, with wide parafacials and jowlsand the eyes reduced. It is also reminiscent of theendemic New Zealand genus X enocalliphora, andwas placed as a subgenus of it by Kurahashi (19 71).I prefer to keep Ptilonesia as a distinct genus onthe basis of its lack of ocellar and superior orbitalsetae, modifications to the fifth tergite, hairing onthe parafacialia and squamae, facial shape, and malegenitalia. The similarity to the Neotropical genusToxotarsus I believe to be due to convergence,rather than an indication of any phylogenetic rela-

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tionship, because both genera are seashore flies.Miller (1939b) says that the single species of Pti-lonesia breeds in decaying seaweed. This may be amistaken observation, and it may be that the flieswere breeding in dead animals tangled in the sea-weed masses.Outside New Zealand Ptilonesia is apparentlyrestricted to beaches north and south of SydneyHeads and at Seaford in Victoria (K. R. Norris,pers. comm). The q uestion arises as to whether thisdisjunct and local distribution represents an intro-duction or is a relic of a previously wider distri-bution. Introduction on floating masses of seaweedis unlikely, as ocean currents run away from Aus-tralia throughout the year, and prevailing windsblow either towards New Zealand or northwardsalong the eastern coast of Australia. The other pos-sibility is of dispersal by boats travelling betweenNew Zealand and Australia, but it is strange that,once established, the individual colonies did notdisperse rapidly, as is usual with other seashoreDiptera. It seems most likely, therefore, that theAustralian populations are a relic of a previouslywider distribution.Genus XenocalliphoraThis genus is endemic, and contains ten species,four of which are described here as new. Three spe-cies occur on the main islands, but the remainingseven are restricted to offshore islands.X enocalliphora is reminiscent of Onesia an d Pti-lonesia in general habitus, having dichoptic males,wide jowls and parafacialia, and in being larvipa-rous. The species can be divided into two distinctgroups which, however, are not distinguishable bytheir general appearance. The first group has twoposterior ia setae, no pd setae on the fore tibia, andan aedeagus with a bifurcate harpes that is curvedfor its entire length (Figures 94 and 100). Thesecond group has one posterior ia seta, a pd setaon the fore tibia, and a non-bifurcate aedeagus thatis broad and curved at the tip (Figures 92 and 96).The first group is restricted to the North Island,northern South Island, and central offshore islands,and the second is found only in the southern islands a distribution suggesting sister-group status.Unfortunately, Hennig (1966) does not mention thisgenus or any of its species in his work.The phylogenetic relationships of X enocalli-phora within the Calliphoridae are not clear. Kura-hashi (1971 ) suggests that the genus is derived fromthe same ancestral stock as the old Calliphora-groupbut has evolved in a way differing from modernCalliphora. Certainly it is of striking appearancewhen com pared with other genera from the South-ern H emisphere. Kurahashi also suggests that it hasaffinities with some H olarctic genera such as Cyno-

mya, Cyanus, Onesiomima, and Abago. This I finddoubtful, and prefer to consider X enocalliphora asa totally isolated group that is now found only inthe New Zealand subregion, but which derives ulti-mately from the common ancestor of the Calli-phora-group.Malloch (1930) keyed four species correctly, usingthe name X enocalliphora and describing one spe-cies as new. Miller (1939b) used Calliphora for allhis blowfly species, including five X enocalliphoraspecies, two of w hich he described as new. Murray(1954) gave a very good key to all the species ofCalliphora known to occur in New Zealand; thisincluded six species of X enocalliphora, one of whichhe described as new. Kurahashi (1971) included fivespecies in his key to X enocalliphora but did not seespecimens of mo st of the species. Harrison (197 6)gave a key to Calliphora species which includedfive Xenocalliphora species. In general, adequatekeys have been given to the know n species, but inmany instances the nomenclature is incorrect,names being confused and assigned to the wrongspecies. Misidentifications are discussed u nder therelevant species.Genus ChrysomyaThis genus is represented in New Zealand by C.rufifacies (Macq uart), which, like other Chrysomyaspecies, is a tropical or subtropical blowfly adopt-ing the ecological role of Lucilia in temperateregions. It is, however, unusual in being able toextend its geographical range to warm temperateregions (cf. C. albiceps in southern Europe). Themale genitalia (Figure 19) are not typical of theChrysomyinae, and the larva is unusual in havingfleshy protuberances on the segments (see Figure116).The earliest recorded occurrence of rufifacies inNew Zealand is 1911 (Miller 1939b, p. 56, andspecimens in BM NH ). Malloch (193 0, p. 315) refersto this species as Chrysomya albiceps (Wiede-mann), with which it may be conspecific (see Zumpt195 6, p. 192).I have recently received from the Common-wealth Institute of Entomology two samples ofChrysomy a m egacephala (Fabricius) found abo ardaircraft arriving in New Zealand from Australia. Ialso have a record o f this species from ro tting fruitat Springston, M C (possibly incorrectly labelled; R.A. H arrison, pers. comm.). This species is tropical,and is unlikely to become established in NewZealand, but I have included it in a key (p. 57) sothat it can be identified should it turn up in sea-ports or at airports.

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IMMATURE STAGESOF N EW ZEALAND CALLIPHO RIDAE

Contributed by . A. Holloway* comprehensive systematic work on NewZealand's Calliphoridae would include a substan-tial section on the morphology and biology of theimmature life stages. Lack of material of the earlystages of the many endemic species of blowfliesprecludes such a treatment in this contribution. Themo st that can be offered are simple keys and shortdescriptions that will enable the user to recogniseeach of the three larval instars and to identify tospecies the third-instar larvae of the six common-est calliphorids of urban and rural areas.Five of these species are of Northern Hemi-sphere and Australian origin. Their larvae are theconspicuous maggots that may be seen feeding ondead animals, and are either primary or secondaryinvaders in sheep strike (see Heath 1985, p. 15).The immature stages of some of them m ay also bediscovered inside meat pies, hambu rgers, and simi-lar meat-based foods that have been exposed to egg-laying adult females.The sixth sp ecies is the large endemic bluebottleCalliphora quadrimaculata. Although its larvaefrequently occur on dead animals, they are also ableto reach maturity without feeding on carrion. Ona recent collecting trip based at Dundas hut in thenorthern Tararua Range (WN-WA; altitude 1250m) I found a mature larva and numerous puparia from which adults subsequently emerged about 2 cm below the surface in gritty soil sur-rounding clumps of the broad-leaved snow tussock,Chionochloa flavescens. The most likely source oflarval food was the deep, wet layer of fermentingleaf sheaths in the base of this species of tussock.Unlike adult calliphorids, which are easily recog-nised as bluebottles, greenbottles, and golden-hairedblowflies, most larvae -- even of quite distantlyrelated species look very similar to the nakedeye. How ever, at 50 magnification under a stereo-microscop e the distinctive features of the three lar-val instars are clearly discernible, and it is notd i f f i c u l t t o i d e n t i f y t h i r d - i n s t a r l a r v a e t o s p e c i e s .Calliphorid larvae which do not key out to one ofthe six common species probably belong to theendemic fauna. If sufficient live specimens areavailable, some should be preserved and theremainder should, if possible, be reared to adults.A well documented collection of correctly asso-ciated larvae and adults would be immensely valu-*Entomology Division, Department of Scientific an dIndustrial Research, Private Bag, Auckland, New Z ealand

ableable to specialists studying the affinities of Ne wZealand Calliphoridae.M aterials and techniquesThe material studied, including associated adults,is in the forensic and general collections of the NewZealand Arthropod Collection (NZAC), held byEntomology Division, DSIR, Auckland. Most ofthe specimens were obtained between 1981 and1983 during a study of the fauna associated withhuman corpses in Auckland (Smeeton et al. 1984).They w ere collected by Drs W . M. I. Sm eeton andT. D. Koelm eyer, senior lecturers in forensic med-icine at the School of Medicine, University ofAuckland, from corpses which, because of thecircumstances of death, were received for post-mortem examination at the Auckland CityMortuary.The characters used for identification are mosteasily seen in fully extended specimens w hich havebeen killed by brief immersion in almost boilingwater. Freshly killed specimens can either be fixedin PEA solution (1 part petroleum ether, 7-10 parts95% ethanol, 2 parts glacial acetic acid) or placeddirectly in 70% ethanol. To examine the structuresused in the keys it is not necessary to clear or slide-mount the larvae. They can be viewed adequatelyin a dish of ethanol under a stereomicroscope at amagnification of about 50.The illustrations were made with the aid of acamera lucida. Structures that are not obvious atlow magnifications and which have not been usedin the key e.g., the perispiracular bristles asso-ciated with the posterior spiracular slits -- areomitted from the drawings. Measurements weretaken from fully extended larvae.M orphology and terminologyThe body of a blowfly maggot (Figures 115 and116) consists of a head, three thoracic segments(T1-T3), and eight abdominal segments (A1-A8) .The head is very small, and is often partiallyretracted into the thoracic segments; it is bilobed,and bears a pair of antennae and a pair of maxil-lary palps (Figures 117 -119 ). The mo uth opens onthe ventral surface of the head, and has associatedwith it a pair of strongly sclerotised mouth hooksand a variably developed oral sclerite (see Figure126). Numerous oral grooves are present on theintegumental surface on either side of the mouth.The mouth hooks are part of a complex cephalo-pharyngeal skeleton which is not easily seen inuncleared specimens.The first thoracic segment of larvae in the secondand third instars bears a pair of conspicuous, lobed,anterior spiracles (Figures 118 and 1 19 ). First-instarlarvae lack lobed spiracles on this segment, but have

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instead a pair of minute spiracular slits which arebarely discernible except und er a scanning electronm i c r o s c o p e ( K i t c h in g 1 9 7 6 a ) . T h e b o d y s e g m e n t sare separated by narrow spine bands (Figures 115 -119) which are always well developed on the ven-tral surface but which may be weak or absent dor-sally and laterally. The size, shape, arrangement,and density of spines in the bands of third-instarlarvae are important characters for identifying dif-ferent species. The posterior end of the lastabdom inal segment bears six pairs of papillae (Fig-ure 115) which are arranged approximately in a ring.In some species paired papillae are present on thethird thoracic segment and all abdom inal segments(Figure 116). The papillae on the last abdominalsegment surround a pair of posterior spiracles (Fig-ures 116 and 123-125). Each spiracle consists of asessile, crescentic to circular peritreme enclosing twoor three obliquely arranged slits. The number ofslits and the shape of the peritreme are easily vis-ible characters from which the different larvalinstars can be identified (Figures 120-122).Various authors (e.g., Miller 1939, Zumpt 1965)have attached considerable taxonomic imp ortanceto the num ber of lobes in the anterior spiracles andthe size, shape, and proximity of the p osterior spir-acles of third-instar larvae. In the species dealt withhere these characters are too variable to be tax-onom ically useful. The most reliable and easily seencharacters for identifying these species occur in theoral sclerite and in the spine bands. Illustrations ofthe posterior spiracles have been included mainlyto show how much these structures may vary insimilar-sized specimens of the same species (Fig-ures 137 cf. 138, 142 c f. 143, and 147 cf. 148).Diagn ostic characters of larvaeCalliphorid larvae can be distinguished from thoseof Anthomyiidae, Drosophilidae, Fanniidae, Mus-cidae, Phoridae, Sarcophagidae, and S ciadoceridae,which m ay occupy sim ilar ecological niches, by thefollowing combination of character states: spinebands present dorsally on at least some segments;posterior spiracles always sessile (never on stalksor m ounds), not located inside a deep p it, encircledby 6 pairs of conspicuous, isolated papillae; peri-treme develop ed as a ring or crescent (never in theform of a uniformly sclerotised disc); slits straight(not coiled or strongly bent), converging towardsventral midline of last abdominal segment.

K ey to larv al instars1 1 without lobed spiracles; posteriorspiracle with a fragmentary, usuallycrescentic peritreme and 2 partiallyjoined slits; length 1.5-4.5 mm... 1st instar

1 with a pair of lobed spiracles;posterior spiracle with a semicircularto circular peritreme and 2 or 3 dis-crete slits; length 10-23.0 mm... 22Posterior spiracle with 2 slits; length3.0-9.0 mm... 2nd instar-- Posterior spiracle with 3 slits; length8.0-23.0 mm... 3rd instarK ey to th i r d -i n s t a r l a r v a e

1- 7 each bearing several pairs offinger-like papillae (Fig. 116); somespine bands with bifurcate- or trifur-cate-tipped spines dorsally...Chrysomya rufifacies 1- 7 without papillae (Fig. 115);none of the spine bands with bifur-

cate- or trifurcate-tipped spines... 22Dorsal part of spine band 1/ 2comprising mainly large spines whichare evenly distributed in somewhatoblique rows (Fig. 127, 131, and 135)... 3 Dorsal part of spine band 1/ 2comprising minute spines which areunevenly distributed in broken, undu-lating, transverse rows (Fig. 140 and145)... 53Dorsom edian part of spine band 7/ 8 with close-set spines arrangeduniformly in about 7 oblique rows(Fig. 128 )... Calliphora hilli Dorsomedian part of spine band 7/ 8 with widely separated spinesarranged irregularly in at most 4oblique rows (Fig. 132 and 136)... 44Oral sclerite short, notched basally,pointed apically (Fig. 130); spinebands containing very large spines(Fig. 131 and 132)

...Calliphora quadrimaculata Oral sclerite long, conspicuouslybifurcate basally, truncate apically(Fig. 134); spine bands containingmoderately large spines (Fig. 135 and136)... Calliphor a sty gia5Oral sclerite well developed (Fig. 139)...Calliphor a vicina Oral sclerite vestigial or absent (Fig.144)... Lucilia sericata-13-


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Descriptive notes on speciesCall iphora h i l liTHIRD-INSTAR LARVA. Oral sclerite (Figure 126)with apex pointed, base bifurcate. Spine bands dor-sally with dense, relatively large, simple seines thatare evenly spaced in somewhat oblique rows (Fig-ures 127 and 1 28). Numbers of spine rows in vicin-ity of dorsal midline as follows: spine band betweenhead and , about 10; 1 / 2, 2/ 3, 3/ 1, 9-11; 1/ 2, 7-9; 2/ 3, 5-7; 3/ 4, 2 or 3; 4/ 5, 5/ 6, 0; 6/ 7, 0 or 1; 7/ 8, 6-8. Anterior spir-acle with 9-11 lobes; posterior spiracle, Figure 129.Miller (1939b) described the third-instar larva ofthis species (as C. rufipes).Call iphora quadrimaculataTHIRD-INSTAR LARVA. Oral sclerite (Figure 130)small, with apex pointed, base notched. Spine bandsdorsally with sparse to moderately close-set, simple,mostly very large seines that are evenly spaced insomewhat oblique rows (Figures 131 and 132).Numbers of spine rows in vicinity of dorsal mid-line as follows: spine band between head and ,7-9; 1/ 2, 6 or 7; 2/ 3, 6-8; 3/ 1, 1/ 2, 5-7; 2/ 3, 5 or 6; 3/ 4, 4-6; 4/ 5, 2-4; 5/ 6,0 or 1; 6/ 7, 1-3; 7/ 8, 3 or 4 . Anterior spiraclewith 11-16 lobes; posterior spiracle, Figure 133.The morphology of the third-instar larva has beendescribed by Miller (1939b); his habitus drawing(Figure 42) incorrectly shows the larva as havingonly ten post-cephalic segments.Call iphora stygiaTHIRD-INSTAR LARVA. Oral sclerite (Figure 134)with apex truncate, base bifurcate. Spine bandsdorsally with sparse to m oderately dense, relativelylarge, simple spines that are evenly spaced in some-what oblique rows (Figures 135 and 136). Numbersof spine rows in vicinity of dorsal midline asfollows: spine bands between head and 1 an d 1/ 2, 7 or 8; 2/ 3, 3/ 1, 4-6; 1/ 2, 3-5; 2/ 3, 2-4; 3/ 4, 4/ 5, 0-4; 5/ 6, 6/ 7, 0or 1; 7/ 8, 0-4. Anterior spiracle with 9-14 lobes;posterior spiracle, Figures 137 and 138.Additional information on the morphology of thethird-instar larva is given by Fuller (1932), Miller(1939b; as C. laem ica), and Zumpt (1965). O'Flynn& Moorhouse (1980) have described the egg andearly-instar larvae.Call iphora vicinaTHIRD-INSTAR LARVA. Oral sclerite (Figure 139)with apex po inted, base not bifurcate. Spine band sdorsally with minute, simple spines that arearranged unevenly in undulating, broken, trans-verse rows (Figures 140 and 141 ). Numbers of spinerows in vicinity of dorsal midline as follows: spine

band between head and , about 8; 1/ 2, T2/ 3, 3/ 1, 7-9; 1/ 2, 2/ 3, 6-8; 3/ 4, 3-6; 4/ 5,0-5; 5/ 6, 0-2; 6/ 7, 0-5; 7/ 8, 4-8. Anteriorspiracle with 8-11 lobes; posterior spiracle, Figures142 and 143.Further information on the morphology of third-instar larvae is given by Miller (1939b; as C. ery-throcephala), Schumann (1954), and Zumpt (1965).The egg has been described by Z umpt (19 65), andearly-instar larvae are described by Schumann(1954 ) and Zumpt (1965).Chrysomya rufifaciesTHIRD-INSTAR LARVA. (Figure 116). Oral scleritevestigial or absent. Segmental papillae very con-spicuous. Spine bands dorsally with moderatelydense, large, scale-like spines, some with bifurcateor trifurcate tips, that are evenly distributed insomew hat oblique rows. Numbers of spine rows invicinity of dorsal midline as follows: spine bandbetween head and 1, 9-11; 1/ 2, 6-8; 2/ 3,5-8; spine bands indistinguishable dorsally onremainder of body. Anterior spiracle with 9-11lobes; posterior spiracle with a broad peritreme.Additional information on the morphology of thethird-instar larva is available in Fuller (1932), andscanning electron micrographs of the egg and third-instar larva are published in Kitching (19 76b). Themorp hology of early-instar larvae has been studiedby O'Flynn & Moorhouse (1980). Zumpt (1965)considered the immature stages of C. rufifacies tobe indistinguishable from those of C. albiceps(Wiedemann).Luc il ia sericataTHIRD-INSTAR LARVA. Oral sclerite (Figure 144)vestigial or absent. Spine band s dorsally with min-ute, simple spines that are unevenly distributed inundulating, broken, transverse rows (Figures 145and 1 46). Numbers of spine rows in vicinity of dor-sal midline as follows: spine band between headand 1, about 6; 1/ 2, 5-7; 2/ 3, 6-8; 3/ 1, 1/ 2, 6 or 7; 2/ 3, 5 or 6; 3/ 4, 2-4; 4/ 5, 5/ 6, 6/ 7, 0; 7/ 8, 0-3. Anterior spiracle with7-9 lobes; posterior spiracle, Figures 147 and 148.Detailed descriptions of larvae of all three instarshave been given by Schumann (19 54) and by Zump t(1965), who includes a description of the egg.

I am grateful to D.W. Helmore, biological illus-trator at Entomology Division, for preparing Fig-ures 115 and 116.References cited in this brief account are listedin the main bibliography, which commences on p .59. -s--14-


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FLY -STR IKE IN N EW ZEALANDContributed by A. C. G. H eath*

Calliphorid flies are able to feed on a wide varietyof animal and plant substances. The adults arestrongly attracted to moisture, and feed mainly onsweet substances such as nectar and honeydew an don the liquid products of organic decomposition,which prov ide the proteins essential for egg matur-ation. The imm ature stages (larvae or m aggots) candevelop on various substrates, ranging from kitchenoffal and faecal material to carrion and living ani-mals. The ability of the larvae of certain genera ofCalliphoridae to attack living animals causes a con-dition known as myiasis.The term myiasis is applied to a particular modeof feeding by dipterous larvae, on or in the bodyof a living vertebrate. These larvae feed on thehost's dead o r living tissue, liquid bod y-substances,or ingested food, and are able to complete or atleast for a certain period continue their normaldevelopment in this manner (Zumpt 1965).A specific type of myiasis, commonly known as`fly-blow' or `fly-strike', is a perennial problem tothe New Zealand farmer. Fly-strike is an extensionof the carrion-feeding habit, and is initiated by onlya few species; these are commonly called `primaryflies' (Colless & McAlpine 1970). Gravid femalesare attracted to certain areas of the sheep's body,and may be stimulated to deposit eggs or live lar-vae there.Before a strike can be initiated, a number o f con-ditions must be met. First, a female fly must be atleast 3 days old and have ha d sufficient protein forher eggs to have reached a certain stage in develop-ment before she will mate (Webber 1958, Applin19 79, C rystal 1983 ). Subsequently a diet consistingof adequate carbohydrate, water, and protein isnecessary before eggs can mature fully (Webber19 57). H aving reached the stage of full egg matur-ation, the female fly is stimulated to lay eggs ordeposit larvae under another circumscribed set ofconditions. First, the site must be moist in orderthat eggs do not becom e desiccated (Vogt & Wood -burn 1980). In fact, flies will not lay eggs unlesstheir tarsi are in contact with water (Barton Brow ne1962). It is also helpful if the chosen ovipositionsite receives low illumination (Barton Browne1958). Lastly, the site the female chooses must beproteinaceous, thus offering a food supply for larvae.History. The first official record o f fly-strike as aproblem in New Zealand appears in the 1896*Wallaceville A nimal Research C entre, Ministry of Agri-culture and Fisheries, Private Bag, Upper Hutt, NewZealand

Annual Report of the Department of Agriculture(pp. 19-20). There is some inconsistency of opin-ion, however, concerning when the flies causingstrike first came to public notice (Hu tton 19 01, G il-ruth 1907); the 1870s are most frequently cited inreports. In the first years of the twentieth centurythe flies apparently were established nationwide. Afarmer with a long memory recalled that "maggotflies have been troub lesome for the past 60 years"(Miller 1921). The problem was severe enough bythe 1920s for a survey of its prevalence and dis-tribution to be undertaken (Miller 1921).Species involved. (Unless otherwise indicated, thedata presented here and in subsequent sections arefrom unpu blished observations and experiments byA. C. G. Heath and J. D. Tenquist.) Calliphoridsidentified from cases of fly-strike in New Zealandare the endemic species Calliphora quadrimaculataand Xenocalliphora hortona and the adventive C.hilli, C. stygia (both possibly self-introduced), C.vicina, Lucilia sericata, and Chrysomya rufifacies(the three last-named most likely introduced byman). The species most comm only involved in fly-strike in New Zealand are C. stygia an d L . s ericata.Material collected over the period 1976-1984 from91 cases of fly-strike showed C. stygia to be presentin 54.9% of cases and L. sericata in 37.4%. Chry-somya rufifacies was found in 5.5% of strikes andCalliphora vicina in 1.1%. In 16.7% o f cases C. sty-gia an d L . sericata occurred together in a single fly-strike lesion. Although the actual pro portions vary,these figures substantiate the findings of earlierauthors (Miller 1922, 19 39a; M acfarlane 1941 ) thatC. s tygia is the species most commonly involvedin fly-strike. By contrast, catches of flies from liver-baited fly-traps show proportions of the main strike-fly species that do not correspond with the pro-portions from cases of fly-strike. Over the period1978-1981, C. stygia constituted 17.7% of the totalcatch of 146,901 flies from 17 traps (14 in the NorthIsland, 3 in the South Island), whereas L. sericataand C. rufifacies accounted for 49.1% and 33.1% ofthe catch respectively.Prevalence. A num ber of surveys of varying scaleand com plexity have investigated the extent of thefly-strike problem in New Zealand (Miller 1921,193 9a; Macfarlane 1941 ; Tenquist & Wright 1976).The largest and most com prehensive. survey (Ten-quist & Wright 1976) indicated that there was anaverage of 1.2% of the national flock affected byfly-strike each year, with differences by region rang-ing from 0.4% to 2.1%. A survey in Marlboroughalone (Macfarlane 1941) arrived at a figure of 6%prevalence, although up to 50% of small flocks havebeen affected in other parts of the coun try at othertimes. These figures indicate the lack of uniform ity

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Number of months>5.9L.0-5.92.0-3.9


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cases of fly-strike in spring. Otherwise, L. sericatawas trapped more frequently in spring than inautumn between 1978 and 1981 . Furthermore, bothC. stygia and L. sericata were nearly equally rep-resented in trap catches in autumn.Geog raphical dist r ibution. Blowflies may be foundthroughout New Zealand, although there is a changefrom no rth to south in the propo rtion of each spe-cies in trapped samples and in cases of fly-strike.Miller (1939a) concluded that C. s tygia was thecommonest fly (as reared from cases of fly-strike)in the North Island. In the South Island C. stygiawas likewise the commonest fly in many cases offly-strike, except on the east coast, where L. seri-cata predominated. In general, C. stygia becameless common as the annual mean temperaturebecame lower, i.e., moving from north to south.Data obtained between 1978 and 1981 from 17fly-traps (14 in the N orth Island and 3 in the Sou thIsland) show a somewhat different picture (seemaps, above). C. s tyg ia appears to be most abun-dant in the north, north-east, and east of the NorthIsland, and also in the Nelson area, but uncommo nin Taranaki and the central North Island. L. seri-cata is most prevalent on the east coast of the No rthIsland, with pockets of abundance in South A uck-land and the K ing Country. These distributions canbe related partly to temperature, but are likely tobe influenced by rainfall as well. Rainfall patternsmay explain the marked west-to-east change inprevalence of the flies, and particularly L. sericata.Predispos it ion to f ly-strike. When blowfly larvaeare deposited or hatch from eggs laid in the damp,dark recesses of a fleece, they are faced with theneed to find food. This can be present in the formof blood from a wound, or serum exuded fromdame skin with scald, or the bacterial decompo -sition products causing fleece rot (Merritt 1980,Watts & Merritt 1981). As the larvae grow theybecome able to attack living host tissues by a com-bination of the tearing action of their mouth-hooksand the digestive prop erties of their saliva (Green -berg 1973).Once an active lesion (fly-strike) is establishedby the primary flies, other species of flies (`super-numerary' or `secondary') may be attracted to thesite. A `tertiary' group of flies, the larvae of whichoccur in healing scabs, is also recognised (Miller1939b, Colless & McAlpine 1970). This hierarchi-cal classification is not rigid, however; prim ary flieshave been found as secondary invaders, and so-called tertiary flies have been know n to initiate fly-strike.The combination of factors that results in a sheepbecoming attractive to flies is still not completely

understoo d. In general, flies depo sit eggs or larvaeon areas of the sheep's body where persistent wet-ting of the wool by urine, sweat, or rain has dis-rupted the skin's waxy layer, allowing serum toexude or bacteria to grow. In addition, other odor-iferous materials such as faeces, pus, blood , or thebacterial decomposition products of footrot willattract a female fly with an urgent n eed to lay eggs.Circumstantial evidence points to lambs sheepless than 1 2 months old perhaps being attractiveto flies for different reasons than are older sheep.From 1 88 cases of fly-strike recorded between 197 6and 1984, 78.7% of the animals affected were lambs.Lambs are more frequently fly-struck around theposterior end (tail, perineum, crutch n early 80%of cases), whereas adult animals are about equallyinfested on these areas or other parts of the bod y,generally the back or side.These differences may be explicable in terms ofwoo l fibre density and length, diet or the ability toutilise food, and parasitism or other diseases. Thefleece of older animals is denser and longer thanthat of lambs except after shearing, and will holdsufficient moisture to encourage growth of micro-organisms and hence fleece rot. Faecal stainingcaused by scouring is accepted as a major predis-posing factor leading to fly-strike in lambs.Pathology. There are numerous pathologicaleffects of fly-strike. First, the host's skin is sub-jected to both mechanical and chemical attack bythe maggots' feeding activity. The skin becomesswollen, inflamed, and weeping much like aburn, hence the term `scald'. The combination offluid loss and release of toxic products into the cir-culatory system is reflected in a change in bloodparameters, such as depressed haemoglobin levelsand elevated white cell numbers (Broadm eadow etal. 1984). The toxins arise from bacterial invasionof wounds or toxic products from the maggots,either in their saliva or excreta. The skin damageis such that w ool fibres break or can be lifted easilyand cleanly from th eir follicles. A pro longed strike(as a consequence of numerous superimposed ovi-positions) can result in larvae penetrating thesuperficial muscle layers and, on occasion, into thebody cavity, with fatal results.If the infestation ha s not been large an d there isno reinfestation nor sup erimposed strikes, the skinwill heal once the maggots leave the site. Healingtakes about 25-30 days, and new wool growthbecomes apparent about 10 days later. This woolis always shorter than the surrounding, untouchedareas of fleece even many months after the initialdama ge. The skin from animals slaughtered beforehealing has occurred will show evidence of maggotdamage when tanned.

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In addition to suffering direct skin damage, theinfested sheep starts to lose its appetite within thefirst 24 hours o f a strike becoming established. Nofurther food is eaten while the maggots are present.When they leave to pu pate, normal appetite returnsin about 21 days in lambs and often about 3 daysin older sheep. The anorexia is accompanied by areduction in liveweight, apparent from 3 to 5 daysafter the strike has started, and amounting finallyto as much as 5 .5 kg. The time taken to regain pre-infestation liveweight can range up to 4 2 days, butmost affected animals tend to remain lighter thanuninfested sheep.Other likely effects of fly-strike are the exacer-bation of footrot throu gh infestation of the affecteddigits, and diminished mating or lamb ing perform-ance due to loss of vigour.Economies of fly-strike. The earliest estimate ofeconomic loss attributable to fly-strike in NewZealand (Miller 1934) was equivalent to about$NZ 15 million a year in present-day terms. Morerecently Tenquist & Wright (1976) calculated thetotal annual cost of fly-strike, on the basis of 1-2%of the national flock being affected, to be $5.3 m il-lion (in 1984 dollars, post devaluation). Thisamount was divided among stock losses ($3.6 mil-lion), labour costs ($1.0 million), and treatmentcosts ($0.7 million), but did not take into accountthe value of production losses from affected ani-mals that survived.The most recent estimate of the current cost offly-strike to the New Zealand sheep industry is$13.75 million (Heath et al. 1983). This comprises$4.25 million in stock deaths, $2.0 million in woollosses, and $7.5 million for labour and materialsassociated with dipp ing. The last figure includes thecost of single statutory dipping for all sheep,together with an add itional treatment for one-thirdof the national flock. This is based on the Tenquist& Wright (1976) survey, in which 35% of farmerswere found to specifically treat for fly-strike inaddition to the statutory dipping. In the total costof fly-strike, no estimate has been made of asso-ciated handling costs such as crutching andadditional shearing, nor has the cap ital cost of dip-ping plant and other equipment been taken intoconsideration. Further costs can result from delaysin sending for slaughter lambs which have lostweight owing to fly-strike, since prices may fall asthe season progresses.Because of the periodic fluctuations in the preva-lence of fly-strike, due largely to year-to-yearweather variations, the costs will also vary year byyear. Nevertheless, the disease remains todayalmost as severe in terms of prevalence andeconomic impact as it has been over the past 100

or m ore years, despite the availability of potent andsupposedly long-persisting insecticides. The mainreason for the continuing success o f blowflies is thatchemical control measures affect only a small pro-portion o f the total population, there being a largenon-parasitic reservoir of maggots. In addition,although today's insecticides are more potent and,in general, more persistent than those in use beforethe Second World W ar, none is capable of provid-ing protection for sheep over the entire period offly activity unless regularly spaced dippings are car-ried out. It is unlikely that many farmers wouldcontemplate such a course of action, so there willalways be unp rotected sheep vulnerable to fly-strike.I am indebted to J.D. Tenquist, who providedtechnical assistance during collection of data, andto A. Barku s, who prepa red the text-figure.The references cited here are listed in the mainbibliography, which comm ences on p. 59

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REARING CALLIPHO RIDAEContributed by Pritam Singh*

Blow flies are relatively easy to rear throughou t theyear in the laboratory with relatively simple equip-ment. They have consequently been used as experi-mental insects in the study of behav iour, nutrition,vision, neurobiology, physiology, and toxicology.They are also used as tools in forensic entomo logy,particularly in establishing the time of death(Smeeton et al. 19 84), and specially reared maggotsare occasionally employed in the treatment of sup-purating wounds and osteomyelitis. Blowflies arealso commercially produced as a bait for anglers,for pollination of greenhouse crops, for rearing ofparasites, and as food for captive birds, amphibi-ans, and reptiles.The life cycle of blow flies is short, and is similarin most species. A female oviposits in a medium,usually carrion, producing clusters of eggs whichhatch in 24 hours. There are three larval instars,lasting in total about 8 days. Larvae feed on liqui-fled tissues which have been dissolved partly byproteolytic bacteria and partly by enzymes theysecrete themselves. When mature, the third-instarlarva ceases feeding and wanders away from thefood supp ly to a suitable pupariation site gener-ally dry, and under the decomposing carcass orslightly away from it about 3 cm below the soilsurface. It empties its gut, acquires fat deposits, andcontracts to form a barrel-shaped puparium. Withinthis structure the larva moults into a pupa, fromwhich the ad ult fly emerges, usually within 2 weeks.Adult females have a p re-oviposition period lasting4-7 days, and in a total lifespan of 2-3 weeks lay600-800 eggs, depending on the species. At 25Con meat, the total life cycle from egg to egg takes20-25 days. All stages are influenced by tempera-ture, humidity, photoperiod, and the type of foodavailable.Choice of rearing methodThis depends on the pu rpose for which the speciesselected for rearing is required, and the numbers,life stage, and supply interval needed. It is recom-mended that blowflies be reared in a well venti-lated room, preferably fitted with an exhaust fanto discharge the foul sm ell outside the building. Theroom should be maintained at 25-28C, 50-60%relative humidity, and a 12-hour light:dark cycle.It should be fitted with a door screen to preventthe entry of wild flies from outside.*Entomology Division, Department of Scientific andIndustrial Research, Private Bag, Auckland, New Z ealand

Too start a colony, insects can be obtained froma well established laboratory, from a fish farm, orfrom the field. Adults can be trapped in meat-baitedcages or with a net near a slaughterhouse o r stock-yards, on a farm, or even in a back garden. Eggscan be obtained using liver, beef, or fish-heads asoviposition substrates. Whichever bait is used, itis possible that more than one sp ecies will be caughtin the trap, and therefore precise identification ofthe adult will be necessary.Generally three rearing methods are employed.REARING ON ANIMAL TISSUE. This method is eas-iest, and is recommended for laboratory colonisa-tion where insects are required for researchpurposes. About 200 adults are sufficient to pro-vide a continuous supply of the various life stagesfor experimental use. Com mercial production is anextension of this method, and is done on a largescale whereby thousands are pro duced daily. Meat,liver, or fish is used as food; the rearing tempera-ture is kept high for fast development and quickturnover of the life cycle. These methods are eco-nomical to o perate.REARING ON ARTIFICIAL DIET. This method is usedfor nutritional studies, the numbers involved usu-ally being only a few hundred. Several diets thathave been compounded are reviewed by Singh(1977). One successful diet is: casein 10.0 g; yeastextract 1.0 g; cholesterol (dissolved in 1 ml ethanol)0.2 g; McCo llum-Da vis salts 0.4 g; l-cysteine 0.1 g;agar 3.0 g; mould inhibitor 2.0 ml; and water 150ml.REARING AXENIC INSECTS. This technique requiresfamiliarity with aseptic techniques and microbiol-ogical routines that are not a usual part of ento-mological training. Aseptic rearing is undertaken toproduce maggots required for surgical use, nutri-tional studies, and in v itro tissue or organ cultures.Rearing can be done either on defined diets ofknown composition or on sterilised meat, chickembryos, or piglets. The methods are com plex, andare given in White (1937), Greenberg (1973), andGreenberg & George (1985).B i o n o m i c sOf the dozen o r so species generally recognised asblowflies in New Zealand, the European greenbot-tle (Lucilia sericata) and the E uropean bluebottle(Calliphora vicina) are common exam ples that maybe reared easily, and about which a good deal isknown . The bionomics of these two species rearedon beef liver at 271 C and 502% RH with con-tinuous light (ordinary incandescent bulbs) aretabulated below from details given by Kamal(1958).

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STAGED U R A T I O Npupariation of mature larvae. The rearing con-aphora vicnatainers are kept covered so that mature larvae doEgg24 (20-28) hoursnot escape. Puparia are collected by tipping the1 st instar24 (18-34) hourscontents of the larval rearing box over a screen,2nd instar20 (16-28) hoursand are transferred to adult cages for emergence.3rd instar48 (30-68) hoursRearing on various foods at 15, 20, 25, and 30C,Prepupa128 (70-290) hoursabout 60% RH , and a 12-hour photoperiod showsPupa11 (9-15) daysthat rearing temperature can have a profound effectTotal immature18 (14-25) dayson the rate of development and the size of C. vicina.mergence success31-43 %The best rearing temperatu re is 25C so far as lar-Time to copulation5-9 daysval establishment, pupariation, puparial weights,Time to oviposition8-15 daysadult emergence, and rate of development are con-Oviposition period6 (12-15) dayscemed. At 15C and 20C development is muchMating activitynot statedslower, and at 30C larval establishment is poor,Adult lifespan25 (24-35) dayspuparial weights are greatly reduced, and no adultsemerge. At 25C larval development (from egg hatchuc asericatato pupariation) on bovine muscle and liver takesEgg18 (12-38) hours6-9 days, puparial weights average respectively 70and 79 mg, and adult emergence exceeds 70%. It1 st instar20 (12-28) hourstakes 17 days for 50% of fl ies to emerge, and these2nd instar12 (9-26) hoursflies lay eggs 4-5 days after emergence.3rd instar40 (24-72) hoursPrepupa90 (48-192) hoursNOTE. The references cited in this supplemen-Pupa7 (5-11) daystary contribution are listed in the main R eferencesTotal immature12 (12-15) dayssection (page 59).mergence success69-91%Time to copulation3-8 daysTime to oviposition5-14 daysOviposition period20 (15-29) daysMating activity15 (9-19) daysAdult lifespan46 (40-59) daysRearing C. vicina i n t h e l a bo r a t u r yRecause Calliphora vicina is the most common flyassociated with animal and hum an corpses in N ewZealand, it has been chosen here to dem onstrate atypical blowfly rearing method , and to illustrate thatthe rate of development is influenced by temper-ature. The rearing is done in a w ell ventilated roommaintained at 25 1 C, about 60% RH, and a 12-hour light:dark cycle.

MAINTENANCE OF ADULTS. Adults can be heldin cages made of galvanised iron with a 30 15cm rectangular base and covered with a nylon net-ting sleeve secured with a rubber ban d at one end.They are fed dry sugar crystals ad libitum in a Petridish, and water via a cotton wick. Bovine liver isprovided for about 12 hours during the dark period,for oviposition. Some 50-60 flies are kept in eachcage, and 3 or 4 cages are ma intained so as to pro-vide for a continuous supply of eggs. As the eggbatches are obtained the, can be transferred to lar-val rearing cages.

REARING OF LARVAE. The larvae can be rearedon beef liver replenished as needed in ventilatedclear plastic boxes (19 12 6 cm). Vermiculiteor untreated sawdust 1-2 cm deep is provided for

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TEXT CONVE NT IONSRepos i t o r i e sAbbreviations used for museums and other insti-tutions where m aterial is located are as follows (afterWatt 1979) :Australian National Insect Collection, CSIRO,C a n b e r r aB r i ti sh M u s e u m ( N a t u ra l H i s to r y ) , L o n d o nC a n t e r b u r y M u s e u m , C h r is t ch u r c h , N e w Z e a la n dNew Zealand Arthropod Collection, DSIR,A u c k l a n dHope Entomological Collections, UniversityM u s eu m , O x f o rdL i n c o l n C o l le g e , C a n t e r b u r y , N e w Z e a l a n d

M u s u m National d ' H i s t o ir e N a t u r e l l e , P a r i sN a t u r h i s t o r i s c h e s Museum, ViennaU n i te d S t a te s N a t i o n a l M u s e u m , W a s h i n g to n , D . C .Z o o lo g is k M u s e u m , C o p e n h a g e n

M o r p h o l o g yThe abbreviations for structural features used inthe keys, descriptions, and figures are those mostcomm only used by workers on calyptrate Diptera,and are as follows.

HEAD: ori, inferior orbital setae; ors, superiororbital setae; vte, external vertical setae; vti, inter-nal vertical setae.THORAX (mesonotum Figure 1): ac, acrostichalsetae; dc, dorsocentral setae; h, humeral setae; ia,intra-alar setae; pa, postalar setae; ph, posthumeralsetae; pra, pre-alar setae; sa, supra-alar setae; stpl,sternopleural setae. The term "pleurotergite" hasbeen used in preference to "supra-spiracular con-vexity", and refers to the convex pleuron immed-iately below the squama.WING: R5, first posterior cell; rm, discal cross-vein.ABDOMEN: St, sternite; T, tergite.Measurements of parafacial and jowlar widthsused for various ratios referred to in the descrip-tions and keys are of the narrow est part, measuredwith the head in profile. Genitalia drawings havebeen made w ith the hypopygium intact and restingin a natural position on cotton wool in glycerineunder a coverslip.

M a t e ri a l e x a m i n e dCollection data for non-type specimens are sum-marised, for the sake of brevity. The full data, intypescript form, are held at NZAC, and may beseen on request. --

KEY T O G ENE RA OF CALLIPHOR IDAEKN OW N FROM N EW ZEALAN D

01Stem-vein setulose dorsally (Fig. 21);head of characteristic shape (Fig. 14);squamae haired to margin. Metallicblue-green flies...(p. 57) .. CHRYSOMYINAE(Chrysomya)-- Stem-vein bare dorsally; head notshaped as in Fig. 14; squamae hairedor bare. Metallic or non-metallic flies... CALLIPHORINAE .. 2

02(01) Lower squama haired above3Lower squama bare503(02) Subcostal sclerite setulose (Fig. 22)... 4 Subcostal sclerite pubescent; headshape as in Fig. 10.. (p. 21) .. Calliphora04(03) Eyes densely haired; ocellar setaeabsent; palpi brown to black; orsabsent; male with tergites 5 and 7greatly enlarged; female tergite 5 withstrong marginal setae and a dorsalmarginal incision; head shape as inFig. 11... (p. 47) .. Ptilonesia Eyes bare or indistinctly haired; ocel-lar setae present; palpi yellow toorange; ors present; abdominal ter-gites normal in both sexes; headshape as in Fig. 13... (p. 48) .. Xenocalliphora05(02) Prosternum, propleuron, and supra-squamal ridge bare; head shape as inFig. 12 ... (p. 28) .. Pollenia Prosternum, propleuron, and supra-squamal ridge setulose (Fig. 23)... 606(05) Pleurotergite haired... (p. 27) .. Hemipyrellia Pleurotergite bare... (p. 27) .. LuciliaDESCRIPTIONS

S u b f a m i ly CA L L I P H O R I N A EGenus Calliphora Robineau-Desvoidy

Calliphora Robineau-Desvoidy, 1830: 433. Type-speciesMusca vomitoria Linnaeus, 1758, by originaldesignation.Neocalliphora Brauer & Bergenstamm, 1891: 87. Type-species Calliphora dasyphthalma Macquart, 1843(= Musca quadrimaculata Swederus, 1787), byoriginal designation.

A N I CB N HC M N ZN Z A CH C O EL C N ZM N H NN H M WU S N MZ M K D

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ParacalliphoraTownsend, 1916: 151. Type-species Cal-liphora oceaniae Robineau-Desvoidy, 1830 (=Musca augur Fabricius, 1775), by originaldesignation.Head of male holoptic or narrowly dichoptic. Aristalong plumose. Outer ph setae present. Suprasqua-mal ridge bare. Prosternum an d propleuron haired.Subcostal sclerite pubescent. Stem -vein bare. Lowersquama haired but bare marginally. Male genitaliaof the normal calliphorid type; female ovipositortelescopic. Oviparo us.Remarks. Calliphora can be distinguised from allother New Zealand Calliphoridae by the bare stem-vein, haired lower squama, and pubescent subcos-tal sclerite.Aspects of general morphology are illustrated inFigures 2, 8, 9, 19, and 24.

K E Y T O S P E C I E S O F CALLIPHORAK N O W N F RO M N E W Z E A L A N D01Abdomen brassy, with a dense, tesse- late golden dusting and bright yellowhairs ventrally; pleura covered withdense yellow hairs; femora and tibiaeorangy yellow2 Abdom en metallic blue -green orviolet, undusted or with a thin, tessel-late silvery dusting; pleura with darkhairs; femora and tibiae blackish-brown302(01) Acrostichal setae 3 + 3; stpl 2(3) +1; head of male holoptic, the eyeswith large facets anteriorly; femalewith frons wider than an eye width;interfrontalia expanded centrally ... stygia Acrostichal setae 2 + 3; stpl 1 + 1 ;eyes in male separated by 3 widthof anterior ocellus, the facets notenlarged anteriorly; female with fronsas wide as eye; interfrontalia parallel-sided...hilli03(01) Abdomen metallic violet or blue-

green and undusted; eyes denselyhaired; thoracic spiracles very large,orange... quadrimaculata Abdomen metallic blue with a tessel-late silvery dusting; eyes bare; thor-acic spiracles normal, the posteriorone brown... vicina Calliphora hilli PattonFigures 10 and 14Calliphora hilli Patton, 1925: 400. Lectotype male, Aus-tralia, Bamawm, Victoria (BMNH); see designa-tion below.Pollenia rufipes of authors, not Macquart [misiden-tifications].Calliphora rufipes of authors, not Macquart [misiden-tifications].Calliphora milleri Hardy, 1 937: 22. Type m aterial eitherlost or never designated.Calliphora varifrons of authors, not Malloch [misiden-tifications].HEAD dichoptic in female. Eyes bare, separated inmale by 3 w idth of anterior ocellus, in female byan eye width. Ground colour black. Parafacialia andjowls orangy-yellow. Interfrontalia matt reddish-brown. Parafrontalia densely dusted brassy yellowwith darker shifting spots and numero us fine, darkhairs. Parafacialia with a brassy dusting and a fewdark hairs above. Jowls golden-dusted anteriorly,brassy yellow posteriorly and with fine black setu-lae and golden-yellow hairs. Interfacial membranewith some dark hairs basally. Occiput dusted brassyyellow, with golden hairs. Male with 10 or 11setae and a pair of vti, female with 6 or 7 ori, 2ors, and a pair each of preverticals, vti, and vte.Vibrissae strong, crossed. Facial ridge with shortsetulae for two-thirds of its length. Antennal seg-ments orangy, brow nish apically; arista plumose fortwo-thirds of its length. Palpi yellow, flattened anddilated apically. Mentum glossy brown.THORAX. Ground colour black. Mesonotumwith dense silvery-grey dusting and a shifting pat-tern of spots and streaks. Ac 2 + 3, dc 3 + 3, 4h, 2 ph, ia 3 + 2, 2 pra, 2 sa, 2 pa. Pleura dustedgrey and brassy yellow, and covered with golden-yellow hairs. Propleural depression haired. Meso-pleuron with a complete row of setae and a fewblack setulae above. Pleurotergite with a pale, shortpile. Spiracles yellow. Stpl 1 + 1. Scutellum witha pair each of apical, basal, and discal setae.WINGS. Veins yellowish-brown. Epaulet andbasicosta yellow. Subco stal sclerite with yellow pile.Squamae infuscated yellow; lower lobe with long,yellow hairs on disc, one or two black ones api-cally, and yellow marginal hairs; upper lobe witha dark margin and dark marginal hairs posteriorly.LEGS. Coxae brow n, with a dense grey dustingand yellow hairs. Trochanters, femora, and tibiaeyellow; fore tibia with 2 or 3 weak ad setae and 1pv ; middle tibia with 2 or 3 ad setae, 1 pd, 2 p, and1 v; hind tibia with a row of short ad setae, 3 pd ,and 2 or 3 av . Tarsi dark brown.ABDOMEN. Ground colour black, with metallicgreen and brassy reflections and a dense, tessellategolden dusting. Dorsal ground setulae black; ven- 2 2


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tral surfaces with long, golden hairs. Tl + 2 withorangy hairs laterally; 3 with a row of fine mar-gial setulae; T 4 with a row of strong marginal setae; 5 with a row of m arginal setae and a few strongerorange and black discal setulae. Sternites dustedgrey and yellow, with long, golden setulae.GENITALIA. Male hypopygium, Figures 10 and14.DIMENSIONS. Length of body 7.5-10.0 mm;length of wing 6.5-8.5 mm.

Type data. Patton described Calliphora hilli fromfour male specimens, three from Bam awm and on efrom Seafo rd, both localities in Victoria. Under thisname in the BMNH I have found two males withtype data. One is labelled "Bamawm, Vic., W.F.Hill", "ex coll. W. S. Patton", "Pres. by LiverpoolSch. Trop. M ed.", "Calliphora hilli Patton, Det. W.S. Patton", in good condition but with the abdo-men m ounted separately and the left hind leg miss-ing. The second is labelled "Seaford, V. W . F. Hill","ex. coll. W. S. Patton", "Pres by Liverpool Sch.Trop. Med", "Calliphora hilli Patton det. W. S.Patton", and is in poor condition the abdomenand right hind and middle legs are missing. I havelabelled, and here designate, the Bamawm speci-men as lectotype and the Seaford specimen as para-lectotype. The lectotype agrees with the most recentinterpretation of this species (Kurahashi 1971; asC. milleri), but the paralectotype is a specimen ofCalliphora bezzii Hardy, according to Dr K.R.Norris.Mater ia l examined. Type specimens, plus 32 non-type examples (3 males, 29 females; ANIC, BMN H,LCNZ, NZAC).Three Kings Is / ND (Fanal I.), TO , TK, WN /NN, KA / SI.Collected at around sea level up to 1500 m(Mount Owen, NN) and even 2500 m (MountEgmont summ it, TK).Taken in January, March, August, November,and D ecember.H abitat records: "open scrub" (Fan al I.). Collec-tion methods noted: "swept".Remarks. Calliphora hilli occurs on sheep withthe more common species C. stygia, but appearsto be of little significance as an agent of fly-strike.Zumpt (1965, p. 63) says that only a few cases ofsheep m yiasis have been recorded for this species.It has been reared from human corpses (Smeetonet al. 19 84), carrion, dead crayfish, and dead moths.Diagnostic characters of the larvae are describedand illustrated by Holloway (1985,p. 14; Figures126-128).This species is known only from Australia andNew Zealand.

CalliphoraCalliphora quadrimaculata (Swederus)Figures 8, 9, 19, 24, 27, 28, and frontispieceMusca quadrimaculata Swederus, 1787: 289. Holotypefemale, New Zealand (Banks Collection, BMNH).Musca sacra Fabricius, 1805: 291. Holotype female, Capeof Good Hope [error] (ZMKD).Calliphora dasyphthalma Le Guillou, 1842: 315. Lecto-type female, New Zealand, Auckland Islands(MNHN); see designation below.Calliphora dasyphtalma Macquart, 1843: 287 [reprint 130].Lectotype female, New Zealand, Auckland Islands(MNHN); see designation below.Musca violacea Walker, 1853: 335. Holotype female, NewZealand (BMNH).Calliphora cockay nei Hutton, 1904: 155. Holotype female,New Zealand, Campbell Island (CMNZ).HEAD holoptic in male, dichoptic in female. Eyesdensely haired, separated by 1.4 width of eye;anterior facets enlarged in male. Ground colourreddish-brown except on occiput, which is black.Interfrontalia matt, with long, fine, black hairs.Parafrontalia thinly grey-dusted, with a shifting sil-very spot centrally, densely haired. Parafacialiadensely haired in upper p art, with an indistinct banddusted silvery-yellow. Jowls b rownish-dusted ante-riorly, grey-dusted posteriorly, with long, darksetulae. Occiput grey-dusted, with yellowish hairs.Male with 7 or 8 on setae and a pair of vti. Femalewith 12-14 on setae, a prevertical, vti, and vte, an d2 pairs of ors. Vibrissae strong, crossed with asecond, slightly weaker pair. Facial ridge with short,strong setulae over its entire length. Antennal seg-ments brown; 3rd segment orangy basally; aristaplumose on basal two-thirds. Palpi yellow, flat-tened and dilated apically. Mentum dusted blackand grey.

THORAX. Ground colour black except on scu-tellum, which is brownish. Mesonotum evenly grey-dusted except for a small shiny patch at humeri.Some specimens less densely dusted between setalrows ac and dc. A c 2 + 3, dc 3 + 3, 3 h, 2 ph, ia2 + 2, 1 or 2 pra, 2 or 3 sa, 2 pa. Pleura grey-dusted, with dark hairs. Spiracles very swollen,bright orange. Small knob above infra-alar bullaorange, with orangy hairs. Pleurotergite pubescent.Stpl 1 + 1. Scutellum with a pair of apical setae,4 pairs of laterals, 2 pairs of basals, and 2 pairs ofdiscals.WINGS. Bases slightly infuscated. Veins brow n.Basicosta and epaulet orangy-yellow. Squamaebrown, with brown marginal hairs; lower lobe withbrown hairs on disc.LEGS. Coxae, trochanters, and femora black-ish-brown w ith a thin grey dusting. Tibiae reddish-brown; fore tibia with a row of short ad setae anda pv seta; middle tibia with 3-5 ad setae, a row ofpd, and 2 v setae in female, and 2 ad, 1 pd, 2 p,and 2 v in male; hind tibia with a row each of ad

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and pd setae, including one or two stronger setaein the rows, and 2 av setae which are stronger inthe female. Tarsi blackish-brown.ABDOMEN. Ground colour black, with metallicviolet or blue-green reflections; without dusting. Alltergites punctured by numerous large setal pores; 3 with lateral marginal setae; 4 with numerousfine marginal setae; 5 with a row of marginal setaeand so me lon ger discal setae. Sternites thinly grey-

dusted on margins. Campbell Island and AucklandIslands material: abdomen less punctured, moregreenish-blue; male parafrontalia more silvery-dusted.GENITALIA. Male hypopygium, Figures 27 and28 .DIMENSIONS. Length of body 9.5-15.0 mm;length of wing 8.5-13.5 mm.

Type data. Musca quadrimaculata.: Swederusdescribed this species from New Zealand materialin the Banks Collection (BMNH), where I havefound a single female under this name. It is ratherdirty and dusty, the right foreleg is missing, andsetae are abraded from the mesonotum. I havelabelled it holotype, and it agrees with the mostrecent interpretation of this sp ecies (e.g., Harrison1976).Musca sacra: Fabricius described this speciesfrom an unknown number of specimens, of unstatedsex, supposedly collected at the Cape of Good H ope.From ZMKD I have received for study a singlefemale found under this name. Although the spec-imen jumped its pin in transit, and was loose inthe box, it is still in fair condition. It is labelled byFabricius "M. sacra e Cap. b. sp.". This specimenis undoubtedly the holotype; Fabricius must havemade a mistake with the locality, as it is a speci-men of Calliphora quadrimaculata.Calliphora dasyphthalma: Le Guillou and Mac-quart described this species independently, frommaterial collected by the A strolabe and Zele expe-dition an unstated number of female specimensfrom the Auckland Islands. From MNHN I havereceived for study three females found under thisname. One is labelled "9 Calliphora dasyphthalmanobis hi les iles Auckland", "130", and apart frombeing dusty is in excellent condition. Another islabelled `Type de Leguillon, Macq. 2,3,130,8" andon a small red disc "1136.41", and is in fair con-dition but has the distal half of the abdom en miss-ing. I consider all three specimens to be syntypes,and here designate the specimen labelled "9 Cal-liphora dasyphthalma" as lectotype and the othersas paralectotypes. All specimens agree with the mostrecent interpretation o f Calliphora quadrimaculata(e.g., Harrison 1976).Calliphora cockaynei: Hutton used this name

when describing a single female from CampbellIsland, which I have received for study from theCanterbury Museum. It is rather dirty but other-wise in good condition. It is labelled "1.84 4", "Cal-liphora cockaynei Hutton Holotype", "Camp. Is.","Camp bell Is. Dr. L. Cockayn e", "Calliphora cock-aynei Hu tt, F.W. H utton det.", and agrees with themost recent interpretation of C. quadrimaculata(e.g., Harrison 1 97 6). The distinguishing charactergiven by Hutton the colour of the abdomen is no more than a colour variation of quadrima-culata found on the Auckland Islands and Camp-bell Island.Material examined. Type specimens, plus 222non-type examples (73 males, 149 females; ANIC,BMNH , LCNZ, NZAC).WO, BP, O, TK, HB, WN / NN, BR, KA, NC,MC, SC, , WD, CO, DN, SL / SI / AucklandIs / Campbell I.Collected at around sea level up to 1250 m (Dun-das Hut, Tararua Range, WN).

Taken in all months.Habitat records: several from "house" and "win-dow"; also noted from "dung", "rotting matter",and "dead elephant seal" (Venus Cove, CampbellIsland). Collection methods noted: "swept" and"malaise trap".Remarks. Calliphora quadrimaculata is restrictedto New Zealand, where it is very common, espe-cially in houses during summer and autumn. Tol-erance of cooler conditions by the adult fly issuggested by winter records, especially one fromTucker Cove, Campbell Island, dated 6 August 1962(LCN Z). It will readily "blow" bedd ing, clothes, andmeat, and will develop in various kinds of decay-ing matter. Holloway (1985,p. 12) found pup ariaand a mature larva under tussock, where they haddeveloped in the absence of carrion. It has occa-sionally been recorded as causing strike in sheep,but as a secondary agent and no t a primary invader.Holloway (1985 , p. 14; Fig. 130-133) describes andfigures the diagnostic characters of the larvae.Calliphora stygia (Fabricius)Figures 10, 29, and 30Musca stygia Fabricius, 1781: 438. Holotype female,Am erica [error] (Banks Collection, BMNH).?Calliphora villosa Robineau-Desvoidy, 1830: 437. Typematerial Australia, Port Jackson, N.S.W. (not inMNHN).Pollenia rufipes Macquart, 183 5: 271. Lectotype female,Australia, Port Western ( ); see designationbelow.Musca laemica White, 1843: 291. Lectotype female, NewZealand (M); see designation below.

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HEAD holoptic in male, dichoptic in female. Frons1.2 5 as wide as eye. Eye in male with enlargedfacets anteriorly. Ground colour reddish-brown.Interfrontalia matt, with num erous fine, black setu-lae. Parafrontalia grey-dusted above, golden bro wnbelow, with numerous black setulae. Parafacialiadensely dusted golden brown, with darker shiftingspots, and with black setulae in the upper part.Jowls dusted golden brow n anteriorly, greyish pos-teriorly, with black setulae and golden hairs. Facedarkened centrally. Occiput grey-dusted, withnum erous yellow hairs. Interfacial membrane b are,coppery-dusted. Male with 9-11 on setae and a pairof vti; female with 8-12 on setae, a prevertical, apair each of vti and vte, and 2 pairs of ors. Vibrissaestrong, crossed. Facial ridge with short setulae fortwo-thirds of its length. Antennal segments orangy;3rd segment browned apically; arista plumose forits entire length. Palpi yellow, flattened an d dilatedapically. Mentum glossy brown.THORAX. Ground colour black. Mesonotum

dusted steel-grey with indistinct, shifting, dustedstreaks; ground setulae black except for a fewgolden-yellow hairs anteriorly. Ac 3 + 3, dc 3 +3, 3 h, 2 ph, ia 2 + 1, 1 pra, 2 sa, 2 pa. Pleuradusted grey and brown, with long, golden-yellowhairs. Pleurotergite with yellow pile. Hypopleuralsetae yellow. Spiracles large, pale orangy-brown. Sipl2(3) + 1. Scutellum with a pair each of apical, lat-eral, and discal setae and 2 pairs of basals.WINGS. Veins yellow basally, brownish api-cally. Basicosta and ep aulet yellow. Squ amae infus-cated brownish-yellow, with yellow hairs basallyand dark hairs apically on disc. Marginal hairs pale

yellow to golden brown.LEGS. Coxae yellow, with a dense grey dustingand yellow setulae. Trochanters and base of femo rawith yellow setulae. Femora and tibiae brightorangy-yellow. Fore tibia with a well spaced rowof short ad setae and 1 pv seta; middle tibia with3 ad setae, 1 pd, 2 p, and 1 v; hind tibia with 1 or2 stronger setae in an ad row, 2 or 3 pd, and 2 av.ABDOMEN. Ground colour black, with brassygreen reflections and a dense, tessellate goldendusting. Dorsal ground setulae black, laterally andventrally golden yellow. T 1 + 2more greyish; T 3 (viewed from beh ind) with a dis-

tinct, undusted median vitta and some long, fine,lateral marginal setae; T4 with a row of marginalsetae; T5 with ground setulae longer and sparser,some of them

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